CONTENTS
The genus Cybianthus (Myrsinaceae) in Ecuador and Pern
John J. Pipolylll
1
New flowering plants from southern New Ireland, Papua New Guinea
W. Takeuchi andjohnj. Pipolylll
161
Three new species of Crataegus (Rosaceae) from western North America:
C. okennonii, C. okanaganensis and C. phippsii
J.B. Phipps and R.J. 'Kennon
169
Taxonomic clarification of Atripkx nxit^llii (Chenopodiaceae) and its near relatives
Howard C. Stutz and Stewart C. Sanderson
193
Flavonoids in Strophostyles species and the related genus Dollchopsis
(Phaseolinae, Fabaceae): Distribution and phylogenetic significance
Juan Pablo Pelotto and Maria A. Del Pero Martinez
213
Ruellia jimnlcensis (Acanthaceae), a new species from the Chihuahuan
Desert area, Mexico
Jose A. VillarrealQ.
223
Una nueva especie de Agave, subgenero Littaea (Agavaceae) de Guerrero y
Oaxaca, Mexico
Abisai Garcia-Mendoza and Esteban Martinez Salas
111
A new species of Mandevilla (Apocynaceae) from Jalisco, Mexico
Justin Williams
231 ^^ ^^^K
A new combination in Mexican Mandevilla (Apocynaceae)
Justin Williams
237
Trachypogon mayaensis (Poaceae: Andropogoneae): A new speaes from Belize
Joseph K. Wipffand Stanley D.Jones
241
Taxonomy of the Sporobolus floridanus complex (Poaceae: Sporobolinae)
Alan S. Weakley and Paul M. Peterson
lAl
Caryopsis morphology of Leptochloa sensu iato (Poaceae, Chloridoideae)
Neil Snow
111
Xanthostemon fruticosus (Myrtaceae), a new species from the Philippines
Peter G. Wilson and Leonardo L. Co
283
(continued on back cover)
CONTRIBUTIONS
TO BOTANY
Volume 18
Number I
October, 1998
SiDA
Contributions to Botany
FOUNDED BY
Lloyd H. Shinners
1962
Wm. F. Mahler
Publisher 1971-1992
Barney L, Lipscomb
Editor
Botanical Research Institute of Texas, Inc.
509 Pecan Street
Fort Worth, Texas 76l02-4060, USA
817 332-4441 / 817 332-4112 FAX
Electronic mail: sida@brit.org
Home page at the URL: http://www.brit.org/sida/
John W. Thieret Prof. Dr. Felix Llamas
Associate Editor Contributing Spanish Editor
Biological Sciences Dept. Dpto. de Botanica, Facultad de Biologia
Northern Kentucky University Universidad de Leon
Highland Heights, Kentucky 41076, USA E-24071 Leon, Spain
Guidelines for contributors are available upon request
and on the inside back cover of the last issue of each volume.
Subscription per year: $25. Individual, $45. USA Institutions, $50. Outside USA;
numbers issued twice a year
®SiDA, Contributions to Botany; volume i8, Number i, pages 1-376.
Copyright 1998
Botanical Research Institute of Texas, Inc.
Printed in the United States of America
ISSN 0036-1488
THE GENUS CYBIANTHUS (MYRSINACEAE)
IN ECUADOR AND PERU
JOHNJ. PIPOLYIII *
Botanical Research Institute of Texas
309 Pecan Street
Fort Worth. TX 76102-4060. U.S.A.
jpipoly@brit. org: clusia@latinrnail.com
ABSTRACT
The genus Cybianthus was revised to provide taxonomic treatments of tlie Myrsinaceae
for Flora of Peru, Catalogue of the Vascular Plants of Ecuador, and flora of Ecuador. Eight of
Cybianthus ten subgenera are represented, and an updated description of the genus, keys
to its subgenera and emended descriptions for each are provided. Detailed descriptions of
the morphology, anatomy and ecology of the genus are presented. Cybianthi/s subgenus Iteoides
is relegated to synonymy under subgenus Microconomorpha . Within each subgenus, keys,
full descriptions, synonymy, distribution, ecology and conservation statuses, local names
and uses are given for each species. In addition, specimens are cited for each species, in-
cluding extralimital ones to show extremes in morphological variation. Thirteen species
are relegated to synonymy, nine names are lector)'pified, and one, Conoino^fha rigida, is neotypified.
One new combination, Cybianthus guyanensis suhsp. pse//doicacoreus, is made, and Cybianth/ts
poeppigit is transferred from subgenus Cybianthus to subgenus Weigeltia. The following 15
new species are described, illustrated and their phylogenetic relationships are discussed:
Cybianthus anthuriophyllus. C. cenepensis, C. coniperuvianus. C. croatii. C. flavovirens, C. fosteri,
C. grandezii. C. granulosus. C. huampamiensis. C. incognitus, C. jensonii. C. nestorii. C. pseudolongifolius,
C. timanae. C. vasquezii.
RESUMEN
Al preparar tratamientos taxonomicos sobre la familia IMyrsinaceae para los proyectos
Flora del Peru. Catdlago de las Plantas Vasculares del Ecuador, y Flora del Ecuador, se llevo a
cabo una revision del genero Cybianthus. Se encuentra en la region ocho de los diez subgeneros,
y se proveen tanto una descripcion actualizada para el genero como para cada subgenero
tambien. Se presenta descripciones detalladas tratando la morfologia, anatomi'a y ecologfa
del genero. Se relega Cybianthus subgenero Iteoides a la sinonimia bajo subgenero Microconomorpha,
y se transfiere Cybianthus poeppigii del subgenero Cybianthus al subgenero Weigeltia. Para
cada subgenero, se proveen claves, descripciones completas, sinonimia, distribucion geografica
y estado actual de su conservacion, nombres locales y usos. Tambien se cita colecciones revisadas
para cada especie, incluyendo ellas fuera de la region delimitada cuando se muestran variacion
morfologica significativa. Se relegan 13 especies a la sinonimia, se lectotipifica 9 binomiales
y uno, Cononwrpha rigida, se neotipifica. Se publica la nueva combinacon, Cybianthus guyanensis
iuhsp. pseudoicacore//s. Se describen, se ilustran y se discuten el parentezco para 15 especies
nuevas, listadas a continuacion: Cybianthus anthuriophyllus, C. cenepensis. C. coniperuvianus,
C. croatii, C. flavovirens. C. fosteri, C. grandezii, C. granulosus, C. huampamiensis, C. incognitus.
C. jensonii, C. nestorii, C. pseudolongifolius, C. timanae, C. vasquezii.
SiDA 18(1): 1-160. 1998
2 SiDA 18(1)
INTRODUCTION
The genus Cybianthus Marti us was cladistically defined by the presence
ot unique glandular granules at the junction of the corolla tube and lobes
by Pipoly (1987). In addition, the unique combination of lateral (axillary)
inflorescences, gamosepalous and gamopetalous flowers, and stamens con-
nate by their filaments at least one-fourth their length, and adnate to the
corolla rube at least one-third its length (Pipoly 19<S7, 1992a) allows for
practical recognition from herbarium specimens. With this contribution,
Cybianthus now contains 167 species in ten subgenera. The genus includes
species formerly included in the genera Comomyrsnn: Hook, f, Couontorpha
A. DC, Correlliana D'Arcy, Cyh'utnthopin Lundell, GraninuukMict Benth.,
Mkroconomorpha Lundell, and Wetgeltui A. DC. (Pipoly 1 987, 1 992a). In Ecuador
and Peru, 43 species in 8 subgenera are known. Among the species, none
are endemic to Ecuador, while 1 1 are endemic to Peru.
The broad generic concept for Cyhianthi/s was first proposed by Agostini
(1970) as a precursor to his dissertation (1972); that was followed by the
formal transfers and a key to the subgenera (Agostini 1 980). Subsequently,
while revising subgenus Gra)ui>iadenia, Pipoly (1987) determined that the
paleotropical genus Embelia Burman f. was the sister group to the entire
genus and that it was most parsimonius to include Gramviadenia within
Cybianthus. Preparation of taxonomic treatments for the Myrsinaceae in Flora
of Per//, Catalog//e of the Vasc//lar Plants of Ec//ador and flora oj Ec//aclor re-
vealed that many taxa were new, and much confusion had arisen among the
concepts prevalent for taxa already described. Among adjacent countries
with significant Amazonian regions, Ecuador and Peru share more species
oi Cybianthus in common than do any other pair of countries. While it would
have been desirable to include Colombia to provide a treatment for the entire
northwestern South America, the number of additional species endemic to
Colombia, or known only from Colombia and Venezuela, would have in-
creased the size of the treatment by fifty percent. Given that it will be some
time until a revision of the entire genus for Flora Neotrop/ca is complete, the
present treatment is mtended to serve as a precursor to that monograph
and to make the names available for the other aforementioned projects. A
revised description of the genus Cybianth//s follows, including keys and emended
descriptions for the eight subgenera occurring in Ecuador and Peru, along
with keys to the taxa and full descriptions for each. For each of the fifteen
new species described, illustrations are provided, while for all species, full
descriptions and synonymy, discussions of distribution, ecology and con-
servation status, etymology, and when known, local names and uses are elucidated.
MORPHOLOGY
Morphological terms in this treatment follow Lindley ( 1 848) and Pipoly
PiPOLY, Cybianthus in Ecuador and Peru 3
(1987, 1992a) for the inflorescence, rachis pedicels and floral parts. Description
of leaf morphology follows Hickey (1984), trichome description follows Theobald
et al. (1984) and basic cell and tissue terminology follow Metcalfe (1984).
Habit and Architecture
The majority o^ Cybianthus species in Ecuador and Peru are large shrubs
or small trees to 18 meters tall. Four species are small shrubs or subshrubs
{Cybianthus croatii. C. huniilis. C. lineatus. C. nanayensis) usually under one
meter tall. With the exception of two species in subgenus Grammadenia, all
species are terrestrial. Cybianthm marginatum is an obligate epiphyte, while
C. magnus is a facultative epiphyte, frequently found on large tree trunks,
but also known from large rock outcrops.
Most members g{ Cybianthus exhibit Rauh's Model of architecture (Halle
et al. 1978), characterized by a polyaxial, monopodial, rhythmically grow-
ing, readily distinguishable trunk, that develops tiers of branches morpho-
genetically identical to itself. All branches are orthotropic and monopo-
dial, with lateral (axillary) inflorescences that do not affect shoot development.
However, two subgenera {Comomyrsine and Triadophora) exhibit Corner's Model
(Halle et al. 1978), characterized by a single aerial meristem that produces
a monoaxial (unbranched) axis on which inflorescences are lateral (axillary).
The resulting monocaulous tree is pleionanthic (not hapaxanthic, or mono-
carpic) and growth is indeterminate. The trunk may grow rhythmically or
continuously. Sporadic occurences of this architectural model occur in members
of subgenus Weigeltia, from the Guianas and eastern Amazonia, however,
none of those species are known from Ecuador and Peru.
While conducting tieldwork in Peru, a collection o1 Cybianthus kayapii
(Lundell) Pipoly was observed reiterating. In the population observed at
Explorama Lodge near Yanamono, (Loreto, Peru) one individual {Pipoly et
al. 12383), exhibited bayonet reiteration, caused by destruction of the api-
cal meristem of the trunk. The individual bore a reduced staminate inflorescence
with flowers slightly larger and more maroon in color than normal, and smaller
leaves with shorter petioles. As I have noted elsewhere (Pipoly 1987, 1992a),
leaves on the reiterative shoots resemble those of saplings. Variation in
inflorescence and floral morphology seen in this Peruvian population is consistent
with variation reported in Pipoly (1992a). Therefore, it appears that changes
in sex expression are consequent to a reiteration phenomenon, rather than
circumstantially associated with it. I postulate that reiterative branches may
be hormonally juvenile and iis such, are morphogenetically incapable of producing
bisexual or pistillate flowers, and produce only sapling leaves until flowering
occurs. This process may account for the great variation in leaf morphology
and apparent sexual lability often associated with many species of Cybianthus.
Unfortunately, for individuals of normally dioecious species with monoaxial
4 Si DA 1<S(1)
models, it is not possible to determine if sex change has occurred because
no inflorescences are usually observed before the original apical meristem
is destroyed, and no inflorescences on other branches are available for com-
paris{)n. Hov^^ever, precociously flowering individuals o'iCybtanthiis incognitiis
(P. Barbour 2403) stipport the hypothesis. Clearly, pruning experiments /'//
.r//// will be necessary to test this hypothesis.
Branchlets
A branchiet is here defined as the distal 10 cm oi any branch. Branchlets
may be flexuous, or straight; terete or angular; smooth, verructflose, or ver-
rucose-papillate (Fig. lA, IB), glabrous, glandular-papillate (Figs. ID, 3F,
4F), ferrugineous stipitate-lepidote (Fig. 2A— F), ferrugineous or rufous dendroid-
and/or stellate-tomentose (Fig. 3A), rufous sessile furfuraceous lepidote, with
a tomentum of malpighiaceous trichomes (Fig. 3D), glandular-granulose,
with hydropotes (Fig. 3C), epunctate or rubiginous punctate-lineate, bear-
ing lenticels or not.
Leaves
Species ofCyi?/a}ith//.s\ like all Myrsinaceae, are exstipulate and have simple
leaves. The leaves are usually alternate, but may be subopposite or
[:)seudoverticillate, especially in subgenera. Mkroaj/wf/mpha, Co//umyrs/m, Weigeltia
and Cybianthus . In the majority of species, the leaves are petiolate, but in
subgentis Grammadenia they are sessile. The ptyxis (the form in which one
shigle leaf is folded on itself in btid) is most often supervolute (Cullenl978),
but it has not been adecjuately studied thtis fitr for each subgenus. The leaf
vernation sensu CuUen (1978), is at least analagous with floral aestivation,
or the relationship of one leaf to another; in Cybianthus, it is in fact, un-
known. The leaf blade texttire may be membranaceous, chartaceous or co-
riaceous. The shape is most often elliptic, but may be oblanceolate, lan-
ceolate, obovate, rarely ovate or oblong {Cybianthus nuirginatus). The apex
may be acute, acuminate, long-attenuate, rounded, obtuse, or emarginate,
without mucro or rarely (stibgenera Grammadenia, Comomyrsine, Triadophora)
mucronate (Fig. 21). The base is rarely auriculate (subgenus Grammdenia)
or obtuse approaching auriculate {Cybianthus grandezii. C. kayapii. C.
anthuriophyUus) and is usually cuneate, long-attenuate, acute or obtuse, decurrent
on the petiole or not. The adaxial surface may be nitid, pallid, or sordid,
rarely scrobiculate , pusticulate or pustulate when mature. The blade mar-
gin may be regular or irregular, flat inroUed or revolute, entire, or rarely
undulate, lobate, crenate or dentate {Cybianthus pastaisis), densely and mi-
nutely serrulate {C, anthuriophyUus), or coarsely dentate {C. schlimii. some
populations of C pastensis, C. poeppigii, ), or with a very subtle btit vascular-
ized vein ending (C verticiHoides) opaque, or rarely scariotis (all species in
PiPOLY, Cybianthus in Ecuador and Peru
Fig. 1 . SEM photos ot morphological features in subgenus Grammadenia. A. Cybianthus marginatus
{'Pipoly 6546), showing verrucose papillae. B. Same, close-up. C, D. C. lineatus (Pipoly et al.
7729), showing stem surface. D. Glandular papillae. E. C. costarkanris subsp. morii {Pipoly
7017), pollen, equatorial ant! polar views. (Bars in figs, equal: A. 50 pm, B. 22.2 pm, C, D.
48 pm, E. 12.3 pm) Figure reproduced from Pipoly, 19H7.
6
SlDA US(1)
Fk;. 2. SEM photos of representative stipitacc fcrrugincous lepiclote scales of SLibt;enus Ciiniiiindrlihii.
A. Cyhiduthiis {liii^autiiphylh/s (holocyjie). V>. C. spichii^^eri (holotype). C. C. hittiis (D/nlley lOHOi).
D. C. per/n-hiiii/s (Dodsofi 282 1 ). E. C. comper/iviarms {Killip &. Smith 260'^ 3). F. (2 )^/tytiiiciisis
subsp. p.seiicliiiuiioyei/s {Siiiipsoii & Schinikc 7,S4). (Bars in Figs. A-F ec]ual: 50 pm).
CybicDitlv/s subgenus Gramnmdema; Cyhianthiis h//»/ilis. and C. troatii ot Cyhuniih/is
subgenus Conioriiyrsim).
Leaf punccation and punctate-lineation may be pellucid (translucent), black,
orange red, or rubiginous (light red) in color. Punctations as defined by Pipoly
(19H7) are definded as rounded, lyso-schizogenously formed cavities and
are distinguished from ptmctate-lineations, which are homologous, linear
cavities in the leaves. Among taxa bearing non-pellucid punctations or punctate-
lineations, all young, undexpanded leaves have orange glands, even il ma-
ture leaves have red or black glands. Experiments to determine integrity of
punctation color have revealed that it is not a reliable taxonomic character
PiPOLY, Cybianrhus in Ecuador and Peru
0t "*
Fig. 3. SEM photos ot representative vestiture in subi2,enera Laxiflorus. Mkrocojioinorpha. Triadophora
and Coiitoinyrsine. A. Cyh'uinthin fiilvopiilvtviikntus stibsp. magnoliifolitis {Cowan & Soderstrom
2146), branchlet, showing' stellate trichomcs. B. C, sp/cati/s {Maguire & Puliti 28064), showing
hytlropote in abaxial leaf pit. C. C. pcisttusis (Killip & Garcia 33886), branchlet, showing
developing hydropotes. D. C. scblinui {Fosherg 20148), branchlet, showing malpighiaceous
trichome. E. C. kayapii (holocype) Sraminate flower, showing vestigial pistillode, structure
of androecium. F. C. hitniilh {l.ehmanii s.n.), showing glandular papillae of abaxial leaf sur-
face. (Bars in Figs. ec]ual: A-C. 50 pm, D & F. 25 pm, E. 0.4 mm),
(Pipoly 1987). The other punctation character states include "conspicuous"
versus "inconspicuous," reflecting the ease of viewing the punctation (a reflection
of its relative diameter in transverse section), and "prominent" versus "not
prominent," which refer to whether the punctation is blister-like or planar,
respectively.
The abaxial and/or adaxial surface of the blade may be glabrous, or with
8 Si DA 18(1)
a vestiturc composed of stipirate ferrugineous lepidote scales (subgenus
Cononm-pha, Fig. 2A-2F), sessile fi.irfuraceous lepidote scales (subgenus Weigeltia),
dendroid and/or stellate crichomes, translucent glandular-lepidote scales,
or with scattered glandular papillae (subgenus Coiuoniyrsine, Figs. 3F, 4F ),
maljMghiaceus trichomes (subgenus Triadophora , Fig. 3D), or hydropotes
(subgenera Laxiflorns, Microcoiioiiiorpha, Grannnadenia, Coniomynine^ Figs- 3A,
3C, 4A-E ). The adaxial surface is usually giabrescent.
Hydropotes ("water drinkers") were previously thought to be unique to
subgenus Granivuknia (Pipoly 1987), but have now been found in all spe-
cies of subgenus Comnniyrsine. Described by Mayr (191 5) and Gruss (1927a,
1927b), hydropotes have been reported for numerous submerged aquatic
angiosperms (Fahn 1979; Gessner & Volz 1951). Elegant ultrastructural
and autoradiographic studies undertaken by Liittge (1964) and Liittge and
Krapf (1972) demonstrated the mineral absorptive function of hydropotes.
While both subgenera bear these structures, their morphology is strikingly
different. The hydropotes of subgenus Grammdenia (Fig. 5) consist of five
to seven subsidiary cells, a central foot cell, a basal cell, a stalk cell, and up
to 12 cells forming a scale-like cap, while those of subgenus Comniiiyrsine
consist of five to seven subsidiary cells, a central foot cell, a stalk cell, and
up to 1 2 cells forming a lenticular cap or globose body. In subgenus Conmiiyrsme,
the cap cell is formed before elongation of the stalk cell. In Cybianthus verticilloides
(Fig. 4B), C. spruce! (Fig. 4D), and C. kayapii (Fig. 4E) the cap consists of a
spherical body of cells, while m C. croatii (Fig. 4 A) and C, simplex (Fig. 4C),
the cap is lenticular. It is interesting to note that within subgenus GraniDuideuui,
the shape of the caj-) is the same among all species, while in Comimiyrsiue^
there are 4 types. The occurrence of hydropotes in subgenus Laxijior/zs, in
pits over the abaxial leaf surface, and in subgenus Microconomorpha, under
the papillate tomentum of the branchlets, was unknown before the present
study. Despite these differences, the structures are homologous, based on
position, function and ontogeny. In early ontogeny, a mucilaginous sub-
stance accumulates around the base cell (Fig. 7E). Later, the cap breaks off
(Fig. 7F), leaving a mucilaginous ring around the broken basal cell (Pipoly
1987, unpubl. data).
Subepidermal fibers, visible as numerous, thin, parallel lineate structures
on both surfaces of leaf blades in subgenus Triadophora are unique to it. They
have recently been shown to be homologous with those of Clavija in the
Theophrastaceae (Pipoly, unpubl.).
Petioles are present in most taxa, with the notable exception of subge-
nus Gram/i/deiiia, where the leaves are sessile. The petioles may be canaliculate,
marginate, or canaliculate with margins, trigonal, or rarely subterete, and
may be pulvinate (abruptly swollen basally) or gradtially tapering basipetally.
PiPOLY, Cybiantlius in Ecuador and Peru
Fig. 4. Representative SEM photos ofmorphological features in subgenus Comouiyniut. A-
E, Hydropotes of abaxial leaf surface. A. Cyhianthns croatii (holotype), showing lenticular
cap. B. C. vertidUoides (holotype), showing globose cap. C. C. iimplex {Zak 1350), showing
lenticular cap. D. C. spruce/ (C//atrea/sas 15743), showing globose cap. E. C. kayapii (holo-
type), showing depressed-globose cap (somewhat collapsed). F. C. kayapii (holotype), branchlet,
showing glandular papilla. (Bars in Figs. ec]ual: A. 25 pm. B. 20 pm. C. 10 pm. D— F, 25
pm).
Cataphylls and Pseudocataphylls
Cataphylls (Fig. 9B) and pseudocataphylls are synapomorphic to subgenera
C omomyrs ine 3.nd Triadophora, respectively. Earlier (Pipoly 1987), I had mis-
takenly interpreted them as inflorescence bracts. They may be alternate or
pseudoverticillate, alternating with pseudoverticels of leaves, or apparently
axillary to them, rigid, chartaceous, or membranaceous, linear-subulate to
10
Si DA IcSd
V]i,. 5 . Onrogeny ot hydroporcs in siilit;c-nLis GrciiiiiiUHk)iui. A, B, & D. Cyhuintbiis aistciyiunim
subsp. cintciricciHiii {Pipfily 76(AS'), SHM photos. A. MatLire hyclropore, cap with at least 8
cells, anti with 5 subsidiary cells. B. Younger hydropote, cap with 1 cells, ami with 6 sub-
sidiary cells. C. C CDSliiyncimis stibsp. iiinri'i (Pipiily 701 7), longisection of yoting hydropote,
showing sLibsidiary cells (sc), loot cell (fc), stalk cell (st), cap cell (ca), basal cell not dis-
cernible. D. Young hydro|-iote, with 2-celled cap, showing miicilagmous SLibstance (ms).
E, K C. coitarkaniis subsp. [hii/iiiiifiisis (Pipuly 7036), paradermal sections. E. Showing ma-
ture hydropote, with 1 2-celled-cap. V. Showing hydropote after cap has broken off. (Bars
in Figs. A-F equal: A & C 28 pm, B. 20 |_im, D. 10 pm, E. 36 pm, F. 5.i pm. Figure
reprodticcd from Pipoly, 1987.
PiPOLY, Cybianthus in Ecuador and Peru 11
acicLilar, rigid to membranous, keeled or flat, apically mucronate or not,
prominently punctate or punctate-lineate, glabrom, aifous pubemlent, glandular-
papillate, bearing hydropotes, or orange lepidote scales, sessile or on a peti-
ole to 2 mm long. Pending further developmental studies, I am distinguishing
cataphylls from pseudocataphylls based on the absence of a petiole in the
former and its presence in the latter. Within Cybianthus, pseudocataphylls
are unique to subgenus Triadophora, while cataphylls are unique to subge-
nus Comomyrsine. Both pseudocataphylls and cataphylls may be distinguished
from an inflorescence bract by the fact that neither of these structures are
axillant to an inflorescence, neither are ephemeral, but both occur in pseudowhorls.
Inflorescence and Flowers
The inflorescence in Cybianthus is always lateral (axillary), and it may consist
of a simple raceme (erect or kix), a subpyramidal or pyramidal panicle (sometimes
thyrsoid), a spike, or rarely, a pleiochasium. At times, species with essen-
tially racemose inflorescences may produce a panicle consisting of a pseudoverticel
of racemes on a reduced peduncle. In this treatment, an inflorescence is considered
spicate if the pedicels are obsolete or less than 0.3 mm long and subspicate
if the pedicels are from 0.4-0.8 mm long. In most species, the staminate
inflorescence tends to be longer, slightly more lax, and bears greater num-
bers of flowers. In those species bearing panicles, the staminate ones bear
secondary branches of the inflorescence that are longer, more floriferous, and
at times, more branched than the pistillate and bisexual panicles.
The inflorescence bracts are early caducous and are rarely seen in the field
or on specimens. The secondary branch bracts of panicles are also rarely collected,
with the exceptions of subgenera Comomyrsine and Triadophora. The floral
bracts may be membranaceous or chartaceous and are usually persistent in
staminate inflorescences, but are at times caducous in pistillate ones. The
floral bracts may be glabrous or bear a tomentum either adaxially, abaxially,
or on both surfaces. The floral bract apices are acute, attenuate, or acicular,
the margins entire or undulate.
The pedicels are cylindrical, clavate, or obconic, and at times accrescent
or crassate in fruit. In most species, the pedicels are subtended by an axillant
floral bract, but in Cybianthus kayapii, it is inserted at about the middle of
the pedicel.
Figure 6 illustrates the tremendous variation in floral morphology among
members of the genus, along with representative staminate and pistillate
flowers from Embelia, the paleotropical sister genus o{ Cybianthus. The flowers
may be functionally staminate, pistillate or bisexual. Consequently, the plants
are normally functionally dioecious, but may also be bisexual or polyga-
mous. Flowers are normally perfect, but in some species of subgenera Weigeltia,
Comomyrsine and Cybianthus, the pistillode is often obsolete in the stami-
12
SlDA IS(I)
M'-
il}{::^^;.-- .:^W]
^m
p. wVi
1 mm
Fi(i. 6. Rc-|Tr(.-sc-nranvf flowers of C)'/'/f/«//v/.f subgenera antl sister genus, £w/W/;/, open flowers
in anriiesis above, calyx lobes below. A-B, Subgenus A\/cri)aji/o///orpha, {Cy/?/ai/fh/fs /),/.\-/ei/s/s),
note monomorphic flowers. A. Srammate flower {Killiji & Cuira'ct }}SS(^). B. Piscillare flower
PiPOLY, Cybianthus in Ecuador and Peru 13
nate flowers. The staminate flowers are usually larger than the pistillate ones.
The flowers are usually 4 or 5(— 6)-merous, but they are 3-merous in subge-
nus Triadophora (Fig. 6K, 6L). The majority of species are homomerous, but
Cybianthus kayapii has heteromerous flowers, (the calyx 4- and the corolla
5-merous), and in C. anthuriophyllus, the calyx 6- and the corolla 5-merous.
Either or both whorls of the perianth may be membranaceous, chartaceous,
coriaceous or carnose, epunctate, punctate or punctate-lineate. The punc-
tations may be inconspicuous or conspicuous, prominent (raised) or not,
pellucid, brown, orange or red. The calyx may be valvate or imbricate, and
may be cotyliform, cupuliform or urceolate. The corolla is valvate or im-
bricate, and may be campanulate, cupuliform, tubiform, infundibuliform,
salverform, rotate or subrotate. The stamens and staminodes are similar in
morphology, but the staminodes are smaller. The filaments of the stamens
and staminodes are partially united at the base to form a conspicuous or
inconspicuous, membranaceous, chartaceous or carnose tube, except in subgenus
Cybianthus, which has a terete or rarely, angulate staminal tube develop-
mentally fused with the corolla tube, the stamens thus appearing epipetalous.
In some species, the tube bears lobes (sterile projections of tissue) which
alternate with the apically free portion of the filaments. The apically free
portions of the filaments may be terete, flat, or absent. The anthers may be
dorsifixed, basifixed or versatile. Anthers may be lanceolate, ovate, cuadrate,
or deltate, apiculate, acute, truncate or emarginate, the apiculate ones may
have the apiculum erect, distally or proximally recurved, glabrous or glan-
dular-papillate. Antherodes of the pistillate flowers are similar to the an-
thers but reduced insize, and normally devoid of pollen, but occasionally,
they may produce copious amounts of abortive pollen (Pipoly 1983a). The
connective may be epunctate or prominenrly punctate, or rarely glandular-
papillate. The pollen is tricolporate and psilate (Fig. IE). The pistillode
may be lageniform, conic or obturbinate. The pistil may be obnapiform,
{Schidtes 3226). C-D, Subgenus Stapfia, {Cybianthus stapjii). C. Staminate flower {Fassett 25 706).
D. Pistillate flower {Cuatrecasas 1229.5). E-F, Subgenus Laxiflori/s {Cybianthus spicatus). E.
Staminate flower (Al^^/^'/rf 33523). F. Pistillate ^ov'/ts: {Prance et al. 3339). G-H, Subgenus
Conomorpha {Cybianthus laurifoUus) . G. Staminate flower {Garcia-Barriga &Jaramillo 19841).
H. Pistillate flower {Grubb etal. 744). I-J, Subgenus Comomyrsine. L Staminate flower, Cybianthus
sprucei {Cuatrecasas 1 6272). J. Pistillate flower, Cybianthus kayapii {Acosta-Solis 5544). K-L,
Subgenus Triadophora, (Cybianthus schlimii). K. Staminate flower {Haught 1502). L. Pistil-
late flower (Herrera 1 720). JM-N, Subgenus Weige/tia, Cybianthus sp. nov. iVI. Staminate flower
{Maas & Westra 4459). N. Pistillate flower {Maguire & fansljawe 405 90). O. Subgenus Cybianthopsis,
bisexual flower, {Cybianthus sintenisii, P. Wilson s.n.). P. Subgenus Grammadenia, bisexual
flower, {Cybianthus piresii, Maguire et al. 37052). Q-R, Subgenus Cybianthus, {Cybianthus
goyazensis). Q. Staminate flower {Irwin & Soderstrom 5378). R. Pistillate flower {Herringer
45). S-T. Embelia sp. S. Staminate flower {Petelot 3887). T. Pistillate flower (C. Wang 76408).
Drawing prepared by Peggy Duke.
14 SiDA 18(1)
conic, ellipsoid, or obturbinate, with a punctiform, or capitate stigma. The
stigma is large and capitate, with erose-fimbriate lobes and is caducous in
subgenus Coniomyrslne. The ovary in species from Ecuador and Peru are ter-
ete. The placenta may be cotyliform, cupuliform or globose, with 2—4 uniseriate
ovules naked or partially immersed. The fruit is a globose or depressed-globose
drtipe and is one-seeded, with a thick or thin exocarp.
ANATOMY
This treatment is not intended to serve as a monograph of the genus, but
a few salient anatomical features may prove useful in identifying sterile material
or wood samples. Druses (Pipoly 1987-Fig. 8B— C) are ubiquitous in iVLyrsinaceae,
as are pericyclic fibers (Pipoly 1987, Fig. 8A). All Myrsinaceae have resin
ducts (canals) at least in the cortex, and in the field, copious amounts of
resin are visible in the canals of the pith and secondary phloem, in freshly
cut branchlets, in species of subgenera Grctuimadenta and Laxiflorns. Aeren-
chyma in the cortex of the primary stem is unique to subgenus Grammdenia
and is found in all species (Fig. 7F), except in Cybianthns lineatm (Fig. 7E).
The pith in primary stems of subgenus Weigeltia is parenchymatotis with
large, rcninded starch grains, while that of Cyhianthiis ))icigniis has angular
coUenchyma (Pipoly 1987). It is interesting to note that the collencyma of
the outer cortex m CyhuinthHS magniis subsp. asynimctrici/s is tangential rather
than angular, while that of C. inapun subsp. magnns is angular (Fig. 8 D—
F). All species of Cybianthiis have cortical vascular bundles, that may be
amphicribal (Fig. 7C) or hemiamphicribal (Fig. 7D), with (Fig. 7C) or without
(Fig. 7D) accompanying perivascular fibers. Wood of subgenus Comoniyrsine
is notable for its thin-walled vessels (Fig. 9D), while the fiber-tracheids of
subgenus Grcimmadenui have walls so thick as to significantly occlude the
lumina (Pipoly 1987- Fig. 7C). Also, starch is present in the phloem fibers
of subgenus Comoniynnn: (Fig. 9F, 9F) but not in Graninukuut. In leaf anatomy,
it is notable that subgenus Grcn)Wicule>iui has functionally acrodromous ve-
nation. This is due to its unique leaf-node continuum, and consequent pri-
mary vascular system (Pipoly 1987-Figs. 7-9; 1 1-12) where the cathodic
and anodic leaf traces are autonomous from each other and from the rel-
evant midrib trace, a system thus far unique among angiosperms. Leaves of
subgenus Tnadophora may be easily recognized by their subepidermal fibers,
while those o'iCybicnithiis Inimtits are unicjue for their bifacial palisade layer
(Pipoly 1987- Fig. 14b).
FXX)1.(K.Y
In Ecuador and Peru, species of Cyb'nnith/is occur in wet or moist, tall
terra firme foKst on laterite, limestone or white sand, seasonally inundated
igapo or varzea, premontane humid, wet or pluvial forest on laterite or sandstone.
PiPOi.Y, Cybianthus in Ecuador and Peru
15
Fk;. 7. Primary stem iiistological featLires in subgenus Gninnnackiiid . A— B. Cybianthus ptariensis,
(Pipoiy et al. 7133), periderm formation, showing epidermal cork development. C. C. marginatum
{Pipoly 6346), amphicribal corical bundle without periv iscular fibers. D. C. ptariensis {Pipoiy
et al. 7133), hemiampliicribal bundle with perivascu'ar fibers (pf). E. C, limatiis {Pipoiy
7229), section showing parenchymatous pith, mnct and outer cortex. F. C. marginatiis (Pipoiy
6546), section showing aerenchymatous mner cortex; parenchymatous pith and outer cor-
tex. (Bars in Figs. A-F equal to: A, 30 [.mi, B. 1 20 pm, C. 48 pm, D. 60 pm, E. 465 pm,
F. 120 pm. Figure reproduced from Pi|-)oly, \'-)H7.
cloud forest, elfin forest, montane or subparamo thickets, or sandstone scrub
at high elevation.
The terra firme and premontane forest life zones are the richest in endemics,
with six and five species, respectively. Terra firme is here divided into two
16
SiDA 18(1)
Fig. 8. Primary stem histological leacures of subgenus Grammadenia . A. C. costarkanm subsp.
morit {Pipoly 7017), showing pericyclic fibers (arrow). B. C. costaricat/j/s subsp. costaricanus
{Pipoly 7068), showing druses in pith. C. C. costaricanus suhsp. pa>2ai//ens/s {Pipoly 7056),
showing druses in pith, using polarized light. D, E. C. magniis %\}i0^p.magnus {Pipoly 6453),
showing angular pith collenchyma. F. C. magnits subsp. asyiiimetricus {huteyn, Pipoly et al.
10415), showing transitional tangential pith collenchyma. (Bars in Figs. A-F equal to: A.
195 pm, B. 45 pm, C. 48 pm, D. 230 pm, E. 74 pm, F. 60 pm.) Figure reproduced from
Pipoly, 1987.
edaphic categories, lateritic and white sand (often referred to as "variUal"
by Peruvians) and is defined as lowland tall moist or wet forest which is not
inundated. The canopy normally reaches .35 meters, with very few emer-
gent individuals. In the lateritic terra firme, Cybianthm kayapti, C. schlhnn,
C. pseuJoIong/folius, C. vasquezii, C. cenepensis, C. grandezh, C. fuscus, C. jemonii,
PiPOLY, Cybianthus in Ecuador and Peru
17
y E
\[
?i*\
Fig. 9. Morphological and anatomical features of subgenus Coniornyrsine {Cybianthus kayapii,
Pjpoly et al. 12490). A. Habit, showing bayonet reiteration. B. Habit, showing pyramidal,
bipinnate panicle and leaf tapering to obtuse base. C. Stem apex, showing cataphylls. D.
Transverse wood section, showing thin-walled vessels. E. Tangential wood section, show-
ing rays and phloem fibers-tracheids. F. Phloem fiber-tracheids, showing starch grains. (Bars
in Figs. A-F, A. 12.6 cm, B. 2 cm, C. 1 cm, D. 400 pm, E. 100 pm,
18 SiDA 18(1)
and C. venez/ielanus arti found. Cyhianthm kayapii, C. schlifiiii, and C. grandezii
are found on steep banks of small creekbeds, C. cenepensis and C. venezuelanus
are ridgetop species, and C. cenepensis, C. jensonii, C. vasqi/ezii and C. pseudolongifolius
grow along terraces above larger rivers. Vasquez ( 1 997) has described "Irapayal,"
"Yarinal" and "Supay Chacras" associations within the lateritic terra firme, but
with so few comparative, quantitative forest inventories completed for both
countries, I am not able to precisely list the Cybianthus species known from
each. The terra firme forests on white sand (varillal) are shorter in stature than
those on lateritic soils, and generally support lower numbers of lianas. Among
the varillal forest types described for Peru, Vasquez (1 997) lists "varillal seco,"
"varillal humedo" and "chamizal" or "ojo de varillal" as.sociations, only some
of which have been noted on label data. However, Cybianthus perinnanus , C.
spichigeri, and C. gigantophyllus occur on rolling hills in the varillal forest
type. Among those species which occur on steep hillsides near light gaps,
Cybianthus gigantopyllns is most notable, as it occurs in the ecotonal area
between forest gaps and mature forest. Cybianthus nanayensis, a subshrub, is
frequently found in gaps left by large treefalls in overmature forests, and
along trailsides, where it occurs in the rather dense herbaceous stratum. Cybianthus
resinosus, another inhabitant of the terra finite forest on white sand, occurs on
terraces above black water rivers in the forest, while C. nestorii is found in
the more open shrubby "varillal seco" transitional area near the riverbank.
Cybianthus Julvopulverulentus suhs\^. Diagnoliifolius typically occurs in campina,
or campinarana formations m Brazil, but in Peru it has been collected once
in the "varillal seco" an open shrubby area on white sand several hundred
meters from a black water riverbank. Unfortunately, no literature directly
addressing this forest type is known for Ecuador.
The two forest types subject to inundation have been divided mto varzea,
flooded by white water, and igapo, flooded by black or black and white water.
The other significant difference between these two forest type is that in varzea,
the forest is indundated for a much shorter time than that of the igapo. To
date, I know of no documentation for occurrences of forests inundated by
clear water in Ecuador or Peru, as they are found elsewhere in Amazonia
(Brazil, Colombia, Venezuela), but are best developed in Colombia. Three
species are known from igapo, with no endemics. Cybianthus guyanensis subsp.
pseudoicacoreus is found in the igapo at its limit with terra firme, while C.
penduliflorus is found well within the igapo and is frequently found in standing
water. Cybianthus spicatus is found both in igapo and varzea, and exhibits
both staminate and pistillate ecotypes, with some exhibiting apparent ran-
dom variation. It is a broadly ranging polymorphic ochlospecies (sensu Pipoly
1983a) with great morphological variation. The known Peruvian popula-
tions, from Huanuco and San Martfn, are identical to those found in cen-
tral Guyana (Pipoly 1 9H3a). The other varzea species, Cybianthus cydopetalus.
PiPOLY, Cybianchus in Ecuador and Peru 19
is known only from Madre de Dios in Peru, and from the Jurua area in Brazil.
It grows in the margin of varzea near its junction with the terra firme forest.
Forest associations within the vazea have been described by Vasquez (1997),
but I have been unable to match the corresponding complement oiCybianthus
species, owing to inadequate label data. The three forest associations present
in varzea for Peru include, "barrillal," "restinga" and "bajial." Clearly, more
fieldwork is needed to discern foristic differences among these association
types.
Premontane forest habitats are found from scarcely above 200 m to nearly
1,000 m. The forest is lower in stature than the terra fir me, have a greater
epiphyte load, and a larger number of lianas. Those on sandstone are dis-
tinguished here from those on other soils. On sandstone, three species are
found of which one, Cylnanthus timanae, is endemic. The other species, Cybianthus
comperuvianus , a new taxon described herein, is known from these forests in
Peru, Bolivia and adjacent Brazil. The other premontane forest formation
on lateritic soils hosts 9 species, Cyhtanthm minutiflorm, C. huanipamiemis,
C. granulosus, C. poepptgii, C. schlmiii, C. peruvianus, C. fosteri, and C. venezuelanus
and C. flavovirens. Among these, only Cybianthus venezuelanus, C. poeppigii,
C. schliniit and C peruvianus are not endemic. A surprising new distribu-
tion record for Cybianthus lepidotus, from Bagua Province, Imaza District,
of Amazonas Department, Peru, is recorded here. Cybianthus lepidotus was
once thought to be a Guayana Higland endemic, but was found in Bolivia
in the Maipiri region on the sandstone "laja" formations there (Pipoly 1992a).
This kind of disJLinction, concomitant with those for species such as Cybianthus
spicatus, and Cybianthus lineatus (see below), support recent thematic map
data at NASA, showing that the Ecuadorean/Peruvian area north, slightly
east, and immediately south of the Cordillera del Condor, contains significant
sandstone formations that may constitute "tepui satellites" (sensu Maguire
1979). Clearly, much more exploration in southeastern Ecuador and north-
ern Peru, should be a high research priority.
There are six species oiCybianthus known from cloud forests. Cybianthus
pastensts and C. incognitus are found in areas of high shade and moisture within
these forests. Along the margins to the leeward side, in that portion of the
Choco Floristic Province that extends into Ecuador, Cybianthus cuatrecasasii
may be found, while on the eastern Andean slopes of Peru, C. laetus grows
in exposed, wind-swept margins of the forest. The lack of records for Cybianthus
laetus in Ecuador is more likely a collection artifact than a reflection of its
rarity, given its occurrence in the Department of Boyaca, Colombia. It is
notable that Cybianthus patensis and C. incognitus, when growing at eleva-
tions below 1 ,000 m, are ridgetop species. Cybianthus magnus subsp. ?nagnus
is an obligate epiphyte in closed cloud forests, growing in the forks of trees
or on deep detritis, and shielded from winds.
20 Sum 18(1)
Elfin forests and montane thickets are transitional formations below the
jalca ( a formation like a paramo but without species of the Asteraceae subtribe
Espletiniae). Elfin forests host considerable numbers of trees and usually
have trees up to 5 m tall. The montane thicket (subparamo) grows in more
exposed areas at higher elevations and is dominated by low shrubs and small
trees to 2 m, with broad crowns. Cyhianthus viarginatiis is found in both of
these habitats, but the leaves and stature of the plants are much smaller in
the thicket formation. Also, the verrucose papillae of the stem are much
more pronotmced in the thicket habitats than in plants growing in the elfin
forest. Cyhianthus, viagnus subsp. asymmetries grows in open montane for-
ests or subparamo thickets, and elfin forests, where it is subject to exposure
to the high winds and rain. Its leal variation is significant, but it is easily
recognized from subspecies magnus by the hydropotes of the adaxial leaf surface
and orange punctations of leaf, inflorescence and perianth parts and the unique
white, then lavender fruits.
Finally, sandstone scrub, called "pajonal," is known thus far only from
Peru. It is the formation growing at the highest elevations where JVIyrsinaceae
occur, mostly well over 3,000 m. In these habitats, there are few, small shrubs
which rarely exceed 1.5 m tall. Cyhianthus I meatus is found in this habitat,
the first locality for this species outside the contiguous Guayana Highland.
No similar habitat has been described in Ecuador, but it may be present in
the Cordillera del Condor.
TAXONOMK; concepts, NOlliS ON KEYS AND SPECIMI-N CITA'l'IONS
JVty species concept follows that of Wiley ( 1 978, 1 98 1 ), who defined a spe-
cies as follows: "An evolutionary species is a single lineage of ancestor-descen-
dant populations which maintains its identity from other such lineages and
which has its own evolutionary tendencies and historical fate." My subspecies
concept (Pipoly 1987), defines a subspecies as follows: "groups of populations
within a single lineage of ancestor-descendant populations that show varia-
tion by unic|ue combinations of plesiomorphies, or homoplasic apomorphies,
correlated with biogeography and/or ecology. This rank is primarily used to
convey information regarding variation in the life histories of these populations
and character state differences hypothesized to be the result of this variation.
The subspecific rank in no way attempts to predict speciation events."
The keys are artificial and designed to expedite identification of herbarium
specimens. An attempt has been made to emphasize vegetative characters to
increase the keys' usefulness with sterile material. The numbers appearing be-
fore the taxa refer to their respective position in the key; any correlations with
phylogenetic relationships are coincidental. Quantitative and cjualitative
data presented in keys and descriptions for floral parts and bracts were taken
from organs rehydrated from herbarium specimens by boiling in water.
PiPOLY, Cybianthus in Ecuador and Peru 21
Measurements from these range from 10% to 15% greater than those mea-
surements taken directly from dried material. Data regarding stem diam-
eters, inflorescence rachises, pedicels, leaf, and fruit shape were taken from
dried herbarium specimens. Extra-Ecuadorean and -Peruvian specimens are
cited for all new species and for recent collections of other species used to
significantly amplify previously pubHshed morphological descriptions (Pipoly
1981, 1983a, 1983b, 1987, 1988, 1991, 1992a, 1993, 1994, 1995, 1996).
A description of the genus Cybianthus and a key to its subgenera in "Ecua-
dor and Peru are provided below. This description, along with that of the
subgenera and species that follow include features found in each taxon as a
whole, including those populations and species occurring outside Ecuador
and Peru. Phylogenetic studies applicable to species in this treatment may
be found in Pipoly (1987) and in the forthcoming Flora Neotropica treatment.
TAXONOMIC TREATMENT
Cybianthus Mart., Nov. Gen. Sp. PI. 3:87. 1831. nam. ettypnsc(ms.\G. Agoscini,
Acta Biol. Venez. 10:l4l. 1980.; Pipoly, Mem. New York Bot. Gard. 43:46. 1987;
Pipoly, Ann. iVIissouri Boc. Gard. 79:9 13.1 992. Type specii-s: C. penduUflorus Martius.
Terrestrial or epiphytic, monoaxial or polyaxial, dioecious, monoecious
or polygamous shrubs or trees to 1 5 m tall. Roots positively geotropic or dia-
geotropic. Branchlets glabrous, glandular-granulose, dendroid- and stellate-
tomentose, furfuraceous- or ferrugineous-stipitate-lepidote. Leaves sessile or
petiolate, alternate, subopposite, or pseudoverticillate, the venation
camptodromous or rarely acrodromous; petioles obsolete or when present,
canaliculate or marginate, tapering gradually to the base, or abruptly swol-
len toward the base, here termed "pulvinate." Inflorescence: staminate, pistil-
late, bisexual or polygamous, lateral (axillary), a simple raceme, panicle of
racemose or spicate (rarely corymbose) branches, a pleiochasium, or an in-
determinate umbel appearing racemose. Flowers functionally unisexual or
bisexual, 3-6(-7)-merous; calyx cotyliform to cupuliform, the lobes im-
bricate, valvate or aberrantly contorted, basally connate 1/5-2/3 their length,
abaxially glabrous, glandular-granulose, ferrugineous stipitate-lepidote, or
translucent-lepidote, adaxially glabrous, epunctate or prominently orange,
red or black punctate, the lobes entire to erose-fimbriate, glabrous or glan-
dular-ciliate; corolla rotate, subrotate, cupuliform or campanulate, rarely
infundibuliform or salverform, the lobes imbricate or valvate, basally con-
nate 1/5-3/4 their length, abaxially glabrous, glandular-granulose, or fer-
rugineous stipitate-lepidote, adaxially glandular-granulose at least at the
junction of the tube and lobe, the margin entire to erose-denticulate, gla-
brous, glandular-granulose or rufous glandular-papillate; stamens and staminodes
adnate to corolla tube at least 2/3 their length, the filaments variously con-
nate to form a tube, the staminal tube adnate to the corolla tube or at times
22 SiDA 18(1)
developmentally fused with it (thus the stamens appearing epipetalous),
bearing fleshy lobes alternate with the apically free portions of the filaments
or not, the anthers erect or distally curved, ovate, widely ovate, or triangu-
lar, basifixed or dorsifixed, apically acute, rounded, truncate, emarginate or
minutely apiculate, the apiculum erect, proximally or distally curved, ba-
sally truncate, cordate, or rarely hastate, deshiscent by apicaly pores, confluent
apical pores (birimose), or by wide or narrow longitudinal slits; pollen tricolporate,
psilate; staminodes morophologically similar to the stamens but greatly reduced
in size, the antherodes at times producing abortive pollen; pistl obnapiform,
ellipsoid, umbonate or obturbinate, the ovary sparsely to densely translu-
cent glandular-lepidote, the style glabrous, the stigma capitate, capitate-
lobate, or punctiform, persistent or early caducous, the placenta free-cen-
tral, carnose, umbonate or globose, the ovules campyiotropous ( l-)2-5(-7),
uni- or biseriate; pistillode conic, lageniform, obturbinate or irregularly shaped
vestigial pistillode, the pistillode hollow or bearing a sterile placenta, rarely
absent. Fruit drupaceous, 1(— 2)-seeded, the endosperm translucent, non-
starchy, the embryo small, linear, flexuous, erect or curved, longitudinal or
transverse, the cotyledons not well-developed.
Distribution. — One hundred sixty-seven species; Nicaragua, Costa Rica
south through Panama to the Andes southward to Bolivia, from Colombia
eastward across Venezuela and Brazil and the Guianas, then southeastward
to the Atlantic coastal forests of SE Brazil.
Ecology. — Members o'i Cybianthus '<iK principally riparian, occurring only
in primary forests or rarely in somewhat disturbed ones, and thus, may serve
as indicators of environmental quality. Throughout the range of the genus,
its members are known from wet tepuf savannas, moist scrub, cloud and
elfin forests (including "ceja de selva"), subparamo thickets, montane, premontane,
pluvial, wet and moist forests, paramo, jalca, igapo, varzea, varillal, campinas,
campo rupestre, restinga, cerrado, and caatinga vegetation types. In Ecua-
dor and Peru, the majority of the species occur in lowland and premontane
forests at the junction of Hylaea and the eastern slopes of the Andean Cor-
dillera (see ECOLOGY section).
Cybianthus is most closely related to the paleotropical lianous genus Embelia
(Pipoly 1 987), and cladistically defined by the unique glandular-granules at
the corolla lobe and tube junction. For practical purposes of identification, the
combination of lateral racemes or spikes, or racemose or spicate panicles, and
filaments which are shorter than the corolla, connate at least 1/4 their length,
and adnate to the corolla at least 1/3 its length, allows for easy recognition.
KEY TO StfHCFNERA Of CYBIANTHUS IN ECUADOR AND PliRl)
1. Corolla corylitorm, cupLiliform, campanulate, or rarely salverForm; anthers
longer than wide, disrally recurved, apically acure or minnrely apiculate.
PiPOLY, Cybianthus in Ecuador and Peru 23
2. Branchlets glabrous or glandular-granulose; anthers dehiscent by narrow
longitudinal slits I. Microconomorpha
2. Branchlets ferrugineous romentose or stipitate-lepidote; anthers dehis-
cent by wide longitudinal slits.
3. Branchlets and calyx ferrugineous stipitate-lepidote; abaxial corolla
surface glabrous near margin II. Conomorpha
3. Branchlets ferrugineous romentose, calyx glandular-granulose or gla-
brous, rarely ferrugineous romentose; abaxial corolla surface glandu-
lar-granulose near margm III. Laxiflorus
1 . Corolla rotate to subrotate; anthers wider than long, erect, apically rounded
to truncareor emarginate.
4. Petioles abruptly swollen basally; anthers dorsifixed, longitudinally
dehiscent.
5. Plants monoaxial; anthers erect, not versatile.
6. Stem glandular papillate, at times with hydropotes, bearing cataphylls
apically and at times, alternating with the leaves; leaves without
subepidermal fibers; flowers 4- or 5 -merous; corolla lobes epunctate
or inconspicuously pellucid or orange punctate, glandular-granulose
within; stigma large, capitare-lobate, early caducous, the lobe mar-
gins fimbriate IV. Comomyrsine
6. Stem with malpighiaceous hairs, without cataphylls, but at times
with aborted leaves (pseudocataphylls); leaves with numerous paallel
subepidermal fibers (most easily seen adaxially); flowers 3-merous;
corolla lobes prominently black punctate, maculate, glabrous within
except at lobe and tube junction; stigma small, capitate-lobate, per-
sistent, the lobe margins entire V. Triadophora
5. Plants polyaxial; anthers versatile VI. Weigeltia
4. Petioles obsolete or not abruptly swollen basally; anthers basifixed, poricidally
dehiscent.
7. Leaves sessile, apically mucronate, basally auriculate, the margins
scarious; staminal tube merely adnate to corolla tube VII. Grammadenia
7. Leaves petiolate, apically acute, acuminate or caudate, basally acute,
attenuate or cuneate, the margins opaque; staminal tube development-
ally fused to corolla tube, the stamens thus appearing epipetalous
VIII. Cybianthus
I. Cybianthus subgenus Microconomorpha (Mez) G. Agostini, Acta Biol.
Venez. 10:150. 1980; Pipoly, Wrightia 7:235. 1983. Conomorpha A. DC. subgenus
Microconomorpha Mez inEngl., Pllanzenr. IV. 236(Heft 9):251. 1902. Microconomorpha
(Mez) Lundell, Wrightia 5:349. 1977. Type Spi'cii^s: Conomorpha verticillata Zahlbr.,
Ann. K.K. Naturhist. Hofmus. 7:3. 1892, non Mez (1902). = Cybianthus pastensis
(Mez) G. Agostini (lectot'i-pi,: Agostini, Acta f^iol. Venez. 10:150. 1980).
Cybianthus Mart, subgenus Iteoides G. Agostini, syn. nov.. Acta Biol. Venez. 10:148.
1980. Type Species: BadiiLi itcoicks Benth., PI. Hartw. 217. 1896. Conomorpha iteoides
(Benth.) Mez in Engl., Pflanzenr. IV. 236(Hef't 9): 254: 1 902. Type Spiiciiis: Cybianthus
iteoides (Benth.) G. Agostini, Acta Biol. Venez. 10:149. 1980.
Terrestrial monoecious, clioecous or polygamous j7:??7/^i or sttiall trees. Bark
smooth to slightly fissured, light brown, thin. Root positively geotropic.
Trunk distinguishable, leptocaulous, the growtli dynamics following Rauh's
24 SiDA 18(1)
Architectural Model (Halle et al. 1978). Branchlets thin, terete, densely ter-
rugineous glandular-granulose, the granules often stipitate. Cataphylls and
pseudocataphylh absent. Leaves pseudoverticillate; blades petiolate, often with
translucent glandular lepidote scales. l}ifloresce)ice a simple raceme or bipin-
nate panicle, staminate, pistillate, or polygamous, the peduncle 1—4 cm long,
densely glandular-granulose; mflorescence and floral bracts, perianth and
pistil bearing prominently raised red or black punctations; infforescence bracts
large, often foliaceous and persistent; fioral bracts linear-lanceolate, the margins
glandular-ciliate, caducous, the pedicels erect, accrescent in fruit. Stami-
nate, pistillate and bisexual flowers monomorphic (similar in shape), the staminate
the largest, the pistillate the smallest in size, white to yellowish-green, (4—)
5(— 6)-merous; calyx cotyliform, the lobes valvate, the margins densely glandular-
ciliate; corolla cotyliform to campanulate, the lobes imbricate, glabrous without
except glandular-granulose near the margin, glandular-granulose over the
entire surface within; stamens and staminodes with a conspicuous staminal
tube, the apically free portions one to three times longer than the anthers,
the anthers elongate-triangular to ovate, prominently curved distally, apically
obtuse to apiculate, basally cordate to hastate, clorsifixed 1/3 to more than
1/2 length from base, dehiscent by narrow longitudinal slits, the staminodes
producing abortive pollen grains; pistil in pistillate and bisexual flowers
obturbinate, the ovary densely translucent-lepidote, the style thick, trun-
cate, the style punctiform, the placenta umbonate, bearing 3—4 uniseriate
ovules immersed in placental tissue, but exposed apically by placental pores;
pistillode similar to pistil but reduced in size, hollow or bearing 2 abortive
ovules. Fn/it drupaceous, 1 -seeded, the exocarp thin, prominently black punctate.
Subgenus Mtcroconomorpha contains 5 species, of which one, Cybianthus
pastensis (Mez) G. Agostini, is known from Ecuador and Peru. I earlier indi-
cated that there was no evidence to support subgenus Iteoides as a separate
entity (Pipoly 1987), so it is treated here in synonymy under subgenus
M icroconomorpha.
1. Cybianthus pastensis (Mez) G. Agostini (Fig. 3c), Acta Biol. Venez.
10:151. 1 980. Conomorpha pastensis Me/ in Engl., Pflanzenr. IV. 236(Hefc 9):252.
1902. Al/rmw/mor/;A<^/wmTO/.v(Mez)LLinclell,Wrigluia 5:349. 1977. Type: COLOMBIA.
Narino: "Paramo de Purugai, Prov. cie Pasco," 2,500 m, 1866 (stam. fl),_/. Triana
2585 (i.F.CTOTypK (Pipoly 1983a): W; isc)lectotypi;s: C, COL, G, P).
Aiyrsine verticillcita C Prcsl, Reliq. Haenk. 2:64. 1835. Conomoyphci vtrthillata {C. Presl)
Mez in Engl., Pflanzenr. IV. 236(Hcfc 9):252. 1902, mm Zahlbr. (1892). Conomorpha
prest//).[\ Macbr., Cantlollea 5:398. I 934. Mkrnconomoipljci verticillcita (C. Presl) Lundell,
Wrighria 5:349. 1977. Tyhi;: PERU. HcAnuco: without further locality, without date,
(stam. fl.), T. Haerike 98 (i.hctotyph (Pipoly 1983a): PR; isolectotypes: HAL, W). Non
Cybianth//s verticillatns (Veil.) G. Agostini, Acta Biol. Venez. 1():16S. 1980.
Coiiomorpija verticillata Zahlbr., Ann. K. K. Naturhist. Hofmus. 7:3. 1892, non C. Presl
PiPOLY, Cybianthus in Ecuador and Peru 25
(1835) Cmwnwrpha jehkii Mez in Engl., Pflanzenr. IV. 236(Heft 9):25l. 1902, nom.
superfl. Corioinorpha preslii ].V. Macbr. \-c\t. jelskii (Mez) J.F. Macbr., Field Mus. Nat.
Hist., Bot. Ser. 13:201. 1959- Muroconomorpha jehkii {Mfi) Lundell, Wrightia 5:349.
1977. Type: PERU. CajaxMARCa: Cutervo, Apr. 1879 (stam. fl.), C vonjelski 11 (ho-
lotypk: W; piioto and fragment, F, F Neg. 31980).
Conoiiwrpha (jeiitalei Mez in Engl., Pflanzenr. IV. 236(Heft 9):252. 1902. Microcono?Horpha
dentata (Mez) Lundell, Wriginia 5:3-49. 1977. Typi:: ECUADOR. Pichincha: In cor-
dillera from Quito to Tungurahua, 2, 000-3, 000 m, 1857-9 (stam. & bisex. fl.), R.
Spruce 3 173 (lpctotype (Pipoly 1983a): K; lsolec:t()TYPF,s: BM, BP, C, CGE, GH, GOET,
LD, LE, F Neg. 22956).
Conoiiwrpha q/ieni folia Mez in Engl., Pllanzenr. IV. 236(FIeft 9):253. 1902. Microamnniorpha
quercifoiia (Mez) Lundell, Wrightia 5:349. 1 977. Type: PERU: without locality, without
date (stam. \\.),J. Pavon s.n. (lectotype (Pipoly 1983a): G).
Conomorpha panamensis Lundell, Wrightia 5:290. 1976. Microconomorpha panamensis (Lundell)
Lundell, Wrightia 5:349. 1977. Cyhianthus morn G. Agostini, Acta Biol. Venez. 10: 154.
1980. Type: PANAMA. Chiriqui: Cerro Pando, on continental divide and Panama-
Costa Rica border, ca. 16 km W of Hato del Volcan, 2,000-2,482 m, 20 Jul 1975
(stam. fl.), S. Mon & A. Bnlten 7292 (hoeotype: LL-TEX; Isotype: MO).
Shrub or tree to 6 m tall. Branchlets and inflorescence densely ferrugineous
glandular-papillose, the branchlets angulate to prominently ridged, (1.5-)
2-3 mm diam. Leaves pseudoverticillate; blades membranaceous to coria-
ceous, narrowly oblanceolate to oblong or obovate, (3.0— )6. 5—14. 5(— 21.0)
cm long, (1.2-)2-4.5(-6.8) cm wide, apically attenutate, acute or acumi-
nate, basally cuneate, not decurrent on the petiole, prominently punctate
and minutely ferrugineous stipitate-papillose above and below, the midrib
impressed above, raised below, the secondary veins 7-15 pairs, prominently
raised below, the margin undulate, lobate, crenate or dentate, rarely subentire;
petioles marginate, (0.2-)0.5-2(-2.7) cm long, densely glandular-papillose.
Staminate, pistillate or polygamous inflorescence: monomorphic, erect or lax, a
simple raceme, 1.8-5.5 cm long, the rachis thin to thick, densely glandu-
lar-papillose; inflorescence bracts chartaceous, obovate to elliptic, (4.3— )6-
11 mm long, 3-7 mm wide, apically acute to acuminate, basally cuneate,
densely and prominently red punctate; floral bracts chartaceous, linear-lan-
ceolate, (0.8-)1.4-2.2(-7) mm long, 0.6-0.8 mm wide, apically attenuate,
caducous; pedicels cylindrical, (1 .3-)2-7(-7.5) mm long, densely glandu-
lar-papillose. Floivers (4-)5-merous, white to yellowish-green; calj^ chartaceous,
shallowly cotyliform, (0.6-)0. 8-1.1 (-1.5) mm long, unequally divided, the
tube 0.2-0.5 mm long, the lobes suborbicular to very widely ovate, (0.4-)
0.6-1 mm long and wide, rounded to acute apically, glabrous, densely and
very prominently orange or black punctate, the margin subentire to erose-
dentate, densely glandular-ciliate; corolla chartaceous, cotyliform, 2-2. 6(-
3.6) mm long, the tube 0.2-0.3 mm long, the lobes ovate to narrowly ovate,
1.7-2.3(-2.8) mm long, 0.8-1.3(-1.5) mm wide, highly reflexed at anthesis,
apically rounded to obtuse, prominently orange or black punctate; stamens
26 SiDA 18(1)
and staminodes 1-1.6(— 2.5) mm long, the staminal and staminodial tube
0.7—1.8 mm long, the apically free portions of the filaments 0.3—0.7 mm
long, the anthers elongate-triangular, 0.6-1.2 mm long, apically obtuse,
basally hastate, the connective red punctate ventrally and dorsally, dorsifixed
ca. 1/3 to slightly less than 1/2 from base; pistil and pistillode 1.2—1.8 mm
long, the ovary (0.6-)0.8— 1 mm long, 1—1.3 mm diam., densely translu-
cent glandular-lepidote, the style thick, 0.5-0.8 mm long, the stigma punctiform,
the pistillode hollow or containmg one abortive ovule. Fruit globose, green,
then red, then black at maturity, 3—4 mm diam. when dried exocarp thin,
prominently pellucid punctate.
Distribution. — Costa Rica to Colombia, southward to Peru, from 1,500-
3,200 m elevation.
Ecology CI nd conservation status. — Cyhianthus pastensis is known from premontane
and montane pluvial and cloud forests, and at elfin forest margins. Popula-
tions in areas exposed to winds have more coriaceous leaves and shorter stature,
frequently as small as one meter in height. The wetter the habitat, the more
membranaceous the leaves become, and the longer the inflorescences. Fieldwork
in Colombia has shown that populations may contain six individuals per
hectare, and that the population rapidly dwindles in areas of disturbance.
Owing to population dynamics thus far observed, Cybianthiis pastensis should
be considered threatened.
Etymology. — The specific epithet refers to the area from which the type speci-
men was collected, near the city of Pasto, Department of Nariiio, Colombia.
Representative specimens examined. COLOMBIA. Antioquia: Mpio. Urrao, Parqiie
National Natural "Las Orqui'deas," Vereda Calles, Permanent Premontane Rainforest In-
ventory Plot, right bank of Rio Calles, 06" 32' N, 76° 19' W, 1 ,450-1 ,=)()() m, 29 Nov
1993 (fr),./. Pipoly. A. Cogolloei al. 17322 (BRIT, COL, JAUM, MO), limits of Parqtie Las
Orqui'deas, left bank of Ri'o Calles, 1 ,450- 1 ,500 m, 30 Nov 1 993 (ster.),/. P/po/y. A. Coi^iillo
et al. 17376 (BRIT, JAUM, MO); near limit of Parque Las Orqui'deas, Alto de Palmitas,
ca. 1 km from INDLRENA C;abana Calles, 1,300-1,400 m, I Dec 1993 (ster.),^. P//?()/y.
A. Cogollodcd. I75()X n323 (BRIT,JAUM, MO), 2 Dec 1993 (fl bud),./. P/poIy. A. Cogdlo
etal. 17334 (BRIT, COL, JAUM, MO), Right bank of Rio Calles, 1,350-1,450 m, 7 Dec
1993 (stam. fl), /. Pipoly et al. 17881 (BRIT, COL, JALJM, MO); Along trail to Finca La
Quince, above LJrrao, 06° 30' N, 76° 10' W, 2,500-2,800 m, 21 Nov 1988 (stam. H), G.
McPhersoii cY al. 13212 (BRIT, HUA, MO); Mpio. Frontino, Region de Murrf, ca. 1 3 km
from Nutibara, 06" 40' N, 76" 20' W, 2,000 m, 9 Dec 1 988 (pist. fl, fr), G. McPher.m/ d al.
13397 (BRIT, HLJA, MO). Norte de Santander: San Antonio, W of Cali, near summit of
Cordillera Occidental, 1 ,9800- 2,350 m, 26 I'eb-2 Mar 1 939 (stam. fl), E. Klll/p&A. Gania
33886 (A, S, US). ECUADOR. Azuay: Chiguinda, on E slopes of cordiUera E of Sigsig,
03° 12' S, 78° 36' W, 1 ,600-1 ,800 m, 1889 (stam. fl), /■ Lehmarni 5 143 (K-2 sheets). Carchi:
Paramo de Achupallas, 01° 46' S, 78° 33' W, 2, 000-3, 000 m, 1899 (stam. fl), F. Lehmaiin
6202 (K-2 sheets); From Prima Vera about 6 hrs. hike up Ri'o Gualchan Drainage to Nilo
Ortiz' shelter, 00" 50' N, 77° 72' W, 1 ,9_^0-2,200 m, 7-8 Jun 1993 (fl bud),/. Bradford et
al. 53 (BRIT, MO, QC;NE). Loja; Cerro Bangala, ca. 10 km E of Yangana, 2,500-2,700 m,
I80ct 1988(pist. ri,fr),(7. H./r//w(,'253 /.)' (GB), (stam. fl), G7 Harling 23334 {GB)\CAmu'm
PiPOLY, Cybianthus in Ecuador and Peru 27
Loja, Carretera Loja-Zamora, at high point, 03° 58' S, 79° 04' W, 2,400-2,600 m, 23 Dec
1991 (pist. fl), D. Ri/hioetal. 2232 (BRIT, xMO, QCNE); Loja, 3,500 m, 1 Dec 1876 (stam.
fl), E. Andre 433 1 (F, K, NY); Divide between Quebrada Jipiru and E fork of Rfo Zamora,
W slope of Cordillera de Zamora (El Condor), 9 km E of Loja, 04° 00' S, 79° 06' W, 2,700
m, 19 Feb 1945 (stam. fl), F.R. Fosherfi & M. Giler 2.3119 (NY, US); Loma de Loro, 6 km S
of Saraguro, on Rd. to Loja, 3,200 m, 1 1 Feb 1985 (stam. fl), G. Harling & L. Andersson
21594 (AAU, S); Saraguro-Loja Rd., km 1 2.4, turnoff toward Fierro Urco, Km 3.8-7.1,
03° 42' 33" S, 79° 18' 03" W, 3,120-3,390 m, 7 Dec 1994 (pist. fl), P.Jorgenseii et al. 1297
(BRIT, LOJA, QCA, QCNE); W slope of Nudo de SabaniUa, ca. 8 km above Yangana on
Rd. to Valladolid, 2,300-2,500 m, 2 Apr 1985 (bisex. fl, fr), G. Harling & L. Andersson
23540 (GB); Cerro Toledo, Rd. to La Torre, ca. 7 km SE of Yangana, 2,500 m, 7 Apr 1985
(stam. fl), G. Harling & L. Andersson 23842 (GB). Morona-Santiago: Between Campanas
and Arenillas, along Ri'o Tintas, 10 leagues SE of EI Pan, 2,195 m, 13 Jul 1943 (stam. fl),
_/. Steyermark 33642 (F, NY); Above Mirador, 2,375 m, 9 Sep 1943 (stam. fl),J. Steyermark
'53897 (F, NY). Napo: 10 km W of Cuyuja, along Quito-Lago Agrio Rd., 00° 25' S, 78°
00' W, 2,700 m, 3 I Apr 1983 (stam. fl), H. Balslev4293 (AAU, QCA); Salcedo-Napo Rd.,
2,390-2,590 m, 7 Feb 1977 (fr),J, Brandbyge 42093 (AAU, QCA); E of Borja, Cerro Antisana,
28 Jul I960 (stam. fl), P. Grubb et ul. H)7 3 (OXF, NY); Sta. Barbara Scumbios, 00" 22' S,
77° 10' W, 2,700 m, 10-15 Feb 1959 (stam. fl), /.. Holm-Nielsen 681 8 {AAU ,QCh);^-l2
km ESE of Sta. Barbara, 00° 40' N, 77° 30' W, 2,780-2,880 m, 1 1 Jan 1985 (stam. fl),/,
Luteyn & E. Cotton llOll (GB, QCA, NY, VEN); Paso de Guamani, Rio Chalpi, at bridge
on Papallacta-Baeza Rd.., 2,800 m, 6 May 1967 (stam. fl), B. Sparre 13940 (GB); Canton
Quijos, Sierra Azul (Agri'cola Industrial Rfo Aragon), 00° 40' S, 77° 55' W, 2,300 m, 2
May 1992 (fr), A. Alvarez et al. 381 (BRIT, MO, QCNE), (fr), A. Alvarez et al. 412 (BRIT,
MO, QCNE), Campamento Estero Chico, 00° 41' S, 77° 56' W, 2,500 m, 18 Jun 1992
(fr), A. Alvarez et al. 490 (BRIT, MO, QCNE); Sierra Azul, Cordillera de Huacamayos, 00°
41' S, 77° 54' W, 2,500-2,700 m. 10 Feb 1994 (fr), A. Alvarez et al.l330 (BRIT, MO,
QCNE). Tungurahua: On Patate-Triunfo Rd., 01° 18' S, 78° 25' W, 2,950 m, 5 Nov
1983 (stam. fl),./, Brandbyge & A. Barford 42306 (AAU, QCA, QNA, S). Zamora-Chinchipe:
Rd. from Loja "to Zamora, km 14, 00° 04' S, 79° 09' W, 2,750-2,770 m, 19-20 Apr 1973
(stam. fl), L. Holm-Nielsen et al. 3965 (AAU, QCA). PERU. Amazonas: Prov. Luya,
Camporredondo-TuUanya, trail to Cerro Huicsocunga, 2,350 m, 3 Sep. 1989 (A), C Diaz
&J. Campos 37 11 (MO, USM); Parte aita de las Montaiias de Galeras, 2,000-2,500 m, 20
Jun 1991 (fl bud), C. Diaz et al. 4448 (BRIT, MO, USM). Ayacucho: Prov. La Mar, E
massif of Cordillera Central, opposing the Cordillera Vilcabamba between Tambo San Miguel,
Ayna and Hacienda Ltusiana, 12° 45' S, 73° 53' W, ca. 30 km SW of Hacienda and Rio
Apurimac, 21 Aug 1968 (bisex. fl), T. Dudley 119 15 (F, NA, US). Cajamarca: Prov. Cutervo,
10 km NW of Socota, 3,200 m, 10 Dec 1938 (stam. fl.), H. Stork & X. Horton 10134 (F);
San Andres de Cutervo, Parque Nacional de Cutervo, "Jalca," trail to Laguna "EI Pileo,"
2,680 m, 15 Mar 1989 (fl, fr), C. Diaz et al. 3330 (AMAZ, MO, USM); Prov. Jaen, E side
of Cordillera E of Huancabamba, 2,400-2,600 m, Apr 1942 (fr), A. Weherbai/er 6099 (F,
GH, US). Huanuco: Prov. Pachitea, region of Pucallpa, W part of Sira Mountains and
adjacent lowland, ca. 26-28 km ESE from Puerto Inca, 09° 25' S, 74° 43' W, 2,210 m, 15
Aug 1988 (stam. fl), B. Wallnbfer 1 1-16888 (BRIT, MO, W, WU, USM); SW slope of Rfo
LluUa Pichfs Watershed, on the ascent of Cerros del Sira, top of first cumbre between camp
4 (Peligroso) and camp 5 (Tabano), 1,680 m, 31 Jul 1969 (ster.), T. Dudley 13513 (NA).
Lima: Prov. Lima, Lima, without date (fr),y, Pavon s.n. (K). Madre de Dies: Prov. Manii,
Cerro dePanciacolla, RfoPlotoa, 10-15 km NNWof Shintuya, 12°35' S, 71°18' W, 1,000-
1,400 m, 15 Dec. 1985 (fr), R. Poster et al. 10860 (F, MO, USM).
Cybianthus pastens is may be easily recognized by its pseudoverticillate leaves
28 SiDA 18(1)
with variously serrate or incised margins, the very fine inflorescence rachis
and minute flowers, and tlie stipitate papillae of the branchlets, leaves, petioles
and inflorescence rachises. The prominent ridges of older branchlets and
the swollen pseudoverticels of leaf scars are also distinctive.
II. Cybianthus subgenus Conomorpha (A. DC.) G. Agostini, Acta Biol.
Venez. 10:150. 1980; Pipoly, Ann. Missouri Bot. Gard. 79:908-957.
1 992. CoNoi/iorphj A. DC, Trans. Linn. Soc. Loncli)n, Bor., 17:1 02. 1 83-1; Co)!(ini(>rphii
sect. Eiimnniwiylihct Mit|., Stirp. Surinam. Select. 111. 1850; Couoiiiorpha subgenus
Eucommorpha Mcz in Engl., Pflanzenr. IV. 236(Heft 9):254. 1902. Typi- Sppcirs: Cononnnphc)
ohlongifolia A. DC. = Cybianthus ohlovgijoliiis (A. DC.) G. Agostini (lkcioiype: by
Agostini, Acta Biol. Venez. 10:1 VI. 1980).
CoiiDDioyphii sect. Aai)ifiinorpl:ki Miq. in Mart., I'l. Bras. 1 0:304. 1856. Type Spi;(:ii;s: Conoinoypbu
heteranthd Benth. (lec;t()[YPe: Agostini, Acta Biol. Venez. 10:151. 1 980)= Cybhnithus
y^/iyam-Hsis (A. DC^. ) Mic]. subsp. i^//yai/ti/s/s.
Terrestrial dioecious, bisexual, polygamous, or rarely, monoecius shrubs
or trees. Roots positively geotropic. Bark smooth or fissured, brown, or beige,
rarely with significant amounts of cork. Ih/nk distinguishable, leptocaulous,
the growth dynamics following Rauh's or rarely, Aubreville's Architectural
Model (Halle et al. 1978). Branchlets thin to moderately thick, terete or ridged,
densely to moderately covered with ferrtigineous stipitate-lepidote scales,
the scales at times appressed, rarely glabrescent. Cataphylls and pseudocataphylls
absent. Leaves alternate, rarely approaching pseudoverticillate (C peruvianus),
petiolate, covered with ferrugineous stipitate lepidote scales, often glabrescent
above; petioles canaliculate, marginate, or rarely winged. Inflorescence race-
mose, spicate or paniculate, the panicles with racemose branches, rarely a
solitary flower; inflorescence bract small, lanceolate, early caducous; rachis
erect or lax, straight or rarely tortuous, ferrugineous stipitate-lepidote; floral
bracts deltate, lanceolate or ovate, ferrugineous stipitate-lepidote, inserted
at the base of the pedicel; pedicels cylindrical, at times clavate in frtiit or
absent, erect, apically recurved, pendent, or nodding, at times accrescent
in fruit. Flowers unisexual, rarely bisexual, dimorphic, (3-)4-5(-6)-merous;
calyx cotyliform, cupuliform, crateriform, urceolate or patelliform, the lobes
valvate, epunctate or with prominent (raised and blisterlike), conspicuous
(readily visible buy flat), or inconspicuous brown, red or black punctations,
the margin entire, rarely crenulate or erose; corolla campanulate to cupuliform,
rarely salverform or tubiform, the lobes erect or reflexed, rarely cucuUate,
valvate or imbricate, ferrugineous stipitate-lepidote or glabrous and epunctate
or prominently, conspicuously or inconspicuously pellucid, brown, or black
punctate without, at times with a narrow line of glandular-granules along
the margin, gland ular-granulose within, the margins enrire or rarely crenulate,
glabrous or rarely glandular-granulose; staminodes resembling stamens but
reduced in size, the tube conspicuous or inconspicuous, adnatc to the co-
PiPOLY, Cybianthus in Ecuador and Peru 29
rolla, lobate or elobate, the anthers ovate or triangular-ovate, rarely linear-
lanceolate, rarely deltate, usually recurved distally, rarely erect, apically acute,
or apiculate, rarely rounded, the apiculum dorsally, rarely proximally re-
curved or erect, the base cordate, dorsifixed from near base to subversatile,
the connective punctate or not; pistillode conic to lageniform, rarely ab-
sent, translucent-lepidote or glabrous, hollow; pistil obnapiform, rarely conic,
the ovary globose, lobed or with an apical apophysis, the style short, the
stigma capitate-lobate, 2-3-lobed or punctiform; placenta cupuliform or
cotyliform, the ovules 2-4. Fri/it subglobose, one(-two)-seeded.
Cybianthus subgenus Conomorpha contains 44 species, 10 of which have
been recorded from Ecuador and Peru.
KEY TO SPECIES OF CYBIANTHUS SUBGENUS CONOMORPHA
1. Branchlets with erect stipitate ferrugineous lepidote scales, their margins
not appressed; leaf blades subbullate to bullate, the secondary veins some-
what to deeply impressed above, prominently raised below; corolla inlundibu-
liform or tubiiorm, or appearing so in bud.
2. Branchlets flexuous, 4-5 mm diam.; leaf blades perpuncticulose above,
sparsely lepidote below, the secondary veins 22-26 pairs, the margin ir-
regular; inflorescence tortuous, pinnarely to bipinnately paniculate; co-
rolla lobes prominently keeled, rugose without; anthers ventrally
recurved 2. C. gigantophyllus
2. Branchlets straight, 2-3 mm diam.; leaf blades not perpuncticulose above,
ensely lepidote below, the secondary veins 8-19 pairs, the margin regu-
lar; inflorescence erect, a simple raceme or poorly formed panicle consist-
ing of basally clustered racemes; corolla lobes flat, smooth or verrruculose
without; anthers dorsally recurved.
3- Secondary veins 12-16; staminal tube epunctate; pedicels cylindric;
calyx cotyliform; corolla membranaceous, infundibuliform, verruculose
without; fruit smooth, 3.5-4.5 mm diam.; plants of premontane plu-
vial forests, subparamo thickets and upper pluvial cloud forests, 1,000
1,960 m elevation 3. C. occigranatensis
3. Secondary veins 8-12; staminal tube punctate; pedicels obconic; calyx
urceolate; corolla carnose, tubiform, smooth without; fruit costate, 7-
15 mm diam.; plants of white sands or on sandstone, 150-180 (-1,500)
m elevation 4. C. spichigeri
1 . Branchlets with appressed ferrugineous lepidote scales, the margins appressed;
leaf blades not subbullate or bullate, the secondary veins planar or slightly
raised above, barely discernible or slightly raised below; corolla campanu-
late to cupuliform.
4. Leaf blades coriaceous, rarely chartaceous, the margins subrevolute to
revolute.
5. Leaf blades densely and prominently pustulate at maturity above, the
secondary veins 24-28 pairs, inconspicuous below; inflorescence spi-
cate (2-)6-l6 cm long; flowers subsessile, the pedicels 0.2-0.6 mm
long; calyx deeply cupuliform 5. C. lepidotus
5. Leaf blades essentially smooth or sparsely pusticulate at maturity above,
30 Si DA 18(1)
thf secondary veins 14—25 pairs, prominently raised below; inllorescence
a raceme or jianicle witli 2-4 racemes branching from base, (2-)3-7 (-
8) cm long; Bowers pedicellate, the pedicels ((>.')-)(). 9- 1 .5 mm lon^;
calyx cocylitorm.
6. Branchlets stibterete, 2-3 mm diam; leaf blades 1 .2-2(-2.5) cm wide,
smooth above; petioles '3~7(-l()) mm long; staminate calyx carnose,
1 .2—1.8 mm long; staminate corolla carnose, densely lepidote with-
out, the scales overlapping, 3.2-3.'1 mm long, the tube equal to the
staminal cube, the lobes symmetric; anthers ovate; fruit with fleshy
exocarp; plants of montane and clotid forests on sandstone 6. C. laetus
6. Branchlets terete, 3~4 mm diam.; leaf blades (2.6-)3.5-3 cm wide,
pusticulate above; jietioles 10-15 mm long; staminate calyx
chartaceoLis, 0.8-1.2 mm long; staminate corolla chartaceous,
glabrous or sparsely lepidote without, the scales not overlapping,
2.0-2.6 mm long, the tube shortet than the stammal tube,
the lobes asymmetric; anthers linear-lanceolate; Iruic with thin
exocarp; plants of lowland and lower montane forests on white
sands 7. C. peruvianus
4. Leaf blades chartaceous to membranaceous, the margins flat.
7. Leaf blades membranaceous to subchartaceous; petioles 5-1 0(-l 2) mm
long; inflorescence a simple raceme or rarely 2-branched at base, 1-3
cm long; corolla salverform or campanulate; fruit globose.
8. Branchlets angulate, 1 .5-2 mm diam.; corolla salverform, the stami-
nate 2,2-2.6 mm; plants of premontane forests on sandstone and
limestone, (244-)400-l ,200 m elevation 8. C. comperuvianus
8. Branchlets terete, 2-] mm diam.; corolla campanulate, the stami-
nate, 2.8-S.2 mm; plants of lowland igapo forests, 90-240(-700)
m elevation 9- C. guyanensis stibsp. pseudoicacoreus
7. Leaf blades chartaceous; petioles ( 1 0-) 1 3- I 7(-22) mm long;
mflorescene a panicle with 2-8 racemes branched from base, 4-8 cm
long; corolla chartaceous, infundibuliform or c(Jtyliform; fruit de-
pressed-globose.
9. Branchlets, petioles, abaxial leaf blades, inflorescence antl calyx lobes
moderately to densely lepidote, but the not scales overlapping; leaf
blades smooth above at matutity; corolla cotyliform, the lobes ob-
long to oblanceolate, flat, smooth without, conspicuously black
l^tinctate, apically acLiminate; staminal and staminodial ttibe charta-
ceous, conspicuous; anthers and antherodes obcordate, the apiculum
distally recurved; pistillode conic; pistil lagenitorm 10. C. tinianae
9. Branchlets, petioles, abaxial leaf blades, inflorescence and calyx lobes
moderately to densely lepidote, the scales overlapping; leaf blades
pustulate above at maturity; corolla infundibuliform, the lobes ovate,
verruculose without, iiicons|Ticuotisly brown |uinctate, apically
rounded; staminal and staminodial cube membranaceous, inconspicu-
otis; anthers ovate, antherodes subdeltate, the apiculum proximally
inllexed; pisiillode lageniform; |-iistil obnapiform I I . C. cuatrecasasii
2. Cybianthus gigantophyllus Pipoly, (Fig. 2 A, 10). CandoUea 46:4 1 . 199 1 •
Bafsapuertoand Moyobamba, 600-1,200
PiPOLY, Cybianthus in Ecuador and Peru
31
Fig. 10. CybicmthHi gigantophylliis Pipoly. A. Habit, siiowing flexuous branchlet, paniculate
inflorescences. B. Pistillate flower, showing urceolate calyx and crenulate corolla lobe mar-
gins. C. Pistdlate flower with one corolla lobe removed, showing cucullate corolla lobe
apices, proximally recurved anrherodes, and capitate, lobed stigma. A— D, drawn from ho-
lorype, by Peggy Duke. Figure reproduced from Pipoly, 1991-
32 SioA 18(1)
m, Aug-Sep 1933 (pist. tl), G. King 3165 (noi.OTYPE: US; isotypf.s; l\ G-2 sheets,
GH-2 sheets, MO, NY, US).
Tree to 4 m tail. Branchlets flexuous, prominently ribbed, 4-5 mm diam.,
moderately lepidote. Leaves alternate; blades chartaceous, elliptic, (15.5)17-
27 cm long, (5.9— )7-9.1 cm wide, apically long-acuminate, the acumen
1 .2-3.5(-4) cm long, pustulate, perpuncticulose and glabrous above, sparsely
lepidote below, midrib slightly depressed above, prominently raised below,
the secondary veins 22—26 pairs, slightly depressed above, prominently raised
below, the margin irregular, flat, entire; petioles canaliculate, thick, (1.6—)
2.7—4 cm long, ca. 3 mm diam., sparsely lepidote, prominently ridged below.
Staminate inflorescence: unknown. Pistillate inflorescence: a pinnate to bipinnate
panicle, 1.5-2.5 cm long, tortuous, the branches spicate, moderately lepi-
dote; peduncle 0.3-0.5 cm long, floral bracts carnose, deltate, 0.8-0.9 mm
long and wide, apically acute, margin crentdate basally, densely lepidote
above and below; pedicels obsolete. Pistillate flowers 4-merous; calyx carnose,
urceolate, 1.6—1.8 mm long, the tube 0.9—1 rnm long, the lobes widely
triangular, 0.5—0.7 mm long, 1-1.2 mm wide, apically acuminate-apicu-
late, the margin regular, entire, lepidote; corolla carnose, campantdate, 2.7—
3.1 mm long, the tube 0.2—0.3 mm long, the lobes erect, 2.5-3 mm long,
apically rounded to obtuse, prominently cucuUate, abaxially carinate, apically
rugose and glandular-granulose along the margins without, inconspicuously
black punctate, the margin glandular-gran tdose, erose-crenulate; staminodes
2.3—2.5 mm long, the staminodial tube membranous, inconspicuous, 0.2—
0.3 mm long, elobate, glabrous, the apical free portions of the filaments
1.2—1.3 mm long, flat, the anthers ovate, 0.8-1 mm long, 0.6—0.8 mm wide,
apiculate, the apiculum ventrally recurved, basally cordate, the connective
epunctate; pistil obturbinate, 1.8—2 mm long, 1—1.3 mm diam., the ovary
1.4—1.6 mm long, the stigma capitate, 3— 5-lobed, the placenta cupuliform,
ovules 3, erect, the upper portions exposed. Fruit globose, 4—5 mm long
and in diam., exocarp thin, black, inconspicuously pellticid punctate.
Distribution. — Cybianthus gigantophyllus is known from the headwaters of
the rios Marafion and Huallaga in San JMartin, and the Iquitos area, along
the Ri'os Napo, Nanay and Amazonas in Loreto, at 130—500 m elevation.
Ecology and conservation status. — Cybianthus gigantophyllus occurs in pri-
mary terra flrme forests, and on white sands (varillal) of lowland Peruvian
Amazonia. Given increasing pressure from deforestation, it should be con-
sidered threatened.
Etymology. — The specific epithet refers to the leaf size, one of the largest
known for the subgenus.
Local names. — Peru: "ukushnum," "wewe," "yakCisnum," "yakushnum"
(Aguaruna).
PiPOLY, Cybianthus in Ecuador and Peru 33
Representarive specimens examined. PERU. Amazonas: Prov. Bagua, Dtto. Imaza,
Comunidad Aguarana de Putuim (CAMPOU), anexo Yamayakat, Monte Alco de Putuim,
450 m, 25 Aug 1994 (fl bud), C. D/az et al. 7007 (BRIT, HUT, MO, USM); Quebrada
ChichijamEncsa, RioCenepa, 130 m, 7 Jun 1973(fr), £. Ancuash 580 {AMAZ, MO, NY);
Vicinity Huampami, 5 km E of Valdivia, 04° 30' S, 73° 30' W, 200-500 m, 12 Aug 1978
(fr), £. Ana/ash 1437 (AMAZ, BRIT, MO, NY, US). Loreto: Maquisapa, Upper Rio Nanay,
Jul 1929 (fr), U. Willuims 1 182 (F); Prov. Maynas, Dtto. Sta. Man'a de Nanay, Caseri'a Mishana,
halfway between Iquitos and Sea. Maria de Nanay, 03° 50' S, 73° 30' W, 130 m, 25 Feb
1979 (ster.), A. Gentry &J. Aronson 25044 (AMAZ, MO); Dtto. Las Amazonas, Quebrada
Yanamono, Explornapo Tourist Camp, above mouth of Ri'o Napo on Rio Amazonas, 9 Nov
1979 (ster.), A. Gentry et al. 27952 (AMAZ, MO), 25 km NE of Iquitos, along Ri'o Amazonas,
southern perimeter path, 1 10 m, 27 Sep 1990 (ster.),/ Fipolyetal. 72^97 (AMAZ, BRIT,
MO, US, USM), 03° 20' S, 72° 55' W, 100-140 m", 15 Feb 1991 (ster.),/ Pipoly et al.
13028 (AMAZ, BRIT, MO, USM), 03° 28' S, 72° 50' W, 106 m, 15 May 1989 (ster.), R.
Vdsqiiez et al. 12 108 (AMAZ, MO, USM); Explornapo Tourist Camp, near Sucusari, along
Ri'o Napo, 03°20' S, 72°55' W, 100-140 m, 22 Feb 1991 (ster.), J. Ptpoly et al. 13284
(AMAZ, BRIT, MO, USM), 23 Feb 1991 (ster.),/ Ptpolyetal. 13423 (AMAZ, BRIT, MO),
(ster.),/ Ptpolyetal. 13426 (AMAZ, BRIT, MO, USM), 1 Mar 1991 (ster.),/ Ptpolyetal.
13931 (AMAZ, BRIT, MO, USM).
Cybianthus gigantophyllus is most closely related to C. occigranatensh (Cuatrec.)
G. Agostini and C. spichigeri Pipoly. However, the large, flat leaves, long
petioles, and tortuous panicles allow for easy recognition. In the original
description (Pipoly 1991), I described the pistillate corolla as tubular, when
it is, in fact, campanulate. When the flower is in bud, the corolla appears
tubular as it longitudinally extends above the calyx, then it gradually opens,
with cucuUate apices. Within the tall terra firiiie forests on lateritic soils, it
may be found above the flood line along small creekbeds.
3. Cybianthus occigranatensis (Cuatrec.) G. Agostini, Acta Biol. Venez.
10:155. 1 980. Conomorpha occigranatensis Cuatrec, Revista Acad. Colomb. Ci . Exact.
8(31):320. 1951. Type: COLOMBIA. Valle Del Cauca: Cordillera Occidental, W
slope, Ri'o Digua River Basin, left bank of Rfo San Juan, around Queremal region,
small stream at km 51, 1,540-1,650 m, (stam. fl),/ C/iatrecasas 237 34 (iiolotypi;:
F; isotype: COL).
Shrub or small tree to 4 m tall. Branchlets straight, subterete, 2—3 mm diam.,
densely lepidote. Leaves alternate; blades membranaceous, elliptic to obo-
vate, (4-)7.5-l4(-21) cm long, (2.5-)4-5(-7) cm wide, apically acuminate,
the acumen 1 .2— 1 .5(— 3.0) cm long, basally acute, decurrent on the petiole,
the midrib impressed above, prominently raised below, the secondary veins
8—12 pairs, deeply impressed above, prominently raised below, the leaf strongly
bullate, adaxial surface smooth, densely lepidote when young, becoming
pusticulate and sparsely lepidote or glabrous with age, abaxial surface densely
lepidote, but the scales not overlapping; petioles canaliculate, 1.0— 1.5(—
1.8) cm long, densely lepidote, persistent. StaniDiate inflorescence a raceme or
a panicle with 1-3 branches from the base, 4—8 cm long; peduncle, rachis,
34 Si DA 18(1)
branches and pedicels densely lepidote; peduncle 0.1-0.4 mm long; floral
bracts membranceous, ovate, shorter than the pedicels, 0.7-1.1 mm long;
0.4-0.5 mm wide, apically acute, densely lepidote abaxially, the margin
entire; pedicels cylindric, thin, 1.5-6 mm long. Stamincite flowers 4-merous;
calyx carnose, cotyliform, 0.8-1.0 mm long, the tube 0.2-0.3 mm long,
the lobes triangular to deltate, 0.5-0.9 mm long, 0.4-0.8 mm wide, apically
attenuate to an acute or round tip, sparsely lepidote without, glabrous within,
conspicuously brown punctate, the margin lepidote; corolla membranaceous,
campanulate, 2.4-2.7 mm long, the tube 0.7-0.8 mm long, the lobes ovate,
1 .6—1 .8 mm long, 1 .0-1.3 mm wide, apically attenuate to a round tip, veraiculose
without, smooth within, sparsely lepidote without toward iipex, apically
glanduiar-granulose within and along margins, conspicuously brown punctate,
the margins entire; stamens 1.8-1.9 mm long, the filaments 2.6-2.8 mm
long, the tube membranaceous, inconspicuous, adnate to the corolla tube,
elobate, the apically free portions 0.2-0.3 mm long, the anthers triangular,
0.8-1.0 mm long, 0.5—0.6 mm wide, apically attenuate to an acute , dor-
sally reflexed tip, basally cordate, dorsifixed just above base, the connective
dark, prominently brown punctate; pistiUode lageniform, 1.3-1.5 mm long,
densely translucent glandular-lepidote near the base. Bisexual and pistillate
inflorescence: as in staminate but only rarely branched from base, 4—6 cm long;
peduncle 0.1—0.3 cm long; floral bracts 0.5-0.8 mm long; pedicels 1.5—
3.5 mm long. Bisexual and pistillate flowers as in staminate but calyx 0.8—
1 .1 mm long, the tube 0.3—0.6 mm long, the lobes deltate to oblate, 0.4—
0.6 mm long, 0.8—1 mm wide; the margin irregular, entire; corolla as in
staminate but 2.4-2.6 mm long, the tube 1 .0-1 . 1 mm long, the lobes ovate,
1.4—1.6 mm long, 0.8—1.1 mm wide. Bisexual flowers with stamens 1.6—
1.8 mm long, the tube 1.0-1 .2 mm long, the apically free portions of fila-
ments 0.2-0.3 mm long, the anthers 0.5—0.8 mm long, pistillode 1.5—1.8
mm long. Pistillate flowers with staminodes 1 .6—1 .8 mm long, the tube 1 .0-
1.2 mm long, the apically free portion of filaments 0.1-0.2 mm long, the
antherodes 0.5-0.7 mm long; pistil obnapiform, 2.4-2.6 mm long, the ovary
1.1 — 1 .2 mm long, 1. 1-1.2 mm diam., the style 1 .0 mm long, the stigma
capitate, 2-lobed, to 0.2 mm long, the ovules 2-4, buried in the placenta
below apical pores. Fr///> globose, 2.5-4 mm long, 3.5—4.5 mm diam., the
endocarp smooth, the aril scanty and adnate to both seed and endocarp, the
embryo straight, ca. 3 mm long.
Distribution. — Panama (Darien), Colombia (Cordillera Occidental) and
Ecuador (Esmeralda, Napo, Santiago-Zamora, Sucumbios), at 1 ,000— 1 ,960
m elevation.
Ecology aiul conservatu))! status. — Cyinaiitbus occigraiiatensis occurs in premontane
pluvial forests, subparamo thickets and in upper pluvial cloud forests. Based
on my observations of populations in subparamo thickets at the Antioquia/
PiPOLY, Cybianthus in Ecuador and Peru 35
Choco interface in tiie Cordillera Occidental of Colombia, this species tol-
erates disturbance well as long as the soil is not compacted. It is retricted
to areas where rainfall exceeds 5,000 mm annually. At this time, the spe-
cies does not seem to be threatened.
Etymology. — The specific epithet refers to its principal range of distribu-
tion, the Cordillera Occidental of Colombia and adjacent Ecuador.
Specimens examined. PANAMA. Darien: S slojie of westermost summit of Cerro Tacaracuna,
massif between Pucro base camp and Tacaracuna summit camj-i, 1,4()0— 1,600 m, 21 Jul
1976 (stam. fl), A. Gentry et a l. 16867 (COL, LL-TEX, MO, PMA). COLOMBIA. Antioquia:
Mcpio Frontino, km 13 Nutibara-La Blanquita Rd., Region de Mum', Alto de Cuevas,
06° 44' N, 76° 23' W, 1 ,990 m, 6 Nov 19<S8 (fl bud),./. Zarucchi et al. 1201 (BRIT, HUA,
MO); Mpio. Frontino, Vereda Venados, Parque Nacional Las Orquideas, sector Dos Bocas,
confluence of Rio Venados and Rio Calles, ()6° 34' N, 76° 30' W, 29 Oc 1986 (stam. fl), R.
Calkjas et al. 2737 (HUA, MO); M]iio. San Lui's, Autopisra Medelli'n-Santafe de Bogota,
sector Rfo Samana, Rd. toward Vereda La Josefina, I (S Dec 1982 (stam. fl), A. Cogollo & C.
Estrada 296 (COL, JAUM, MO); Mpio. Urrao, Parque Nacional Las Orqufdeas, Vereda
Calles, Permanent Premontane Rainforest Inventory Plot, right bank of Rfo Calles, 06°
32' N, 76° 19' W, 1,450 m, 26 Nov 1993 (ster.),^, Pipoly. A. Cogollo et al. 17 1 59 (BRIT,
JAUM, MO), 27 Nov 1993 (ster.),^. Pi/mly, A. Cogollo et al. 17182 (BRIT, JAUM, MO),
Range NW of Cabana de Calles, 1 ,450 m, 28 Nov 1993 (ster.), A. Cogollo etal. 7529 (BRIT,
JAUM, MO), 1,450-1,500 m, 28 Nov 1993 (ster.),/ Pipoly etal. 17253 (BRIT, JAUM,
MO), (ster.),/ Pipoly et al. 17281 (BRIT,JALJM, MO), 7 Dec 1993 (.ster.),/ Pipoly et al.
17871 (BRIT, JAUM, MO), 9 Dec 1993 (fl. bud),/ Pipoly et al. 17979 (BRIT, JAUM,
MO), Vereda Calles, Alto de Palmitas, ca. 1 km from Cabana de Calles, 1 ,700-1 ,750 m, 2
Dec 1993 (ster.),/ Pipoly etal. 1 7542 (BRIT, JAUM, MO). Choco: Mpio. Itsmina, Quebrada
Raspadura, between Raspadura and Quibdo, split of Rfo Atrato and Rfo San Juan drainage
basins, ca. 05° 15' N, 76° 38'W, 18 Apr 1979 (fr), E. Forero & R. Jaramillo 5307 (COL,
MO); Serranfa del Darien, along Colombian/Panamanian border, 1,400 m, 20 Jul 1976
(stam. fl, bisex. fl), A. Gentry. H. Leon & L. Porero 16842 (COL, MO); without locality, 1866
(fr),/ Triana 2589 (G). Huila: Rfo Suaza, SW of Alejandrfa, 1 ,670 m, 23 Aug 1 944 (stam.
fl), E. Little 8532 (COL, US). Quindi'o: Mariquita, 1866 (stam. fl),^, Triami 2562 (P).
Valle Del Cauca: Cordillera Central, 5 km N of Darien along Rd. toward La Guajira, Upper
Rfo Calima, 03° 58' N, 76° 28' W, 1,550-1,700 m, 24 Jan 1986 (fl bud), B. Stein & L.
McDacle 3284 (BRIT, HUA, MO); Finca Zungara, Corregimiento La Divisora, crest of Cordillera
Occidental, W of Cali, 6 km N of Cali-Buenaventura Hwy, 03° 32' N, 76° 35' W, 1,960
m, 12 Dec 1985 (ster.), A. Gentry etal. 53167 (COL, MO, US), 24 Mar 1986 (fr), A. Gentry
etal. 53551 (COL, MO, US); Rfo Digua Drainage Basin, Piedra de Moler, 900-1,180 m,
20 Oct 1943 (pist. fl, fr),/ Cuatrecasas 14918 (COL-2 sheets, F); Rfo Sanquininf, La La-
guna, 1,250-1,400 m, 10 Dec 1943 (stam. fl),/ Cuatrecasas 15658 (COL, F, US); Monte
La Guardia, La Carbonera Range, between Las Brisas and Alban, 1,950—2,000 m, 16 Ocr
1946 (stam. fl),7. Cuatrecasas 2213 1 (COL, F, US, YEN); San Antonio, W of Cali, 1 ,900-
2,350 m, 26 Feb 1 939 (stam. fl), E. KilUp&A. Garcia 33898 (A, BM, COL, F, NY, US); La
Cumbre, 7 May 1 922 (stam. fl), P. Penmll5l47 (GH, K, NY, US). ECUADOR. Esmeraldas:
San Lorenzo Canton, Reserva Etnica Awa, Parroquia Alto Tambo, Centro de la LJnion, Cation
del Rfo Mira, 00° 52' N, 78° 26' W, 250 m, 22 Mar 1993 (fr), C. & M. Ai/lestia 1313
(BRIT. MO, QCNE), Napo: Carrerera Nueva, Cotundo-Coca, 1 ,130 m, 5 Aug 1984 (pist.
fl, fr), C. Doclson et al. 15115 (MO); Canton Archidona, 1 50 m NE of Caserio of Huamanf,
right side of Carretera Hollfn-Loreto, 00° 43' S, 77° 36' W, 1,200 m, 9 Sep 1988 (fr), R
Hurtado & D. Neill 235 (MO, QCNE), Cordillera de Guacamayos, Rd. to Archidona, Rfo
36 SlDA 18(1)
HoUin PeqLicno, primary forest on 90° slopes, 00° 38' S, 77" 48' W. 1 ,900 m, Aug I 990
(stam. fl), W. Pcilacnis & E. fveire 4S99 (BRIT, MO, QCA); Canton El Chaco, Right margin
of Rio Quijos, Finca "La Ave Brava," of Scgundo Pacheco, 00° 12' S, 77° 39' W, 1,800-
1 ,900 m, 7- 1 Sep 1990 (fr), W PaLnws 5394 (BRIT, MO, QCNE); S slope of Volcan Reventador,
left bank of Ri'o Reventador, between Rd. and trail to crater, 00° 07' S, 77° 30' W, 1 ,600-
1 ,850 m, 1 1 Oct 1990 (stam. H), W. Paiacios 61 76 (BRIT, MO, QCNE), (stam. tl), W. Pa/aaos
6187 (BRIT, MO, QCNE), (fr), W. Pa/aaos 6218 (BRIT, MO, QCNE); Proyecto Hidroelectnco
(]oca, Punto ST4; right margm of Rfo Quijos, ca. 10 km S of Reventador, 00° 11' S, 77°
39' W, 1 ,500 m, 3-5 Oct 1990 (pist. fl), W. PaU/os 58 H (BRIT, MO, QCNE), 08° 08' S,
77° 30' W, 1,450 m, 6-10 Oct 1990 (pist. Il), W. Paiacios 6040 (BRIT, MO, QCNE); Yasum'
National Park, Maxus Rd and pipeline construction project, km 15, 00° 3 1 ' S, 76° 32' W,
250 m, 30 Jtin 1 994 (bud) N. P/tma/i 461 (BRIT, MO). Santiago-Zamora: Between Campanas
and Arenillas, along Ri'o Tmtas, 10 leagues SE of El Pan, 2,195 m, 1 3 Jul 1913 (stam. fl),
J. Steyer/fiark 53550 (NY). Sucumbios: Sendero toward Volcan el Reventador from km
100 of Baeza-Eago Agrio Elwy, 1,900 m, 7 Oct 1990 (stam. ?i),J. Jaramillo & E. Grijalva
12988 (QCA).
Cybianth?/s occigranatensis is most closely related to C timanae Pipoly, but
is easily distinguished by the fewer secondary veins of the coriaceous leaf
blades, the campanulate corolla with verrucose, prominently black punc-
tate lobes and attenuate apices, and obnapiform pistil. The population from
Alto de Cuevas in Antioquia, Colombia, has by far the largest leaves of any
population of this species known thus iar. Further study oi the population
biologies of Cybunithus montciniis ( Lundell) G. Agostini from Panama, C
oaigranatensis, and C. timanae will be necessary to fully resolve the precise
relationships and microecological roles each plays in montane wet and plu-
vial forests.
4. Cybianthus spichigeri Pipoly, Candollea 46:43. 1991- (Fig. 2B, 1 1).
Typf.: PERU. LoREit); Prov. Rec]uena, Trocha al Ajuajal, 2 km Irom Centro Eorestal
Jenaro Jerrera, right bank of Rfo Ucayali, I 5 Eeb 1982 (stam. fl, tr), \i. Sptchiger & E.
Encarniic'iihi 1224 (hoi.otypi;: US; is()rvpi;s: AMAZ, G, MO).
Tree to 1 5 m tall. BraHchlets thin, straight, terete, 2—3 mm diam., densely
lepidote. Leaves alternate; blades chartaceous, elliptic to narrowly oblanceolate,
( 1 0— )1 5—20 cm long, (3— )5.2— 6.5(— 7.2) cm wide, apically caudate-acumi-
nate, the actmien 1 .9—2.3 cm long, basally acutish to obtuse, not decurrent
on the petiole, bullate, the midrib and secondary veuis strongly impressed
above, prominently raised below, smooth and inconspicuously to promiently
pellucid punctate above, moderately lepidote below, the margin essentially
flat, but very slightly inroUed at the very margin; petioles canaliculate, (I—)
1.2—2 cm long, densely lepidote. Staminate inflorescence: a pyramidal pinnate
pannicle, 1-4.5 cm long, 1-3 cm wide, peduncle 0.3-1 cm long; branch
bracts chartaceous, linear-subulate, 0.6—1 mm long, 0.1—0.2 mm wide, apically
attenuate, densely lepidote; pedicels cylindrical, (0.8— )1— 1 .5 mm long. Staminate
flowers 4— 5-merous, carnose; calyx suburceolate, 1.3—1.5 mm long, the tube
0.3—0.5 mm long, the lobes deltate, ca. 1 mm long and wide, apically actite.
PiPOLY, Cybianthus in Ecuador and Peru
37
F](;. 11. Cybianthus spichigeri Pipoly. A. Habit, showin^q minute inflorescences and large,
costate fruits. B. Staminate flower in bud, showing suburceolate calyx and tubiform co-
rolla. C. Staminate flower, showing long, prominently lobate staminal tube, cuculiate co-
rolla lobes, proximally recurved anthers. A-C, drawn from holotype, by Peggy Duke. Fig-
ure reproduced from Pipoly, 1991-
38 SiDA 1S(I)
sparsely lepidote, prominently ru^M)se, with one prominent brown pancta-
tion per lobe, the margins irregular, entire, sparsely lepidote; corolla tubiform,
2.4—2.8 mm long, the tube ca. 0.5 mm long, the lobes oblong, 1.9-2.9
mm long, 0.9-1.1 mm wide, apically acute, prominently cucuUate, with
only a few, scattered scales without, glandular-granulose within, the mar-
gin entire, glandular-granulose; stamens 2.2-2.4 mm long, the tube carnose,
conspicuous, 1—1.4 mm long, lobate, the lobes 0.1-0.2 mm long alternat-
ing with the filaments, the apex of the tube and lobes punctate, the apically
free filaments, 0.9-1 3 mm long, the anthers deltate, 0.5-0.7 mm long and
wide, apically apiculate, the apiculum slightly proximally recurved, basally
cordate, the connective dorsally punctate with small brown dots forming a
triangle along connective margin; pistillode conic, 0.5—0.7 mm long, 0.2—
0.3 mm wide, hollow, glabrous, the style conspicuously brown punctate,
the stigma punctiform. Pistillate and poly^ai//.()//s inflorescence: a raceme, occa-
sionally a poorly formed panicle of 1—3 racemes branched from base, 0.4—
1.5 cm long, densely lepidote, tardily glabrescent; peduncle 0.1-0.4 mm
long; floral bracts chartaceous, ovate, 0.8-1 mm long, 0.3-0.5 mm wide,
apically attenuate, densely lepidote; pedicels obconic, (0.8-)1.5-2.5(-3) mm
long, to 1.2 mm diam. apically in fruit, densely lepidote. Pistillate flowers as
in staminate but calyx 1.0-1.2 mm long, the tube 0.4-0.5 mm long, the
lobes 0.6-0.7 mm long and wide, staminodes and ]Tistil unknown. Fr/nt
depressed-globose, 0.7-0.8 cm long, 0.7-1.5 cm wide, prominently cos-
tate longitudinally, the exocarp costate, inconspicuously pellucid punctate.
Distribution. — Principally known from forests along the rfos Ucayali, Maranon
and Napo Drainage Basin complex, Loreto, Peru, with one disjunct popu-
lation in nearby Morona-Santiago, Ecuador, 150-1 80 (-1 ,500) m elevation.
Ecology and conservation status. — Cybianthus spicbigeri is known from only
a handful of specimens, but is locally quite common. Not enough is known
of the population biology to categorize its conservation status, but its fre-
quency in forest study plots of the Jenaro Herrera Reserve in Peru suggest
it is reproducing and may not be in imminent danger. Cybianthus spicbigeri
is a varillal or premontante sandstone species in Peru, and in Ecuador it is
known only from premontane sandstones. Occurrence of this species in the
Cerros del Sira, Peru, reinforces the concept that those mountains contain
many unusual populations of otherwise lowland Amazonian plants.
Etymology. — The epithet commemorates Rudolphe Spichiger, Director
of the Conservatoire et Jardin Botaniques, Chambesy, Geneve, Switzerland.
Dr. Spichiger has devoted much of his career to study of global change,
conservation of biodiversity, and systematics of the genus Ilex. Under his
leadership, the Jardin has maintained active research programs in Paraguay,
Peru, Madagascar, and throughout Europe.
PiPOLY, Cybianthus in Ecuador and Peru 39
Representative specimens examined. ECUADOR. Morona-Santiago: Cordillera del Condor,
Cuangos, 20 km E of Gualaquiza, near disputed Peru-Ecuador border, 03° 29' S, 78° 14'
W, 1,500 m, 18 Jul 1993 (ster.), A. Gentry 80096 (BRIT, MO, QCNE), 1,470 m, 19 Jul
1993 (infl. bud), A. Gentry 80179 (BRIT, MO, QCNE). PERU. Huanuco: Prov. Pachitea,
region of Pucallpa, W part of "Sira Mountains," and adjacent lowland, ca. 24 km SE to 26
km ESE of Puerto Inca, from beginning of rainforest to Campamento Pato Rojo, 09° 27' S,
74° 46' W, 1,380 m, 31 Jan 1988 (fl bud), W. Moniwetz & B. Wallnbfer 14-31188 (BRIT,
W, WU). Loreto: Prov. Maynas, Allpahuayo, IIAP Station, 04°10' S, 73°30' W, 150 m,
13 Nov 1984 (fr), R. Vdsquez et al. 5911 (AMAZ, MO, NY), 6jun 1985 (fr), R. Vchquezet
al. 6588 (AMAZ, MO, NY); Rio Nanay, Mishana, 30 km SW of Iquitos, 03° 55' S, 73°
35' W, 150 m, 19 Aug 1978 (fr), R. Foster 4226 (MO, NY, USM), 16 May 1981 (fr), R.
Va.u/uez &J. Crtollo 1801 (AMAZ, MO, NY), 20 Jan 1985 (fr), R. Vasquez & N.Jaramillo
6/37 (AMAZ, MO, NY). Prov. Requena, Reserva Forestal JenaroHerrera, ()4°55' S, 73°45'
W, along Rio Ucayali, 120 m, 1980 (stam. fl), R. Marimllod 9-R-137 (G, US), 1980 (bud),
R. Manrnllod 4-R-90, 10 Jun. 1982 (fr), R. Sphhiner et al. 1973 (G, US), 24 Feb. 1987
(ster.), A. Gentry et al. 5653 1 (AMAZ, MO); Aguajal, 3 km from Centro Forestal Jenaro
Herrera, right margin Rfo Ucayali, 18 May 1982 (fr), F. Fncarnaaon 26105 (AMAZ, MO),
22 May 1982 (fr), F Fncarnaaon 26200 (AMAZ, MO, NY, US); Arboretum, Centro For-
estal Jenaro Herrera, 180 m, 13 Nov 1981 (fr, stam. fl), R. Spichiger & F. Fmarnacim 1027
(AMAZ, G, MO, US).
Cybianthus spichigeri is unique within subgenus Conomorpha because of its
costate fruits. In addition, the subbuUate leaf blades, punctate staminal tube,
obconic pedicels, and polygamous inflorescences are also exceedingly rare
characters that allow for easy recognition. Since its description (Pipoly 1991),
an entirely staminate specimen {R. Marmillod 9-R-137) has been located.
While the staminate inflorescence structure is quite different from that of
the polygamous one, flowers of both are identical. The occurrence of this
taxon three times in one inventory conducted at the Jenaro Herrera Re-
serve indicate that the relative frequency of reproductive individuals would
permit a study of the breeding system and population biology for this most
unusual taxon.
5. Cybianthus lepidotus (Gleason) G. Agostini, Bol. Soc. Venez. Ci. Nat.
22:388. 1976. Conomorpha lepidota Gleason, Bull. Torrey Bot. Club 58:146. 1931.
Type: VENEZUELA. Tf.rritorio Fi^dhkal Ama/onas: Summit of Mt. Duida, 1,500
m, Aug 1928-^Mar 1 929 (pist. fl), G. H. H. Tate 741 (hoi.otype: NY, F Neg. 040832;
lsotype: US).
Conomorpha curvii'enia Gleason, Bull. Torrey Bot. Club 58:444. 1931. Typp: VENEZU-
ELA. Territorio Federal Amazonas: Mt. Duida, 1 ,260 m, Aug 1928-Mar 1929 (stam.
fl), G. H. H. Fate 927 (holotype: NY; isotypi;: US).
Conomorpha lepidota Gleason f. ac/itata Steyerm., Fieldiana, Bot. 28:465. 1953- Type:
VENEZUELA. Territorio Federal Amazonas: Cerro Duida, 2 Dec 1 944 (fr),/. Steyermark
58265 (holotype: F; isotype: NY).
Shrub or small tree to 6 m tall. Branchlets straight, terete, 2-2.5 mm diam.,
densely lepidote. Leaves alternate; blades elliptic to narrowly elliptic, chartaceous
to coriaceous, (3.1-)5-15 cm long, 1.6-6 cm wide, apically acuminate, the
40 SiDA 18(1)
acumen 0.3—2.0 cm long, basally acute to obtuse, midrib depressed above,
prominently raised below, the secondary veins 24-28, inconspicuous above
and below, pustulate and densely lepidote above at first, glabrescent, densely
lepidote below, the scales not overlapping, inconspicuously pellucid punc-
tate, the margin entire, subrevolute to revolute; petioles thin, marginate,
1 .0-2.5 cm long, densely lepidote. S tarn mate inflorescence a spike, rarely two
subsessile spikes, (2— )6-l 6 cm long, 8— 15-flowered, peduncle, pedicels, and
axis densely lepidote, the scales not overlapping; peduncle (0.2-)0. 5-0.8
mm long; floral bracts ovate to widely ovate, chartaceous, 0.5-0.6 mm long,
0.3—0.4 mm wide, apically acute, densely lepidote adaxially, the margin
entire, glabrous. Stcimincite flowers (4-)5-merous; calyx cupuliform, carnose,
0.9-1.2 mm long, the tube 0.2-0.4 mm long, the lobes deltate to ovate-
triangular, 0.6—0.8 mm long and wide, apically acute or acuminate, rarely
obtuse, attenuate to a rounded tip, conspicuously brown punctate, the margin
entire, lepidote; corolla cupuliform, carnose, 1.5-2.5 mm long, the tube
0.5—1.0 mm long, the lobes ovate to broadly ovate, 1.2—1.7 mm long, 0.7-
0.9 mm wide, apically attenuate to a round, cucullate tip, at times with a
few, scattered lepidote scales without, glabrous without, glandular-granulose
within over the entire surface, punctations brown, submarginal, the mar-
gin entire, glabrous; stamens 1.5-1 .6 mm long, adnate 0.5-1.0 mm to corolla
tube, the staminal tube 0.3—0.4 mm long, carnose, bearing lobes alternat-
ing with the apically h-ee portions of the filaments 0.1-0.2 mm long, the
filaments fiat, 0.3-0.4 mm long, erect, glabrous, the anthers dorsifixed less
than 1/4 from base, ovate-triangular, 0.6—0.7 mm long, 0.4-0.5 mm wide,
apically attenuate to a rounded tip, basally cordate, slightly dorsally reflexed,
the connective epunctate; pistillode lageniform, 1.1-1.2 mm long, hollow,
costate basally, sparingly translucent lepidote, pellucid-punctate. Pistillate
inflorescence as in staminate but a spike, (2-)6-l6 cm long, 6-10-flowered;
peduncle 0.6-0.8 mm long; fioral bracts 0.5-0.6 mm long, 0.3-0.4 mm
wide. Pistillate floivers as in staminate, but staminodes 1.2—1.3 mm long,
adnate 0.5-1 .0 mm to corolla tube, the staminodial tube 0.5-0.6 mm long,
carnose, bearing lobes alternating with the apically free portions of the fila-
ments 0. 1-0.2 mm long, those apical portions flat, 0.3-0.4 mm long, erect,
glabrous, the antherodes dorsifixed less than 1/4 from base, deltate, 0.7-
0.8 mm long and wide, apically attenuate to a rounded tip, basally cordate,
slightly dorsally reflexed, the connective epunctate; pistil pyriform, 1.4-
1.5 mm long, the ovary 1-1 .2 diam., the style not differentiated, the stigma
punctiform, the placenta patelliform, bearing 2(-3) naked ovules. Fr/«V globose,
purple at maturity, 0.5- 1.0 cm long, 0.6-1.0 cm diam., the endocarp smooth,
the embryo curved, 3.5-4.0 mm long.
Distribution. — Guayana Highland of Venezuela and Brazil, and sandstone
PiPOi.Y, Cybianthus in Ecuador and Peru 41
formations in Bolivia and Peru (reported for the first time here), 600—2,300
m in Venezuela, 850-950 m in Bolivia, and 760-850 m in Peru.
Ecology and conservation status. — Cybianthus lepidotm is restricted to large
cloud forest formations in transition zones between sandstone and diabasic
intrusions. It is often associated with species o^Erythroxylum, which are also
edaphic endemics. It is a widespread, but locally infrequent species and therefore,
should be considered threatened.
Etymology. — The epithet refers to the densely lepidote vestiture of the
vegetative and floral parts of the plant.
Specimens examined. PERU. Amazonas; Prov. Bagua, Dtto. Imaza, Comunidad Aguaruna
Putuim, anexo Yamayakat, SW of Putuim, 760-850 m, 26 Sep 1994 (stam. fl), C. Diaz et
al. 7252 (BRIT, HUT, MO, USM). BOLIVIA. La Paz: Prov. Nor Yungas, valley of Rfo
Coroico, Sacramento, 10 km NE of C^huspipaca on Coroico Rd., 27 Jan 1984 (seer.), A.
Gentry & J. Solomon 44668 (MO), 4 km NE (above) Inabuara, 13.5 km above San Pedro,
1,530-1,560 m, 22 Jan 1984 (stam. ti), A. Gentry &J. Solomon 44407 (MO); Prov. Larecaja,
Maipiri, 6 Nov 1926-28 Feb 1 927 (stam. fl), 0. Buchtten 1 738 (HBG, NY, US), Copacabana,
10 km S of Maipiri, 850-950 m, 8 Oct-1 5 Nov 1 939 (fr), B. Krnkoff 10987 (A, K, MICH,
MO, NY, UC, US).
Cybianthus lepidott/s, restricted to sizeable cloud forests in transition zones
between sandstone and diabasic intrusions, is most easily recognized by its
long, lax spikes. When sterile, it may be confused with Cybianthus roraimae
(Steyerm.) G. Agost., but may be easily distinguished by the branchlets 2—
2.5 (not 3.5—4.5) mm in diameter. It may also be confused with Cybianthus
punctatus (Mez) G. Agost. and C. cardonae G. Agost. For a discussion of the
differences between these taxa in sterile condition, see Pipoly (1992a).
The area of Bolivia in which this species has been collected is of biogeo-
graphic interest because it also supports several other Guayana Highland
taxa in the Ericaceae and Clusiaceae. Its new discovery in Bagua Province
of Amazonas, Peru, reinforces thematic map data from satellite imagery that
indicated tepui-like vegetation could be expected in the area. It is interest-
ing that C. Diaz et al. 7232 from Peru, and B. Krukoff 10987 from Bolivia,
are qualitatively and quantitatively identical to specimens diCybianthus lepidotus
from Cerro Duida, Amazonas, Venezuela.
6. Cybianthus laetus (Mez ) G. Agostini (Fig. 2C), Acta Biol. Venez. 10:153.
1980. Conomorpha laeta Mez m Engl., Pflanzenr. IV. 236(Heft 9):259. 1902. Type:
PERU. Amazona.s: Taulia, without elevation or date (stam. fl.), A. Matthews ]y6l
(hoi.otype: K; isotype: K).
Shridb orstnalltree to 2 m tall. Branchlets straight, subterete, 2—3 mm diam.,
densely lepidote. Leaves alternate; blades coriaceous, obovate, 3-8(-12) cm
long, 1.2-2(— 5.0) cm wide, apically acute or short-acuminate, basally cu-
neate, decurrent on the petiole, midrib depressed above, prominently raised
below, the secondary veins 14—18 pairs, planar above, conspicuous below,
42 SiHA lcS(l)
glabrous and smooth above at maturity, densely lepidote below, the mar-
gin revolute; petioles marginate, 0.5—1 cm long, densely lepidote. Stami-
nate inflorescence: a raceme or panicle with 2 racemose branches from base,
2—3 cm long; rachis, and pedicels densely lepidote; peduncle ().1"{).3 cm
long; pedicels cylindrical, 0.9—1 -5 mm long; floral bracts chartaceous, nar-
rowly ovate, 1—2 mm long, densely lepidote adaxially. Staminate floiven 4-
merons; calyx carnose, cotyliform, 1.2—1.8 mm long, sparsely to densely
lepidote without, glabrous within, the tube 0.2-0.3 mm long, the lobes
triangular to deltate, 0.8—1 mm long, 0.6—0.8 mm wide, apically attenu-
ate to a rounded tip, inconspicuously pellucid punctate, the margin entire,
lepidote; corolla carnose, campanulate, 3.2—3.4 mm long, the tube 1.8—2
mm long, densely lepidote without, the scales overlapping, glabrous within,
the lobes ovate or ovate-triangular, 0.8—0.9 mm long, 0.3—0.4 mm wide,
symmetric, apically attenuate to a rounded tip, inconspicuosly pellucid punctate,
densely lepidote without, glandular-granulose within, the margin entire,
glandular-granulose; stamens 2.6—2.7 mm long, the tube carnose, conspicuous,
equalling the corolla tube, 1.8-2 mm long, lobate, the lobes alternating
with the filaments ca. 0.1 mm long, the apically free portions of the fila-
ments flat, 0.1—0.2 mm long, the anthers dorsifixed, ovate, 0.6—0.9 mm
long, 1.8—1.9 mm wide, somewhat curved distally, apically apiculate to
subapicLilate, basally cordate, the connective inconspicuously brown punc-
tate; pistillode conic, 1.5—2 mm long, translucent glandular-lepidote ba-
sally, hollow. Pistillate inflorescence: as in staminate but 3—3.5 cm long; pe-
dtmcle 0. 1-0.4 cm long; pedicels cylindrical, 0.9-1 .5 mm long; floral bracts
chartaceous, linear-lanceolate, 1.3-1.5 mm long, densely lepidote adaxially.
Pistillate flowers as in staminate but calyx 1 .2—1 .4 mm long, sparsely to densely
lepidote without, glabrous within, the tube ca. 0.2 mm long, the lobes deltate,
1 — 1.2 mm long and wide, apically attenuate to a rounded tip, inconspicu-
ously pellucid punctate, the margin entire, lepidote; corolla, staminodes
and pistil unknown. Fr//i/ globose, 5—8 mm long and indiam., exocarp carnose,
black.
Distribution. — Eastern slopes of the Andes, Colombia, Peru and Bolivia,
1,980-2,850 m.
Ecology and conservation status. — The species occurs in primary cloud for-
est, a life zone being cleared rapidly for cultivation throughout the Andes,
which may account for the paucity of collections. Within subgenus Conoviorpha,
Cyhianthi/s laetiis is the species most in danger of extinction.
Etymology. — The specific epithet is Latin for "cheerful or bright," and probably
refers to the plant's aesthetically pleasmg appearance, having the same general
form as many species of Waccinium, Myrsine dependens, other Ericaceae, and
other diminutive Andean shrubs. The thick juicy exocarp is said to be very
tasty although slightly acidic (T. Dudley, pers. comm.).
PiPOLY, Cybianthus in Ecuador and Peru 43
Representative specimens examined. COLOMBIA. Boyaca: Arcabuco, NE of town, 2,650
m, 1 1 Nov 1965 (stam. fl), L. Uri/K s.n. (COL); Sierra Nevada del Cocuy, path from Laguna
to Cobugon, near Alto del Oso, 2,900 m, 27 Aug 1958 (stam. fl), P. Gr/ihh et al. 744 (K).
PERU. Amazonas: Prov. Luya, Dtto. Camporredondo, Anexo TuUanya, between Pajaco
Tigreand Palma,06°04' 35"S,78° 21' 45" W, 2,500-2,600 m, 9 Dec I996(fr),J. Ciwpos
et dl. 5UiI (BRIT, HUT, MO, USM); Along Rd. E of Chachapoyas between Pipos and
Molinopampa, 06°15' S, 77°40' W, 1,980-2,340 m, 14 Feb 1985 (pist. fl, iv)J. Liiteyn &
E. Cotton 11414 (NY, TEX, US, USM); E of Chachapoyas, 2,000 m, without date (stam.
fl), A. Weberbaiier 43^4 (G). Cusco: Prov. La Convencion, Cordillera Vilcabamba, 12° 37'
S, 73° 32' W, ceja and cumbre, 2,550 m, 3 Jul 1968 (pist. bud), T. Dudley 10690 (F, NA,
USM), 5 Jul 1968 (pist. fl, fr), T. Dudley 10803 (F, NA, USM). Huanuco: Prov. Huanuco,
Carpish Hills, trail to summit from W entrance, 2,700-2,850 m, 09° 42' S, 76° 05' W, 2
Mar 1985 (stam. fl.), B. Stem & C. Todz/a 2292 (MO, USM).
Cybianthus laet/ts is easily recognized by its small leaves, short petioles,
subsessile anthers and thick, juicy exocarp. It is known outside of Peru from
only two other collections, one from the department of Boyaca, Colombia
and the other from Nor Yungas, Bolivia. A recent collection from Luya Province
in western Peru (J. Campos et al. 3161) is referred here because of its thin,
angulate branchlets, and the thick exocarp of its fruits, despite the larger,
thinner leaves. It is expected in Ecuador, in either Napo or Santiago-Zamora
Provinces.
7. Cybianthus peruvianus (A. DC. ) Miq. (Fig. 2D) in Mart., Fl. Bras.
10:298. 1856. Conomorpba peruviana A. DC. , Ann. Sci. Nat., Hot. set 2, 16:92.
\M\.Peckia peruviana (A. DC. ) Kuntze, Revis. Gen. PI. 402. 1891. Type: PERU.
Amazonas: Moyobamba, 1838 (stam. fl), A. Mathews s.n. (holotype: G-DC; lsotypes:
G, G-BOIS, GH, K).
Conomorpha weherhaueri Mez, Repert. Spec. Nov. Regni Vcg. 3:101. 1906. Type: PERU.
LoRirro: In mountains near Moyobamba, 1 ,300 m, 28 Aug 1904 (stam. fl), A. Weberbauer
4668 (hoi.otypi:: B-destroyed; Lectotype, here designated: F).
Shrub or small tree to 16 m tall. Branchlets straight, terete, 3-5 mm diam.,
densely lepidote. Leaves alternate, at times approaching pseudoverticiUate
at some nodes; blades coriaceous, elliptic to obovate, 7-12.7 cm long, (2.6—)
3.5—5 cm wide, apically acuminate, the acumen 1.2—1.6 cm long, basally
acute to obtuse, decurrent on the petiole, midrib depressed above, promi-
nently raised below, the secondary veins 10—25 pairs, prominently raised
below, densely lepidote at first, then pusticulate above at maturity, sparsely
lepidote below, the margin revolute; petioles canaliculate, 1—1.5 cm long,
densely lepidote. Staminate inflorescence: a panicle with 1—4 racemes branch-
ing from the base, (2— )3— 8 cm long; peduncle, rachis and pedicels densely
lepidote; peduncle 0.1—0.2 cm long; floral bracts chartaceous, ovate, 0.7—1
mm long, 0.5—0.6 mm wide, apically attenuate, densely lepdote abaxially;
pedicels cylindrical, 0.5— 1(— 1.2) mm long. Staminate floivers 4— 5-mcrous;
calyx green, cotyliform, 0.8—1 .2 mm long, lepidote without, glabrous within,
the tube 0.3-0.4 mm long, the lobes ovate to deltate, 0.7—0.8 mm long,
44 Sum 18(1)
0.5—0.8 mm wide, apically artenuate ro a rounded tip, the margin lepi-
dote, inconspicuously brown punctate; corolla translucent green, chartaceous,
campanulate, 2—2.6 mm long, the tube 0.6— 1 .2 mm long, glabrous or sparsely
lepidote externally, glabrous internally, the lobes ovate to narrowly ovate,
1.2—1.8 mm long, O.")— 1.1 mm wide, asymmetric, apically rounded or at-
tenuate to a roimded tip, sparsely lepidote externally, the scales not over-
lapping, sparsely gland ular-granulose internally, inconspicuously brown punctate,
the margin glandular-granulose; stamens 1 .4—2 mm long, the tube chartaceous,
conspicuous, longer than the corolla tube, 0.8—1.2 mm long, elobate, the
apically free portions of the filaments 0.2—0.4 mm long, the anthers linear-
lanceolate, 0.7—0.9 mm long, 0.2—0.3 mm wide, somewhat recurved dis-
tally, attenuate to a round or acute tip, the connective conspicuously brown
punctate; pistiUode obclaviform, 1 — 1.5 mm long, translucent glandular-
lepidote basally. Pistillate inflorescence: like the staminate but, 2—7 cm long;
peduncle 0. 1-0.2 cm long; floral bracts 0.7-1 mm long, 0.4-0.5 mm wide;
pedicels 0.2—0.5 mm long. Pistillate liowers as in staminate but calyx 0.9—
1.1 mm long, the tube 0.3-0.7 mm long, the lobes deltate, 0.5-0.6 mm
long, 0.4—0.5 mm wide, apically subacute to obtuse; corolla 1.3—1-7 mm
long, the tube 0.2-0.3 mm long, the lobes ovate, 1-1.4 mm long, 0.7-0.9
mm wide, apically obtuse; staminodes 1—1 .2 mm long, the tube longer than
the corolla tube, 0.3—0.4 mm long, the apically free portions of the fila-
ments 0.2—0.3 rnm long, the antherodes 0.6—0.7 mm long, 0,2—0.3 nim
wide, pistil obnapiform, 1.3— 1.4mm long, 1-1.1 mm diam., the ovary 0.6—
0.7 mm long, translucent glandular-lepidote basally, the style 0.5—0.6 mm
long, conspicuously brown punctate, the stigma punctiform, the placenta
globose, ovules 3, apically exposed. Fruit subglobose, 0.3—0.8 mm long,
0.4—0.9 mm diam., the exocarp thin, black.
Distribution. — Amazonian Ecuador through Peru to Bolivia, at 122—1,500
m elevation.
Ecology and conservation status. — Cyhianthus perui'ianus occurs on the east-
ern slopes of rhe Andes in moist or wet lowland and premontane foresrs on
white sands, especially in transition zones, where brownish sand-clay mix-
tures occur.
Etymology. — The epithet refers to the type locality, in (Moyobamba) Peru.
Eocal navie. — Peru: "Tarrafactispi." (Quichua), "uchi yacushnum" (Aguaruna).
Represencarivc sjiecimens examined. ECUADOR. Napo: CanttMi A^^tiarico, Reserva Fauni'stica
Cuyabeno, Layima Zancutlo Cocha (Iripari), SE side ol Laguna, 00" 33' S, 75° 32' W, 230
m, 28 Sep 1991 (fr), W. Palacios el at. 7761 (BRIT, MO, QCNH); Cant6n Orellana, Sector
Huashito, 20 km N of Coca, PALMORIENTE property, 00" 20' S, 77° 05' W, 250 m, 3~
21 Nov 1989 (fr), E. Gi/uliiio 137 (BRIT, MO, QCNE); Sendero to Palma Roja, 28 Apr
1986 (stam. ^)J.Jc/nimillo H522 (QCA). Zamora-Chinchipe: Canton Nangaritza Campamento
Miazi, along Rfo Nangaritza, 900 m, 1 9 Eeb 1 994 (fr), //, van cler Werffet at. / J2<S'0 (BRIT,
MO, QCNE); 11 ill above military post, 04" 18' S, 78° 40' W, 1,000 m, D. Nnll & W.
PiPOLY, Cybianthus in Ecuador and Peru 45
Palacios 9613 (BRIT, MO, QCNE), 04° 16' S, 78° 42' W, 970 m, 20 Oct 1991 (fr), W.
Pa/acm eta/. 8486 (BRIT, COL, MO, QCNE). PERU. Amazonas: Along Ri'o Maranon,
near confluence with Ri'o Santiago, 1924 (stam. fl.), G. Te.ss>uann 3523 (B, G); Prov. Bagua,
Dtto. Imaza, NE region of Rfo Maraiion Drainage Basin, Comunidad Kampaenza, along
Qucbrada Shimutaz, Rio Maranon, 04° 55' S, 78° 19' W, .320 m, 9 Sep 1994 (pist. fl, Fr),
N.Jaramilloetal. 436 (AMAZ, BRIT, HUT, MO, USM), 09 Oct 1995 (fr), N. Jaramillo &
D. Chamik 813 (AMAZ, BRIT, MO, USM); Comunidad Aguaruna de Putuim (CAMPOU),
anexo Yamayakat, Monte Alto de PutLiim, 450 m, 25 Aug 1994 (fr), C. D/az et al. 6993
(BRIT, HUT, MO, USM), 660-760 m, 21 Sep 1994 (stam. fl), C, D/az et al. 7170 (BRIT,
iMO, USM); Cerrosde Putuim, 05° O.V 20" S, 78° 20' 23" W, 350 m, 13Jun 1996 (fr),/?.
Va'sqt/ez et al. 21131 . Huanuco: Prov. Pachitea, region of Pucallpa, W part of Sira Moun-
tains and adjacent lowland, 20-24 km SE of Puerto Inca, Campamento Oro, 09° 29' S, 74°
50' W, to Campamento Sira, 800 m, 17 Jul 1988 (stam. fl), W. Morawetz & B. Wallmfer
22-19188 (BRIT, W, WU). Loreto: Prov. Loreto, Nauta, 04° 32' S, 73° 35' W, 160 m, 2
Jun 1984 (fr), R. Vdsqiiez & N.Jaramillo 5060 (AMAZ, MO, USM); Ztingaru Cocha, 15
km SE of Iquitos, 9 Sep 1964 (stam. fl), C. Dochon 2821 (AMAZ, MO, US, USM); Prov.
Maynas, Moropon, lower Rio Nanay above Bellavista, 29 Aug 1968 (fr), S. McDaniel 1 0942
(AMAZ, lEB, MO); Tamishiyacu, Quebrada Blanco Biol. Station, Camp II, Quebrada Blanco,
Tahuayo River, 04° 23' S, 73° 17' W, 4 Apr 1 985 (fr),/. Castro 27 (AMAZ, MO, US, USM);
Dtto. Iquitos, Hwy to Sto. Tomas, path in front ol "Chaparal" chicken farm, 140 m, 16
Dec 1 983 (stam. fl), M. Rimachi 7232 (AMAZ, IBE, MO, US); Estacion Expetimental IIAP
Allpahuayo, 21 km S of Iquitos,04° 10' S, 73° 30' W, 160 m, 18 Sep 1990 (ster.),/ P/poly
etal. /2i72(AMAZ,MO,US, USM), 220 m, 19 Sep 1990 (stam. B) J. Pipoly et al. 12210
(AMAZ, MO, NY, USM); Dtto. Sta. Maria de Nanay, 10 km W of Caserio Mishana, in
Cocha Yarama Reserve, 03° 55' S, 73° 35' W, 130 m, 14 Mar 1991 (ster.),J. Pipoly etal.
14994 (AMAZ, MO, US, USM), (ster.),/ Ptpalyetal. 14997 (AMAZ, MO, USM); Mishana,
along Rfo Nanay, 03° SI'S, 73° 32' W, 150 m, 22 Apr 1986 (stam. bud), R. Vaujuez et al.
7503 (AMAZ, MO, US, USM), 8 Sep 1990 (fr), R. Vdsquez etal. 14335 (AMAZ, BISH, F,
MO, NY, TEX, US, USM); Mishana,() 3° 52' S, 73° 30' W, 140 m, 4 Jan 1983 (fr), A.
Genttj etal. 39040 (AMAZ, MO, USM); Dtto. Iquitos, Puerto Almendras, Arboretum Ciencias
de Ingenieria Forestal, UNAP, 03° 48' S, 73° 25' W, 122m, 4 Sep 1992 (ster.), C. Gra'/ulez
et al. 471 1 (AMAZ, BRIT, MO, USM), Puetto Almendras, along Rio Nanay, 03° 45' S,
73° 25' W, 122 m, 30 Oct 1984 (fr), R. Mchquez & N.Jaramillo 5867 (AMAZ, MO, NY,
USM), 29 May 1986 (fr), R. Vazquez & N.Jaramillo 7570 (AMAZ, MO, US, USM), 30
May 1986 (stam. bud), R. YdsqNez& N.Jaramillo 7587 (AMAZ, MO, US, USM); Quistococha,
IMARPE tract, along Rd. 13 km from Iquitos, I4 Jul 1976 (fr), F. Encarnaaon 864 (AMAZ,
US); Prov. Requena, Dtto. Saptiena, Jenaro Herrera, Rio Ucayali, 04° 55' S, 73° 40' W,
160 m, 16 Aug 1994 (stam. fl), R. Ortiz et al. 98 (AMAZ, BRIT, MO, USM). Madre de
Dios: Prov. Tambopata, Santuario Nacional Pampas del Heath, Quebtada Palma Real Grande,
12° 57' 1 1" S, 68° 54' 48" W, 210 m, 21 Apr 1996 (fr), Al. Agu/lar & D. Castro 623 (BRIT,
MO, USM), Puesto Enahuipa, 12° 39' 23" S, 68° 44' 13" W, 2 10 m, 25 Apr 1996 (fr), M.
Agi/ilar & D. Castro 655 (BRIT, MO, USM). San Martin: Prov. San Martin, trail to televi-
sion antenna, km 17.5 of Tarapoto-Yurimaguas Rd., 2.5 km N of Cataratas de Ahuashiyacu,
06° 27' S, 76° 21' W, 850-1,200 m, 7 Sep 1986 (pist. fl, fr), S. Kuapp 8290 (MO, US,
USM).
Cybianthus peruvianiis is most closely related to C. comperuviaJius Pipoly
(herein described), but may be recognized by its longer, canaliculate peti-
oles, revolute leaf margins, shorter pedicels, asymmetric, lepidote corolla
lobes, and linear-lanceolate anthers. Field studies near Iquitos have shown
46
SiDA 18(1)
Fk,. 1 2. Cylnanthiii comperiivianus Pij^oly. A. Habit, showing inflorescence of racemes or malformed
panicles. B. Staminate flower and axillanr bract, showint^ floral densely lepidote floral bract
as long as pedicel, and lepidote calyx margins. C. Opened staminate flower, showing staminal
rube longer than apical tree portions ot the lilamencs, lobes glandular-grannlose adaxially
and crennlate margins. A--C, ch^awn Irom holotype, by Linny licagy.
PiPOLY, Cybianthus in Ecuador and Peru 47
it is a ridgetop species in the lowlands, with a density of approximately 2—
4 individuals per hectare.
8. Cybianthus comperuvianus Pipoly, sp. nov. (Fig. 2E, 12). Type: BRA-
ZIL. Mato Grosso: Sta. Anna da Chapada, 1903 (stam. fl), G. Malme 3483 (holo-
type: S; jsotypes: G, GH, R, S).
Cybtanthus comperuvianus Agostini ex Pipoly in Killeen et al., Guia Arb. Boliv. 570. 1993.
no?n. nud. Agostini (1972) provided the first description of this species in his disserta-
tion, but never published it. I subsequently annotated herbarium specimens with the
name, fully intending to publish it, but it was still not validly published when it
appeared in Guia de Arholes de Bolivia, without Latin description or reference to type.
Presumably, the name was obtained by them based on herbarium determinations, and
thus a nomen nudum and invalid. Validation of the name is effected here,with the cita-
tion of holotype above and the Latin diagnosis, and accompanying description, pro-
vided below.
Species haec cum C. peruviano saepenumero confusus est, sed laminis membranaceis (non
coriaceis), secus margines planis (ncc revolutis), petiolis marginatis (non canaliculatis) 0.5-
1 (nee 1-1.5) longis, pedicellis 1.5-2.5 (non 0.7-1.0) mm longis, coroUis staminaribus
salverformibus (non campanulatis) statim diagnoscenda.
Shrub or small tree to 6 m tall. Branchlets angulate, 1.5-2 mm diam., densely
lepidote. Leaves alternate; membranaceous, elliptic, narrowly elliptic, to obovate,
9-13(-17) cm long, 3-5(-6) cm wide, apically acuminate, basally acute,
decurrent on the petiole, the midrib slightly depressed above, prominently
raised below, pusticulate above, densely lepidote below, the secondary veins
20-30 pairs, the margin flat; petioles marginate, 0.5-1 cm long, densely
lepidote. Staminate inflorescence: a raceme, rarely a malformed panicle with 2
branches from the base, 1-3 cm long; peduncle 0.1-0.5 cm long; floral bracts
membranaceous, ovate, 1 .3-1 ■5(-2.0) mm long, ca. 0.6 mm wide, subglabrous,
sparsely lepidote above and densely lepidote below, apically acute, slightly
shorter than or as long as the pedicels, entire; pedicels cylindrical, 1.2-2.5
mm long, sparsely lepidote. Staminate flowers whitish-green, 4(-5)-merous,
membranaceous; calyx cupuliform, 1-1.2 mm long, the tube 0.3-0.6 mm
long, the lobes triangular to deltate, 0.6-1 mm long, 0.5-0.7 mm wide,
apically attenuate to a rounded tip, sparsely lepidote without, glabrous within,
conspicuously brown punctate, the margin entire, sparsely ferrugineous-
lepidote; corolla salverform, 2.2-2.6 mm long, the tube 1-1.2 mm long,
the lobes ovate to ovate-triangular, 1.2-1 .6 mm long, 0.8-1 mm wide, apically
rounded or attenuate to a rounded tip, glabrous without, glandular-granulose
throughout within, the margin minutely crenulate, glandular-granulose,
conspicuously brown punctate; stamens 1.8-2.2 mm long, adnate 1.2-1.5
mm to corolla tube, staminal tube 1.6-1.9 mm long, elobate, longer than
the apically free portions of the filaments 0.2-0.3 mm long, the anthers
deltate to triangular, slightly distally curved, 0.7-0.8 mm long, 0.5-0.6
mm wide, apically apiculate, basally broadly cordate, the connective incon-
48 SiDA 18(1)
spicuously brown punctate; pistillode lageniform, 1.0-1.2 mm long, 0.3—
0.4 mm wide, the ovary 0.4—0.5 mm long, densely translucent glandular-
lepidote near base, the style 0.5-0.6 mm long, the stigma punctiform. Pis-
tillate inflorescence as in staminate but pedicel 1.5-2.5 mm long. Pistillate
flowers as in staminate but calyx 1.0-1.2 mm long, the tube 0.3-0.6 mm
long, the lobes deltate, 0.6-1 mm long and wide, corolla, staminodes and
pistil unknown. Fruit globose, 5-7 mm long, 6-8 mm diam.
Distribution. — Ecuador southward to Bolivia and adjacent Brazil, 400—
1,200 m elevation.
Ecology a)id conservation status. — Cybianthus coniperuvianus occurs in primary
premontanc forests on sandstone and limestone, in relatively sparse popu-
lations (teste coUectore). Therefore, it should be considered threatened.
Etymology. — The specific epithet refers to the fact that Cybianthus comperuvianus
has long been confused with C. peruvtanus.
Local name. — Peru: "wewe" (Jivaro); "uchi apikna" (Huambisa); "CumaliUa"
(Spanish).
PARAT^-prs. ECUADOR. Morona-Santiago: Sitio La Planada, 01° 46' S, 77° 57' W,
900 m, 2 1 Sep 1993 (stam. H), \V. Pciljcio.s 1 1407 (BRIT, MO, QCNE). PERU. Amazonas:
Near Yucui Entsa, 6 hrs. from Camino tie Kusu, 300 m, 1 1 Mar 1973 (It), R. Kayap 538
(MO, NY, USM), (fr), E. Ancuast) 93 (AM AZ, BRIT, F, LL-TEX, MO, NY, US, USM); Qucbrada
Huampami, Tseasim, near Nayumpim, 244 m, 3 Apr 1973 (fr), E. Anaunlo 141 (AMAZ,
F, LL-TFX, MO, US, USM); Quebracla Yurui Entsa, 305 m, 12 Apr 1973 (fr), E. Aiiauish
220 (AxMAZ, F LL-TEX, MO, US); Prov. Bagua, Drto. Imaza, NE Region of Ri'o Marafion
Drainage Basin, Comunidad de Yamayakac, Rio Maraiion, 04° 55' S, 78° 19' W, 320 m, 8
Aug 1994(stam. H), N.Jaramillaetcil. 321 (BRIT, HUT, MO, USM). Ayacuclio: Tambillo,
Toche Colorado, 27 Jul. 1878 (fr), C.Jelski 360 (PR, W). Huanuco: E of Tingo Man'a, 5
Oct. 1972 (sram. (i), T. Croat 21194 (F, USM, MO); Agua Bianca, trail to Monzon, 9 Feb
1966 (stam. 11.),/, Sitjmike 1049 (AMES, MO, NY, S, USM, VEN); vicinity Rondos, 24
Mar. 1962 (sram. H.),_/. Sc/vnike 388! (F, US, USM); Prov. Pacliirea, region of Pucallpa, W
part of Sira Mountains and adjacent lowland, 20-24 km SE of Puerto Inca, Campamento
Oro, 09° 29' S, 74° 50' W, to C;ampamento Sini, 800 m, I "Jul 1988 (stam. fl), B. Walliwfer
12-17788 (BRIT, MO, W, WU), 30 Aug 1988 (fr), W. Alonnmz & B. Wallnofer 13-30888
(BRIT, W, WU), from Campamento Sira, 09° 28' S, 74° 47' W, SE to valley of Rio Negro,
650 m, 1 I Aug 1988 (fr), W. Morawetz & B. Walinofer 1 13-1 1888 (BRIT, MO, W, WU).
Juni'n: E of Quimiri Bridge, near La Merced, 800-1,300 m, I Mar 1929 (stam. H.), E.
Killip & A. C. Simtlo 2401 1 (F, NY, US); La Merced, Hacienda Schunke, 27 Aug-1 Sep
1923 (stam. fl.),/, Mac/mc/e5677 (F); Pichi's Trail, San Nicolas, 1 ,100 m, 4 Jul 1929 (stam.
fl.), E. Killip & 'a. C. Smitl} 26(P3 (F, NY US), Sta, Rosa. 625-900 m, 6 Jul 1929 (stam.
H.), 26768 (BM, F, NY, US, USM); Puerto Yessup, iOO m, 10 Jul 1929 (stam. fl.), /:. K//tip
& A. C. Smitlj 26286 (NY, US); Puerto Bcrmudez, 375 m, 1 4 Jul 1929 (stam. H.), E Killip
&A. C. Switlj 26464 (NY, US), 26548 (NY, US), 2656.1 (NY, US). Pasco: Prov. Oxapampa,
Pichi's Valley, San Manas Ridge, 10-12 km SW of Puerto Bermudez, above Sta. Rosa de
Chin's, trail to Loma Linda, 10" 20' S, 75° 00' W, 500 m, 29 Sep 1982 (fr), R. Eo.stcr et at.
8962 (F, MO, USM). Puno: Below San Ciabon on Rfo San Ciabon, 500-1,000 m, 17-24
Jul 1978 (stam. bud), Al. Dillomjtjl. 1219 (BRIT, F MO, LJSM). San Martin: Prov. Mariscal
Caceres, Drto. Tocache Nuevo, Palo Blanco near Funtlo de Manuel Aranjo, 700-800 m, 1
PiPOLY, Cybianthus in Ecuador and Peru 49
Mar 1979 (fr) J. Schunk 10893 (AMAZ, BRIT, F, MO, NY, US); without specific locality,
1778-1788 (stam. fl.), Ru/z L. &J. Patm 3/36 (F, MA), BOLIVIA. La Paz: Mapin Re-
gion, 1926 (stam. fl.), 0. Buchnen 1733 (F, GH, HBG, NY, US); Tuiri, near Mapiri, 490-
750 m, Sep 1939 (stam. fl.), B. Knikoff 10930 (A, G, GH, MO, MICH, NY, S, U, UC, US).
Santa Cruz: Velasco; Parque Nacional Noel Kempff Mercado, Campamento las Gamas,
14° 48' 41" S, 60° 23' 45" W, 850 m, 26 Mar 1993 (fr), L. Arroyo & K. Keill 164 (BRIT,
MO, USZ); Campamento Huanchaca, 13° 54' S, 60" 48' W, 650 m, 17 May 1994 (stam.
fl), L. Arroyo et ai. 61 4 (BRIT, MO, USZ). BRAZIL. Mato Grosso: Mpio. Cuiaba, Burity,
NE of Cuiaba, 750 m, Jul 1927 (stam. fl.), B. Collenette 113 (NY); Chapada dos Guimraes,
Cachoeira Veu de Noiva, do Rio Coxipozinho, 15° 30' S, 55° 45' W, 21 Oct. 1985 (fr),/
Prra>2! 1326 (INPA, MG, NY, SP), 720 m, 16 Oct 1973 (fr), G. Prance et a I. 19073 (AAU,
F, INPA, K, MG, NY, K, S, SP, U, US); Sta. Anna da Chapada, 1902 (stam. fl), G. Malme
2048 (S, UPS), 1903 (fr), G. Malme 3483 (G, GH, R, S), 1827 (stam. fl.), L. Riedel 939
(LE, NY, US), 1902 (fr), A. Robert 322 (BM). Rondonia: 1 km NE of Ariquemes, Porto
Vehlo-Cuiaba Hwy., 13 Aug 1968 (fr), £, Porero & L. Wrigley 7033 (MG, MO, NY).
Cybianthus comperuvianus was first recognized as a novelty by Agostini (1972).
Despite the fact that over 25 years has past since its first recognition, pistil-
late flowers are still unknown, reinforcing the concept that within the ge-
nus they are ephemeral (Pipoly 1983a, 1992). Cybianthus comperuvianus is
most closely related to C. peruvianus but is easily distinguished by its long
pedicels, salverform staminate corolla, and membranaceous leaves.
9. Cybianthus guyanensis (A. DC.) Miq. in Mart, subsp. pseudoicacoreus
(Miq. in Mart.) Pipoly, comb, et stat. nov. (Fig. 2F). Ardisia pseadoicacorea
Miq. in Mart., Fl. Bras. 10:284. 1856. Conomorpha psendoicacorea (Miq. in Mart.) Mez
in Engl., Pflanzenr. IV. 236(Heft 9):26l. 1902. Cybianthus pseudoicacoreus (Miq. in
Mart.) G. Agostini, Acta Biol. Venez. 10:155. 1980. TYPE: BRAZIL. Amazonas:
In forest near Rio Japura, Jan 1820 (pist. fl, fr), C. Martins s.n. (lectotype, here desig-
nated: M; isolhctotype: M).
Shrub or tree to 7 m tall. Branchlets terete, 2—3 mm diam., densley lepi-
dote. Leaves alternate; blades membranaceous to chartaceous, narrowly obovate
to elliptic, symmetric, (8-)9-12(-18.5) cm long, 3^.5(-5.7)cm wide, apically
abruptly acuminate to caudate, the acumen 1-2 cm long, basally cuneate,
decurrent on the petiole, midrib flat or slightly depressed on the upper surface,
prominently raised below, the secondary veins 12-25 pairs, pustulate above,
sparsely lepidote below, the margin flat; petioles canaliculate, 0.5— 0.8(— 1.5)
cm long, densely lepidote. Staminate inflorescence: a raceme or panicle with
1—2 branches from base, 1-3 cm long; peduncle, rachis and pedicels densely
lepidote; peduncle 0.1-0.2 mm long; floral bracts chartaceous, narrowly
ovate, longer than the pedicels, 1.3-1-5 mm long, 0.2-0.3 mm wide, apically
attenuate, densely lepidote abaxially; pedicels cylindrical, 1—1.2 mm long.
Staminate flowers 4-merous; calyx carnose, subcupuliform, 1—1.2 mm long,
sparsely lepidote without, glabrous within, the tube 0.3—0.4 mm long, the
lobes triangular, 0.8-l(-1.2) mm long, 0.5-0.7 mm wide, apically attenu-
ate to a rounded tip, conspicuously brown ptmctate, the margin lepidote;
50 SiDA 18(1)
corolla carnose, camjoanulate, 2.8-3-2 mm long, the tube 1.2-1.8 mm long,
glabrous, the lobes ovate, 1.3-1.8 mm long, 0.8-1 mm wide, erect to spreading,
apically attenuate to a round tip, prominently or conspicously brown punctate,
sparsely lepidote without, densely glandular-granulose on the upper half
within, the margin entire, glandular-granulose; stamens 2.2-2.6 mm long,
the staminal tube consj^icuous, carnose, 1.2—1.6 mm long, lobate, the lobes
to 0.2 mm long, the apically free portions oi the filaments ().4-().6(-7) mm
long, the anthers triangular, 0.7-0.9 mm long, 0.4-0.5 mm wide, slightly
distally recurved, dehiscent by narrow introrse slits, apically attentuate to
an apiculate tip, basally cordate, the connective inconspicuously brown punctate;
pistillode elongate, conic, 1.2-1.8 mm long, densely translucent glandu-
lar-lepidote basally. Pistillate inflorescence as in staminate but 1—3 cm long;
lioral bracts 1—1 .3 mm long, ca. 0.2 mm wide; pedicels 0.6—0.9 mm long.
Pistillate flowers as in staminate but calyx ca. 1 mm long, the tube 0. 1—0.2
mm long, the lobes 0.8—1 mm long, 0.-1— 0.6 mm wide; corolla, staminodes
and pistil unknown. Print subglobose, 4—6 mm long, 5—7 mm diam., exo-
carp thin, pellucid punctate.
Distribution. — Venezuela, Ecuador, Peru, Amazonian Brazil, and reported
here for the first time from Bolivia, from 70—700 m.
Ecology and conservation status. — Cybianthus guyanensis subsp. pseudoicacoreiis
inhabits igapo forests of South-Central and Western Amazonia. It occurs
in these forests on deep white sands just bek)w the floodline. It is periodi-
cally inundated, but not for long periods. Quantitative fieldwork in Perti
has shown it occurs in jiopulations of 8—1 individuals > 2.5 cm DBfi per
hectare.
Etymology. — The subspccific epithet refers to the growth habit of the plant,
somewhat reminiscent oi Ardisia (subgenus I cacorea) guyanensis (Aublet) Mez.
Representative specimens examined. ECUADOR. Napo: Canton Aguarico Reserva Etnica
Huaorani, jVIaxus Oil Ilwy., km 60-61, S of Rfo Tivacuno, 00° 51' S, 76° 26' W, 250 m,
21-25 Oct 1993 Or), /\l. A/ilcslia &J. Audi 923: Maxus Petroleum pipeline RcL, km 68,
10 km SW of Ri'o Tivactino, 00" 49' S, 76" 2& W, 2fO m, 1 3 Dec 1993 (Ir), D. Matt a a/.
10303 (BRIT, MO, QCNE). (BRIT, MO, QCNE); Estacion Experimental INIAP-Payamino,
5 km NE of Coca, 00" 26' S, 77" 01' W, 250 m, 18-26 Feb 1986 (fr), W. Palmias d cil.
I()4(> (MO, NY, QAME), (stam. fl), W. Palacios ct dl. lOSi) (MO, NY, QAME), 00° 25' S,
77" 00' W, 250 m, 29 Nov 1 986 (fr), D. Nail 7494 (MO, QAME); Parqiie Nacional Yasiim',
Lagunas de Garza Cocha, shore of Ri'o Garza, TO 1' S, 75 ' 17' W, 200 m, 22 Sep ! 997
(fr), C. Ceroii &N. dllo "tOfij (MO, QCNE). trail hehind the house, 850 m, 27 Apr 1986
{(v),J.Jdn/M/llo 8301 (QCA). PERU. Loreto: Prov. Maynas, Iquiros, G. Tessiiunin ,j65(;
(NY), "l 00 m, 3-1 f Aug 1 929 (fr), II. Killip & A. C. Smith 2^005 (F, NY, US); Misluiyacu,
near Iquitos, 100 m, 1930 (stam. il), G. Kl//;^ 1-412 (I'), I'eh 1932 (fr), G. Kl/zt^ 256^ (F,
NY), 24 Sep 1 929 (stam. 11), I:. Killip & A. C. Smith 2<AS'" / ; Dtto. Alto Nanay, trail leading
N from N end of Sta. Maria ck- Nanay, 5 Mar 1968 (stam. Ik), D. Simpson &J. Schittikt '^S4
(F, US); Dtto. Indiana, Ex|ilorama Lodge, hallway between Indiana and mouth ol Rio Najio,
03° 28' S, 72" 50' W, I sO m, 26 jun 1983 Or), /). Gt/itry et al. 42IH3 (AMAZ, MO), Far
PiPOLY, Cybianthus in Ecuador and Peru 51
end of Bushmascer Trail, l40 m, 5 Jan 1991 (ster.), A. Gentry et al. 12129 (AMAZ, MO,
US, USM), Explorama Lodge, near Yanamono, 25 km NE of Iquitos, 03" 30' S, 72° 50' W,
106 m, 24 Nov 1981 (fr), R. Vasquez & N. Jaramillo 2703 (AMAZ, MO, US, USM), Pe-
rimeter path at southern boundary o^ reserve, 110 m, 27 Sep 1990 (ster.), J. Pipoly ct a!.
12492 (AMAZ, MO, US, USM), 03" 28' S, 72° 52' W, 106 m, 15 Apr 1992 (fl bud), R.
Vchq/wz & N.Jannmllo 18240 (AMAZ, BRIT, MO,USM); Dtto. Iquitos, Allpahuayo, Estacion
Experimental del IIAP, 04° 10' S, 73° 30' W, 150-180 m, 3 Nov 1990 (fr), R. Vdsquez &
N.Jarannllo 14545 (AMAZ, BRIT, MO, USM)8 Nov 1990 (ster.), R. Vasq//ez & N.Jarc/wilb
15016 (AMAZ, MO, US, USM); Puerto Almendras, 2 Jul 1992 (fr), R. RimL, '&J. Ruiz
597 (AMAZ, BRIT, MO); PuertoAlmendras, Arboretum Colegion Ingenerfa Forestal UNAP,
03° 48' S, 73° 25' W, 122 m, 4 Sep 1992 (ster.), C. Grandez et cd. 4487 (AMAZ, BRIT,
MO, USM); Dtto, Las Amazonas, Quebrada Sucusari, 03° 15' S, 72° 55' W, l40 m, 11
Aug 1994 (stam. fl), R. Ortiz tt cd. 7-4 (AMAZ, BRIT, MO, USM); Dtto. Sta. Marfa de
Nanay, Mishana, 03° 55' S, 73° 35' W, 90 m, 1 Ocr 1990 (ster.), J, Pipuly et al. 12706
(AMAZ, MO, USM). Madre de Dies: Prov. Manu, Cerro de Pantiacolla, Rio Palotoa, 1 0-
15 km NNW of Shintuya, 12" 35' S, 7 1 " 18' W, 650-700 m, 13 Dec 1985 (ster.), R. Poster
et ill. 10993 (F, NY, USM); Prov. Tambopata, Santuario Nacional Pampas del Heath, Rio
Heath, 12° 39' 23" S, 68° 44' 1 3" W, 2 1 m, 5 Jun 1 996 (stam. B), M. Aguilar & D. Castrn
805 (BRIT, MO, USM). BOLIVIA. Santa Cruz: Dtto. Velasco, Parque Nacional Noel
Kempff Mercado, Campamento La Torre, 13" 39' 20" S, 60° 49' 08" W, 200 m, 24 Nov
1993 (fr), L Arroyo etcil. 510 (BRIT, MO, USZ).
Cybianthus gNyanerisis s\.\hs\) . lisei/c/oiatcore/ts is one of three subspecies. The
species is defined by the autapomorphic contorted anthers of the stamens
and staminodes (Pipoly 1992a). The three subspecies may be separated in
the following key.
1. Leaf blades symmetric; calyx subcu|Tulilorm; corolla chartaceous or carnose,
rhe lobes erect to spreading; apically free portions ot the filaments shorter
than the anthers; anthers narrowly triangular or ovate-triangular, apically
apiculate, dehiscent by narrow, introrse slits.
2. Petioles canaliculate and winged, (1-)1 .5-1 .9(-2.3) cm long; staminate
peduncle 0.2-0.5 cm long; floral bracts shorter than the pedicels, 0.7-
0.8 mm long; pedicels 0.5 — 1 mm long; corolla chartaceous, 2.3—2.6 mm
long, the lobes elliptic; staminal tube 0.9-1 ■ 1 mm long; apically Iree |X)rtions
of the filaments 0.2-0.4 mm long subsp. guyancnsis
2. Petioles canaliculate, 0.5-0.8(-1.5) cm long; staminate peduncle 0. 1-0.2
cm long; floral bractslonger than the pedicels, 1 .3-1 .5 mm long; pedicels
1-1.2 mm lonir; corolla carnose, 2.H-3.2 mm lont;, the lobes ovate; staminal
tube 1.2—1.6 mm long; apically Iree |-)()rtions of the filaments 0.4—0.6
(7) mm long subsji. pseudoicacoreus
1. Leaf blades asymmetric; calyx cotylihirm; corolla membranaceous, the
lobes reflexed-recurved; apically free portions ol the filaments longer than
the anthers; anthers ovate, apically acute, dehiscent by wide, sublatrorse slits.
subsp. multipunctatus
Cybianthus subspecies multipunctatus (A. DC.) Pipoly is distributed in eastern
Amazonia and the Guianas in premontane forests on lateritic and white sands
of the Roraima Superimposed Sediments, while subsp. guyanensis is located
principally in central Amazonia in igapo forests (Pipoly 1992a). Subspe-
52
Si DA 18(1)
Fic. 1 3. (.yhiciii!l.i/is tiDhDhiL l^ipoly. A. I labic, showint; paniculate iiiHoresccnces. B. Branchler
apex. C. Adaxial leaf surlacc, showing midrib slightly raised but canaliculate. D. Abaxial
leaf SLirhice, showing lepidote scales and prominently raised midrib. E. Section of raceme,
showing coriaceous, delrate floral bracts, cotyliform calices, obcordate anthers with dis-
tally recurved apiculae. F. Pistillate flower bud. G. Pistillate corolla with one lobe removed,
showing lagenilorm pistil and siibsessile antherodes. A— F, drawn Irom holorype. F— (i, drawn
Irom 'iinunsci l()4'^. Figure drawn h\ Finda Fllis.
PfPOLY, Cybianthus in Ecuador and Peru 53
c'les, pseudokacoreus is distributed south-central and western Amazonia, along
banks of smaller streams on deep white sands. S\xhs]i&cies pseudokacoreiis and
guyanensis are sympatric only in central Brazilian Amazonia, and differences
in their ecology are not known in sufficient detail. In Ecuador and Peru,
subspecies pseudokacoreus may be most easily confused with C comperuvianus
Pipoly, but may be recognized by the thicker branchlets, longer floral bracts,
and carnose perianth.
10. Cybianthus timanae Pipoly, sp. nov. (Fig. 13). Type: PERU. Junin; Prov. Satipo,
Gran Pajonal, Mapari, ca. 12 km SW of Chequitavo, 10° 45' S, 74° 23' W, 1,300 m,
7 Apr 1984 (stam, fl), D, Smith 6782 (iiolotype: MO; isotypes: BRIT, US, USM).
Propter ramulos graciles angulatos, laminas ellipticas vel oblanceolatas, ad apices acuminatas
ad bases acutas, inflorescentiam paniculatam, calycem cotyliformem, necnon tubum staininarem
staminodiaremque inconspicuo, C. occigranatens i a.Tcte affinis, sed ab ea nerviis secundariis
16-40 (non 8-12)-]ugis, perianthiis chartaceis (non coriaceis), corolla cotylilormi (non
campanulata), lobis corollinis laevibus (non verrucosis), ad apices acuminatis (nee attenuatis),
conspicue (nee incounspicue) atro-punctaris, pistilo lageniformi (nee obnapiformi) pertacile
separabilis.
Subshrub to 1 m tall. Branchlets angulate, 2-2.5 mm diam., densely lepi-
dote. heaves alternate; blades chartaceous, elliptic to oblanceolate, (8—)
9-5-1 5(-l 9) cm long, (2.7-)3-5.5(-6.5) cm wide, apically abruptly acuminate,
caudate, the acumen 1—2.2 cm long, basally acute, decurrent on the peti-
ole, smooth and nitid above, pallid and moderately lepidote below, midrib
slightly raised and canaliculate above, not decurrent on the petiole, promi-
nently raised below, secondary veins 16-40, brochidodromous, planar to
somewhat impressed above, not bullate, the margin flat, entire; petioles
canaliculate, (1.3-)1.5-2(-2.2) cm long, glabrous above, densely lepidote
below. Staminate inflorescence: a panicle of 2—8 racemes branched from base,
(3-)4-6(-7) cm long; peduncle 3-7 mm long; rachis densely lepidote; floral
bracts coriaceous, deltate, 0.6-0.8 mm long and wide, apically acute, somewhat
cucullate, densely lepidote above and below, the margin entire; pedicels
cylindrical, 1-2.5 mm long, densely lepidote. Staminate flowers 4-merous,
cream; calyx chartaceous, cotyliform, 0.7-1 mm long, the tube ca. 0.2-0.3
mm long, the lobes ovate-triangular, 0.5-0.7 mm long, 0.3-0.4 mm wide,
apically acute, moderately lepidote, the margin entire, somewhat involute;
corolla chartaceous, cotyliform, 2—2.4 mm long, the tube ca. 0.2 mm long,
the lobes oblanceolate, 1 .8—2.2 mm long, 0.8—1 mm wide, apically subacuminate,
sparsely lepidote apically near margin, prominently black punctate and punctate-
lineate without, densely glandular-granulose throughout within, the mar-
gin glabrous, entire; stamens 1.4-1.6 mm long, the tube conspicuous,
chartaceous, 0.1—0.2 mm long, elobate, the apically free portions of the filaments
terete, 0.4-0.6 mm long, the anthers obcordate, 0.6-0.7 mm long, 0.3-
0.4 mm wide, apically apiculate, anther and apiculum distally recurved,
54 SioA LS(1)
the connective prominently black punctate dorsally; pistillodc conic, 1.2—
1.4 mm long, 0.3—0.4 mm wide, hollow, densely translucent glandular-
lepidote. Pistillate inflorescence a^ in staminate, but (1— )1.5— 3 cm long; pe-
duncle 2—4 mm long; floral bracts 0.4—0.6 mm long and wide; pedicels
obconic, 0.6—0.9 mm long, sparsely iepiclote, conspicuously black ptmc-
tate. PistilL/te flowers as in staminate but translucent green; calyx 0.7-0.9
mm long, the tube ca. 0.2 mm long, the lobes 0. "3-0.7 mm long, 0.3—0.4
mm wide, sparsely lepidote, corolla 1.2-1 .5 mm long, the tube ca. 0.2 mm
long, the lobes oblong to oblanceolate, 0.9-1 -3 mm long, 0.4—0.6 mm wide,
apically obtuse to subacuminate; staminodes 0.6-0.8 mm long, the tube
conspicuous, chartaceotis, ca. 0. 1 mm long, the antherodes subsessile, 0.6—
0.7 mm long, 0.3-0.4 mm wide; pistil lageniform, 1 .2-1.4 mm long, 0.3-
0.4 mm wide, hollow, densely translucent glandular-lepidote, the ovules
2-3, partially immersed on the placenta. Fr//il slightly depressed-globose,
4.5—5.5 mm long, 5.5-7.5 mm diam., the exocar)-) thin, black at maturity.
Distrih//ti())i. — Southeastern Ecuador to (4isco Peru, at 720—1,300 m
elevation.
Ecology and conservation status. — Cybianthiis t'nnanae usually occurs in wet
premontane forest on sandstone soils. The restricted distribution oi Cyhianthus
timanae indicates it should be considered a threatened species.
Etymology. — The species is named for Martin Timana de la Flor, former
Peruvian Field Associate ol the Missouri Botanical Garden, and currently a
graduate student at the University of Texas at Austin. Martin is specializ-
ing in the systematics of high altitude Caryophyllaceae.
PARATYPF.S. ECUADOR. Zamora-Chinchipe: Nans^aruza Canton; lower slopes of
Cordillera del Condor, above PachicLirza, Rfo Nangaritza Valley, 04° 07' S, 78" 3<S' W,
1,000-1,200 m, 6 Dec 1990 (fr), D. Nell & W. Falacios 9536 (BRIT, MO, QCNB); Rio
Nangarriza, Sliaime, confluence ol Rios Nangaritza and NLimpacakaime, 04° 20' S, 78°
40' W, 1,000 m, 7 Dec 1990 (fr), /). Nail 9602 (BRIT, MO, QCNE); Pachicutza. Rd. to
Hito, Cordillera del Condor, 04" 07' S, 78° 37' W, 1,000-1 ,100 m, 19 Oct 1991 (pist. fl
bud), W. PciLuios i't cil. 8346 (BRIT, COL, MO, QCNE), 20 Oct 1 991 (stam. fl), W. Palaaos
eta/. 8407 (BRIT, COL, MO, PORT, QCNE, USM); Parroc|uia Pachicutza, NE of military
camp, 900 m, 6 I5ec 1990 (fr), _/. _/</;vw//^ & E. Cn/,/l,u/ 134/9 (COL, QCA). PERU.
Cajamarca: CAitervo National Park, 1 2 km NE of San Andres de Cutervo, Transect 3, 06°
10' S, 78" 40' W, 2,230 m, 10 Sep 1991 (ster.), A. G7;y/n c/ .//. 74630 (BRIT, MO, USM).
Cusco: Pro\\ Quis|iicanchi. Camanci, Manin, along trail parallel to Ri'o Maniri to Quebrada
Garrote, 13°I7'S, 70"48AV, 720 m, r Oct 1990 (pisr. fl), ,\I, T/m/i/a 1047 (BRIT, CUZ,
MO, US, USM).
Cybianthiis timanae is most closely relared to C occigranatensis (Cuatrec.)
G. Agostini, but may be easily separated by its more numerous secondary
veins, chartaceous perianth parrs, cotyliform corolla with smooth, conspicuously
black punctate, actiminate lobes, and lageniform pistil. The involute calyx
lobes and the obconic pedicels of the pistillate flowers are also extremely
rare within the subgenus.
PiPOLY, Cybianthus in Ecuador and Peru
55
2 mm
D
M5)6K)
2 cm
Fic;. 14. Cybianthus c/uitrecasasii Pipoly. A. Habit, showing irregularly shaped, basaliy branched
panicles. B. Scaminate flower, habit, showing cotyliform calyx, infundibuliform corolla. C.
Opened staminate flower, showing inconspicuous staminal tube and lageniform pistillode.
D. Opened pistillate flower, showing obnapiform pistil and subsessile antherodes. A-C,
drawn from holotype. D, drawn from A. Gentry et al. 33960. Figure drawn by Juan Pinzon.
11. Cybianthus cuatrecasasii Pipoly (Fig. 14), Caldasia 18(3):285. 1996.
Type: COLOiVIBlA. Huila: Municipio La Plata, Vereda Agua Bonita, Finca iVIerenberg,
cerca del Cementerio (Jardi'n Botanico), 22 Abr 1982 (stam. fl)J. H. Torres 1034
(iiOLOTYPP,: COL).
Dioecious tree to 6 m tall. Branchlets narrowly angulate, 3-4 mm diam.,
56 Sida18(1)
densely appressed-lepidote, the scale margins frequently overlapping. Leaves
alternate; blades chartaceous, elliptic or oblanceolare, 8—20 cm long, 3-5—
6.5(— 6.9) cm wide, apically acuminate, the acumen (0.5-)l.()-1.5(-2.0) cm
long, basally acute, slightly decurrent on the petiole, midrib impressed above,
prominently raised below, the secondary veins 26-34 pairs, inconspicuous
above, prominently raised below, sparsely lepidote above at first, then pus-
tulate and glabrescent, densely appressed-lepidote below with margins fimbriate
and overlapping, the blade margin flat, entire; petioles canaliculate, (1 .0)1 .3-
1 .7(— 2.0) cm long, densely appressed-lepidote. Staniinate inflorescence: an irregular
panicle with 2—4 basal branches, 4-8 cm long; peduncle, rachis and pedicels
densely lepidote; peduncle 0.1—0.4 cm long; Horal bracts chartaceous, tri-
angular, subequalling the pedicels, 1.2—1.5 mm long, ca. 0.6 mm wide,
apically attentuate, densely lepidote below, the margin entire, glabrous; pedicels
cylindric, 1 .0-1 .5 mm long. S tarn inate flowers 4-merous; calyx carnose, cotyliform,
1.3—1.4 mm long, the tube 0.2—0.4 mm long, the lobes triangular or ovate-
triangular, 1.0—1.2 mm long, 0.6-0.9 mm wide, apically attenuate, con-
spicuously brown punctate, the margin entire, regular, with few scales; corolla
chartaceous, infundibuliform, 3.0—3.5 mm long, the tube 1.2—1.3 mm long
the lobes ovate, 2.0-2.2 mm long, 1.0—1.2 mm wide, apically rounded,
carinate medially, densely lepidote without and along the margins, glan-
duiar-granulose toward the apex and along the margins within, inconspicuously
brown punctate, the margins regular, entire; stamens 2.5—3 mm long, the
staminal tube membranaceous, inconspicuous, 1 .2-1 .3 mm long, hyaline,
elobate, glabrous, the apically distinct hlaments adnate to the corolla lobe
above the staminal tube 0.3-0.5 mm and apically free 0.1-0.3 mm, gla-
brous, the anthers ovate, 0.8-1 .0 mm long, 0.5-0.6 mm wide, distally reflexed,
apically apiculate, the apicuium slightly inHexed, basally cordate, the con-
nective dark, eglandular; pistillode lageniform, 1 .6-1 .8 mm long, densely
glandular-lepidote basally. Pistillate inllorescone: as in the staminate except
a simple raceme, 4.4—10 cm long; peduncle 0.2-0.5 cm long; floral bracts
lanceolate, 1 .2—1 .5 mm long, 0.3—0.4 mm wide; pedicels 0.7—1.3 mm long,
incrassate in fruit. Pistillate flowers similar to the staminate except calyx 1 .4—
1 .7 mm long, the tube 0.2—0.4 mm long, the lobes ovate, 1 .3—1.5 mm long,
0.6-1.2 mm wide; corolla 3.5—4.0 mm long, the tube 0.9-1.1 mm long,
the lobes elliptic, 2.7—3.1 mm long, 0.9-1 -3 nim wide, apically acute; staminodes
similar to stamens but 3.0—3.2 mm long, the staminodial tube 1.0 mm
long, the apically free portion of the filaments adnate to corolla lobes 1.3—
1.5 mm and apically free ca. 0.1 mm, the antherodes subdeltate, 0.8-1.1
mm long and wide, {always longer than wide), apically acute, basally sag-
ittate; pistil obnapiform, 1.8-2.0 mm long, the ovary 0.9-1.0 mm long,
1.2-1 .5 mm diam., densely translucent glandular-lepidote, the style 0.9-
1 . 1 mm long, the stigma bilobate, not capitate, the jilacenta deeply cupuliform.
PiPOLY, Cybianthus in Ecuador and Peru 57
the ovules 2, deeply embedded in the placenta below apical pores. Fruit
drupaceous, depressed-globose, 4-6 mm long, 6-8 mm diam., the endocarp
smooth, the embryo erect, 3 mm long.
Distribution. — Cybianthus cuatrecasasii occurs in the Western Cordillera
of Colombia in the Department of Antioquia, in the Central Cordillera near
the Macizo Colombiano, in the Departments of Huiia and Cauca, and in
the Western Cordillera of adjacent Ecuador, with disjunct populations in
the Province of Santiago-Zamora, and Zamora-Chinchipe, Ecuador, from
(l,200-)l,960-2,850 m elevation.
Ecology and conservation status. — Cybianthus cuatrecasasii occurs along the
margins of primary cloud forests and along small watercourses. The popu-
lation from Antioquia, Colombia is rare because it is from premontane plu-
vial forest, and has leaves much shorter than normal. Because it is known
only from primary forests, it should be considered threatened.
Etymology. — ^This species is named to honor the late Jose Cuatrecasas, prodigious
field botanist, monographer of many plant families, and ardent student of
the Colombian flora. Don Jose freely offered his advice and assistance to all
who asked, despite his limited time and numerous projects.
Specimens examined. COLOMBIA. Antiocjuia: Mpio. San Luis, Piedra de Casrrillon,
3-4 hours by foot S of town, 06° 01' N, 75° 01' W, 1,500-1,700 m, 8 May 1989 (A bud),
D. Dalyetal. 3926 (HUA, MO, NY, US). Huila: Cordillera Central, E slope, Finca Merenberg,
km 101 of La Plata-Popayan Rd., 13 km E ofSta. Letfcia, 02° 15' N, 76° 12' W, 2,300 m,
24 Mar 1986 (fl bud), 6. Stein 3721 (BRIT, MO); Cordillera entre cuencasde los rios Guarapas
y Guachicas, arriba de Palestma, al S\V de Pitalito, 2,000-2,300 m, 6 Feb 1943 (stam. fl),
r.R. Pasherg 19969('NY, US); FincaMerenberg, E de Volcan Purace, ccrcade lazona limi'trofe
con Cauca, 02° 26' N, 76° 12' W, 2,300 m, 1 Apr 1986 (bud), A. Gentry et al. 53881
(COL, MO, US), 3 Apr 1986 (pist. fl, fr), A. Gentry et al. 53960 (COL, MO, US); 15 km
NE de Algeciras, cerca del Campamento La Gironda, 2,400-2,850 m, 26 Mar 1944 (stam.
fl)£. Uttle7481 (COL, NY, US);Mpio. Sanjosedelsnos, VeredaElHornito, 1,960-2,000
m, 23 Jul 1980 (stam. fl), G. Lozano 3366 (COL); Mpio. La Argentina, arriba de Finca
Palmira, 2,100 m, 26 Sep 1984 (fl bud), G. Lozauo et al. 4133 (COL). Cauca: Moscopan,
margen del Rio San Jose, La Chorrera de Candelaria, 2,100-2,350 m, 1 Feb 1947 (fr),./.
C//atrecasas 23583 (COL, F, NY, US); Cucnca del Rio La Plata, 43 km SE de Popayan, 2,1 60
m, 26 Nov 1944 (stam. fl), F.R. Fosberg 22376 (NY, US). ECUADOR, Santiago-Zamora:
along Quebrada Fionda, vicinity of Rancho Achupallas, 2,500-2,700 m, 1 Oct 1 943 (stam.
fl),J. Steyermark 54552 (NY). Zamora-Chinchipe: Fiill ca. 2 km downstream from Campamento
Shaime along Rio Nangaritza, 900 m, 15 Feb 1994 (fr), H. van ckr Werff 13074 (BRIT,
MO, QCNE).
Cybianthus cuatrescasasii is unique within subgenus Conomorpha because
of its infundibuliform corolla and subapical portions of the filaments ad-
nate to the corolla lobe. The overlapping covering scales of the abaxial leaf
surface, branchlets and inflorescence rachis is found otherwise only in Cybianthus
crotonoides (Mez) G. Agostini, a species endemic to the Guayana Highland.
This species has been confused with Cybianthus peruvianus (Mez) G. Agostini,
58 SiDA 18(1)
but is easily separated from it by rhe infundibiiliform corolla, long pedicels
and eglandiilar connectives. CybuDtthiis ciuitreccncisi'i is closely related to C
Ictetiii (Mez) G. Agostini and C. oaigrciiiciteusis ((Aiatrec.) (3. Agostini, from
which it is easily distinguished by its inftindibulitorm corolla, apical por-
tions of the filaments adnate to the corolla and lack of lobes on the staminal
and staminodial tribes.
III. Cybianthus subgenus LaxifllorusG. Agostini, Acta Biol. Venez. 10:144.
198{); Pipoly, Bnccoma 35:61-80. 1983. Cww^w^r/'^v secnon C<iiio)iii)ypbnhi W\l\.,
Stirp. Surinam Selc'c:. 111. Y'A'^Y^pyii pdvte. Coiioinorphc/ subgenus Eiaonouwrphu Mez
in Engl., Pflanzenr. IV. 236(Heft 9):25-1 . 1 902. proper te nimore. Tyhi; Spiicii-s. Covoiiiorpha
lax/Jlora (Mixrz.) A. DC. = Cybitnitlu/.s spiaiU/s {\\. B. K.)G. Agostini. (lecto'iype: by
Agosrini, Acta Biol. Venez. 10:11'). 1980).
Terrestrial dioecious shrubs or sniaU trees. Roots jiositively geotropic. Trunk
distinguishable, leptocatiloiis, rhe growth dynamics following Rauh's Ar-
chitectural Model (Halle et al. 1978). Brauehlets tomentose, with dendroid
or stellate ferrugineous trichomes or both. Cataphylls and jiseudocataphylls
absent. Leaves alternate, petiolate, tomentose, glabrescent. Inflorescence a simple
raceme or rarely a spike, the peduncle 0.1-0.5 cm long, the staminate ra-
chis tortuous or lax, the pistillate rigid and erect or rarely lax. Flowers A- or
5-merous; calyx cotyliform, valvate, epunctate, inconspicuously pellucid or
conspicuously brown or orange punctate, glabrous, tomentose, or glandu-
lar-granulose, rarely bearing translucent glandular scales, the margins glandular-
ciliate; corolla campanulare, rhe lobes valvate, glandular-granuiose only along
margins without, but over the entire surface within, epunctate or incon-
spicuously pellucid or conspicuously brown or orange punctate, and punc-
tate-lineate, medially and basally glabrous, glandular-granuiose or with a
few stellate trichomes; stamens and staminodes with filaments basally con-
nate into a conspicuous or inconspicuous tube, the tube elobate or bearing
small lobes alternate with the apically free filaments, the anthers elongate,
triangular, distally curved, rarely erect, dorsifixed 1/4—1/2 from base, apically
acute, basally cordate, dehiscent by wide longitutHnal slits; staminodes re-
sembling sramens but reduced in size, the sterile anthers wirhout pollen or
at times producing abortive pollen; pistil obnapiform, the ovary translu-
cent glandular-lepidote, the style short, truncate with apunctiform stigma;
pistillode lageniform, the ovary translucent glandular-lepidote, hollow the
style elongate and curved apically, the nonfunctional stigma capitate. Fn/it
drupaceous, one-seeded, the exocarp thin.
Cybianthus subgenus Laxifloriis contains 6 species, 3 of which have been
recorded from Peru. To date, none are known from Ecuador, but we may
expect Cybianthus spicatiis (Kunth) G. Agostini in sandstone are^LS in the Cordillera
del CcSndor area.
PiPOLY, Cybianrhus in Ecuador and Peru 59
KEY TO SPECIES OF CYBIANTIIUS SUBGENUS LAXIFLORl'S
1. Inflorescence rachis black |-iunctate; calyx lobes coriaceous, delcace, medi-
ally carinate, basally rugose; fruit depressed-^ulobose, broader than long; large
trees to 18 m tall, terra finne "varillal seco" [dry, on white sand] forests.
12. C. nestor i i
I. Inflorescence rachis epunctate; calyx lobes chartaceous, widely to narrowly
ovate-triangular, medially thicked but flat; basally smooth; fruit globose, as
broad as long; shrubs or small trees to 1()(-1 2) m tall; riparian gallery for-
ests, either periodically flooded (varzea or iga|i6) or "varillal humedo" {wet,
white or brown sand] forests.
2. Leaf blades chartaceous, the tertiary veins prominently raised above; ca-
lyx lobes longer than broad; staminal and staminodial tubes inconspicu-
ous; receptacle or pedicel or both enlgarged in fruit; seasonally inundated
forests (varzea or igapo) 1 3. C. spicatus
2. Leaf blades coriaceous, the tertiary veins not visible from above; calyx
lobes broader than long; staminal and staminodial tubes consjiicuous;
neither receptacle nor pedicel enlarged in fruit; forests with
poor drainage on sand, but non-intmdating, ("varillal humedo")
M. C. fulvopulverulentus subsp. magnoliifolius
12. Cybianthus nestorii Pipoly, sp. nov. (Fig. 15). Type: PERU. Lorpto: Prov.
Maynas, Mishana, on Rfo Nanay, 03° 55' S, 73" 35' W, 150 m, 20 Jan 1985,
R. Vcfsqmz & N. JaniwilU, 6122 (iiolotypi,: MO; istrtYPEs: AMAZ, F, NY, LJS, USM).
Ob folia coriacea, rhachides inflorescentiares atro-punctato-lineatos, lobis calycinis deltatos
inter se aequilatos coriaceosque, C. Jeltato valde arete afhnis sed ab ea habitu arboreo (non
fruticoso) usque ad 18 (non 3) m, laminis anguste ellipticis vel anguste oblanceolatis vel
anguste oblongis (non obovatis) 9.5-1 8(-28), (nee 3.2-4,4) cm longis, 3.2-4.6 (nee 1.7-
2.8) cm latis, inflorescentiis 5-7 (non 1-4.2) cm longis, lobis calycinis 1.8-2 (non 1.3-
1.7) mm longis, carinatisque (nee planisque) denique secus marginem minute erosis (nee
undulatis), statim separabilis.
Tree to 18 m tall. Branchlets terete, 5-7(-15) mm diam., appressed fer-
rtigineous dendroid tomentose. Leaves alternate; blades thinly coriaceous,
narrowly elliptic to narrowly oblanceolate, rarely narrowly oblong, (9-5-)
11.5-14(-18) cm long, (3.2-)4-6 cm wide, apically obtuse, emarginate,
rounded or acute, basally acute to attenuate, decurrent on the petiole, gla-
brous and nitid above, glabrous and nitid but pallid below, midrib slightly
raised above, prominently raised below, decurrent through length of peti-
ole above and below, the secondary veins (l4-)20-47 pairs, prominulous
(slightly raised) above and below, inconspicuously pellucid punctate and
punctate lineate below, the margin entire, glabrous, revolute; petioles mar-
ginate, (2.2— )2.5— 3.2(— 3.7) cm long, glabrous. Staminate inflorescence: un-
known. Pistillate inflorescence: an erect raceme, 5-7 cm long, the rachis black
punctate-lineate, ferrugineous glandular-granulose, glabrescent; floral bracts
coriaceous, linear, minute, 1-1.1 mm long, 0.2-0.3 mm wide, apically acute,
densely glandular-granulose below, glabrescent, the margin glandular-cili-
60
SioA 18(i)
Fk;. 1 5. Cyhuinthiii nestorii Pipoly. A. Habir, showini; depressed-^ lobosc fruits and leaf blades
with prominuloLis secondary veins and revokice margins. B. Portion of infructescence, showing
depressed-globose fruit and carinate, rugose calyx lobes with erose, sparsely glandular-ciliolate
margins. (;. Abaxial leaf stirface, showing prominently raised midrib and promintilotis secondary
veins. D. Branchlet apex, showing appressed dendroid tomentum. A-D, drawn from holo-
type, by Linda Ellis.
PiPOLY, Cybianthus in Ecuador and Peru 61
ate; fruiting pedicels cylindrical, 2—4 mm long, densely glandular-granulose,
minutely black punctate apically. Pistillate flowers unknown; fruiting calyx
coriaceous, 1.8—2 mm long, the tube 0.3—0.5 mm long, the lobes deltate,
1.3—1 .6 mm long, 1 .4—1.7 mm wide, apically acute, medially carintae, rugose
basally, the margin minutely erose, sparsely glandular-ciliolate; corolla, staminodes
and pistil unknown. Fr/zzV depressed-globose, 4-6 mm long, 6-8 mm diam.,
black at maturity, inconspicuously pellucid punctate.
Distribution. — Known only from the Rio Nanay Drainage Basin, Maynas
Province, Department of Loreto, Peru, at 150 m elevation.
Ecology and conservation status. — Cybianthus nestorii occurs in "varillal seco"
habitats, consisting of tall terra firme (non-inudating), relatively dry forest
on deep white sands. It is a rare species and thus, should be considered threatened.
Etymology. — It is a pleasure to dedicate this species to Nestor Jaramillo,
of Iquitos, Peru, prodigious collector, and authority on plant collecting in
tropical forests. Nestor, with his supervisor, Rodolfo Vasquez, form one of
the most formidable botanical exploration teams in South American botany.
Paratypes: PERU. Loreto: Prov. Maynas, Dtco. Iquitos, Puerto Almendras, UNAP, Tree
No. 324, Tree inventory, 03° 48' S, 73° 25' W, 122 m, 17 Jan 1993 (ster.), C. Grdndez. N.
Jaramil/o et al. 5321 (BRIT, MO, UNAP), Tree No. 373 (ster.), C. Grdndez, N.Jaramtllo et
al. ')570 (BRIT, MO, UNAP, USM); Tree No. 651 (ster.), C. Grdndez. N. Jaramillo et al.
5642 (BRIT, MO, UNAP).
Cybianthus nestorii is unique within the subgenus because of its autapomorphic
depressed-globose fruits and carinate calyx lobes. Other characters which
readily seaparate it from its closest relative, Cybianthus deltatus Pipoly, of
the Rio Guainia drainage basin of Venezuela, include its much larger ar-
borescent habit, larger elliptic to narrowly oblanceolate leaves, longer
inflorescences, and longer, minutely erose calyx lobes.
13- Cybianthus spicatus (Kunth) G. Agostini (Fig. 3B), Acta Biol. Venez.
10: 146. 1980. Mymm ipicata Kunth in H.B.K., Nov. Gen. Sp. 3:250. 1818. Omomorpha
spicata (Kunth) Me2 in Engl., Pflanzenr. IV. 236(Heft 9):259. 1902. Tvpi;: VEN-
EZUELA. Thiuutorio Fp.DBRAL Amazonas: In inundated area along the Ri'o Atabapo,
without date (stam. fl), A. von Humboldt & A. Bonpland 1096 (iiolotypi;: P-BON;
isotyph: B-destr.).
Wallenui laxiflora Mart., Nov. Gen. Sp. PI. 3:89- 1829- Conomorpha laxiflora (Mart.) A.
DC, Trans. Linn. Soc. London, Bot. 17:102. 1834, Typp: BRAZIL. Amazonas: Prov.
Rio Negro, "In sylvis Japurensibus," Jan 1826 (stam. fl), C. Martins s.)i. (i.i;c:tot^'fi;
by Pipoly 1983b: M; isolhctotype: CGE).
Conomorpha laxiflora var. longifolia Miq. in Mart., Fl. Bras. 10:302. 1856. TvFi;: BRA-
ZIL. Amazoxas: Prov. Rio Negro, vicinity Barra [Manaos], Dec-Mar 1850-5 1 (stam.
fi), R. Spruce !()4() (LEf;i()TYPi"., here designated: M; isoleci-otypp.s, CGE, GH, GOET,
K, LD, LE, OXF, U).
Conomorpha laxiflora var. latijolia Miq. in Mart., Fl. Bras. 10:303. 1 856. Conomorpha tatijolia
(Miq. in Mart.) Mez in Engl., Pflanzenr. IV. 236(HeIt 9):255. I 902. Typi;: VENEZUELA.
62 SiDA 18(1)
[CUYANA]. 1839 (scam. H), R. Sdmiiihiiriik 1002 (uicrorvPH by Pipoly 1983b: G;
isoi.EC'iorYPr,: C^GE).
Conomorpha catululleciiui Mcz in En^M., Pflanzenr. IV. 236(Heft 9): 23(3. 1 902. Typi': VEN-
EZUELA. [GUYANA}. 1840 (scam, fl), R. Schowhuriik 88') (hopotypi:: B-dcstroyed;
fragment at F; li-,c:ix)TYPi;, here clesignatetl: G; isoleci'otypi'.s: CGE, GH, K-2 .sheets,
US, W).
CdJKiiuorlibci grcitultjlorci Mez in Engl., Pflanzenr. IV. 236(HeI't 9):258. 1902. Typp.:
HRAZIL.Amazonas: Rio Negro, above Moureira, Dec 1 85 1 (stam. fl.), R. Spr/ia- 1946
(iioloiypp: B- destroyed; i.HC-roTYPP,, here designated: K; i.s()i.i;( totypps: CCtE, GH,
GOET, LD, LE, NY, OXF).
Coimiumfbciglam-onihoisMfi m Engl., Pflanzenr. IV. 236(HeFt 9):260. 1 902. Type: BRAZIL.
Ama/()\'as: Rio Negro, about 00° 30' S, (ri° 00' W, 21 Jun 18"?" 4 (stam. fl), J. Trail
)()8 (hoi.otypp,: K).
Coi/ijworplh/ >/k/(/c/m/si.\ A.C Smith, J. Arnold Arbor. 20:300. 1 93 I . Typi;: BRAZIL. Ama/.oxas:
Mtinicipality oi: F^tmiayta, between Monte Cristo and Sta. Victoria on Rio Ipixtma,
1 5-17 Nov 1934 (stam, fl), B. Krukoff724U (holotype: NY; isotypics: A, R M, MAD-
Y, MICH, MO, S, US).
Co>/oworp/ja gnni/is A.G. Smith, Bull. Torrey Bot. Glub 6'7:295. 1940. Tyi>i:: GUYANA.
Artibaru River, Kako Territory, Upjier Mazaruni drainage, 600 m, 2 Feb 1939 (stam.
fl), A. Pnikiis 181 (hou)type: NY; i,s()typi;s: BR, F, GH, M, MO, NY, S, US).
Shi7/b or sijiall tree to 12m tall. Braiichlets thin to moderately thick, (.^.5-)
4-8(-20) mm diam., appressed to floccose-dendroid and stellate ferrugin-
eous tomentose, the stellate hairs often appearing lurfuraceous, glabrescent.
Leaves alternate; blades tliinly coriaceous to chartaceous, to membranaceous,
ovate, elliptic, obovate or rarely oblanceolate, (4.5— )6— 17.5(— 2 1 ) cm long,
2.9— iS(— 1 1 .5) cm wide, apically actite, rounded, obtuse or rarely emargin-
ate, basally acute to cuneate, glabrotis, pustulate and often nitid above, sparsely
lerrugineous puberulent and minutely glandular-lepidote below, the mid-
rib, secondary and tertiary veins prominently raised above and below, the
midrib decurrent to the base of the petiole, pellucid or black punctations
obscure below, the margin slightly revolute, entire; petioles distinctly marginate,
(1— )1 .5— .3.2(— 4) cm long, stellate and dendroid ferrugineous tomentose, early
glabrescent. Slciiiii)hite 'inflorescence, a simple raceme, lax, (3— )6-9(-l 2.5) cm
long, the rachis epunctate, glandular-granuk)se with scattered stellate and
dendroid ferrugineous trichomes, glabrate; floral bracts chartaceous, linear-
lanceolate, ().7-L.7(-2.2) mm long, densely ferrugineous tomentose, early
caducous; pedicels cylindrical, 0.9— 2.1 (-3) mm long, glabrous to densely
glandular-granulose. Staimnate flowers (4— )5-merous, chartaceous, cream to
yellow, with a sweet odor; calyx shallowly cotyliform, 1.5—2.5 mm long,
the tube 0.3— 0.6 mm long, glandular-granulose at first, glabrescent, the
lobes ovate or ovate-triangular, (1 .2-)l .4-2 mm long, 0.9—1.3 mm wide,
acute to abruptly acuminate apically, somewhat cordate basally, prominently
thickened medially, conspicuously punctate, the margins mostly entire, but
occasionally erose apically, glandular-ciliate; corolla campanulate, 3.6—4.8
mm long, the rube 1-1.2 mm long, the lobes ovate, 2. 1-3.4 mm long, 1.2—
PiPOLY, Cybianthus in Ecuador and Peru 63
2.6 mm wide, slightly reflexed at anthcsis, obtuse to acute apically, epunctate
or conspicuously brown punctate; stamens 1.6—2.2 mm long, the tube
membranaceous, inconspicuous, 1—1.5 mm long, the apically free portions
of the filaments 0.3-0.55 mm long, the anthers elongate-triangular, 0.7-
1 .2 mm long, 0.2—0.3 mm wide, dorsifixed ca. 1/3 from base, apically acute,
basally cordate, distally recurved, the connective inconspicuously brown punctate;
pistillode lageniform, 1.3-1.8 mm long, the ovary 0.6—0.9 mrn long, 0.5—
0.75 mm diam., densely translucent glandular-lepidote, the style elongate,
0.8-1.3 mm long, slightly curved apically, the stigma capitate, 0.1-0.2 mm
long. Pistillate inflorescence as in staminate, erect, (3.3-)4.5-13(-17.5) cm
long; floral bracts 0.5-2.4 mm long; pedicels 1.3-1.8 mm long, accrescent
in fruit to 3-1 mm long, the receptacle or pedicel or both incrassate in fruit.
Pistillate flowers as in staminate but chartaceous, yellow; calyx cotyliform,
1.3—1-9 mm long, the tube 0.6—0.9 mm long, the lobes widely to narrowly
ovate, 1.1-1.7 mm long, 0.9-1.5 mm wide, erect, sparsely brov/n punc-
tate, prominently thickened below, the margins entire or occasionally erose
apically; corolla 2.7—3.5 mm long, the tube 0.7—0.9 mm long, the lobes
1.6—2.7 mm long; staminodes as in stamens but 1 .6—2.1 mm long, the tube
chartaceous, conspicuous, 0.7-1.1 mm long, the apically free portions of
the filaments 0.3-0.5 mm long, the antherodes 0.6-0.9 mm long, 0.2—0.3
mm wide, at times producing abortive pollen; pistil obnapiform, 1.6—2.2
mm long, the ovary 0.8-1.2 mm long, 0.9-1.2 mm diam., densely trans-
lucent glandular-lepidote, the placenta cupuliform, ovules 4, partially im-
bedded, the stigma capitate, 0.1-0.2 mm long. Fruit globose, 2.5-6 mm
long and in diam., prominently black punctate-lineate, with a tew persis-
tent lepidote glandular scales below the usually persistent style base, green
then purple, then black.
Distribution. — Venezuela, Colombia, Peru, Brazil and Guyana, in inun-
dated forests (varzea and igapo), from 100—400 m.
Ecology and conservation status. — Cybianthus spicatus is restricted to primary
riparian habitats on white sandy soils. It is common in igapo and varzea,
but the latter only when sufficient c^uantities of sand exist, a mixed varzea
type. While it is a widespread species, it is sensitive to soil compaction,
and should be considered threatened.
Etymology. — The epithet refers to the inflorescence shape, which is a raceme
bearing flowers on short pedicels, thus appearing spicate.
Represenrarive specimens examined. PERU. Huanuco: Cerros del Sira, SW slope of
the Rfo Lulla Pichis, 1 190 m, I 2 Jul. 1 969 (fr),_/, Wolffe 12295 (F, NA); 100 m, 22 Jul
1969 (fr), 7.' D//cllty H 1 24 (F, NA); Prov. Pachicea, region of Pucallpa, W part of Sira Mountains
and adjacent lowland, ca 24 km SE ro 26 km ESE of Puerto Inca, next to Campamento
Pato Rojo, 09° 27' S, 74" 46' W, 1 ,000 m, 27 Jan 1 988 (pist. fl), W. Momwetz & B. Wallnofcr
72-27788 (BRIT, MO, W,W[]), 1,320 m, 1 May 1988(fr),/3. WW/«r//tT 7 7 /-/.5<S\S' (BRIT,
W, WU), 1 ,230 m, 1 3 Jun 1 988 (fr), B. Wcillndfer I I2-I368ti (BRIT, W, WU). San Martin:
64 Si DA 18(1)
Lamas, on old trail from San Antonio ck- Cumbasa, S oi Shapajilla, upper slopes of Cerro
Isco, 06" 22' S, 76° 23' W, 6()()-S()0 m, 5 Oct 19H6 (btul), S. Knapp et al. 85 14. (fr), S.
Kncipp €t al. 8") 11 (MO, US, USM).
Cybtanthus spicatiis is a polymorphic ochlospecies, sensu White (1962),
Prance (1972) and Pipoly (1983a), wirh many semi-isolated populations
throughout the Amazon and eastern Guayana Floristic Province (Maguire
1979). These localized populations have produced several seemingly dis-
tinct ecotypes, resulting in overdescription. Collections oiCybianthus spkatiis
from Peru match the type oi Conomorpha gracilis. A. C. Smith, described from
Guyana, in every detail. Fieldwork conducted in Guyana, Venezuela, Bra-
zil and Peru since my earlier treatment (Pipoly 1983a), where I discuss the
four ecotypes present within the species, and the synonymy rationale, has
confirmed the concept that Cyhianthus spicatns is a polymorphic ochlospecies.
Steyermark (1981 ) discussed the fact that Richard Schomburgk's collec-
tions attributed to Guyana are now known to be from Venezuela.
14. Cyhianthus fulvopulverulentus (Mez)G. Agostini subsp. magnoliifolius
(Mez) Pipoly (Fig. 3 A), Brittonia 35:72. 1983. Commwrpha magmliifolia
Mez in Engl., Pflanzenr. IV. 236(Heft 9):258. 1902. Cybunithm »ui\^u,iliifdiin (Mez)
G. Agostini, Acta Biol. Venez. 10:146. I 980. Type: SURINAME. without locality,
(scam. II), H. Kt',v.e/ 244 (i.ectotyph by Pipoly 1983a: GOET).
CoiniiiKirphi! rigicLi Mez, Repert. Spec. Nov. Regni Veg. 16:420. 1920. syn. nov. Type:
BRAZIL. Amazona.s: "Hylaea," without date (Fr), E. Ilk 8722 (iiolotype: B-desrr., F
Neg. 4831). Despite searches of herbaria housing significant Ule collections, no du-
plicates of the type have been located. Therefore, I select a neotype herewith: Type:
C;LJYANA [BRITISH GUIANA]. Orealla Savanna, Corenryne River, Sep 1879 (fr),
/;. Imth/irn 6/9 (neotype here designated: K; KSONEOIyte: BRG). The neotype was annotated
by Mez in 1901, bur was not mentioned in the protologue.
Shr/ib or tree to 10 m tall. Branchlets mostly thick, (3-)5.2-12 mm diam.,
glabrate or densely stellate and dendroid ferrugineous tomentose and glan-
dular-granulose, at times glabrescent. Leaves alternate, at times clustered
and approaching pseudoverticillate; blades coriaceous, elliptic, oblanceolate,
ovate or obovate, 7.2-21 .7 cm long, (2.2-)3.6-9.5(-17.1) cm wide, apically
obtuse, acute, rounded or emarginate, basally obtuse to acute (rarely acuminate),
sparsely pitted with superimposed glandular lepidote scales and glabrous
or glandular-granulose, at times sparsely ferrugineous tomentose above and
below, glabrescent, eptmctate, the margin entire and revolute; petioles marginate,
(1 .3—)! .6— 4.2(— 4.6) cm long, ferrugineous tomentose, and glandular-granulose,
glabrescent. Staininate inflorescence: a simple, tortuous raceme, rarely lax, (3.6-)
4.5-1 5 cm long, the rachis maroon, opaque, glandular-granulose or with a
few scattered dendroid ferrugineous trichomes, glabrescent; floral bracts
chartaceous, linear-lanceolate, 0.8—3.6 mm long, ca. 0. 1—0.2 mm wide, densely
ferrugineous tomentose, caducous; pedicels cylindrical, ({).2-)2.9— 3.6 mm
PiPOLY, Cybianthus in Ecuador and Peru 65
long, densely glandular-granulose, glabrescent. Sta77iinate flowers 4- or 5-merous,
chartaceous, white, cream or yellow; calyx deeply cotyliform, 0.9-2.6 mm
long, densely glandular-granulose then glabrescent, the tube 0.3-0.7 mm
long, unequally divided, the lobes chartaceous, ovate to ovate-triangular,
0.6-1.9 mm long, 1.1-2.4 mm wide, obtuse to acute or acuminate apically,
prominently orange punctate, densely ferrugineous tomentose and ferrugi-
neous glandular-granulose or partially glabrescent or ferrugineous glandu-
lar-granulose and then sometimes totally glabrescent, the margins extremely
undulate, entire, densely glandular-ciliate; corolla campanulate, (2.8-)3.3-
4.7(_5.5) mm long, the tube (0.6-)0.9-1.4(-1.7) mm long, the lobes ovate,
(2-)2.2-3.1(-4) mm long, (1.2-)1.5-2.4(-2.8) mm wide, the lobes highly
reflexed at anthesis, obtuse to acute apically, conspicously orange punctate
and punctate-lineate, at times scattered ferrugineous trichomes before an-
thesis without, the margin entire; stamens (2.4-)2, 8-3-4 (-3.8) mm long,
the staminal tube 0.8-1.1 mm long, the apically free portions of the fila-
ments 0.6-0.9 mm long, the anthers elongate-triangular, (0.8-)l-1.4 mm
long, apically acute, basally cordate, dorsifixed ca. 1/4 from base, distally
recurved or rarely and aberrantly erect; pistillode (1.6-)1.9-2.5 mm long,
the ovary 0.7-1.1 mm long, 0.7-1.2(-1.4) mm diam., densely translucent
glandular-lepidote, the style elongate, curved, 1-1.6 mm long, the stigma
0. 1-0.2 mm long. Pistillate inflorescence as in staminate but erect (3-)4-l 1 .4
cm long, the rachis green then red; floral bracts 0.7-2.9 mm long, ca. 0.1-
0.2 mm wide; pedicels cylindrical, (0.2-)0.6-1.5(-2) mm long. Pistillate
flowers as in staminate but dull yellow to brown; calyx 1.2-2.6 mm long,
the tube 0.4-0.7(-l) mm long, the lobes 0.7-2 mm long, 1 .3-2.4 mm wide;
corolla (2.7-)3.2-3.5(-4) mm long, the tube 0.6-1.2 mm long, the lobes
2-2.3(-2.6)mm long, 1.1-1.5 mm wide; staminodes as in stamens but 2.4-
2.9 mm long, the staminodial tube 0.7-0.9 mm long, the apically free portions
of the filaments 0.7 mm long, the antherodes triangular, 0.6-0.9 mm long;
pistil obnapiform, 1. 9-2. 5 (-3) mm long, the ovary (0.7-)0. 9- 1.6 mm long,
(0.7-)1.3-1.7(-2.2) mm diam., densely translucent glandular-lepidote, the
style thick, (0.4-)0.6-0.9(-1.3) mm long, the stigma pseudocapitate, very
minutely 4-lobed, up to 0.3 mm long. Fruit globose, 3-9 mm long and in
diam., green, then red, then black, inconspicuously pellucid punctate.
Distribution. — Venezuela, Guyana, Surinam, French Guiana, Brazil (Amazonas,
Roraima, Para, Mato Grosso) and reported here for the first time from Peru
and Bolivia 50-500(-850) m.
Ecology and conservation status. — Cybianthus fulvopulverulentus subsp . magnoliifolius
is extremely common in wet tepui savannas, but is rare in Peru in varillal
humedo (wet sandy, non-inundating) habitats. It is anticipated, but has not
yet been collected, in Ecuador.
Etymology. — The epithet, 'fulvopulverulentus' refers to the rusty tomen-
66 Si DA 18(1)
turn of the species, forming a powdery like vestiture on ciie adaxial leaf surface,
caducous as the leaf matures. The subspecific epithet refers to the shape and
shiny adaxial leaf surface, giving the general impression of a Magnolia leaf.
Specimens examined. PERU. Loreto: Prov. Maynas, Iquicos, Nma rumi-Ri'o Nanay, 03°
48' S, 73" 25' W, 122 m, 5 Mar 1987 (bud), R. Vau/uez et al. 8905 (AMAZ, MO, US,
USM). BOLIVIA. Santa Cruz: Prov. Velasco, Parque Nacional Noel Kempff Mercado,
Campamenco Huanchaca I, 3.35 km from Rio Pauserna, 13° 56' 01" S, 60° 49' 30" W,
600 m, 22 May 1994 (fr), L. Arroyo et al. 738 (BRIT, MO, USZ), 26 Jul 1995 (pist. H), T.
Killeen et al. 7328 (BRIT, MO, USZ), (stam. ft), T. Ktlleen et al. 7330 (BRIT, MO, USZ),
850 m, 3 Nov 1995 (Ir), A. Rodriguez &J. Suruhi 66'-) (BRIT, MO, USZ).
Specimens cited here are the first known from Peru and Bolivia. Those
collections represent remarkable disjunctions, the closest population known
heretofore occurring approximately one degree north of the Rio Negro in
southern Venezuela near the Brazilian frontier, or halfway across Brazil to
the east, on Serra do Cachimbo, state of Mato Grosso. To date, no speci-
mens are known from Ecuador.
In my previous treatment (Pipoly 1983a), I included Conomorpha rigida
in the synonymy oi Cybianthus fulvopulverulentus (Mez) G. Agostini subsp.
fulvopulverulentm. The type at Berlin had been destroyed and I synonymized
it based on the description and photo. However, the discovery of two du-
plicates of the Imthurn gathering annotated by Mez permitted neorypification
of the name with the Kew specimen and its inclusion here. Populations
represented by this collection differ from the type of subspecies magnoliifolius
only in their diminutive stature and vegetative parts.
IV. Cybianthus subgenus Comomyrsine (Hook, f ) G. Agostini, Acta Biol.
Venez. 1 0:162 1 980. Cuimmyrsme Hook. f. m Bench. & Hook., Gen. PL 2:643. 1 876;
Wetgeltia A. DC. subgcniis Comomyrsine (Hook, f.) Mez in Engl., Pflanzenr. IV. 236(Hefc
9):290. 1 902. Pax /« Engl. &Prantl, Nat. Pflanzenfam. IV, 1:92, 1897;J.F. Macbr., Field
Mus. Nat. Hist., Bot. Ser. 1 3 (5, 1): 175. 1 959- Type Spi;cies: Cybianthus sprucei {Hook.
f.) G. Agostini (m:ct()type: by D'Arcy, Ann. Missouri Bot. Gard. 60:445. 1973.)
Terrestrial, erect, monoaxial suhshriibs or treelets to 2(-5) m tall. Roots positively
geotropic. Bark mostly brown, smooth, longitudinally fissured or transversely
checked. Trunk distinguishable, leptocaulous, monoaxial, following mor-
phogenetic dynamics of Corner's Architectural Model (Halle et al. 1978),
growth rythmic. Stems terete distally, sparingly to densely glandular-papil-
late, at times with rufous hydropotes or orange glandular lepidote scales,
without lenticels. Cataphylls alternate or pseudoverticillate, alternating with
pseudoverticels of leaves, or apparently axillary to them, linear-subulate to
acicular, rigid to membranous, keeled or flat, prominently punctate or punctate-
lineate, glabrous, glandular-papillate, bearing hydropotes, or orange lepi-
dote scales. Leaves pseudoverticillate or alternate, apically acute to rounded,
often mucronulate, basally symmetric or asymmetric, acute, obtuse, or ta-
PiPOLY, Cybianthus in Ecuador and Peru 67
pering, rarely auriculate, the venation camptodromous, to brochidodromous,
apically acute or attenute to rarely rounded, mucronulate or not, basally
acute, cuneate, or rarely obtuse, at times subauriculate, symmetric or asymmetric,
black or pellucid punctate, bearing hydropoten, at times with orange glan-
dular lepidote scales or glandular papillae, the margin opaque or scarious,
entire, subentire, undulate and sparse dentate or pectinate-dentate, flat or
inroUed, the teeth vascularized or not; petioles pulvinate (basally swollen),
canaliculate, except rarely marginate distally. Infioreicence pinnately to tripinnately
paniculate, (rarely reduced to a pseudoraceme), pyramidal to columnar, the
branches spicate or racemose (rarely glomerulate); inflorescence bract lin-
ear-subulate, indistinguishable from the cataphylls; branch bracts membranous,
linear, subulate; floral bracts linear-lanceolate, subulate, subtending or rarely
on the pedicel, variously punctate, glandular-papillate or glabrous; pedicels
terete, thin, or obsolete. Flowers unisexual or bisexual, 4-5(-6)-merous,
homomerous or heteromerous; calyx cotyliform, at times unequally divided,
the lobes valvate, spreading, linear-subulate or rarely subdeltate, apically
acute, attenuate or rarely premorse, pellucid to black punctate, the margin
entire or erose-serrulate, glabrous or glandular-ciliate; corolla rotate to subrotate,
the lobes valvate, linear-subulate, apically acute, rounded or attenuate, es-
sentially glabrous without, densely glandular-granulose throughout within,
pellucid or black punctate, the margin irregular, entire; stamens and staminodes
similar, the staminodes reduced in size, partially connate to form a con-
spicuous or inconspicuous tube, at times bearing lobes alternating with apically
free filaments, the tube adnate to the corolla, at times so as to mimic epipetalous
stamens, the apically free portions of the filaments terete or flat, glabrous
or glandular-granulose, the anthers dorsifixed, as wide or wider than long,
apically rounded, obtuse or emarginate, basally cordate to deeply cordate,
dehiscent by wide longitudinal slites; pistillode lageniform, clavate or conic,
hollow, or entirely absent; pistil obpyriform, subglobose, obturbinate or clavate,
the ovary terete, sparsely to densely translucent glandular lepidote, glan-
dular-papillate or glabrous, the placenta umbonate, 2-3(-4)-ovulate, the
ovules immersed in the placenta ca. 1/2 their length, the style short or ves-
tigial, the stigma large, capitate, the margin lacinate, the lacinae with large
lobes, the stigma early caducous. Fruit globose to depressed-globose, the
exocarp sometimes fleshy, the mesocarp and endocarp stony, black or pellu-
cid punctate, one-seeded, the testa corrugate, the embryo cylindrical, transverse.
Distribution. — From the Darien of Panama southward through the Andes
of Colombia, Ecuador, Peru and Bolivia to Rondonia, Brazil, at elevations
of sea level-2,2()() m elevation.
Ecology. — Members o^ Cybianthus subgenus Comomyrsim occupy the for-
est floor, growing in sheltered areas under cliffs, and in other shady areas.
The monoaxial stems are weak and break easily when trampled. They are
68 Si DA 18(1)
extremely sensitive to soil compaction and occur mostly in areas where leaf
litter accumulates.
Pipoly ( 1 987) demonstrated that Cyhianthus subgenus Comomyrsine is most
closely related to subgenus Triadophora by the monoaxial habit. The cataphylls
of Comomyrsme without petiolar structures, as opposed to the petiolate
pseudocataphylls found in subgenus Triadophora, serve to further distinguish
Comomyrsine. Likewise, the caducous, oversized stigma with lacinate lobes
is unique to subgenus Commnynine. The subgenus contains 8 species, of which
7 occur in Ecuador and Peru.
KEY TO SPECFF.S OF CYBIANTUUS SIIRGF.NUS COMOMYRSINE
1. Leaves 5.7-29 cm long; stems less than 1 cm diameter.
2. Stems apices bearing orange hydropotes; leaf blades linear-lanceolate, 1.6-
4.0 cm wide, the margin repand to undulate; petioles 0.6-1.0 cm long;
inflorescence pseudoracemose, the flowers glomerulate 15. C. verticilloides
2. Stem apices bearing rufotis glandular-papillate or with rufous hydropotes;
leaf- blades ovate, obovate, oblanceolate, elliptic, or oblong, (4.0-), 10-
22(-30) cm wide, the margin regular, entire or minutely straight-serru-
late; petioles (I.O-)1.5-5(-21) cm long; inflorescence pyramidal-bipin-
nate paniculate, or a columnar thyrsoid panicle, never a pscLidoraceme,
the flowers spicate, corymbose, or rarely cymose.
3. Stem apices sparsely glandular-papillate, bearing rtifous hydropotes; leaf
bases broadly rounded, the margins scarious; petioles progressively longer
acropetally along the stem, 2.0-1.5 cm long; inflorescence paniculate,
the flowers corymbose; pethcels 1.0-2.5 mm long; sepals deltate; pet-
als CLicullate apical ly, the margin glandular-granulose; filaments
^1'iIt"()us 1(S. c. croatii
5. Stem apices densely glandular papillate, without hydropotes; leaf bases
acute, to acute with a rounded base, the margins opacjue; petioles subec]ual
along stem, slightly shorter acroi^etally, 1.6-2.0 cm long; inflorescence
pinnately paniculate, the flowers spicate or racemose; |uxlicels absent;
sepals ovate-lanceolate; petals flat, the margin glabrous; filaments glandular
MHinulose 17. c. humilis
1. Leaves (16-)31-105 cm long; stems 1-5 cm diameter.
4. Leaves oblong to elliptic, apically rounded or rarely acute, not mucrontilate,
basally abruptly acute to broadly rounded; petioles ( (2.5-)5 cm long; flowers
homomeroiis, 4— or 5-merous.
5. Shrub or tree to 5 m tall; inflorescence a pyramidal |ianicle, the branches
spicate, the flowers sessile to subsessile; leaf base symmetric; cataphylls
(2.5-) 3.5-5.0 cm long 18. C. sprucei
5. Semi-woody shnfl^is to l(-3.5)m tall; inflorescence a columnar, thyrsoid
panicle, the branches racemose, the flowers on pedicels 2.3-5.0 mm
long; leaf base asymmetric; cataphylls 0.6-2.5 cm long 19. C. simplex
4. Leaves oblanceolate, apically acute, mucronulate, basally gradually tapered
on the petiole, olten subauriculate; petioles 1 .5-2 cm long; flowers het-
eromerous, tlie calyx 5-6-lobed, the corolla 4-5-lobed.
6. Leaves chartaceous, 31-55 cm long, the margin flat, entire; cataphylls
PlPOLY, Cybianthus in Ecuador and Peru 69
1 .5-4.0(-4.5) cm long; calyx lobes deltate, the margin glabrous; co-
rolla carnose 20. C. kayapii
6. Leaves coriaceous, longer than 5 5-1 25cm, rhe margin mroUed, densely
and minutely serrulate; cataphylls 4.0-6.5 cm long; calyx lobes linear-
subulate, the margin densely glandular-granulose; corolla membrana-
ceous 21. C. anthuriophyllus
15. Cybianthus verticilloides (Cuatrec.) G. Agostini (Fig. 4B), Acta Biol.
Venez. 10:165. 1980. Weige/fia vertni/Iozdes Cuatrec, Revisvj. Acad. Colomb. C'\.
Exact. 8(31):327. 1951. Type: COLOMBIA. Valle del Cauca: Rio Call riverbed,
between Quebradade Juntas and El Recreo, 2,070-2,260 m, 7 Jul 1946 (stam. R),J.
Cucitrecasas 2 1981 (holcjtype: F; isotype: COL).
Shrub to 1 m tall. Stem terete, 5-8 mm diam., sparingly rufous glandu-
lar-papillate and bearing orange hydropotes, early glabrescent, the bark
horizontally checked. Cataphylls chartaceous, subulate, (1.2— )1. 5-2.1 mm
long, 0.9-2. 1 mm wide, flat, apically long-attenuate, basally somewhat decurrent,
sparingly glandular-papillate and orange lepidote above and below, black
punctate. Leaves pseud overticiUate; blades chartaceous, linear-lanceolate, 12-
20 cm long, 1.6-4.0 cm wide, apically long-attenuate, basally acute, grey-
ish-green above and pallid green below when fresh (teste Cuatrec), spar-
ingly rufous glandular-papillate and orange lepidote above and below at
first, persistent only below, inconspicuously pellucid-punctate, midrib planar
above, prominent below, the secondary veins 8-12 pairs, prominent below,
the margin inroUed except revolute basally, undulate to a vascularized blunt
tooth at nerve end, translucent throughout its length but not scarious; petioles
canalicualte, 0.6-1.0 cm long, 0.5-1.0 mm diam., glabrous, swollen ba-
sally at point of attachment. Staminate Inflorescence: a pseudoraceme, 10—12
cm long; peduncle ca. 1 cm long, the rachis glandular-papillate, the flowers
glomerulate; inflorescence branch bracts membranous, subulate, 5.5—7.0
mm long, 1.2-2.0 mm wide, minutely glandular-papillose publerulent,
glabrescent, prominently black lineate-punctate, the margin entire; floral
bracts similar to branch bracts except 0.9—2.0 mm long, 0.2—0.5 mm wide;
pedicels obsolete to cylindrical to 1.3 mm long, prominently black punc-
tate. Staminate flowers 5— 6-merous, lilac; calyx membranaceous, subcotyliform,
1.5-2.3 mm long, unequally divided, the tube to 0.2 mm long, the lobes
linear-lanceolate, subulate, 1.3-2.0 mm long, 0.4-0.7 mm wide, apically
long-attenuate, flat, epunctate, glabrous, the margin irregular, entire, gla-
brous; corolla rotate, chartaceous, 2.5-4.0 mm long, the tube opaque, 0.8-
1.5 mm long, the lobes hyaline, ovate, 2.1-3.0 mm long, 1.4-1.9 mm wide,
reflexed 180°, apically acute to rounded, moderately glandular-granulose
throughout within except densely so along margin, glabrous without, epunctate,
the margin irregular, entire; stamens 3.0-3.5 mm long, the tube 0.8-1.5
mm long,epunctate, elobate, glabrous, the apically free portions of the fila-
70 SiDA 18(1)
menrs chick (base of filament as wide as anther), terete, glabrous, 1.5-2.0
mm long, ventrally reHexed apically, the anthers subglobose, wider than
long, 0.3-0.5 mm long and wide, apically rounded to obtuse, basally barely
cordulate, the connective prominately black punctate dorsally; pistillode
ovoid, ca. 0.9-1. .3 mm long, 0.6-0.8 mm diam. P ntillate infloresctna resem-
bling staminate, but 4.5-6.5 cm long, the glomerules mostly reduced or a
single flower; branch bracts and floral bracts identical, membranous, sublate,
2.8-4.0 mm long, 0.7-1.0 mm wide, sparsely glandular-papillate, promi-
nently black punctate, the margin erose; pedicels (1.5-)1.8-3.2 mm long.
Pistillate flowers as in staminate but white; calyx chartaceous, cocyliform,
2.2-2.7 mm long, hyaline, the tube 0.3-0.4 mm long, the lobes 1.9-2.3
mm long, 0.7-1.1 mm wide, apically attenuate, at times rufous-papillate
apically; corolla rotate, 2.5-3.2 mm long, the tube 0.9-1 .0 mm long, the
lobes widely ovate, 1.6-2.2 mm long, 1.2-1 .5 mm wide, apically rounded
to acutish, glandular-granulose within and along margin, the margin regular;
staminodes similar to stamens but 1.6-1.9 mm long, the tube 0.9—1.0 mm
long, the apically free portions of the filaments terete, 0.7-0.9 mm long,
the antherodes globose, often malformed, 0.2-0.3 mm long, 0.3-0.4 mm
wide, apically obtuse, basally obtuse to rounded or cordulate, the connec-
tive prominently black punctate ventrally; pistil obpyriform, 2. 1-2.5 mm
long, 1.2-1 .6 mm diam, the ovary 1.6-1.8 mm long, translucent-lepidote,
glandular-papillate, prominently black punctate, style 0.5-0.7 nim long,
the stigma capitate, 4-5-lobed, each lobe lacinate, to 0.6 mm long, the placenta
deeply cupuliform, 2 ovulate. Pr/zit globose, 7-7.5 mm long and diam.,
prominently black punctate, exocarp thin.
Distribution. — Previously considered endemic to the Western Cordillera
of the Andes, on the cliffs (Farallones) of Call, in the Department of Valle,
Colombia, at 1,900-2,600 m, but reported for the first time here from Napo,
Ecuador, at 200 m elevation.
Ecology and consen'ation status. — Cybianthus verticilloides is a rare species in
lowland and premontane forest, growing near watercourses. Given that it
has well-known uses, C. verticilloides might be suitable for cultivation. However,
at present nothing is known regarding its natural population dynamics. Given
the rapid urban development around Call, and its apparent rarity in Ecua-
dor, this species should be considered endangered.
Etymology. — The specific epithet refers to the strikingly pseudoverticillate
phyllotaxis.
Local names and uses. — Colombia: "verticilado" (Spanish); Ecuador:
"Carnerupachapanga," "Yanacarneru" (Quichua), "Carnero Negro" (Spanish).
Used to get rid of small fish in the urinary tract chat cause bleeding and pain.
Specimens examineti. COLOMBIA, Valle del Cauca: Ri'o C^ali rivcrbetl, above Ri'o Pichinde,
El Rnbal, IM^") m, 25 Jul 1 946 (fr),_/, Ciuitmciscis 2 1 12 1 (COL, F); Penas Blancas, 2,200
PiPOi.Y, Cybianthus in Ecuador and Peru 71
m, 9 May 1940 (stam. fl), A. Figueroa 873 (COL, US); Rfo Tulua, 1,200 in, without date
(pist. fl),./. Duque-Jaramilk 4022 (COL); El Silcncio, Yanaconas, 1,900-2,200 m, 28 Feb
1939 (pist. fl), E. Killip & H. Garcia 33802 (COL, US); Farallones de Cali, 1,800 m, Oct
1883 (pist. fl), F. Lehmann 3027 (K, US); km 18-20, Cali-Buenaventura Hwy, entering
near Finca Zingara, summit of the Cordillera Occidental, 1,500-2,000 m, 28 Feb 1988
(fr), H. van derWerff& I. Cabrera 15786 (COL, MO, VALLE). ECUADOR. Napo: Rfo
Napo, S bank a few km below Itaya, 00° 28' S, 76° 33' W, 200 m, 20 Aug 1982 (ster.) H.
Balslev & Santas Dea 2850 (QCA).
Cybianthus vertkilloides appears to be most closely related to C. goudotianus,
by the synapomorphic vascularized leaf teeth. However, C. verticilloides is
defined by the autapomorphic horizontally checked bark, the orange hydropotes
of the branchlets, and the linear-lanceolate leaf shape.
16. Cybianthus croatii Pipoly, sp. nov. (Fig. 4A, 16). Type: ECUADOR. Pastaza:
Along Rd. between Diez dc Agosto and Arajuno, 18 km NE of main Puyo-Macas
Rd., 8.2 km NE of Diez de Agosto, 01° 27' S, 77° 51' W, 970 m, 4 May 1984 (stam.
fl), r. Croat 59009 (hoi.otyfp: MO).
Quoad habitum deminutum petioles brevistipitatos et laminas chartaceas, C. hiimilem
valde cognatum, sed ab ea basibus laminaribus obtusis vel rotundatis (non acutis) inflorescentiis
paniculatis cum ramulis floriferis corymbosis (nee tripinnati- paniculatis cum ramulis floriferis
spicatis vel racemosis) petalis ad apicem cucullatis (nee planis) ad marginem glandulari-
papillosis (nee glabris) filamentis glabris (nee glandulari-granulosis) antherarum connectivis
manifeste punctatis (nee epunctatis) prompte cognoscitur.
Suhshrub to ca. 15 cm tall. Stem terete, ca. 4 mm diam., bearing rufous
hydropotes and sparingly glandular-papillate. L&ives alternate; blades chartaceous,
ovate to elliptic, 5.7-14 cm long, 4.0-7.5 cm wide, apically acute, the tip
mucronulate, basally obtuse to rounded slightly decurrent on the petiole,
bearing rufous hydroporen above and below, midrib slightly impressed distally,
slightly raised proximally above, prominent below, the secondary veins 4-
9 pairs, barely visible above, prominent below, without collecting vein, the
margin scarious, subentire or entire; petiole deeply canaliculate, decurrent
on stem, at times appearing to form a small sheath, (1.5-)2.0-4.5 cm long,
0.2-0.3 cm diam., increasing in length acropetally along stem. Cataphylls
membranaceous, alternate, subulate, 6-12 mm long, 0.6-1.2 mm wide,
located just below center of internode, psuedoverticillate, apically acicular,
keeled, conspicuously black punctate-lineate, bearing hydroporen and glandular
papillae. Inflorescence bract subulate, 3.9-4.0 mm long, 2.0-3.0 mm wide.
Staminate inflorescence: supraaxillary, paniculate, 1.2-3.5 cm long, appearing
succulent, the branches racemose, the rachis densely glandular-papillate;
peduncle 0.5-1.0 cm long; inflorescence branch bracts linear-lanceolate, 1.8-
2.2 mm long, 0.4-0.6 mm wide, conspicuously punctate, glandular-papil-
late; floral bracts subtending and equal to the pedicels, 1.0-2.5 mm long.
Staminate jhwers homomerous, 4-merous, green; calyx cotyliform, membranaceous,
1.2-1.5 mm long, more or less equally divided, hyaline, the tube 0.2-0.3
72
SiDA 18(1)
Fig. 16. Cybianthiis crvattt Pipoly. A. Habit, showing stem with small, acicular cataphylls,
atropetally longet petioles, and supraaxillary inflorescences. B. Staminate flower bud, showing
sparsely glandular-papillate, cotyliform calyx. C. Open staminate corolla, showing cucul-
late lobe apices, suborbicular anthers and conic pistillode. D. Open pistillate corolla, showing
oblate antherodes, subglobose pistil. A-C, drawn from holotype. D, drawn from L. Albert
ck Escobar il 44- Figtire drawn by Peggy Duke.
PiPOLY, Cybianchus in Ecuador and Peru 73
mm long, the lobes deltate, 1.0—1.2 mm long and wide, apically acute to
somewhat acuminate, prominately black punctate, sparsely glandular-papillate
without, the margin irregular, entire, glandular-papillate at first, glabres-
cent; corolla subrotate, carnose, 2.4—2.7 mm long, the tube 0.7—0.8 mm
long, translucent, the lobes opaque, oblong, 1 .7—2.1 mm long, 0.9— l.O mm
wide, apically rounded to obtuse, cucullate, sparingly glandular-papillate
without, densely glandular-granulose within, prominently black punctate
without, especially at apex, the margin entire, glandular-granulose; stamens
2.5—2.6 mm long, the tube conspcuous, carnose, 0.7—0.8 mm long, subtruncate,
the apically free portions of the filaments terete, 0.9-1-0 mm long, epunctate,
glabrous, erect except slightly recurved ventrally at point of attachment to
anther, the anthers suborbicular, 0.7—0.8 mm long and wide, apically rounded,
deeply cordate basally, the connective epunctate ventrally, prominently black
punctate dorsally; pistillocle conic, 1.3 mm long, 0.7 mm wide, densely
translucent-lepidote, conspicuously black punctate, hollow. Pistillate inflorescence:
as staminate, but 2.5—3.5 cm long; pedicels 0.6—1.1 mm long. Pistillate
flowers as in staminate but calyx 1 .3-1-5 mm long, equally divided, the tube
0.2—0.3 mm long, the lobes triangular, 0.9— 1-2 mm long, 0.5—0.7 mm wide,
apically acuminate, sparsely glandular-papillate along margin without; corolla
subcampanulate, the lobes 1.3-1-5 mm long, the tube to 0.2 mm long, the
lobes suborbicular, 1.1—1.3 mm long, 1.0—1.1 mm wide, apically rounded
to obtuse, minutely glandular-granulose along margin without and throughout;
staminodes as in stamens but 0.9—1-0 mm long, the tube, ca. 0.2 mm long,
the apically free portions of the filaments 0.2—0.3 mm long, the antherodes
oblate, 0.4—0.5 mm long, 0.5—0.6 mm wide, apically truncate, basally obtuse,
the connective prominently black punctate ventrally; pistil subglobosel, 0.6—
0.7 mm long and diam., densely translucent-lepidote, conspicuously black
punctate, the placenta deeply cupuliform, bearing 2 ovules, the style barely
discernible, to 0.1 mm long, stignma subcapitate, the margin lacinate, early
caducous. Fruit unknown.
Distribution. — Known only from the type (Ecuador) and from Antioquia,
Colombia, at 970—2,440 m elevation..
Ecology and conservation status. — Cybiantbus croatii occurs in premontane
and montane wet forest, where it is locally common in protected areas near
forest margins. Because of its restricted habitat, it is presumed to be threatened.
Etymology. — It is with great pleasure that 1 dedicate this species to Tho-
mas B. Croat, pre-eminent authority on the systematics of Neottopical Araceae,
indefatigable collector, gentleman, scholar, and the P. A. Schuize Curator
of Botany at the Missouri Botanical Garden. During the tropical botany
course I took from him in Costa Rica in 1977, 1 was first shown and be-
came intrigued with the systematics and populaton biology of the Myrsinaceae.
74 SiDA 1<S(1)
ParA'i YFi:: COLOMBIA. Antioquia: Mpio. Caklas, above town, Finca La 7:drz.d, 2, 140
m, 2 Feb 1984 (pist. H), L. Albert cle EsaAir et al. 3744 (HLJA, US).
Cyhiaiitb//s crocit'a is the smallest myrsinaceous shrub chat exhibits Corner's
Model (Halle et al. 1978). It is interesting to note that with each successive
pseudowhorl of cataphylls, an inflorescence is produced, followed by a larger
leaf with a longer petiole. This species appears to be most closely related to
another diminutive plant, C. humilh, known only from the Department of
Antiocjuia, Colombia and adjacent Choco, and from Ecuador. The synapomorphy
which defines the Cybtanthus humllis-C. croatii clade is the scarious leaf margin,
known elsewhere only in subgenus Grammadenia (Pipoly 1 987, unpubl. data).
However, becatise that subgenus occurs on the other side of the generic cla-
dogram presented by Pipoly (1987), it is most parsimonius to hypothesize
that the margin has arisen independently in these distant lineages. The pistillode
is present in both of these species, and I have chosen to postulate that this
represents a reversal because it is the most parsimonius conclusion. Cybianthns
croatii is defined by the autapomorphic rufous stem hydropotes and the ac-
ropetally longer petioles. Despite the fact that the distribution of this spe-
cies entirely overlaps that of C, hnniilis, it appears that C. croatii is restricted
to montane and cloud forests, where C. h/nnilis is restricted to premontane
pluvial forests and subparamo thickets. This is yet another example of sis-
ter species in alritudinally adjacent liabitats, already reported in Cybia)!tb//s
subgenus Laxiflor//s (Pipoly 1983) C. subgenus /\i/rmY;;/'r;wr;r/7/:^^ (Pipoly 1983b)
and C. subgenus Conormrpha (Pipoly 1992a). Whether this supports the conce[it
of speciation by peripheral isolation (parapatric) is unknown. Futher stud-
ies of the respective population biologies of the subgenus are needed.
17. Cybianthus humilis (Mez ) G. Agostini (Fig. 3F), Acta. Biol. Venez.
10:163. 1980. \Veigelthih/(mil,sUi:z in EngL, Pflanzenr. IV. 236(Hetr 9):291 . 1902.
Typh: ECUADOR. Without locaHty, 1896 (stam. fl), A. Sodiro 100/14 (holotyph:
B-destr., F Neg. 4856; i.nc'icnYPE, here designated: COLOMBIA. Antioquia: 2,650-
2,800 m, 1 Apr 1880 (stam, H), W: Kdlhreyer n34 (K). Altliotigh Mez (1902) did
not specifically mention the Sodiro collection as the t\pe, he cited the Sodiro and
the Kalbreyer specimens in the protologtie. However, the F photograph clearly shows
that a drawing of a dissection accompanies the Sodiro specimen, while that of Kalbreyer
does not, stiggesting that the Sodiro specimen (ormed the |irincipal element upon
which the description was based. LJnfortimately, no duplicates of that Sodiro collecton
have been located. Mez also annotated the Kalbreyer sheet at K, and thus I designate
it as the lectotyjie.
CoiitoD/yr.iiiii' soclirihnni Mez, Btill. Fierb. f^oissier, 2 ser 5:535. 1905. syn. nov. Cyb'uinthin
mimkiniis (Mez) C. Agostini, Acta Biol. Venez. 1 0: 1 63. 1 980. Tyi'p: ECUADOR. Atacat/o.
Jan 1902 (stam. fi), A. Sodivn 100/2 (iioi.otypf.: B-destr.; i.H;r()rYPP., here designated: P).
Sabshri/b to 25 cm tall. StcDi terete, 3.5-4.0 mm diam., densely glandu-
lar-papillate, early glabrescent. Cataphylls in alternating nodes with leaves,
membranaceous, subulate, 1 0-13 mm long, 1 .7-2.6 mm wide, apically long-
PiPOLY, CybianchLis in Ecuador and Peru 75
attenuate, keeled, midrib prominent below, prominently black punctate and
lineate-ptmctate, glandular-papillate, glabrescent, the margin opaque, flat,
entire. Leaves subopposite; blades chartaceous, elliptic to narrowly elliptic,
1 5— 20(— 27) mm long, (4.0— )7. 0—8. 5 cm wide, apically acute to subacuminate,
terminating in an inconspicuous mucro, basally acute, bearing hydropoten
above and below, inconspicuously pellucid punctate, midrib somewhat impressed
above, prominent below, the secondary veins 6—1 1 pairs, slightly impressed
abive or not, prominently raised below, not united by a submarginal col-
lecting vein, the margin entire, opaque, subrevolute; petioles canaliculate,
thin, 1.0—2.0 cm long, 2.0—2.5 mm diam., glabrous, not decurrent on the
stem. Infiorescence bracts similar to cataphylls, but 1.2—1.3 mm long, 1.5—
2.2 mm wide. Staminate inflorescence: a bipinnate panicle, 3-5— 15(— 19) cm
long, the rachis densely glandular-papillate, the branches spicate or rarely
racemose, appearing subglomerulate apically; inflorescence branch bracts
membranaceous, subulate, 3—4.5 mm long, 0.3—0.5 mm wide, densely and
prominently black pimctate-lineate, sparingly papillate; pedicels essentially
obsolete, to 0.3(— 2) mm long; floral bract membranaceous, ovate, asym-
metric, 2.3—2.8 mm long, 0.9-1 -2 mm wide, apically abruptly acuminate,
medially keeled, epunctate, the margin erose, stipitate glandular-papillate.
Staminate flowers 4— 5-merous; calyx membranaceous, cotyliform, 1.5—2.0 mm
long, tube 0.2—0.3 mm long, the lobes ovate to lanceolate, 1.3—1.7 mm
long, 0.5—1.3 mm wide, unequally divided, apically acuminate to acute and
often erose or premorse, glandular papillose-puberulent without, the mar-
gin erose, glabrous; corolla chartaceous, appearing subrotate, 3-7-4.5 mm
long, the tube 1.1—1.5 mm long, the lobes linear-lanceolate to oblong, 2.4—
3.2 mm long, 1.0—1.5 mm wide, often unequal, apically long-attenuate to
obtuse, moderately rufous gladular-granulose throughout within, sparsely
glandular-papillate without, hyaline, the margin entire, glabrous; stamens
3-0-3.9 nnm long, the tube 1.1—1 .5 mm long, coriaceous, conspicuous, sparsely
glandular-grandulose, elobate, the apically free portions of the filaments basally
as wide or wider than anther, then tapering apically, 1 .9— 2.4 mm long, terete,
glandular-granulose, the anthers oblate, 0.4—0.5 mm long, 0.6—0.7 mm wide,
apically truncate, basally subcordate, the connective epunctate; pistillode
glabrous lagenform, 1.4—1.6 mm long, 0.8-1.0 mm wide, the stigma punctiform.
Pistillate inflorescence: as in the staminate but 7—10 cm long, 4—6 cm wide;
inflorescence branch bracts 2.5-3.5 mm long, 0.2—0.3 mm wide; peduncle
1—2.5 cm long; floral bracts 1 — 1.5 mm long; pedicels obsolete. Pistillate
flowers unknown; fruiting calyx as in staminate but 1.3-1.5 mm long, the
tube ca. 0.2—0.3 mm long, the lobes 1 .1 — 1 .2 mm long, 0.2—0.3 rnm wide.
Fruit globose, reddish-purple when fresh, 4—6 mm long and diam. when
dried, the exocarp thin, conspicuously pellucid punctate.
76 SiDA 18(1)
Distribution. — Known only from Colombia and Ecuador, at 1,450—2,700
m elevation.
Ecology and conservation status. — Cybianthus hinnilis grows in deep shade
among rocks at the margins of premontane pluvial forests. Populations I
observed in Colombia grew only in undisturbed areas with deep shade, leaf
litter and organic humus. Because of the apparently restricted habitat, this
species should be considered threatened.
Etymology. — The specific epithet refers to the low habit of the plant.
Local names and uses. — Colombia (Choco): "Hierba del palo grande." Ground
to make crude syrups; syrup applied externally to cure cuts, taken inter-
nally to cure internal infections, clean the stomach and against chills. Given
that it is a shade species, the common name probably refers to its frequency
in sliaded areas near large trees.
Si^cci mens examined. COLOMBIA. Antioquia: Mjiio. Urrao, i-'arque Nacional Uis Orqui'deas,
VeredaCalles, permanent inventory tor premontane pluvial iorest, right bankof Ri'o Calles,
06° 32' N, 76" 19' W, 1,450 m, 27 Nov 1993 (fr),./. P,poly et at. 17186 (COL, JAUM,
MO), 1,450-1,500 m, 29 Nov 1993 (lr),_/. Pi/w/) via/. 17361 (COL, JAUM, MO); with-
out locality and date, (.stam. fl), F. Lehiiiatni s.u. (L, K). Choco: Mpio. de Quibdo, C^orregimiento
San Francisco Icho, Quebrada Caledonia along Caledonia Rd., 9 Apr I 9H7 (ster.), F. Garcni
& J. Ecl:uirary/d 259-A. (COL, CHOCO, MO). Nariiio: Mpio. Barbacoas, Corrcgimiento
Orri'z y Zamora, Vereda El Barro, Reserva Natural Rfo Nambf, ca. 5 km W de Altaquer,
taldas occidentales de la Cordillera Occidental, 01" 18' N, 78° 08' W, 1,350-1,100 m, 3
Sep 1997 (ster.),/ P//W). A. Cngollo. et ul. 21240 (BRIT, rMB,JAUM, PSO). Quindio:
Mpio. De Salento, Estacion Navarco, Alto San Ignacio, 2,850 m, 23 Nov 1990 (stam. fl),
P. Franco et ul. 3204 (COL, MO). Risaralda: Mpio. Sta. Rosa, Camino de LIcrradura etre
Termales y Paramo Sta. Rosa, Cordillera Central, vertiente Occidental, Hacienda El Margariral,
2,500 m, 18 Atig 1980 (fr),/ hlrolmetal. 967/ (COL, MO). ECUADOR. Napo: Canton
El Chaco, Proyecto Hidroelectrico Coca, Punto ST3, right bank of Ri'o Quijos, ca. 10 km
S of Reventador, 1,500 m, 3-5 Oct 1990 (fr), W. Pjlacim 5930 (MO, QCNE). Pichincha:
Cant()n Quito; Parroquia Calacah', Reserva Geof-)otanica Pukiiahua, 00° 01' N, 78° 35' W,
1,800-2,000 m, 29 Jul I 989 (stam. fl), C. Cerm 7 1 H4 (MO, QCNA); Mindo, 26 Jun 1876
(stam. fl), £. Andre 3819 (K).
Cybianthus bumilis is most closely related to C. croatii, by virtue of its
synapomorphic scarious leaf margin. The autapomorphies that distinguish
C. humilis from all other species of the subgenus include the premorse apices
of the calyx lobes, the unecjual corolla division, and the oblate anther shape.
18. Cybianthus sprucei (Hook, f.) Cj. Agostini, Acta Biol. Venez. 10(2): 164.
1980. Couioniyrsine iprncei Hook, I. ni Benth. et Hook., Cien. PI . 2:614. 1876. Weigeltia
spriicei (Hook, f.) Mez in Engl., Pflanzenr. IV. 236(Heft 9):291 . I 902. Type: ECUA-
DOR. [CiiiMBORA/.o: W slopes of Volcan Chimborazo, 17 Jun 1860] (stam. fl), 11.
Spruce 6)144 (houxitpe, K-2 sheets). Note: label on specimen does not indicate place
or date. Information was derived by comments regarding habit, etc. on the label which
matched data gi\'en in Sprtice ( I 880).
Weigeltia pitncimensis Standi., Publ. Field Mus. Nat. Hist, Bot. Set. 22:l6'l. 1940. syn.
nov. Cyhicnitljus paiuiinensis (Standi.) CL Agostini, Acta Biol. Venez. 1 0: 163- 1980. Type:
PiFOLY, Cybianthus in Ecuador and Peru 77
PANAMA. Darien: Cana, Cuasi Trail, Dtto. Cheijana, 1000 m, 10 Mar 1940 (stam.
fl), M E. Terry & R. A. Terry 1490 (holotype: F-2 sheets; isotypes: A, MO).
Weigeltia purpurea Cuatrec, Revista Acad. Colomb. Ci. Exact. 8(31):326. 1951. Type:
COLOMBIA. Valle del cauca: Bahia de Buenaventura, Quebrada de San Joaqui'n,
0-10 m, 21 Feb 1946 (stam. fl),J. Cuatrecasas 19892 (holotype: F; isotype: COL).
Shrub to tree to 5 m, flowering from less than 1 m. Stem terete, 1.3-2.5
cm diam. below uppermost leaves, swollen at nodes, semi-woody, glandu-
lar-papillate-pLiberulent, glabrescent. Cataphylls few, alternate, coriaceous,
subulate, (2.5-)3.0-5.0 cm long, 2-4 mm wide, strongly keeled, densely
glandular-papillose-puberulent, glabrescent, consiptiously black punctate,
the margin opaque, regular, entire. Letft'(?j-pseLidoverticillate; blades chartaceous,
widely oblong to elliptic, rarely widely obovate, (26— )3 1—7 5 cm long, (6.5—)
10—30 cm wide, apically rounded, obttise or rarely acutish, not mucronulare,
basally abruptly subcuneate, asymetric, slightly decurrent on petiole, bear-
ing a few hydropoten above, moderately rufous glandular-papillate and with
a moderate number of hydropoten below, prominently red or black puctate,
the margin regular, opaque, entire; petioles stiff, (2.5-)5.0-l4(-23) cm long,
0.3-0.9 cm diam., slightly to moderately canaliculate, swollen basally, sparingly
glandular-papillate, glabrescent. Staminate Inflorescence: a pyramidal bipin-
natethyrsoid panicle, 9-l6.5(-19)cm long, 9-17(-22)cmwide, the branches
subspicate, the rachis densely glandular-papillose-puberulent; peduncle 0.5-
2.0 cm long; inflorescence branch bracts chartaceotis, linear-lanceolate, 8.5—
12.9 mm long, 0.6—0.9 mm wide, apically narrowly acute, conspicuously
black punctate, densely glandular-papillate, gabrescent, flat, the margin opaque,
regular, entire; floral bracts membraneous, linear, 1.5— 2.0(-2.5) mm long,
0.5—0.6 mm wide, subulate, hyaline, densely glandular-papillose puberu-
lent, the margin entire, glandular-papillate; pedicels obsolete to 0.2 mm
long. Staminate flouers homomerous, 4-merous, coriaceous, subscssile, race-
mose, 4— 5-merous, white in btid, green in anthesis, then crimson; calyx
coriaceous, subcotyliform, unequally divided, 0.9—1-1 mm long, the tube
0.1—0.2 mm long, the lobes linear-lanceolate, 0.7—0.9 mm long, 0.2-0.5
mm wide, apically subulate, keeled, brown punctate-lineate or punctate medially,
sparsely glandular-papillate, the margin irregular, subentire to erose, densely
glandular-ciliate; corolla subrotate, 2.0-2.9 mm long, the tube 0.2— 0.3mm
long, the lobes narrowly ovate to lanceolate, (1.7— )1.9— 2.3(-2.6) mm long,
0.9-l-0(-1.5) mm wide, reflexed 135° from tube at anthesis, apically at-
tenuate, densely glandular-granulose throughout within and along margin
within and without, inconspicuously pellucid punctate, the margin entire,
somewhat irregular, densely glandular-granulose; stamens exserted to slightly
shorter than the corolla lobe, 1.6—2.2 mm long, the tube ca. 0.5 mm long,
conspicuous, coriaceous, taller than the corolla tube, elobate, opaque, epunctate,
glabrous, the apically free portions of the filaments terete, thicker than the
78 SiDA 18(1)
anthers, 0.8-1.6 mm long, ventrally recurved at anthesis, epunctate, gla-
brous, the anthers subglobose, ca. 0.3 mm long, 0.4-0.5 mm wide, apically
rounded, basally cordulate, dorsifixed near base so as to appear basihxed,
the connective prominently red or black punctate dorsally; pistiUode nor-
mally absent, occasionally conic, to 1 mm long, 0.3 mm wide, densely translucent-
lepidote. Pistillate hiflorescence resembling staminate in all features, but smaller,
2. 5-3. 5(-l 3) cm long, 3.5-8. 0(-l 2.5) cm wide; peduncle 0.5-1 .5 cm long;
inflorescence branch bracts 3.0-3.5(-5.0) mm long, to 0.6 mm wide, at
times somewhat cucuUate; floral bracts 2.0-2.6 mm long, ca. 0.5 mm wide,
pedicels virtually obsolete, or cylindrical to 0.1 mm long in flower, accrescent
and incrassate to 2.0 mm long, 1.0-1 .5 mm diam. in fruit. Pistillate flowers
as in staminate, forming a condensed spike on the inflorescence branches;
calyx subcupuliform, 1.2—1.4 mm long; the tube to 0.1-0.2 mm long, the
lobes 0.9-1.1 mm long, 0.9-1-0 mm wide, widely ovate, apically acute;
corolla rotate, 2.7-2.9 mm long, the tube 0.6—0.7 mm long, the lobes ob-
long to elliptic, 2.0—2.2 mm long, 0.9—1 .0 mm wide, apically acute; staminodes
1.5—1 .6 mm long, the tube 1 .0—1. 1 mm long, the apically free portions of
the filaments 0.4-0.5 mm long, the antherodes malformed, 0.2-0.3 mm
long, 0.3—0.4 mm wide; pistil obturbinate, 1.5-1 .6 mm long, densely trans-
lucent-lepidote and prominently pellucid punctate, the style obsolete, the
stigma capitate, 0.2—0.3 mm long, subsessile, 4— many-lobed, the lobes 1.3—
2.3 mm long, viscid, bright crimson, the placenta ovoid, the ovules 2, born
on side of placenta. Fr//it subglobose, 5.0—9-0 mm long, 9-14 mm diam.,
fleshy, the exocarj:> thick, orange at maturity, prominently black punctate.
Bisexual Inflorenscence resembling staminate in all features except: 3-7-14.5
cm long, 2.5-12.5 cm wide; inflorescence branch bracts and floral bracts
and pedicels as in pistillate. Bisexual flotuers spicate, less crowded than in
the pistillate, more crowded than staminate; calyx cotyliform, (4-)5 -parted,
1.0—1.2 mm long, the tube ca. 0.2 mm long, the lobes narrowly ovate to
narrowly triangular, 0.8-1.0 mm long, 0.3—0.4 mm wide, apically acute
to narrowly acute; corolla 4-lobed, rotate, 2.0—2.7 mm long, the tube ca.
0.3 mm long, the lobes oblong, 1 .7—2.4 mm long, 0.9—1 -2 mm wide, apically
acute; stamens 1.2-1.5 mm long, the tube ca. 0.5 mm long, the apically
free portions of tlie filaments 0.6—0.7 mm long, the anthers oblate, 0.3—
0.4 mm long, ca. 0.5 mm wide; pistil as in pistillate flowers except 1.2—1.4
mm long, the ovary 1.0—1.1 mm long, 0.7—0.9 nim diam., the style short,
0.1—0.2 mm long, the stigma capitate, to 0.2 mm long, the lobes 0.9-1-1
mm long, the placenta ellipsoid, the ovules 2—3, borne on side of placenta.
Bisexual fruit s\xh^\ohose, as in pistillate, but 4-5 mm long, 5-7 mm diam.
Distribution. — From Darien, Panama to Loreto, Peru, from 0—1,700 m
elevation.
Ecology and conservatifm status. — Cybianthus sprucei is a ridgetop species.
PiPOi.Y, Cybianrhus in Ecuador and Peru 79
occurring in premontane wet and rainforests and also in lowland forests on
forest margins of exposed hilltops. This species, as opposed to its closest
congener, Cybianthus simplex, occurs in areas of high incident light for at
least part of the day. Because Cybianthus sprucei continues to reproduce in
spite of mild forest intervention, it is not considered threatened or endan-
gered.
Etymology. — This species was named in honor of Richard Spruce, ardent
collector and student of the Andean and Amazonian flora.
Local names and uses. — Colombia: "margoandre," "Tunda" Spanish (Valle
del Cauca, Colombia); Ecuador: "urcu tahucu" (Quichua); ground and a vapor
bath is taken to "send evil away" and to treat body pains, headaches, inter-
nal colds; Peru: "kurup" (Jfvaro); the root is mashed and boiled, and the
decoction is drunk to "strengthen" the body.
Specimens examined. PANAMA. Darien: Orro Pirre, 10-20 Jul 1977 (bisex. fl, it), J.
PoLuwi 4544 (MO, PMA); on ridge of Cerro Pirre, 08° 00' N, 77° 45' W, 1,000-1,080 m,
14 Sep 1989 (fr), G. McPherson 14066 (BRIT, MO). COLOMBIA. Cauca: Rfo Micay, en
Guayabal, 5-20 m, 25 Feb 1943 (pisr. fr),/. Cua/recasas 1415B (COL, F, US); Distrito Cauca,
El Tambo, 900 m, Apr 19.37 (stam. fl), K. imi Sneidern 1615 (S). Choco: Along Rd. be-
tween Quibdoand Medellin, Km 207.5, 0-200 m, 18 Dec 1980 (pisr. fl, fr), T. Croat & A.
Cogollo 522^7 (COL, JAUM, MO); Alto del Buey, 1,200-1,800 m, 8 Jan 1973 (pist. fl, fr),
A. Gentry & E. Forero 73/7 (COL, F, MO). Narifio: La Guayacana, Funes, 24 Jun 1951
(stam. fl), R. Castatleda 2873 (COL, F); Ricaurte, 1 ,300 m; 18 Apr 1941 (bisex. fl), K. von
Sneidern A61 2 his (S); Reserva Natural La Planada, 7 km above Chucunes on Rd. between
Tuquerres and Ricaurte, along Sendero La Vieja, 01° 06' N, 77° 54' W, 1,780-1850 m, 7
Mar 1990 (fr), T. Croat 71155 (MO, PSO). Valle del Cauca: Pacific coast, Rfo Naya, Puerto
Merizalde, 5-20 m, 22 Feb 1943 (bisex. fl),/. C/u/trecasas 14053 (COL, F); Rfo Yurumangui,
550 m, 28 Jan-10 Feb 1944 (pist. fl, fr),J. C/tatrecasas 15743 (COL, F, US); Rio Calima,
Choco region. La Trojita, 5-50 m, 20 Feb 1 944 (stam., pist. fl-sheets mixed),J. Cuatrecasas
16272 (COL, F, US); Rfo Cajambre, Barco, 5-80 m, 30 Apr 1944 (fr),/ Cuatrecasas 1 7625
(COL, F, US); Bajo Calima, 15 km N of Buenaventura, Carton de Colombia concession,
Juanchacho region, 03° 56' N, 77° 08' W, 500 m, 27 Mar 1986 (stam. fl), A. Gentry et al.
53713 (COL, MO), Concesion Pulpapel/Buenaventura, 03° 55' N, 77° 00' W, 100 m, 7
Mar 1985 (fr), M. Monsa/ve 767 (COL, CUVC, MO), 1 9 Mar 1 985 (stam. fl), M. Monsalve
790 (CUVC, MO), 21 Mar 1985 (sram. fl), M. Monsaive 797 (CUVC, MO), 12 Mar 1986
(stam. bud), M. Monsalve 981 (CUVC, MO), 24 Aug IS'86 (fr), M. Monsaive 1 124 (CUVC,
MO); Bahia de Malaga, near moutii of Quebrada La Sierpe,0 4° 00' N, 77° 1 5' W, 0-20 m,
17 Feb 1983 (stam. fl), A. Gentry et aJ. 40453 (COL, MO); 1 8 km E of Buenaventura, 50
m, 14 Feb 1939 (fr), E. Killip & H. Garcia 33219 (BM, COL, NY, US), Buenaventura, Jun
1901 (pist. fl, fr) F. Lelomann B. T 651 (K, NY). Putumayo: Umbna, 00° 54' N, 76° 10'
W, 325 m,Jan-Feb 1931 (stam. fl), G. Klug 2108 (US). ECUADOR. Bolivar : Along first
15 km of ChiUanes-El Tambo, 2,400 m, 18Jul 1991 (stam. fl), H. van clerWerffet al. 12430
(BRIT, MO, QCNE); along Rd. ChiUanes-San Pablo, 6 km outside Chillanes, 2,600 m, 21
Jul 1991 (fr), H. vander Werffetal.l 12561 (MO, QCNE), Morona-Santiago: 15 km N of
Macas, Rd to Rio Upano, 02° 07' S, 78 °08' W, 1250 m, 20 Feb 1987 (bisex. fl),^. Bohlin
et al. 1493 (GB); Cordillera de Cutucu, W slopes along trail from Logrono to Yaupi, 02°
46' S, 78° 06' W, 1,200 m, 10 Nov 1976 (stam. fl), Al. Maddtson et aL 3204 (US). Napo:
Canton Archidona, Carretera Hollfn-Loreto, Rfo Huataraco, 2 hrs walk from Guagua Sumaco,
80 SiDA 18(1)
00° 43' S, 77° 32' W, 800-1,000 m, 23-30 Aug 1989 (stain, fl), C. Ceron & M. Factos
7648 (MO, QCNE); Canton Orellana, Resetva Flori'stica El Chuncho, 5 km N of Coca, 00°
25' S, 77° or W, 250 m, 23 May 1993 (fr), W. Palados 10680 (MO, QCNE), Ei Chuncho,
el Payamino, Estacion Experimental INIAP-Napo, 5 km NW of Coca, 00° 30' S, 77° 01'
W, 250 m, 12 Oct 1987 (stam. fl), C, Cerd/i 2494 (MO, QCNE); Canton Tena, Mishualli,
junction of Rios Mishualli and Napo,01° 03' S, 77° H ' W, 500 m, 13-14 Aug 1979 (fr),
L. Hohii-Niehoti 19295 (AAU); Estacion Biologica Jatun Sacha, along S bank of Ri'o Napo,
1 km E of Puerto Misahualli, 00° 04' S, 77° 36' W, 450 m, 1 Apr 1992 (fr), T. Croat 73552
(MO, QCNE), 8 km E of Puerto Mishualli, 01° 04' S, 77° 36' W, 400 m, 14 Aug 1989
(ster.), C. Ceron 7409 (MO, QCNE), 22 Sep 1989 (stam. fl), W. Palacios 4471 (MO, QCNE),
16 Mar 1991 (fr), D. Neill98J3 (BRIT, F, MO, QCNE); Ri'o Blanco Comunidad, headwa-
ters of Rio Huambuno, 6 km NNW of Ahuano, 01° 44' S, 77° 44' W, 440 m, 18 Jul-9
Aug 1990 (fr), E. Kohn 1311 (MO). PERU. Amazonas: Prov. Bagua, Yamayakat, trail to
Putuim, 04° 55' S, 78° 19' W, 500 m, 17 Oct 1996 (stam. fl), R. Va'squez & N. Jarcmillo
20318 (AMAZ, BRIT, MO). Loreto: Prov. Loreto, Pampa Hermosa and vicinity, Rio Corrientes,
1 kmSofjunctionwithRjoMacusari,03° 15' S, 75" 50' W, l60m, 3-20 Dec 1985 (stam.
fl), W. Lewis etal. 10180 (BRIT, MO, USM).
Cybianthus spr/icei was misinterpreted by Mez, and confused with C. sim-
plex (Mez 1902). From there, Weigeltia panamensh was described based pri-
marily on subtle differences and geography. Finally, Cuatrecasas described
Weigeltia purpurea from the Choco floristic region of Colombia, notable only
for its narrower leaves, the secondary veins more arcuate, and some quanti-
tative floral characteristics.
Cybianthus spnicei is most closely related to C. simplex because of the homomerous
flowers, long petioles and non-mucronulate leaf apices. Flowever, Cybianthus
sprucei may easily be separated from C simplex because of its arborescent
habit, pyramidal panicle with spicate branches, symmetric leaf base and longer
cataphylls.
19. Cybianthus simplex (Hook, f ) G. Agostini (Fig. 4C), Acta Biol. Venez.
10: 163. 19H(). Comomyrsine simplex Hook, f /« Benth. & Hook., Gen, PI. 2:644.
1876. Weigeltni simplex (Hook, f ) Mez in Engl., Pflanzenr IV. 236(Hefr 9): 290. 1 902.
Tvpt: ECUADOR. Chimborazo: At foot of Volcan Chimborazo, 760 m, Aug I860
(pist, fl, fr), /?, Spri/ee 61 43 (holotyph: K-2 sheets).
Weigeltia chamaephytci Diels, Nonzbl. Bot. Gart. Berlin-Dahlem 15:383. 1941. syn. nov.
Cybianthus chamaephyta (Diels) G. Agostini, Acta Biol, Venez. 10:163. 1980. Type:
ECUADOR. Pasta/.a: Mera, 1,200 m, 15 Nov 1938 (stam. fl), H. Schnltze-Rhonhof
2983 (HOLOTYPH: B-destr. ; no isotype found). I defer neotypification until material
has been regathered from the type locality or an adjacent one.
Senii-woody siibshrub to 1 (—3.5) m tall. Stem terete, 1—5 cm diam., sparsely
glandular-papillate, glabrescent. Cataphylls few, spirally arranged in inter-
nodal areas, coriaceous, subulate, 6.5—26.5 mm long, 1.12—1.7 mm wide,
keeled, densely and prominently black punctate and punctate-Iineate, sparsely
glandular-papillate, the margin flat, entire. Lfir;z'^.s"pseudoverticillate; blacies
membranaceous, elliptic to oblong, (34.5— )38— 46. 5(— 80) cm long, (12. 5-)
PiPOi.Y, Cybianthus in Ecuador and Peru 81
13. 5- 18. 5 (-3 2. 5) cm wide, apicaliy broadly acute or rounded to a short
acumen, rarely acute, basally asymmetric, broadly rounded or rarely abruptly
acute, slightly decurrent on the petiole, hydropotes few above, numerous
below, often sparsely glandular-papillate below, conspicuously black punc-
tate, the margin irregular, hyaline when juvenile, opaque at maturity, flat,
entire; petioles rigid, deeply canaliculate, 7-1 7 (-2 1) cm long, 4-6 mm diam.,
slightly marginate at junctions of the blade, expanded basally and slightly
decurrent on stem, sparsely glandular-papillate, glabrescent. biflorescence bracts
membranaceous, linear-lanceolate, 17-25 mm long, 2-4 mm wide, apicaliy
long-attenuate, hyaline, densely and prominently black punctate-lineate,
the margin irregular, entire; peduncle ( 1 .2-)2.4— 4.5 cm long. Staminate inflorescence:
a thyrsoid panicle (columnar) 1 1-28(-54) cm long, bi- or tripinnate, the
primary branches subopposite, each branch pyramidal-paniculate, the flowers
racemose; inflorescence branch bracts linear-lanceolate, subulate, (50-)6l-
72 mm long, 0.5-1.3 mm wide, apicaliy long-attenuate, medially keeled,
hyaline, glandular-papillose throughout, the margin entire; floral bracts early
caducous; pedicels 2.3-3(-5) mm long, glandular-papillate and -ciliolate.
Staminate flowers 5-merous, green to greenish-white when fresh; calyx mem-
branaceous, subcotyliform, 1.1-1.4 mm long, equally divided, the tube 0.2
mm long, the lobes lanceolate to lanceolate-subulate, apicaliy narowly acute
to long-attenuate, hyaline, epunctate, the margin erose-dentate; corolla coriaceous,
rotate, 2.2-3.1 mm long, the tube 0.3-0.7 mm long, the lobes linear-lan-
ceolate, 1.5-2.6 mm long, 0.9-1.1 mm wide, reflexed distally 180° from
the tube, the apicaliy subulate, densely glandular-granulose throughout within
and along the margin, epunctate, the margin regular, entire; stamens 1.0-
2.3 mm long, the tube conspicuous, coriaceous, 0.3—0.7 mm long, the apicaliy
free portions of the filaments terete, as wide as the anthers, 1.0-1.1 mm
long, ventrally recurved, the anthers connivent at first, subglobose, 0.4-
0.5 mm long, 0.5-0.6 mm wide, apicaliy rounded, basally cordulate, dorsifixed
just above the base, the connective prominently black punctate dorsally;
pistillode absent. Pistillate inflorescence: as in staminate but 16-30 cm long;
inflorescence branch bracts ovate-subulate, 4.5-7 mm long, 1.5-2 mm wide,
the margin irregular, erose; floral bracts membranaceous, linear-subulate,
1.6-2 mm long, 0.4-0.6 mm wide, apicaliy long-attenuate, densely glan-
dular-papillose puberulent, the margin irregular-entire; pedicel terete, 1-
10 mm long, translucent, glandular-puberulent. Pistillate flowers green; ca-
lyx coriaceous, 0.6-1 mm long, unequally divided, the tube 0.2-0.3 mm
long, the lobes ovate to widely ovate, 0.4-0.7 mm long and wide, apicaliy
acute, often moderately glandular-papillate without, the margin irregular,
erose; corolla 1.5-1.7 mm long, the tube 0.4-0.5 mm long, the lobes ob-
long, 1.0-1.2 mm long, 0.7-0.8 mm wide, apicaliy acute, reflexed 135°
82 SiDA 18(1)
from tube, glandular-papillate without; staminodes 0.8-1.2 mm long, the
tube inconspicuous, membranaceous, 0.4-0.5 mm long, glabrous, elobate,
the apically free portions of the filaments 0.3-0.4 mm long, the antherodes
subglobose, 0.2 mm long, 0.3 mm wide, apically rounded, basally cordulate;
pistil subglobose, 0.8-1.1 mm long, the ovary 0.6-0.8 mm long, 1.0-1.2
mm diam., densely translucent-lepidote, the style short, thick, 0. 1 mm long,
0.7 mm diam., the stigma capitate, the margin lacinate, with numerous
lobules to 0.2 mm long, the placenta umbonate, the ovules 3, exposed apically
1/3 their length on the placenta. Fruit subglobose, 4-5 mm long, 5-8 mm
diam., red, exocarp somewhat thick, juicy, prominently black punctate.
Distribution. — Endemic to the slopes of the Western Cordillera of the Andes
in Colombia and Ecuador, 60-2,200 m.
Ecology and conservation status. — Cybianthus simplex occurs in premontane
wet and rainforests, occasionally in the understory of ridgetops forests. This
species occurs in deep shade under the shrub stratum of the forest. It occurs
more frequently in primary forest wet enough to maintain Chusquea popu-
lations, but can survive in disturbed forests as long as the shrub and Chusquea
populations exist. Because of its habitat flexibility, it is not considered threatened
or endangered.
Etymology. — The epithet "simplex" refers to the monoaxial habit of the
species.
Specimens examined, COLOMBIA. Nariiio: Mpio. Barbacoas, C^orregimiento Orn'z y
Zamora, Vereda El Barro, Reserva Natural Rio Nambi', ca. 5 km W de Aitaquer, faldas
occidentales de la Cordillera Occidental, 01° 18' N, 78° 08' W, 1,350-1,400 m, 1 Sep
1997 (scam, fl), /. Pipoly, A. Cogollo. d al. 2!02{^ . 21046, 2105L 2]055_{BR1T, FMB,
JAUM, PSO), 2 Sep 1997 (bisex. (Dj. P,po/y. A. Co^ollo et al. 2/7(;9_(BRIT, JAUM, PSO),
(ster.)J. Pipoly. A. Coi^ollodal. 21 131. 21148 (BRIT, PSO), (pist. fl, fr),/ Pipoly. A. Co^ollo
etal. 21173 (BRIT, FMB, JAUM, PSO), 1, -190-1, 500 m, 4 Sep 1997 (stam. fl),J. PipoIy,
A. Cogolloetal. 21294 (BRIT, FMB, JAUM, PSO), (pist. fl),/. PipoIy. A. Cogolloetal. 21296
(BRIT, FMB, JAUM, PSO), 1,350-1 ,400 m, 5 Sep 1997 (scam, fl), /. Pipo/y. A. Cogo/Io et
al. 21326 (BRIT, JAUM, PSO), (ster.),J, Pz/Wj. A. Cogollo ,t al. 21328 (BRIT, JAUM,
PSO),l, 350-1, 145 m, 6 Sep 1997 (ster.),J. Pipoly. A. Cogolloetal. 21417 (BRIT. FMB,
JAUM). 1 Sep 1997 (fr),/ Pipoly. A. Cogolloetal. 21469. 21471, 1,450-1,500 m, 8 Sep
1997 (scam, fl),/ Pipoly. A. Cogolloetal. 27,520 (BRIT, FMB, JAUM, PSO), (ster.),/. Pipoly.
A. Cogollo et al. 21524 (BRIT, JAUM, PSO); along trail from main Pasco-Tumaco Rd. to
Rk) Nambi, departing main Rd. ac Escuela Mixta El Mirador, 7 km W of Alcaquer, 01"
18' N, 78° 04' W, 1,100 m, 26 Feb 1992 (fr), T. Croat 72394 (JAUM, MO); Corregimiento
Alcaquer, Vereda el Barro, Reserva Nacural Rfo Nambf, W slope, W Cordillera, 01° 18'
N, 78° 08' W, near Cabana Fundacion FELCA, 1,325 m, 1 1 Dec 1993 (fr),/. Betancur et al.
4857 (COL, MO); La Planada, Finca Salazar, 7 km above Ricaurce, 01° 08' N, 77° 58' W,
1,750 m, 29 Nov 198 1 (pist. fl, fr), A. Ge>/try etal. 35188 (BRIT, COL, MO, US); La Planada,
SofRicaurte, 7 km from Tumaco-Pa.sto Rd., 01° 10' N, 77° 58' W, 1,800 m, 24 Jul 1986
(scam, fl), A. Gentry et al. 55053 (MO, PSO); trail to Hondon, 6-12 km SW of La Planada,
01° 04' N, 78° 02' W, 1,750-1,800 m, 5 Jan 1988 (fl bud), 0. de Benavu/es & R. Keating
6041 1 (MO, PSO); Valley of Rfo Guiza, Rd. from El Espino to Tumaco, ca. 21 km W of
PlPOLY, Cybianthus in Ecuador and Peru 83
Ricaurtc, 01° 1 5' N, 78" 07' W, 1 ,000 m, 7 Dec 1 988 (stam. fl), B. Hamniel 17150 (JAUM,
MO). ECUADOR. Bolivar: Hacienda Changuil, LA 16; 02° 06' S, 79° 10' W, 500 m, 17
Aug 1995 (stam. H), X. Cornejo & C. Bonifaz 4339 (GUAY, MO). Carchi: Prominent hillcrest
directly N of Lita, on N side of Rio Mira, E of Rfo Baboso, W-facing slope, 00° 5.^' N, 78°
27' W, 760 m, 7 Aug 1994 (ster.), B. Boyle 347 3 (MO, QCNE), Steep N-facing slope S of
Baboso, S side of Rio Baboso, 00° 53' N, 78° 27' W, 750 m, 1 1 Aug 1994 (ster), B. Boyle
3399 (MO, QCNE); Ri'o Blanco drainage above Chical, tributary of Rfo San Juan, 12 km
W of Maldonado, 1300-1500 m, 25 Sep 1979 (bisex. fl), A. Get/try & C. Shupp 26363
(MO, QCNE); Canton Tulcan, Parroquia Tobar Donoso, Reserva Indigena Awa, Centro El
Baboso, 00° 53' N, 78° 25' W, 1 ,800 m, I 7-27 Aug 1992 (stam. fl), G. Tipaz et al. 1709
(BRIT, MO, QCNE), (stam. fl), G. Tipciz el al. 1H86 (MO, QCNE), (ir), G. Tipaz et al. 1924
(MO, QCNE); 6 km above Maldonado, just below Puentc de Palo, 00° 54' N, 78° 06' W,
2,275 m, 23 May 1993 (stam. fl), B. Boyle &J. Bradford 1878 (MO, QCNE); Trail from
Pailon to Gualpi Chico, Reserva Indi'gena Awa, 1 .5 km past Rfo Blanco, 00° 5 1 ' N, 78°
16' W, 1,000-1,450 m, 14 Jan 1988 (stam. fl), W. Hoover et al. 2436 (MO, QCNE); SE
Trail, Gualpi Chico area of Awa Reserve, 00° 58' N, 78° l6' W, 1,330 m, 19 Jan 1988
(pist. fl, fr), W. Hoover et al. 2809 (MO, QCNE); Trail along ridge and forest slope to NW
of Awa encampment, Gualpi Chico area near Finca Rodriguez, 00° 58' N, 78° 16' W, 1,258-
1 ,323 m, 19 Jan 1988 (fr), W Hoover et al. 3338 (MO, QCNE). Cotopaxi: Rio Guarapa, ca.
20 km NW of El Corazon, 250 m, 19 Jun 1967 (stam. fl), B. Sparre 17091 (S), 20 Jun
1967 (pist. fl), B. Sparre 17081 (S). El Oro: 1 1 km W of Las Pinas on new Rd. to Sta. Rosa,
850 m, 8 Oct 1979 (stam. fl), C. Dodsoii et al. 9101 (MO, SEE); Hacienda Buenaventura,
12 km W of Las Pinas on Rd. to Machaia, 03° 48' S, 79° 46' W, 1,000 m, 1 Mar 1991
(stam. fl), M. Kessler2601 (GOET, MO); New Rd. Saracay-Balzas-Velacruz, ca. 8 km SE of
Saracay, 400 m, 30 Apr 1980 (stam. fl), G. Harling & L. Andersum 18778 (GB). Guayas:
Cordillera Chogon-Colonche, Cerro Los Pontones; 01° 44' S, 08° 40' W, 500 m, 2 Jul 1994
(stam. fl), X. Cornejo & C. Bonifaz 2979 (GUAY, MO). Loja: Tierra Colorada, I km E of
Landara, 8kmEofMercadillo,04°02'S, 79° 57' W, 1,500 m, 9 Feb 1991 (pist. fl, fr), M.
Kessler 2401 (BRIT, GOET). Manabi: Machalilla National Park, zona de San Sebastian,
01 ° 36' S, 80° 42' W, 600-700 m, 2 1 Jan I 99 1 (fr), A. Gentry et al. 72499 (MO, QCNE).
Pichincha: Quito-San Juan Chiriboga-Sto. Domingo de los Colorados Rd., Branch km
59, 18 km NW of Rd., 1,700-2,000 m, 27 Sep 1 986 (bud), V. Zak 1330 (MO, US); Quiro-
Aloag-Sto. Dominigo de los Colorados, km 94, 10 km S of Rd., W slopes ol Volcan El
Corazon, 00° 21' 30 "S, 78° 51' 15" W, 1,300-1,500 m, 25 Dec 1986 (fr), V. Zak 134')
(MO, US); 15 ha. Patch of forest in Cooperativa Sta. Marta No. 2, along Rio Verde, 2 km
SE of Sto. Domingo de Los Colorados, 530 m, 5 Feb 1979 (fr), C. Dodson et al. 7397 (MO,
SEL); Reserva Flon'stica-Ecologica "Rfo Guajalito," Km 59, Quito-Santo Domingo de los
Colorados, 3.5 km NE of Rd., lower slopes of Volcan Pichincha, 00° 13' 53" S, 78° 48' 10"
W, 1 ,800-2,200 m, 28 Dec 1985 {k)J.Jaraniillo 8298 (MO, QCA); Canton Quito, Parroquia
Nanegal, Reserva Maquipucuna, along Inca Trail to Rfo Tulambf, ca. 5 airline km SE of
Nanegal, 00° 07' N, 78° 38' W, 1 ,350 m. 1 5 Sep I 989 (fr), G. Webster & P. Delprele 27394
(DAV, QCA), along trail between Rfo Umachaca and Rfo Tulambf, 00° 07.5' N, 78° 38.5'
W, 1,200-1,300 m, 7 Jul 1990 (fr), G. Webster et ai 27793 (DAV, QCA); Montarias de
Maqmpucuna, Cerro Sosa, 00° 0.5' N, 78° 37' W, 1,950 m, 3 Jul 1991 (fr), G. Webster
28702 (DAV, QCA), 1,750 m, 3 Jul 1991 (stam. fl), G. Webster et al. 28710 (DAV, QCA),
on ridge between Base Camps 1 & 2, 00° 5.5' N, 78° 37' W, 1,800-1,900 m, 6-7 J til
1991 (fr), G. Webster & B. Castro 28769 (DAV, MO, QCA); along Rfo Umachaca near Ha-
cienda El Carmen, 00° 07-7.5' N, 78° 38' W, 1,250 m, 6-7 Jul 1991 (fr), G. Webster etal.
28796 (DAV, QCA). Quininde: Bilsa Biological Reserve, Montarias de Mache, 35 km W
of Quininde, 5 km W of Sta. Isabela, SE ridge trail, 00° 21' N, 79° 44' W, 400-600 m, 21
84 SiDAlcSd)
Sep 1994 (scam, tl), N. Pitiiuiii c/ al. 688 (MO, QCNE), 5 Dec 1994 (fr), N. P/lm/i/ 993
(MO, QCNE), Alon^ Dogala and Invaders Trails, 00° 21' N, 79° 4i' W, 4()()-6()() m, 2
Jan 1995 (fr), N. P/liinin I I6l (MO, QCNE). Province unknown: withoLir locality, Sep
1896 (pist. fl),./, Soc/iro 1U()/12 (B-descr., F Neg 4859).
This species has often been confused with CyhianthHS ipri/cei, owing to
variation in leaf morphology and inflorescence size. However, recent field
studies conducted at the Rfo Nambi Natural Reserve of Narifio, Colombia,
have revealed that juvenile individuals have obtuse to somewhat broadly
rounded, asymmetric leaf bases with long petioles, while mature individu-
als have tapering, asymmetric leaf bases. The confusion was due, in part, to
precociously flowering individuals, detectable by their extremely small flowers,
or to reiterative shoots, detectable by the renewal shoot visible below the
"bayonet", that bears juvenile leaves and pink flowers. While the largest
individuals of Cybianthus siniphx may approach the size of many C sprncei.
the large pith of the stem in the former renders them extremely weak, and
the stems may easily be snapped by hand, while the pith of C. spriwei is
relatively smaller, and the stems can be bent without snapping in the field.
Cybianthiis sniipkx is most closely related to C. sprncei, but may be sepa-
rated from it by the columnar, thyrsoid panicles with racemose branches,
the pedicellate flowers, asymmetric leaf base and shorter cataphylls. Popu-
lations corresponding to the type of Weigeltia cbcinuiephyta differ from the
type of C. simplex in floral structure, directly attributable to the fact that
the former is based on a pistillate, and the latter a staminate collection. However,
the autapomorphic columnar thyrsoid panicle leaves no doubt that they are
synonymous. Because no further collections have been made in the region
from which the type was collected, I defer neotypification until collections
from that area are available.
20. Cybianthus kayapii (Lundell) Pipoly, comb. nov. (Figs. 3E, 4F, 6J,
9A-F). \Vi7c;(-///)/ kjyapi! Lundell, Writ;htia 6: 118. 1 980. Tvfh: PERU. Amazoxas:
Camino de chichijam, cntsa, 300 m, 2 May 1973 (fr), R. Knyap 723 (i iolotyfi:: MO;
isoTYPi;: LL-TEX).
Monoax'u/l treelet to l(-2) m tall. Stems terete, (O.(S-)l-l .7 cm diam., glandular-
papillate at first, glabrescent. Cataphylls alternate in a high spiral, coria-
ceous, subulate, 1 5—45 mm long, ().5-2(-3.5) mm wide. lu^ives psetidoverticillate;
blades chartaceous, oblanceolate to oblanceolate-oblong, (22-)31.3-55.5
cm long, 8.5-19.6(-23) cm wide, apically acute or broadly rounded to a
small acutish ti]^, mucronate, the mucron to 0.5 mm long, the blade gradually
tapering to an abruptly obtuse base appearing auriculate, to 1.5 cm wide,
midrib slightly raised above, prominently raised below, the secondary veins
12-16 pairs, with prominent marginal and submarginal collecting veins,
slightly sunken above, prommently raised below, glabrous above, with ru-
fous hydropoten below; petiole deeply canaliculate, 1-2 cm long, ca. 3-5-
PiPOi.Y, Cybianrhus in Ecuador and Peru 85
4 mm diam., densely glandular-papillate adaxially. Staminate inflorescence: a
pyramidal, bipinnate panicle, (3-)5.5-29 cm long, 5-15(-26) cm wide, the
branches racemose, densely glandular-papillate, succulent, then drying hyaline;
peduncle 3-5.5 cm long; branch bracts membranaceous, subulate, 6.5-8
mm long, 0.5-1.5 mm wide; pedicels cylindrical, 1.2-1.8 mm long, sparsely
glandular-papillate, glabrescent; floral bracts membranaceous, subulate, inserted
on the pedicel about at middle, longer than the pedicel, 1.5-2.5 mm long,
0.1-0.2 mm wide, hyaline, densely glandular-papillate, the margin entire.
Staminate flouws pink, heteromerous, the calyx 5-merous, the corolla 4-merous;
calyx deeply membranaceous, cupuliform, 0.9-1. 1 mm long, the tube 0.1-
0.2 mm long, unequally divided, the lobes deltate to subdeltate, 0.6-0.9
mm long, 0.2-0.7 mm wide, highly reflexed at anthesis, apically acute, epunctate,
hyaline, densely glandular-papillate, the margin glabrous, entire; corolla
carnose, subrotate to rotate, 2-3 mm long, the tube 0.5-0.8 mm long, the
lobes ovate, 1.5-2 mm long, 1.1-1.6 mm wide, apically acute, distally re-
curved 90° from tube axis at anthesis, opaque, densely glandular-granulose
within and along margin, sparsely glandular-papillate along margins without,
epunctate or sparingly and inconspicuously pellucid punctate, the margin
entire; stamens 2.2-2.9 mm long, subequal to corolla lobe or exserted, the
tube conspicuous, carnose, 0.5-0.8 mm long, hyaline, glabrous, elobate,
the apically free portions of the filaments terete, 1 .6-2.2 mm long, free from
corolla, proximally recurved, the anther oblate, 0.3-0.5 mm long, 0.5-0.8
mm wide, always wider than long, apically emarginate to retuse, basally
widely cordate, the connective prominently black punctate dorsally, con-
spicuously black punctate ventrally; pistillode absent or to 1 mm long, 0.1-
0.3 mm wide, densely glandular-papillate. Pistillate inflorescence as in stami-
nate but 6.5-9(-10.5) cm long, erect, not succulent, opaque, densely
glandular-papillate; peduncle 1-2 cm long; branch bracts 2-3 mm long,
0.2-0.3 mm wide; pedicels subobsolete or cylindrical, to 1 .2 mm long, incrassate
and accrescent in fruit to 1.5 mm long; floral bracts inserted on pedicel,
longer than the pedicel, 1-1.3 mm long, 0.2-0.3 mm wide. Pistillate flowers
as in staminate but pink to pinkish-white; calyx 0.9-1 .2 mm long, the lobes
unequally divided, the smaller linear-lanceolate, 0.8—0.9 mm long, 0.3—
0.4 mm wide, the larger deltate, 1.0-1.1 mm long and wide; corolla ro-
tate, 2.6-2.9 mm long, the tube 0.9-1 mm long, the lobes elliptic, 1.7-
2.0 mm long, 0.6-0.7 mm wide, reflexed at anthesis, distally rectirved 180°
from tube axis, glabrous without, sparsely glandular-granulose within, the
margin slightly irregtilar; staminodes very poorly developed, 1.2—1.5 mm
long, the tube 0.9-1.0 mm long, the apically free portions of the filaments
0.3-0.6 mm long when developed, recurved proximally, the anthers badly
formed, at times consisting of 2-3 prominently punctate glands at filament
apex, or orherwise as in the stamens, ovate to subglobose, 0.2-0.3 mm long,
86 SiDA 18(1)
0.3-0.4 mm wide, apically irregular, obtuse, emarginate or refuse, basaliy
cordate, the connective when distinguishable prominently black punctate
dorsally, conspicuously punctate ventrally; pistil clavate to lageniform, 3-
3.5 mm long, the ovary 0.9-1 . 1 mm long, 1 .2-1 .5 mm diam., densely papillate,
the style 2. 1-2.4 mm long, the stigma large, capitate, with 4 principal lobes,
each irregularly lacinate-lobulate, early caducous, the placenta deeply cupuliform,
the ovules 2, buried for 1/2 their length. Frai/ depressed-globose, 5-6 mm
long, 7-9 mm wide, prominently black punctate, the exocarp thin. Bisex/uil
inflorescence: as in stammate but 4-13 cm long. Bi sex// a I flowers as in stami-
nate flowers but calyx 1.1-1.9 mm long, the tube 0.2-0.3 mm long, the
lobes unequally divided, deltate to elliptic, the smaller 0.6-0.7 mm long
and wide, the larger 0.9-1.6 mm long, 0.5-0.6 mm wide, otherwise as in
pistillate flowers; corolla 2.6-2.8 mm long, the tube ca. 0.6 mm long, the
lobes narrowly ovate, 2.0-2.2 mm long, 1.2-1.3 mm wide, recurved dis-
tally 90° Irom tube, sparsely glandular-granulose within, glandular-papil-
late along the margin; stamens as in staminate flower, but 2.2-2.7 mm long,
always slightly shorter than corolla tube, the tube ca. 0.6 mm long, the
apically free portions of the filaments 2.0-2.2 mm long, the anthers widely
ovate, ca. 0.3 mm long, 0.5-0.6 mm wide, apically obtuse to emarginate,
basaliy widely cordate; pistil 2.7-2.9 mm long, the (wary 0.8-0.9 mm long,
0.6-0.7 mm diam. , densely glandular-papillate, the placenta deeply ctipuliform,
the ovules 2, buried for 1/2 length. B/id^A7/f7//r///y depressed-globose, 5-6
mm long, 6-7 mm wide, prominently black punctate, the exocarp thin.
D/str/b//tion. — Colombia (Amazonas, Choco, Narino, Valle del Cauca, ),
Ecuador (Chimborazo, Napo and Pichincha), Peru (Amazonas, Loreto, San
Martm) and Brazil (Acre), growing at sea level-2,530 m elevation.
Ecology cind couservatio}/ stat/ts. — Cyb/anth/is kayap'// is locally common in
small populations at the high watet line in primary "tahuampa" habitats
(varzea forest), along white water rivers, or rarely in premontane habitats
along the edge of pools led by creeks. The species is not known to be culti-
vated and occurs only in areas where deep leaf litter and alluvial deposits
are left after flooding action. It appears that the species can easily be de-
stroyed by soil compaction as a result of trampling, and thus, it should be
considered threatened.
Etymology. — The epithet commemorates Rubio Kayap, an indigenous
Aguaruna Peruvian plant collector who worked with Brent Berlin, known
for his great knowledge of Amazonian flora and ethnobotany.
Local names and /ises. — Colombia: "Hierba de palo grande"(Spanish). Ground
in crude syrups to cure cuts, internal infections, to clean the uterus and the
stomach. Ecuador: "putush" (Shuar). Used against intestinal parasites and
for chronic rectal bleeding (colo-rectal carcinoma °); 10 lbs. of root boiled
in 8 liters of water, down to one liter; 8 cc given as enema before bed; useful
PiPOLY, Cybianthus in Ecuador and Peru 87
for "dysentery." Peru: "mantaya," "kugkuima muspari" (Aguaruna); used
to disinfect dog and insect bites; "napi tsuake" (Huambisa). Brazil: used in
curare cf. B. Krukoff7663.
Representative specimens examined. COLOMBIA. Amazonas: Mpio. Leticia, Parque
Nacional Natural Amacayacu, Centre Administrativo Mata-mata, trail to Amacayacu, km
4, 03° 47' S, 70° 15' W, 120 m, 25 Sep 1991 (stam. fl), A. Ruclas & A. Pneto5l47 (COL,
FMB, MO), 110-120 m, 28 Oct 1991 (A bud),/ Ptpoly &J. Murillo 15483 (COL, FMB,
MO), Quebrada de Agua Pudre, ca. 1.5 km NE of junction with Ri'o Amacayacu, perma-
nent inventory plot, 200-220 m, 11 Nov 1991 (ster.),/. Pipoly et al. ] 5896 (COL, FMB,
MO), 15 Nov 1991 (stam. fl),/ Ptpoly etal. 16075 (COL, FMB, MO); Rio Loretoyacu, 100
m, Oct 1946 (bisex. fl), R. E. Schultes & G. Black 8427 (US). Choco: Bahia de Solano, IS-
IS Apr 1982 (pist. fl, fr), R. Dressier 6036 (COL, FLAS, MO); Mpio. de Quibdo, Corregimiento
San Francisco Icho, Quebrada Caledonia along Caledonia Rd., 9 Apr 1987 (ster.), F. Garcia
&J. Echavavria 259-A (COL, CHOCO, MO). Narino: Mpio. Ricaurte, Reserva Natural La
Planada, 1,800 m, 13 Nov 1993 (fr), C. Restrepo 723 (BRIT, MO, PSO); La Planada, Finca
Salaazar, 7 km above Ricaurte, 01° 08' N, 77° 58' W, 1,750 m, 27 Nov 1981 (fr), A. Gen-
try et al. 35062 (MO, PSO). Narino: Mpio. Barbacoas, Corregimiento Ortiz y Zamora,
Vereda El Barro, Reserva Natural Rio Nambi, ca. 5 km W de Altaquer, faldas occidentales
de la Cordillera Occidental, 01° 18' N, 78° 08' W, 1,350-1,400 m, 3 Sep 1997 (ster.),./.
Pipoly, A. Cogollo, et al. 21241 (BRIT, FMB, JAUM, PSO). Valle del Cauca: Rfo Calima,
region del Choco, La Trojita, 5-50 m, 20 Feb 1944 (stam. fl),/. Cuatrecasas 16272 (COL,
F, US); Mpio. El Cairo, Cerro del Ingles, summit, Cordillera Occidental, Serrania de los
Paraguas, limit Vallc/Choco, El Cairo-Rio Blanco Hwy, 1 hour in jeep from El Cairo, 2,400
m, 1 Jan 1987 (pist. fl), P. Silventom-Sopktn et al 2871 (CUVC). ECUADOR. Carchi: Canton
Tulcan, Parroquia Tobar Donoso, Reserva Indigena Awa, Centro El Baboso, 00° 53' N,
78° 25' W, 1,800 m, 17-27 Aug 1992(stam. R),G. Tipazetal / 706 (MO, QCNE). Esmeraldas:
Canton San Lorenzo, Parroquia Ricaurte, Reserva Indfgena Awa, Comunidad Balsarefio,
Rfo Palabf, 01° 09' N, 78° 31' W, 100 m, 15-29 Apr 1991 (fl bud), D. Ruhio & C. Quelal
1335 (MO, QCNE). Chimborazo: Cordillera Occidental, "El Carmen," Sibambe, 2,450
m, 22 Aug 1943 (pist. fl), Al, Acosta-Solh 5544 (F, QCNA); on slopes of Chimborazo Vol-
cano, (pist. fl, fr), A. Sodiro 100114 (B, destr., QA?, n.v.). Morona-Santiago: Basin of Ri'o
Morona, Rfo Mangosiza, Nayumbime, 45 km SE of Sucua (by air), 200 m S of Don Lui's
Najamte's house, ca. 02° 43' S, 77° 38' W, 300 m, 27 Feb 1990 (fr), C. Limbach 140 (QCA,
NY). Napo: Antisana, Shinguipino Forest, between Rios Napo and Tena, 8 km SE of Tena,
01° 00' S, 77° 50' W, 450 m, 17 Sep I960 (stam. fl), P. Griibhetal. / 633 (K, NY); Canton
Archidona, Carretera HoUin-Loreto, Rio Huataraco, 2 hrs by foot from Guagua Sumaco,
00° 43' S, 77° 32' W, 800-1 ,000 m, 23-30 Aug 1989 (fl bud), C. Ceron & M. Eactos 7641
(MO, QCNE); Canton Tena, 17 Oct 1939 (stam. fl), E. Asplund9396 (S); 3 km E of Caseri'o
Huamanf, N of Carretera Hollin-Loreto, 00° 43' S, 77° 36' W, 1,200 m, 17 Sep 1988 (fr),
F. Hi/rtado & A. Alvarado 503 (MO, QCNE). Pichincha: Along Rd. from Tandayapa to
Mindo, 10 km from Tandayapa, 2,530 m, 16 Dec 1979 (fr), T. Croat 49361 (MO, QCNA).
PERLJ. Amazonas: Prov. Bagua, Dtto. Imaza, Comunidad Aguaruna de Kampaentza (PUJ AIM),
property of Juan Mayan, 740 m, 6 Oct 1994 (pist, fl), C. Diaz et al. 7265 (BRIT, HUT,
MO, USM); Prov, Condorcanqui, Dtto. El Cenepa, NE region of Marahon Drainage Basin,
Rfo Cenepa, Comunidad Tutino, 04° 33' S, 78° 10' W, 350 m, 21 Nov 1993 (pist. fl, fr),
R. Vasq/tez et al. 18395 (AMAZ, BRIT, HUT, MO, USM); Rfo Cenepa, vicinity of Huampami,
ca, 5 km E of Chavez Valdfvia, 04° 30' S, 78° 30' W, Quebrada Chigkishinuk, 10 Apr
1973 (fr), E. Anaiash 211 (MO), 11 Aug 1978 (pist, fl), E. Ancuash 1405 (MO); Trail one
day walk from Huampami to Shaim, creek running into Nahim, 600 m, 27 Nov 1972
88
SlHA 18(1)
Fic;. 17. Cybianthjis anthnriophyll//s Vipoly. A. Habir, showint; pleiochasium wich sympo-
dial branches. B. Sraminate flower, sliowing lanceolate-SLibulatc corolla lobes, elobare scaminal
tube, and conic, vestigial pistillode. C. Pisrillace flower, showing attenuate-acicular calyx
lobes and obturbinate pistil. A, C, drawn from holofype. B, drawn from Bravo & Gomez 49.
Figure drawn by Peggy Duke.
PiPOLY, Cybianthus in Ecuador and Peru 89
(scam. H), B. Berlin 395 (MO); Rio Santia^-o, W bank, 400 m beyond La Poza, 180 m,
without date (fr), F. Donii'ng/iez 147 (MO); 800 m beyond Caterpiza, 200 m, 4 Sep 1979
(fr), V. Hiiasbikat 3^6 (MO), 10 Sep 1979 (stam. fl), V. Hiiaihikat 507 (MO), 1 2 Sep 1979
(bisex. ID, V. Huashikat 5H1 (MO); 26 Mar 1 980 (fr), S. Tunqiii 1110 (MO). Loreto: Prov.
Maynas, Explorama Lodge, near Yanamono, between. Indiana and mouth of Rfo Napo 03°
28' S, 72° 50' W, 103 m, 27 Jun I 983 (fr), Gei//)y el al. 42247 (AMAZ, MO, NY), 106 m,
15 Apr 1985 (fr), R. Vasquez & N.Janiwillo 6325 (AMAZ, MO), 28 Sep 1988 (stam. fl), R.
Wdsqmz & N.Jaramillo 11100 (AMAZ, MO)l 1 Jul 1990 (fr), R. Vdsqmz & N. Jciraiiiillo
14092 (AMAZ, MO, US), 25 km NE of Iquitos, along Rfo Amazonas, 90 m, 03" 30' S,
72" 50' W, 26 Sep 1990 (stam. fl),./, Pi/m/y vt al. 12383 (AMAZ, MO), along S border
trail, 1 10 m, 27 Sep 1990 (stam. i\),J. P/poly ct al. 12490 (AMAZ, MO, NY, US, USM),
(scam. fl),J. Plpoly et al. 12541 (AMAZ, MO, US, USM); Prov. Alto Amazonas, N slopes of
Cerros Camapaqufz at Pongo de Mansenthe, right bank of Ri'o Maraiion, 300-550 m, I 9-
21 Oct 1962 (stam. fl),J. Wurdack 2324 (US, USM). San Martin: Pongo de Cainarachi,
Rio Cainarachi, tributary of Rio Huallaga, 230 m, Sep-Oct 1962 (stam. fl-except speci-
men at S-bisex. fl), G. King 2691 (A, F, GH, K, MO, NY, S, US). BRAZIL. Acre: Cruzeiro
do Sul, Rios Jurua and Moa, 8 km above Cachoeira Grande, 07° 30' S, 73° 30' W, 27 Apr
1971 (fr), G. Prance etal. PI 2555 (IAN, MG, NY). Amazonas: Basin of Riojavarf, Mpio.
Sao Paulo de Oliven<,-a, near Esperanga, Dec 1935 (fr), B. Kri/koff7663 (NY, U).
Cybianthm kayapii may be confused with C. sprucei (Hook, f.) G. Agostini,
but may be recognized by the shorter petioles, heteromerous flowers and
mucronate leaf apices. However, the leaf blades with mucronulate apices
and subauriculate bases, and heteromerous flowers indicate Cybianthus kayapii
is more closely related to C. anthiiriophyllm. From Cybianthus anthrmophyllus,
C. kayapii may be separated by the much smaller, flat, entire leaves with
entire margins, the deltate calyx lobes with entire margins, and the carnose
corolla.
21. Cybianthus anthuriophyllus Pipoly, sp. nov. (Fig. 17). Type: ECUA-
DOR. Napo: Canton Gonzalo Pizarro, Rfo Tigre, affluent of Rfo Dashino, entering
from 73 km of Rd. from Lumbaque to El Reventador, 10 km S oi Lumbaque, 00°
05' S, 77° 24' W, 900-1,100 m, 18-21 Feb 1 987 (pist. fl, fr), W. Palacios & D. Neill
1584 (hoi.otype: US; i.sotypes: K, MO, QCNE).
Ob folia oblanceolata ab lamina ad pctiolum gradatmi contracco, a primo intuitu cum
C kayapi confusa est, sed ab ea marginibus laminaribtis revolutis serrulatisque (nee integettimis
planisque) inflorescentiis 28 (non 8-15) cm longis, pleiochasia cum ramulis floriteris cymosis
(nee bipinnatipaniculatis cum ramulis floriferis racemosis) lobulis calycinis subulatis (nee
ovatis) praeclare distat.
Monoaxial treelet to 0.6 m tall. Stems terete, weakly woody, ca. 2 cm diam.,
sparsely glandular-papillate, glabrescent. Cataphylls tightly pseudoverticillate,
coriaceous, linear-subulate, 4.0-6.5 cm long, 0.3-0.6 cm wide, densely and
prominently punctate-lineate, sparingly glandular-papillate, glabrescent.
Leaves pseudoverticillate, erect; blades coriaceous, narrowly lanceolate, (56-)
104-1 10cm long, (l4.5-)17-22 cm wide, apically broadly rounded or rounded
to a minute abrupt submucronate acumen 2 mm long, the blade gradually
tapering to the petiole, almost obtusish basally, midrib slightly raised above.
90 SiDA 18(1)
prominently raised below, the secondary veins ca. 42 pairs, impressed above,
prominently raised below, nitid and epunctate above, purple (when fresh),
inconspicuously punctate and papillate-puberulent below, the papillae erect,
rufous, the margin thin, opaque, inroUed, prominently straight-serrulate,
the teeth alternatively larger and smaller (ca. 1 mm and 0.5 mm long, re-
spectively); petiole deeply canaliculate, thick, ca. 1.5 cm long, 0.8 cm diam.,
sparingly glandular-papillate. Staminate inflorescence: a pyramidal thyrsoid
panicle, ca. 16 cm long, 23 cm wide, the branches pseduoracemose (sym-
podial), the rachis densely glandular-papillate, the peduncles longer below,
shorter above; inflorescence branch bracts linear-lanceolate, 7—9 mm long,
2-3 mm wide, apical ly attenuate-acicular, sparsely glandular-papillate, the
margin entire; floral bracts acerose, 2-3 mm long, 0.4-0.8 mm wide; pedicels
1.0-2.5 mm long, angular, thin, the longer pedicels ui the lower portion of
the inflorescence, the shorter ones above, densely glandular-papillate, the
papillae persistent. Staminate flowers heteromerous, membranaceous, hyaline,
the calyx (5-)6-merous, the corolla 5-merous; calyx subcotyliform 3.7-4.7
mm long, 0.4—0.7 mm wide, apically acuminate-acicular, prominently keeled,
the keel thickened hyaline, epunctate, sparsely glandular-papillate except
densely so along the margin, the margin entire; corolla subrotate, 7.4—10
mm long, the tube 0.7-0.9 mm long, the lobes lanceolate-subulate, 6.3-9
mm long, 2.7-3.2 mm wide, the apically long-acuminate, highly reflexed,
sparsely glandular-papillate without, very sparsely glandular-granulose within
above the junction within tube the margins densely glandular-papillate,
entire; stamens 2.2—4.1 mm long, the staminal tube conspicuous, coria-
ceous, 1.4-1 .8 mm long, glabrous, elobate, the apically free portions of the
filaments ventrally recurved, 0.8-0.9 mm long, flat, glabrous, the anthers
ovate, 0.3-0.4 mm long and wide, apically and basally emarginate, dorsifixed
less than _ length, the connectives prominently black punctate ventrally
and dorsally; pistillode conic, vestigial-0.8 mm long or absent. Pistillate
inflorescence a pleiochasium, with branches cymose (sympodial), the rachis
densely glandular-papillate, the peduncles 1-3 cm long, longer below, shorter
above; inflorescence bracts resembling cataphylls but smaller, 1.3-1.6 cm
long, 1 .0-1.3 mm wide, conspicuously punctate-lineate; inflorescence branch
bracts linear-lanceolate, 1 .2-1 .5 mm long, 0.2-0.4 mm wide, apically at-
tenuate-acicular, sparsely glandular-papillate, the margin entire; floral bracts
acerose, longer than the pedicels, 0.5-1.0 cm long; pedicels angular, thin,
1.2—7.5 mm long, the longer pedicels in the lower portion of the inflorescence,
densely glandular-papillate, the papillae persistent. Pistillate flowers as in
staminate but calyx 6-merous, (2.1-)3-4.1 mm long, the tube 0.3-0.7 mm
long, 0.5-1 .0 mm wide, apically long-attenuate-acicular; corolla (from dried
remnants) subrotate, hyaline, 5.0-7. 1 mm long, the tube 1 .0-1 .3 mm long,
the lobes hnear-subulate, 4.0-5.8 mm long, 1.1-1 .5 mm wide at base, apically
PiPOi.Y, Cybianthus in Ecuador and Peru 91
subulate, highly reflexed, twisted and distally recurved at anthesis; stami-
nodia 1.2—1.3 mm long, the staminodial tube 0.9—1.2 mm long bearing
well-developed lobes alternate with the apically free filaments, the filaments
0.3—1.1 mm long, flat, glabrous, the antherodes malfi^rmed, mostly con-
sisting of undifferentiated tissue surrounding prominent black punctations
or suborbicular and 0. 1—0.5 mm long, 0. 1—0.6 mm wide, always wider than
long, apically emarginate, dorsifixed slightly less than _ length, the connectives
prominently black punctate dorsally; pistil obturbinate, 2.5—3.0 mm long,
the ovary 1 .0—1.5 mm long, 0.8-1.3 mm wide, densely papillate and prominently
black punctate, the style 1.3-1.7 mm long, densely glandular-papillate, the
stigma large, capitate, lobes, the lobes lacinate, each lobe to 0.4 mm long,
early caducous, the placenta widely conic, bearing 4 uniseriate, exposed ovules,
the ovules on the periphery of the placenta. Bisexual fruit pink, globose, 4—
6 mm long and diam., the punctation prominent, brown when fresh (teste
coll.), red or black upon drying, the exocarp thin.
Distribution. — Amazonian ("Oriente") Ecuador and adjacent Peru (Loreto),
l60-300(-l,100) m elevation.
Ecology and conservation status. — Cybianthus anthuriophyllus grows in pri-
mary tropical wet forest and premontane wet forest on terra firnie above the
high water contour. It is found in primary forest as well as in secondary, but
it is not known whether the plant is cultivated in secondary forest situa-
tions. At this time, the species can be considered locally common but not
threatened.
Etymology. — The specific epithet refers to the unique shape of the adult
leaves, held erect in vivo and reminiscent o{ Anthurium crassinerviuyn (Araceae).
Local names and uses: — Ecuador: "namakuk" (Achuar JiVaro); "Acuari" (dialect
unknown); "challuo panga" (Quichua). Peru: "kutukupish," "takushia,"
"mutupash," "kurup" (Maynajivaro), "sierra panga" (Quichua). Leaves crushed
and used as a fish poison( W. Leivis et al. 14051); inner stem is scraped and
an infusion given to dogs to drink to improve their hunting abilities and to
enhance their stamina {W. Lewis et al. 1 1 133): stem is scraped and its juice
put into a dog's nose to produce sneezing {W. Lewis et al. }()475), or to "make
it an attack dog against thieves" (Lewis et al. 12833).
Paratypes. ECUADOR. Napo: Cuyabeno-Punta AreniUa; Sep 1981 (stam. fl), E. Bravo
&P. Gamez 49 (QCA); Canton Orellana, Yasunf National Park, Maxus Rd. and pipiineconstmction
project km 10, 00° 29' S, 76° 34' W, 250 m, 29Jun 1994 ( fr), N. Pitwati 448 (JVtO, QCNE).
Pastaza: Kapawi (Amuntaf), rfo Pastaza; Village area, 02° 31' S, 76° 48' W, 235 m, 25-
29 Jul 1989 (ster.), W. Lewis et at. 14051 (MO). PERU. Loreto: Prov. Alto Amazonas,
Puranchim, Rio Sinchiyacu, 02° 50' S, 76° 55' W, 200 m, 3-7 Dec 19<S8 (stcr.), W. Lewis
et al. 14390 (MO); Washintsa and vicinity, Rfo Huasaqa, 03° 20' S, 76° 20' W, 185 m,
16-26 Jun 1986 (ster,), W. Lewis et at. 11153 (MO); Prov. Loreto: Nueva Jerusalem and
vicinity, Rio Macusari, 02° 55' S, 76° 15' W, 220-300 m, 29 Dec 1985-3 Jan 1986 (Fr),
W. Lewis et at. 10475 (MO); Pampa Hermosa and vicinity, Rio Corrientes, 1 km S of junc-
92 SiDA 18(1)
cion with Rio Mucusari, O.V 15' S, 75" 50' W, 16() m, 3-20 Dec 19H5 (sram. H), W. Leins
et cil. lO.UO (MO); Vista Aiegre, Rio 'I'igre, 02° 40' S, 75° 35' W, 210 m, 17 Mar 1987
(stcr.), W. Leir/s ci a/. 1285,^ (MO).
Cy/jh///tb//s ai/thur/ophyII//s is unique within subgenus Cnmoni^niue by its
small, marginal pectinate leaf serrations and pleiochasial inflorescence, ap-
pearing paniculate, but with sympoclial primary and secondary branches,
and thus, cymose. The leaves appear subsessile, with the blade gradually
tapering to the deeply canaliculate petiole, a feature found otherwise only
in its closest congener, Cyhianthus kayapii. However, Cybianthns anthuriphyllin
is clearly distinct from C. kaycipii because of the inroUed, pectinate-serru-
late leaf blade margin, tlie much longer, pleiochasial inflorescence, and subulate
calyx lobes. The extremely long cataphylls are the best developed in the
subgenus.
V. Cyhianthus subgenus Triadophora (iVIez) G. Agostini, Acta Biol. Venez.
1 0: 1 64. 1 9H(). Wasi^dtn-t subgenus TridJophoni Mez in Engl., Pflanzenr. IV. 236(Hei'c
9):291 . I 902. Typi; Species. Wcii^dtid sdtliiiiii (Hook, f.) Mez in Engl. = Cyh'hnithi/s
schl/niii (Hook, i.) G. Agosrini.
Currdiunid D'Arcy, Ann. Missouri Hot. Card 60:442. 1973. Type Species. Corrdlnvui
spectcihilis (Standi.) D'Arcy = Cyhu/iithus .uhliinii (Hooker 1.) Ct. Agostini.
As here interpreted, Cyhictnthns subgenus Tnculophora is monotypic. Its
only species, C. schlni/ii, is easily recognized by its monoaxial habit and
autapomorphic rufous glandular tomentum of malpigiaceous trichomes, leaf
blades with subepidermal fibers and pseudocataphylls (here defined as petiolate
cataphylls). The first full description oi Cyhianthi/s schliiini is provided, along
with complete synonymy and complete exsiccatae for Ecuador and Peru,
and representative ones for other areas.
22. Cyhianthus schlimii (Hook, f.) G. Agostini (Fig. 3D, 6K, 6L), Acta
Biol. Venez. 10:165. 1980. Connwiyrsiiw schliwii Hook. f. in Benth. 6- Hook.,
Gen. Pi. 2:644. 1 876. Weigdtui sddunii (Hook. \.) Mez in Engl., Pflanzenr. IV. 236(Heft
9):29l. 1902. CoynUuina Khlimn {W^oV. f.) D'Arcy, Ann. Missouri Bot. Cjard 60:443.
1973. Type: COLOMBIA. Meta: Llano de San Martin, 300 m, Jan 1836 (stam. fl),
J. 'Wunui 7594 (iioi.oi ype: K; esotypes: COL, LE, MA, P).
Weigeltici multijlora A.C. Smith., Bull. Torrey Bot. Club 60:387. 1 933- syn. nov. CmreUiiinu uiiiU't-
flora {K. C. Sm.) D'Arcy, Ann. Missouri Bot. Gard 60:4 15. 1973. (A. C. Sm.)G. Agostini,
Acta Biol. Venez. 1 0: 165. 1980. Type: BRAZIL. Mato Grosso: Near Tabajara, upper
Rio Machado, 2t Nov 1931 (sram. fl), B. A. Kri/koff 1 .]HS (hoi.otyi'i,: NY; lsotype: A).
Aniisni ipeclahilis SvAnd[., Publ. Eield Mus. Nat. Llist. Bot. Ser. 18:893. 1938. syn. nov.
Weigdtia spectabilLs (Standi.) Lundell, Wright ia 4: 1 69. 1 97 1 . Cotrdliami speclahilis (Standi.)
D'Arcy, Ann. Missouri Bot. Gard. 60:443. 1973- Cyhunith//.s spaUihilis (Standi.) G.
Agostini, Acta Biol. Venez. 10:165. 1980. Type;: COSTA RICA. Aeajuiea: Cataratas
(Los Angeles) de San Ramon, A|-ir 1935 (stani. ti), /\, Bn//a 20"^ iO (molotype: E; esotypes:
CR, F).
Wc'/gdt/j trunidni Aspl., Bot. Not, I9t9:8{)2. 1939. T\m- COLOMBIA. Cauca: Near
Distrito El Tarn bo, 900 m, 3 1 Jul 1936 (stam. II), K. vtm SiiL'/der)i 919 (i ioeo'EYPi:: S).
PiPOi.Y, Cybianthus in Ecuador and Peru 93
Weigeltid schiimii (Hook, f.) Mtv van inteniiecl/n Moklcnkc, Phytokigia 2:242. 1 947. Typi;:
COLOMBIA. Vai.ii; di;i. Calica: Pacific coa.s:, Rfo Cajambre, San Isidro, 5-100 m, 2-
5 May 1944 (stam. fl),_/. Ci/citreaiscis 1 7312 (noioTVi^r,: NY; isotypi-s: COL, F-2 sheets).
Monoaxtal tree to 5 m. Stem terete, 0.8—2.0 cm diam., the wood dense,
minutely rufous glandular appressed tomentose, the trichomes malpighiaceous,
early glabrescent. Pseiidocataphylls produced only irregularly, chartaceous,
subulate, ca. 2.0—3.0 cm long, 0.5—1.0 cm wide, apically acute, mucronate,
densely rufous puberulent, black lineate-punctate, the margin entire; peti-
ole subobsolete, to 0.2 cm long. Leaves tightly pseudoverticillate; blades
chartaceous, elliptic, oblong or oblanceolate, 25—65 cm long, 5.5-20 cm
wide, apically acute or subacuminate, mucronulate, the mucro often sclerified,
the acumen 0.5-3.0 cm long, base long-attenuate, the blade decurrent on
the upper portion of the petiole, midrib slightly elevated above, prominent
below, the secondary veins 9—13 pairs, prominent, the marginal veins loop
connected, conspicuously striolate by subepidermal fibers, these visible above
and below, sparsely rufous puberulent above, moderately puberulent be-
low, at times glabrescent, hydropotes absent, sparsely to densely punctate
or lineate-punctate below, the margin opaque, irregular, entire to roughly
serrate; petioles canaliculate, 1 .0— 3.0(-10) cm long, 0.5—1 .0 cm diam., abruptly
swollen basally, puberulent, glabrescent. Staniinate Inflorescence a pinnate or
bipinnate columnar panicle 13^0 cm long, 3-20 cm wide, the rachis densely
glandular-papillate and rufous puberulent, the flowering branches racemose;
peduncle 8-15 cm long; inflorescence bract chartaceous, ovate, 9—15 mm
long, 2.4—4.5 mm wide, apically acute, densely rufous glandular puberu-
lent, conspicuously black punctate and lineate-punctate, the margin opaque,
entire; inflorescence branch bracts membranaceous, linear, 10—13 mm long,
1.9—2.1 mm wide, apically narrowly acute, mucronulate, minutely rufous
puberulent, orange furfuraceous lepidote, densely and conspicuously black
lineate-punctate, the margin opaque, entire; floral bracts membranaceous,
subulate, 0.8—1.3 mm long, 0.2—0.4 mm wide, sparsely rufous puberulent,
the margin entire; pedicel terete, 2.0^.5 mm long, prominently black punctate,
densely papillate and rufous puberulent. Staminate flowers 3(^)-merous, light
purple, then dull yellow; calyx chartaceous, cupuliform, 0.9-1-8 mm long,
the tube 0.3 mm long, the lobes subdeltate, 0.7—1.6 mm long, 0.9-1-2
mm wide, apically acute, densely rufous puberulent, glabrescent, densely
and prominently black punctate, the margin flat, wide, hyaline, densely ciliolate,
the cilia often caducous; corolla rotate, chartaceous, 3.0-5.0 mm long, the
tube hyaline, 0.8—1.0 mm long, the lobes elliptic or oblong, 2.2—4.0 mm
long, 1.6—2.2 mm wide, apically obtuse to rounded, subcucullate, invo-
lute, distally recurved 180" relative to tube, sparsely rufous puberulent without,
very sparsely glandular-granulose within basally, often glabrescent, very densely
and prominently black punctate except margin hyaline, irregular, glabrous,
94 SiDA 18(1)
entire; stamens 2.4-3.5 mm long, the tube membranaceous, inconspicu-
ous, 0.8—1.0 mm long, hyaline, epunctate, glabrous, elobate, the apically
free portions of the filaments 1 .8—2.3 mm long, sparsely or epunctate, sparsely
rufous puberulent at first, glabresccnt, the connective prominently punc-
tate, or inconspicuously so, the punctation orange, red or black, the anthers
cordate, 0.5-0.6 mm long, 0.7-1.0 mm wide, apically subacute to rounded,
base deeply cordate, dorsifixed at point less than 1/5 distance from apex;
pistillode none or conic, to 1.0 mm long, densely and prominently black
punctate and rufous papillate. Pistillate inflorescence as in staminate but more
columnar, (3.5-)8.0-] 8.5 cm long, (2.0-)3.5-6.() cm wide, the branches
subcorymbose to rarely racemose; peduncle 1.5—6.8 cm long; inflorescence
bract ovate to elliptic, 4.5-12 mm long, 1.9—4.0 mm wide, conspicuously
black punctate; inflorescence branch bracts 5-15 mm long, 0.5-2.5 mm
wide; floral bracts 0.9-2.0 mm long, 0.3-0.6 mm wide; pedicels terete,
2.0— 4.0 mm long, accrescent to 5.0 mm long and incrassate to 2.0 mm
diam. in fruit. Pistillate flowers as in staminate but 3-merous, calyx purple,
corolla yellow; calyx subcotyliform, 1.8-2.0 mm long, the tube 0.2-0.3
mm long, the lobes subdeltate to widely ovate, 1.6-1.8 mm long, 1.6-2.2
mm wide, apically acute, the margin slightly erose, conspicuously long glandular-
ciliate, the cilia often caducous; corolla 4.2—5.0 mm long, the tube 0.3-0.5
mm long, the lobes oblong, 3-2-4.5 mm long, 2.3-2.6 mm wide, apically
obtuse, somewhat cucuUate, reflexed ca. 45" from tube axis, densely pu-
berulent without, sparingly glandular-granulose basally within; staminodes
to 2.5 mm long, the tube membranaceous, inconspicuous, to 0.5 mm long,
the filaments adnate to corolla lobe ca. 0.5 mm long, then apically free to
1.0 mm long, thick, terete, rarely punctate, the antherodes cordate, ca. 0.6
mm long, 0.7—0.8 mm wide, apically acute to apiculate, base widely cor-
date, the connective punctate, prominently or not; pistil clavate to obnapiform,
4.0—5.2 mm long, the ovary 1.8 mm long, 0.5-1.6 mm wide, prominently
black punctate and rufous puberulent, the style to 1.3 mm long, densely
rufous puberulent, the stigma capitate to lobed, 3(— 4)-lobed, the placenta
deeply cupuliform, bearing 2 ovules exposed apically. Print globose, 1.0—
1.3 cm long, 1.0—2.0 cm diam., at maturity, the exocarp bright orange to
red-orange, prominently black punctate.
Distribution. — From the Atlantic Slope (Dpto. Ri'o San Juan) Nicaragua,
to state of Pando, Bolivia and adjacent Amazonia of Brazil, 0-1,800 m.
Ecology and conservation status. — Cyhianthus schlimii occurs in a variety of
habitats, from lowland to premontane tropical moist, wet and pluvial for-
est. It is locally common, but restricted to primary forest. Therefore, it should
be considered threatened.
Etymology. — The species is named for Louis Joseph Schlim, a Belgian plant
collector working for J.J. Linden in Brussels, who collected extensively around
PiPOLY, Cybianthus in Ecuador and Peru 95
Caracas to the Venezuelan Andes, and from there, to the grasslands of Meta
and the Sabana de Santafe de Bogota, Colombia, during the period 1841-
1852. Schlim also collected with Nicolas Funck later in Venezuela.
Local names and uses. — Peru: "napi tsuake" (Huambisa).
Representative specimens examined. NICARAGUA. Rio San Juan: El ReJos, midpoint
between El Castillo and Delta de San Juan, 0-50 m, 23 Mar 1961 (stam. fl), G. Bunting &
L. Lick 77^ (F, NY). COSTA RICA. Alajuela: 15 km NW of Arenal by air, 2 km NW of
Nuevo Arenal on Rd. to Tilaran, then 3 km NE on Rd. to San Rafael de Guatuso, then 2
km W on Rd. to Finca Cote, 10° 34' N, 84" 54' W, 700 m, SE side of Lago Cote, 30 Apr
1983 (fr), R. Liesmretal. 15093 (CR, MO, WIS). Heredia: Zona Protectora, N slopes of
Volcan Barba. betweem Rio Peje and Rio Guacimo, along Quebrada Cantarana, 300-400
m, 18 Jan 1983 (stam. fl), M. Grayum&G. 5/)^/z 3 i 70 (CR, DUKE, MO); Finca La Selva,
Rio Puerto Viejo 2 km E of jet with Rio Sarapaqui, 10° 26' N, 84" 00' W, 100 m, 14-17
Jun 1968 (stam. fl), W. Burger & R. Stolze 5803 (CR, F, MO, NY). Limon: Near Finca
Castilla, 30 m, 24 July 1936 (ster), C. Dodge & V. Goerger9283 (F, MO). San Jose: Estacion
Carrillo, Cafion del Rio Sucio, 450-700 m, 12 Nov 1983 (pist. fl), /. Chacon & G. Herrera
1720 (CAS, CR, MO, NY). PANAMA. Bocas del Toro: Cerro Bonyic, above Quebrada
Huron, 180-400 m, 13 Apr 1968 (fr),/ Kirkhnde &J. Duke 610 (MO). Colon: Base of
Cerro Bruja, along Ri'o Escandaloso, above Mina Boqueron, No. 2, 47.5 km from Tran-
sisthmianHwyonRd. to Salamanca, 09° 50' N, 79° 32 'W, 10-200 m, 18 Mar 1982 (stam.
fl), S. Knapp & W.J. Kress 4282 (MO, NY, PMA). Darien: Rio Tuquesa, Tuquesa Mming
Co. camp, Charco Peje, 250 m, 7 Jul 1975 (stam. fl), 5. Mori 7015 (MO, SCZ). Panama:
Cerro Campana, 800 m, 22 Jun 1967 (fr), T. Croat 17167 (MO). San Bias: Trail from Rio
Estadi to Cerro Banega, 300-530 m, 09" 23' N, 78" 51' W, 21 Dec 1985 (stam. fl), G. de
Nevers & H. Herrera 6642 (CAS, MO, PMA). COLOMBIA. Antioquia: 6 km E of Guapa,
53 km S of Turbo, 240 m, 13 May 1945 (stam. fl), 0. Haught 4660 (US); Mpio. Anori,
Corregimientode Providencia, Buenos Aires, 500-600 m, 4 Feb 1972 (fr), D. Soejarto 3205
(HUA, MO, NY); Vicinity Planta Providencia, 26 km S, 23 km W (by air) of Zaragoza,
07" 13' N, 75" 03' W, valley of Rio Anori between Dos Bocas and Anorf, 1 Jun 1976 (fr),
J. Shepard323 (COL, WIS); Vicinity Medellin, 20 Aug 1927 (stam. fl), R. Toro 356 (MEDEL,
NY); Medellfn-Bogota Hwy, sector Rio Samana-Ri'o Claro-San Luis, 400-1,000 m, 24 Aug
1982 (fr),/ Hernandez & S. Hoyos 483 (COL, HUA); Mpio. Urrao, Boundary of Parque
Nacional Natural Las Orqufdeas, Vereda Calles, Permanent Inventory, Premontane Rainforest,
left bank of Rio Calles, 06° 32' N, 76° 19' W, 1,450-1,500 m, 30 Nov 1993 (ster.),/
Pipo/y et al. 17406 (BRIT, JAUM, MO). Boyaca: Region of Cerro Chapon, extreme W
part of Boyaca, NW of Bogota, 2,300 m, 31 Jul 1932 (fr), A. Laurence 370 (A, NY, S); El
Umbo region, 1,000 m, 12 Oct 1932 (stam. fl), A. Laurence 530 (A, BM, F, G, GH, MO,
NY, S, U, UC, US). Choco: Mpio. Quibdo, Quebrada La Platina, Hwy to Medelli'n, 25 Sep
1983 (fr), L. Arias eta/. 134 (MO); Mpio. San Jose del Palmar, along Rfo Torito (affluent of
Rio Habita), W slopes, 850- 950 m, 15 Mar 1980 (fr), E. Forero et al. 7350 , 16 Mar 1980
(fl bud), E. Forero et al. 7393 (COL, MO), Vereda Portachuelo, Hacienda Barro Blanco, 1 ,350
m, 15 Jan 1983 (fr), P. Franco et al. 1325 (COL); Rio Mecana, ca. 10 km E of Mecana, 06"
15'N, 77"25' W, 100 m, 7 Mar 1983 (stam. fl),A. Gentry & A. J iincosa 41072 {QO\.,U.O,
JAUM), 710-880 m, 8 Jan 1984 (stam. fl), A. Juncosa 1769 (COL, MO, JAUM); Mpio.
Novita, vereda Curundo, left bank, Rio Ingara, 550 m, 1 Dec 1983 (fr), P. Franco et al.
1059 (COL); Rio Nuqui, 400 m, 25 Jan 1947 (stam. fl), 0. Haught 5479 (COL, US); La
Mojarra, upriver from Istmma, 05" 12' N, 76" 37' W, 30-60 m, 5 Nov 1983 (fr), A.J/mcosa
1255 (COL, JAUM, MO, NY); S of Rio Condoto, between Quebrada Guarapo and Mandinga,
120-180 m, 22-28 Apr 1939 (fr), E. Ktlltp 35675 (COL, US). Cundinamarca: Cordillera
96 SiDA 18(1)
Bo^'oni, 2,000 m, Sep 1H55 (Fr),/. 'I'm/i/a 4 (BM), Meta: Near Salitre, 6 Jan 1876 (stam.
ti), E. A/Jc/n 1 151, 7 Jan 1876 (ster), E. Andre s.n. (K); C]ano Tigre, between Cano Aguas
Claras and Cano Grande, 4.5 km SW of ViUavicencio, 04" 07' N, 73" 39' W, 500-550 m,
24 Feb 1943 (pist. H), ER. Eoiherg 20148 (COL, US); Llano de San Martfn, (scam, fl, pisr.
fl mixed), f/. Karstc/i s.n. (LE-2 sheets). Nariiio: Mpio. Barbacoas, C^orreginiiento Orci'z y
Zamora, Vereda El Barro, Rcserva Natural Rfo Nambi, ca. 5 km W de Altaquer, laldas
occidcntales de la Cordillera Occidental, 01° 18' N, 78" 08' W, 1,350-1,400 m, 1 Sep
1997 (stam. fl), /. P/po/y. A. Cogollo. et al. 2/095 (BRIT, FMB,JAUM, PSO), 21144, 21149.
1,450-1,500 m, 2 Sep 1997 (fr),./. P//mIy. A. Cogollo et ul. 21483 1250-1,350 m, 8 Dec
1997 (ir),y, P/po/y. A. Cogollo et cd. 21598 (BRIT, JAUM, PSO). Norte de Santander:
Ocana, I 200 m, May 1846-52 (stam. fl), L. Schlim ()86 (BR-3 sheets, F-2 sheets, G-BOIS-
2 sheets, G-DEL, MA-2 sheets). Santander: 8 km SE ot Barrancabermeja, Ri'o Opon, 200
m, 28 Atig 1954 (stam. fl), R. Cdstiinedci 4~^4(^ (COL), vicinity Barrancabermeja, between
Ri'os Sogamoso and Colorado, 100—500 m, S of Rio Sogamoso, Camp Mesa, 8 Jan 1935
(stam. fl\ 0. Haiight 1502 (A, COL, F, NY, US). Valle del Cauca: Cordillera Occidental,
W slope, along Rio Sanc]Liininf, left side, La Laguna, 1,250-1,400 m, 13 Dec 1943 (stam.
fl),J. Citdtyecdias 15474 (VALLE); Bahfa Malaga, Quebrada Algeria, new Rd. to military
base,0 4" 02' N, 77" 22' W, 50 m, A. Gentry et al. 53326 (COL, MO, US); Bank of Rfo
Digua, Rfo San Juan, below Queremal, to the right ol river between km 52 and 53, 1300—
1500 m, 19 Mar 1947 (fr),/ Ctuttrecma.^ 23855 (COL, F-3 sheets); Calima Dam, 1600-
1700 m, 17 Sep 1966 (stam.).V. Esf>tm/1 2 109 (MO, VALLE). Without locality, date, (stam.
(\)J. Mittis449 (MA). 29/9 (MA), 3907. 5/02^/ (MA, US), Warscewiczs.n. (B-destr., Photo-
F Neg. 4858). ECUADOR. Carchi: Trail along plain above Tovar-Donoso and Rio Guape,
01" 10' N, 78" 18-31' W, 280-450 m, 18 Feb 1984 (stam. fl), W. Hoover 1194 (MO), SE
Trail, (iualpi Chico Area of Awa Reserve, near encampment, 00" 58' N, 78° 16' W, 1 ,330
m, 19 Jan 1988 (stam. fl), W. Hoover et al. 2815 (MO, QCNE), Gualpi Medio Community,
Awa Reserve, 900 m, 2 I rVIay 1992 (bud), C. Quelal et al. 764 (MO, QCNE) El Pailon, 45
km below Maldonado, along path to Tobar Donoso, 800 m, 1 Dec 1979 (fr), AL Madison &
E. Besse 7201 (AAU, F, QCNE, SEE); Canton Tulcan, Reserva Indfgena Awa, Parrocjuia
Tobar Donoso, sector El Baboso, 00° 53' N, 78° 20' W, 1,600 m, 3 Oct 1991 (A, fr), G.
Tipaz et al. 260 (BRIT, MO, QCNE), (fr), 3 / 1 (BRIT, MO, QCNE), Centro El Baboso, 00°
53' N, 78° 25' W, 1,800 m, 17-27 Aug 1992 (fr), G. Tipaz et al. 1950 (BRIT, F, MO,
QCNE). Esmeraldas: Canton San Lorenzo, Reserva Indigena Awa, Canon del Rfo Mira,
10 km W of Alto Tambo, Comunidad "La Union," 01° 02' N, 78° 26' W, 250 m, 16-26
Mar 199 1 (fr), D, Riibto et al. 1262 (MO, QCNE). Sucumbios: Canton Lago Agrio, Reserva
Cuyabeno, Laguna (Trande, Near NEOTROPIC Cabins, 00° 00' S, 76" 1 1' W, 230 m, 15
Nov 1991 (stam. fl), W Palacios et al. 9269 (BRIT, MO, QCNE). Zamora-Chinchipe:
Canton Nangaritza, Valle del Rfo Nangaritza, Miazi, 04° 18' S, 78° 40' W, 1,200 m, 10
Dec 1990 (pist. fl), W. Palacios 67.34 (BRIT, MO, QCNE), Behind military camp, 04° 16'
S, 78° 42' W, 970 m, 20 Oct 1991 (stam. fl), W. Palacios et al. 8483 (BRIT, MO, QCNE).
PERU. Amazonas: Camino de chichijam, entsa, 180-250 m, 2 May 1973 (stam. fl), R.
Kayap 728 (MO); Valle del Rio Santiago, 03° 50' S, 77" 40' W, Quebrada Caterpiza, 2-3
km behind Caterpiza, 200 m, 4 Jan 1980 (stam. fl), S. Tiinqi/i 549 (MO). Loreto: Prov.
Maynas, Guarnicion Pijuayal, near Pebas, 130 m, 7 Sep 1988 9stam. fl), C. Diaz et al. 566
(MO). Ucayali: Prov. Padre Abad, Boqueron del Padre Abad, 400 m, 20 May 1969 (fr),./.
Schunke 3068 (F-2 sheets, US, USM). BRAZIL. Amazonas: Mpio. Humayta, on plateau
between Rio Livramento & Rio Ipixuna, 7-18 Nov 1934 (stam. fl), B. Kr//koff7290 (A,
NY, S, U); Sfu) Paulo de Oliven^;a, near Esperanc;a, Dec 1935 (ster.), B. Kr//koff7(y63 (K,
NY), Behind Sao Paulo de Olivenga, 16 Atig 1 973 (fr), E. Lleras et al. PI 73 1 ^ (GB, INPA,
MG, NY); km 500, Manaus-Humayta Rd., 17 Sep 1980 (stam. fl),5. Loirrieetal. 5.2 (INPA,
PiPOLY, Cybianthus in Ecuador and Peru 97
MG, NY). Mato Grosso: Near Tabajara, upper Rio Machado, Nov-Dec 193 1 (sram. fl), B.
Krukoff 1577 (A, NY). Rondonia: SaoLourenqo, cassiterite mine, 20 km Nof S. Louren^'o
on Rd. to "A Macisa" Mine, 15 Jul 1979 (scam, fl), C. Calderon et al. 2852 (INPA, US).
BOLIVIA. Pando: W bank of Rio Madeira, 3 km above Abuna, 13 Nov 1968 (stam, fl),
G. Prance et al. 8388 (LPB, MG, INPA, NY).
The "pseudocataphylls" referred to in the description are poorly devel-
oped and do not occur in a regular phyllotactic spiral as is found in subge-
nus Comomyrsine. Likewise, their morphology is essentially that of a leaf arrested
at different stages of development, and as such, do not have a distinctive
morphology.
Northern populations from Nicaragua and Costa Rica corresponding to
the type of C. spectabilis (as Ardhia spectabilis) differ in their smaller flowers
and anther connectives sometimes eglandular, a feature which is more a function
of ecotype then anything else. Eglandular anthers and entire leaves may be
found in very lowland wet forest populations in the Darien of Panama, and
the Choco of Colombia.
The type of Weigeltia multifiora A.C. Sm. represents populations whose
inflorescences are less branched (although the duplicates of the type collec-
tion vary in that regard) and the largest flowers of the species. They, like
many of the Panamanian populations, have entire leaves and are otherwise
inseparable from the type of Weigeltia tnandra Asplund.
The collections of Kayap from Amazonian Peru are referred to this spe-
cies, despite their longer petioles,
VI. Cybianthus subgenus Weigeltia (A. DC.) G. Agostini, Acta Biol. Venez.
10:156. 1980. Weigeltia A. DC, Trans. Linn. Soc. London, Bot. 17:102. 1834.
Cybianthus sect. Weigeltia (A. DC.) Miq. /;/ Mart., Fl. Bras. 10:299- 1856. Type Spe-
cies: Salvadora stirnamensis Spreng. Tent 7. 1828. ^Cybianthus siirinaniensis (Spreng.)
G. Agostini.
Polyaxial dioecious or rarely, monoecious shrubs or small trees. Roots posi-
tively geotropic. Trunk distinguishable, growth dynamics following Rauh's
Architectural Model (in ours), rarely Corner's Model (not in Ecuador or Peru)
sensi/ Halle et al. (1978). Bark grey to beige, thick, vertically fissured. Branchlets
sessile furfuraceous lepidote or rarely, rufous tomentose, often glabrescent
Cataphylls and pseudocataphylls absent. Leaves alternate, rarely subopposite,
with minute sessile rufous furfuraceous lepidote scales abaxially. Inflorescence
a pinnate or bipinnate panicle, rarely a simple raceme; peduncle 0.5-2 cm
long. Flowers 4- rarely 5-merous; calyx valvate, crenate or rarely entire, gla-
brous or rarely glandular-ciliolate, punctations red or black, prominent or
not; corolla rotate to subrotate, the lobes mibricate, glandular-granulose
only at the junction with the corolla tube, the punctations red or black,
conspicuous, or rarely, prominent; stamens and staminodes united to form
an inconspicuous or conspicuous tube, the tube without lobes alternating
98 SiDA 18(1)
with the apically free filaments, the filaments one to three times longer than
the anthers, terete, and recurved proximally, the anthers subglobose, or widely
ovoid, versatile, apically acute to emarginate, basally widely cordate, de-
hiscent by wide longitudinal slits, the connectives mostly prominently red
or black punctate; pistil conic, pyriform or obturbinate, the ovary translu-
cent glandular-lepidote or glabrous, the style capitate-lobate, the lobes entire;
pistillode minutely conic, or at times, absent. Fruit globose or depressed-
globose.
Cyhiantbiis subgenus Weigdtia contains approximately 46 species in South
America and the Caribbean. Five species are known from Ecuador and Peru;
they are restricted to the lowlands and premontane forests on the lower slopes
of the Western Cordillera in Ecuador and the Eastern Cordillera in Ecuador
and Peru.
KF.Y '\X) SPF.CinS Oi; CYBIANTHUS SUBGENUS WEIGELTIA
I . Inflorescence bipinnately paniculate; petioles canaliculate; staminate Mowers
with stamens shorter than corolla.
2. Branchlets terete; leaf blades apically long attenuate-subulate, basally long-
attenuate.
3. Branchlets 2.5-4 mm diam., minutely rufous-lepidote; leaves
pseudoverticillate, the blades membranaceous, prominently black punctate-
lineate below, margins subentire to obtusely serrate; petioles 0.5-1.5
cm long; calyx membranaceous, 1-1.3 mm long, the lobes obtuse, promi
nently punctate, glandular-ciliate along the margin 23. C. poeppigii
3. Branchlets 5.5-6 mm diam., minutely ferrugineous tomentellous;
leaves alternate, the blades chartaceous, minutely and prominently
pLinctictdose below, margins entire; petioles 2—3-5 cm long; calyx
chartaceous, 0.7-1 mm long, the lobes acuminate, epunctate, glabrous
along the margm 24. C. pseudolongifolius
2. Branchlets angulate or winged; leaf blades apically acute or short-acumi
nate, basally acute or cimeate.
4. Branchlets angulate, 8-10 mm diam., conspicuously rubiginous punctate-
lineate; leaf blades coriaceous, conspicuously rubiginous punctate-lineate
below; petioles 2.5-3 cm long; staminate calyx membranaceous, 1.4-
1.6 mm long, the lobes apically acute; staminate corolla membrana-
ceous, 2.3-2.5 mm long; stamens 1.8-2 mm long 23. C. vascjuezii
4. Branchlets vvuiged, (2.5-3-)5-6 mm diam., epunctate; leaf blades
chartaceous, incons|Ticuously pellucid punctate; petioles 2-2.5 cm long;
staminate calyx chartaceous, 1-1.2 mm long, the lobes apically
subacuminate; staminate corolla chartaceous, 1.2-1.3 mm long; sta-
mens l-l.l mm long 26. C. cenepensis
I . Inflorescence racemose; petioles marginate; staminate flowers with stamens
exserred or equalling corolla 27. C. nanayensis
23. Cybianthus poeppigii Mez in Engl., Pflanzenr. IV. 236(Heft 9):218.
1902. Tvpi:: PF.RU. Lokito: Prov. Maynas. Tocache, without elevation, without
date, (stam. (i), /:. Poepp/i^ .\.i/. (ikm.otype: W).
PiPOLY, Cybianthus in Ecuador and Peru 99
Wei^dtiii alhiflora A. C. Sm., Amer. J. Bot. 27:546. 1940, syn. nov. Cyhiarithi/s cilhiflori/s
(A.C. Sm.) G. Agostini, Acta Biol. Venez. 1 0:1 57. 1980. Type: COLOMBIA. Choc6:
Andagoya, 70-110 m, 20-30 Apr 19.39 (stam. fl), E. Ktllip 33372 (hoi.otype: NY;
isoTYPES: A, BM, US-2 sheets).
Cybianthus gentry il^undeW, Wright ia 5:195. 1975, syn. nov. Typi;: COLOMBIA. Ch(x;6:
Cerro Mali', on border with Panama, 1,200-1,400 m, 17 Jan 1975 (stam. fl), A. Gm-
try & S. Mori 13709 (holotype: LL; isotypi;: MO).
Shrub or small tree to 4(-7) m tall. Branchlets thin, terete, 2.5-3.5(-4) mm
diam., minutely rufons-lepidote. Leaves in loose pseudoverticels; blades
membranaceous, elliptic, oblanceolate, lanceolate or rarely obovate, (6— )6.8—
18.5(-24) cm long, (2.2-)3.5-6.5(-8.5) cm wide, apically acuminate to
subacuminate-attenuate, basally cuneate to acute, not decurrent on the petiole,
midrib depressed above, prominently raised below, the secondary veins 7—
1 2(-l4) pairs, rufous lepidote above and below early glabrescent above, tardily
glabrescent below, sparsely black punctate and densely black punctate-lineate
below, the margin flat, subentire to obtusely serrate; petioles canaliculate,
0.5—1.5 mm long, minutely rufous-lepidote. Staminate and pistillate inflorescences
monomorphic, bipinnately paniculate, somewhat pyramidal, sometimes
malformed and appearing racemose, 8-15 cm long, 8-10 cm wide, the ra-
chis densely glandular-papillate, the flowers racemose; inflorescence bracts
unknown; floral bracts membranous, linear-lanceolate, 1— 2.5(— 3-5) mm long,
0.3— 0.8(— 1.2) mm wide, apically attenuate, densely glandular-papillate,
epunctate, the margin glandular-ciliate; pedicels cylindrical, (0.7— )1.6— 1.9(—
2.5) mm long, densely glandular-papillate. Staminate floivers 4-merous, yel-
low or yellowish-green; calyx membranaceous, cotyliform, 0.8— 1 .3 mm long,
the tube 0.2-0.3 mm long, glabrous, the lobes widely ovate to suborbicu-
lar, (().6-)l mm long, (0.5-)0.6-l mm wide, apically obtuse, prominently
punctate, the margin hyaline, irregular, erose-serrtilate apically, densely glandular-
ciliate; corolla membranaceous, rotate, (1.7-)1.8-2.2(-2.6) mm long, the
tube (0.3-)0.4-0.6 mm long, glabrous without, glandular-granulose within,
the lobes suborbicular, (0.9-)l .2-1 .6(-2) mm long, (0.8-)l .2-1.7(-2) mm
wide, apically obtuse, prominently punctate, glabrous without, glandular-
granulose medially at stamen base within, the margin hyaline, irregular,
erose; stamens (1-)1.4— 1.6 mm long, the tube inconspicuous, hyaline, (0.3-)
0.4—0.6 mm long, densely glandlar-granulose within, the apically free por-
tions of the filaments (0.3-)0.4-0.5 mm long, the anthers ovate-triangu-
lar, 0.3-0.4 mm long, 0.2-0.3 mm wide, apically rounded, basally obtuse,
the connective epunctate; pistillode come, 0.3-0.4 mm long, densely ru-
fous glandular-papillate. Pistillate flowers as in staminate, but calyx 1—1.3
mm long, the tube 0.2-0.3 mm long, the lobes deltate to widely elliptic,
0.8-1 mm long and wide, the margin opaque; corolla 2-2.3 mm long, the
tube, 0.6-0.7 mm long, the lobes 1.4-1.6 mm long, 0.6-0.7 mm wide,
100 SiDA lcS(l)
glandular-granulose within above filament junction with tube, the margin
often erose; staminodes resembling stamens but 1-1.3 mm long, the staminodial
tube ca. 0.6 mm long, the apically free portions of the filaments to 0.3 mm
long, the antherodes suborbicular, to 0.2 mm long and wide; pistil obnapiform,
1.2—1 .3 mm long, the ovary 0.8—0.9 rnm long, 0.3—0.5 mm wide, densely
translucent-lepidote, the style 0.3—0.4 mm long, the stigma bilobed, the
lobes decurrent, ca. 0.4 mm long. Fruit depressed-globose, 3.5-4.5 mm
long, 4.5—6 mm diam. when dried, red, then black at maturity, inconspicuously
punctate, exocarp thin.
Distribj/iton. — Known from the easternmost Darien of Panama south to
Amazonian Peru and Brazil, 100-1,800 m.
Ecology and conservation status. — Cybianthus poeppigii is a broad ranging
polymorphic ochlospecies, occurring in primary premontane wet and plu-
vial forests, from the transition zone with lowland forests, to the transition
zone to cloud forests. Recent fieldwork in the Cordillera Occidental of Co-
lombia has shown it is a conspicuous element of primary pltivial premontane
forests, with a density of approximately 20 individuals per hectare, clus-
tered mostly along the matgin of the windward side of the forest, and along
streambanks above the high water level. While locally common, its restricted
primary habitat verifies its threatened status.
Etymology. — The specific epithet honors Eduard F. Poeppig (1 798-1868),
professor at Leipzig, explorer and plant collector, who made numerous valuable
contributions to our knowledge of the Peruvian Amazon Basin.
Representativf specimens examined. PANAMA. Darien: Scrrani'a del Darien, Panama/
Colombia frontier, Cerro Tacaracuna, Cerro Mali, summit, 1,400 m, 17 Jan 1975 (sram.
fl), A. Gentry &S. Mon 13665 (LL-TEX, MO-2 sheets), W ridge, Cerro Tacaracuna, 1,800-
1 ,850 m, 3 1 Jan 1 975 (Ir), A. Gentry &S. Mon 1 4023 (COL, LL-TEX, MO-2 sheets); Serranfa
de Pirre, Cerro Pirre, above Cana Gold Mine between Ri'os Cana and Escucha Ruido, 1 ,()()()-
1,310 m, 21 Jul 1976 (stam. fl), T. Croat J 7785 (LL-TEX, MO, NY, PMA), SW ridge
leading to Alturas de Nique, Panama/Colombia border, 1 , 100-1 ,200 m, 30 Dec 1980 (stam.
fl), R. Hiirt)iki>i 12401. 12461 (MO). COLOMBIA. Antiociuia: Mpio. De Anon', Corregimiento
Providencia, Rfo Anori Valley, between Dos Bocas and Anorf, •100-900 m, 24-31 May
1973 Or), D. Soejc/rto et ul. -f()9(} (A, COL, F, LIUA, MO); Mjmo. Sonson, Rfo Verde region,
Hacienda "La Soledad," 1 ,430- 1 ,800 m, 2 1 Jan 1 947 (stam . fl ), G, G//t/envz 1 1 86 (E, MEDEL,
MO, LJC); Mpio. San (;arl()s, Corregimiento Alto de Samana, Vereda Miraflores, 820-900
m, 15 Jun 1989 (stam. fl), R. fomiei^rj et ul. 30^6 (BRIT, IILJA). Mpio. Urrao, Parque
Nacional Natural Las Orquitleas, Vereda Calles, Permanent Inventory, right bank, Rfo Calles,
06° 32' N, 76° 19' W, 1 ,450 m, 27 Nov 1993 (ster.),/ P,poly etui HI 83 (BRIT, JAUM,
MO), 1,450-1,500 m, 29 Nov 1993 (ster. seedling),/ Pipoly et al. / 7360 (BRIT, JAUM,
MO). Choco: Mpio. San Jose del Palmar, vereda "El Tabor," 1, 540 m, 18 Jan 1983 (pist.
fl), P. Prcinco et al. 1469 (COL). Vaupes: Rfo Pacoa (tributary of Rfo Apaporis), 00° 20' N,
71° 20' W, 300 m, 7-12 Feb 1952 (pisr. fl), R. Schi/ltes & I. Cabrera 1^423 (COL, GH,
US). ECUADOR. Morona-Santlago: Macuma, 50 km NofMacas, 21 Mar 1973 (pist. fl),
H. Ugo 3633 (GB); Rfo Cuycs and Boboiza-Gualaquiza Rd., 03° 25' S, 78° 35' W, 800 m,
1 Nov I 986 (stam. fl), U^ Pataaos ! 466 (MO, US). Napo: Between Tena and Napo. 1 Jan
PiPOLY, Cybianchus in Ecuador and Peru 101
1940 (stam. fl), U. Aspli/rid 10209 (S), 7 Jan 19i0 (stam. fl), E. Aspl/nid 10302 (S); Tena,
400 m, 3 Apr 1935 (stam. fl), Y. Alexia 7206 (NY, UC, US); ReservaBiologicaJatun Sacha,
8 km from Puerto Misahuallo, right bank, Rio Napo, 01° 04' S, 77° 36' W, 450 m, C.
Cerdn2585 (MO,QCNE). Pastaza: Mera, 1 ,100 m, (stam. fl), E. Asplund 18717(5), 1,500
m 29 Dec 1958 (stam. fl), G. Hurling et al. 9764 (GB); Loracachi, on path to Lagartococha,
01° 38' S, 75° 58' W, 25 May 1980 (fr) J. Jaramillo et al. 30984 (AAU, QCA), 31 May
1980 (tr), 3 I -5 79 (AAU, QCA). Tungurahua: Ri'o Negro, 1 ,200 m, 1 3 Mar 1980 (pist. fl),
G. Marling & L. Andersson 17255 (GB); 10 km E of Paquisha, 1,400-1,500 m, 13 Apr
1985 (stam. fl), G. Marling & L. Andersson 24097 (GB). PERU. Amazonas: Rio Santiago,
3 km behind Caterpiza, 14 Nov 1 979 (fr), V. Muashikat 1248 (MO, NY); Tingo Maria, 25
Jul 1940 (stam. fl), E. Asplund 12497 (S). Pasco: Prov. Oaxapampa, Cabeza de Mono, Palcazu
Valley, 10° 20' S, 75° 18' W, 320 m, 1 1 Jun 1983 (fr), A. Gentry et al. 41880 (MO); Ozuz
to Rio Lobo, 10° 19' S, 75° 16' W, 400-500 m, 10 May 19H5 (fr), R. Foster & B. d'Achilk
10013 (F). San Martin: Quebrada de Haquisha (right margin Rio Hualiaga), Prov. Mariscal
Caceres, Dtto. Tochache Nuevo, 400-500 m, 1 Jul 1974 (stam. fl),^. Schunke 7106 (MO,
NY). BRAZIL. Amazonas: Near Jurua, Dec 1900 (stam. fl), E. Ule 5160 (HBG).
Cylnanthus poeppigit was mistakenly placed by Agostini (1980) in subge-
nus Cybianthus, but the versatile, dorsifixed anthers clearly place it in sub-
genus Weigeltia. I had reported earlier (Pipoly 1983a) that Cyhtanthus albiflorus ,
its synonym, was closely related to C. lawrencei Moldenke. However, the
bipinnate inflorescences with racemose flowers, stamens shorter than the
corolla and usually long petioles indicate that Cyhianthm poeppigii is more
closey related to C. longtfolius Miq., a vicariant species of the southwestern
Amazon Basin of Brazil and adjacent Bolivia. The glandular grantiles of
the staminal and staminodial tubes represent a unique (autapomorphic) character
state within the genus.
The holotype oi Cylnanthus poeppigii is staminate, as are those of its taxo-
nomic synonyms. The type of Weigeltia albiflora {Cybianthus albiflonts) rep-
resents populations with entire, irregularly margined leaves, but is other-
wise qualitatively identical with that of C. poeppigi. Likewise, the type of
Cybianthus gentryi Lundell represents isolated montane populations of the
Darien/Choco regions, and exhibits more notable lineate-punctations, much
smaller leaves, and abbreviated inflorescences. According to annotations by
Killip at US, description of another taxon was at one time contemplated,
based on the fact that the populations of this species from near Tena, Ecua-
dor, have roughly serrate leaf margins.
24. Cybianthus pseudolongifolius Pipoly, sp. nov. (Fig. 18). Type: PERU.
Pasco: Prov. Oxapampa, Palcazu Valley, Cabeza de Mono, 5-6 km W of Iscosacfn
10° 12' S, 75° 14' W, 325 m, 13-19 Apr 1983 (fr), D. Smith 3808 (holotype; MO;
isoTYPEs: US, USM).
Quoad folia magna chartacea, inflorescentias bipinnatipaniculatas, j^ediceilos cylindricos,
lobos calycinis ovatos, ca. 1/3 connatos, petiolos canaliculatos C. longijolio arete accedens, sed
abearamuiis teretibus (non angulatis), 5.5-6 (non 3.5-4) mm diammetris, laminis anguste
oblanceolatis (non ellipticis vel lanceolatis) desuper sordidis (non nitidis) ad apices longi-
102
SlDA 18(1)
Fic. 18. Lybicnithns p\i:iid(ili)iigijtil}iis Pipoly. A. H;ibi:, showing bipinnate panicles. B. Pe-
duncle and axillant leaf, showmg canaliculate petiole. C;. Detail of prominently puncticulosc
abaxial leal surtace. D. Pedicel, calyx and fruit, showing hyaline and erose calyx lobes. A &
C, drawn from isotype; B ik. D, drawn From holotype. Figure drawn by Linda Ellis.
PrpoLY, Cybianthus in Ecuador and Peru 103
attenuacis ct subulatis (nee acutis vel atuminatis) ad bases longi-attenuatis (nee aeutis),
inflorescentiis 8-13 (non 15-20)cni longis, pedicellis 2.8-3 (nee 0.6-1) mm longis necnon
fructibus depresso-globosis (nee globosis) acque minute cosatis (nee laevibuss) perfacile diseenda.
Treelet to 3 m tall. Branchlets terete, 5.5-6 mm diam., densely and mi-
nutely ferrugineous tomentellous. Leaves alternate; blades chartaceous, nar-
rowly oblanceolate, (22-)26.5-33(-36.5) cm long, (5-)6.5-8.5 cm wide,
apically long attenuate, subulate, basally long-attenuate, decurrent on the
petiole, sordid and glabrous above, pallid, minutely and prominently pel-
lucid puncticulose and minutely ferrugineous puberulent below, midrib slightly
raised above, prominently raised below, the secondary veins 13-18 pairs,
the margin entire, glabrous, flat; petioles canaliculate, (2-)2.5-3(-3.5) cm
long, glabrous above, minutely ferrugineous puberulent below. Staminate
inflorescence: unknown. Pistillate inflorescence: a lax bipinnate panicle, 8—13
cm long, 1.5-4 cm wide, densely feraigineous puberulent, glabrescent; secondary
inflorescence bracts unknown; floral bracts unknown; pedicels cylindrical,
2.8—3 mm long, densely ferrugineous puberulent; Pistillate flowers unknown;
fruiting calyx chartaceous, cotyliform, 0.7-1 mm long, the tube 0.3-0.4
mm long, the lobes ovate, 0.5-0.7 mm long, 0.5-0.6 mm wide, apically
acuminate, densely and prominently red punctate, the margin hyaline, erose,
epunctate, glabrous. Fr///7 depressed-globose, 3—4 mm long, 5-6 mm diam.,
inconspicuously pellucid punctate, minutely longitudinally costate.
Distribution. — Known only from the type.
Ecology and conservation status. — Cybianthus pseudolongifolius appears to be
restricted to the lowland primary forest of the eastern Andean slopes. The
Oxapampa Province of Pasco is home to numerous Peruvian endemics, and
it would not be surprising if the species was of extremely limited distribu-
tion or endemic. With only one specimen known, no determination can be
made of its true conservation status.
Etymology. — The specific epithet refers to its general likeness to Cybianthus
longifolii/s Miq., a lowland black water river species from Amazonian Brazil
and Venezuela.
Cybianthus pseudolongifolius is most closely related to C. longifolius, but
differs by its terete branchlets, 5.5-6 mm in diameter, narrowly oblanceolate
leaf blades that are sordid above, long-attenuate and subulate apically and
long-attenuate basally, longer inflorescences, much longer pedicels and depressed-
globose, minutely costate fruits. The Oxapampa region of Pasco contains a
number of endemic species and disjunct taxa and as such, is one of the most
important underexplored areas in Peru.
25. Cybianthus vasquezii Pipoly, sp. nov. (Fig. 19). Type: PERU. Loreto:
Prov. Alto Amazonas, Capahuari Norte, 02° 45' S, 76° 25' W, 220 m, 7 Jun 1981
(stam. fl), R. Vdsciiiez &N.Jc!nimillo 1993 (hoi.otype: MO; isotypes: AMAZ, BRIT,
F, NY, US, USM).
104
SiDA 18(1)
Fic. 19. Cyb'uinthus vasquezii Pipoly. A. Habit, showing angulate, punctate-lineate branciilet.
B. Inflorescence branch, showing racemose-glomerulatc floral arrangement. C. Open co-
rolla. D. Abaxial calyx lobe surface. E. Abaxial leaFsurface, showing minute furufuraceous
scales and prominent punctate-lineations. R Branchlet apex. A-F, drawn by Linda HUis,
from holotype.
PiPOLY, Cybianrhus in Ecuador and Peru 105
Propter ramulos crtissos manifesce angulatos, fol ia coriacea subter pallida ad apices subaciiminata
ad bases acuta, petioles canaliculatos, inflorescenrias anguste bipinnatipaniculatas, ramulos
inflorescentiares dense spicatos vel glomerulatos lobos calycines grosse crenatos C. potiaeo
valde affinis sed ab ea ramulis aibiginoso-puncrato-lineatis (non epunctatis), folia pseudoverticillata
(non alterna), laminis oblanceolatis (nee ellipticis), (9— )10— 13(— 1 5.3) (nee 6.5—8) cm latis,
petiolis 2.5—3 (nee 1.5—2) cm longi.s, inflorescentiis 12—25 (non 3—6) cm longis, corollae
lobis ovatis (non ellipticis) ad apices acute rottmdatis (nee emarginatis) ad bases abrupte
constrictis (nee rectis), secus margines gros.sc crenatis (nee integerrimis) confeste separabilis.
Terrestrial dioecious tree to 8 m tall. Branchlets angulate, 8—10 mm diam.,
densely and minutely rubiginous furfuraceous-lepidote, conspicuously ru-
biginous punctate-lineate below. Leaves pseudoverticillate; blades coriaceous,
oblanceolate, (21— )26— 34.5 cm long, (6.5-)9— 12.2 cm wide, apically acute
to subacuminate, basally acute, decurrent on the petiole 4—7 mm, glabrous
above, very minutely rubiginous furfuraceous-lepidote below, the midrib
slightly raised above, prominently raised and rubiginous punctate-lineate
below, the secondary veins 7-10 pairs, slightly impressed above, promi-
nently raised below, the margin entire, flat; petioles canaliculate, 2.5—3 cm
long, swollen below to 0.5—0.7 cm diam. basally, rubiginous furfuraceous-
lepidote at first, early glabrescent, conspicuously rubiginous punctate-lineate.
Staminate inflorescence: a bipinnate panicle (12— )14— 17.5(— 25) cm long, 2—3
cm wide; peduncle (l-)2— 3.5 cm long; secondary inflorescence bracts chartaceous,
linear, 2—3 mm long, 0.3—0.6 mm wide, apically attenuate, densely glan-
dular-papillate, the margin entire; branches racemose-glomerulate (0.5— )1 —
1.5 cm long; floral bracts membranaceous, linear, 1.6—1.8 mm long, 0.3—
0.4 mm wide, apically attenuate, densely rubiginous puberulent; pedicels
cylindrical, 0.3—0.5 mm long, glabrescent. Staminate flowers 4-merous, pink;
calyx membranaceous, cotyliform, 1.4—1.6 mm long, the tube ca. 0.2 mm
long, the lobes ovate, 1.2—1.4 mm long, 1—1.1 mm wide, apically acute,
sparsely and inconspicuously orange punctate, glabrous, the margin coarsely
crenulate, glabrous; corolla membranaceous, subrotate, 2.3—2.5 mm long,
the tube 0.2—0.3 mm long, densely glandular-granulose, the lobes ovate,
2-2.3 mm long, 1 .7—1 .8 mm wide, apically acutely rounded, abruptly constricted
basally, inconspicuously orange punctate without, sparsely gladular-granulose
behind the base of the filaments, the margin hyaline, coarsely crenate, gla-
brous; stamens 1.8—2 mm long, the tube carnose, squarrose, 0.2—0.3 mm
long, the filaments terete, 1.1—1.2 mm long, slightly curved proximally,
the anthers ovate, 0.5—0.6 mm long, 0.6—0.7 mm wide, apically rounded,
basally cordate, the connectives inconspicuously orange punctate dorsally;
pistillode obturbinate, 0.5 mm long, 0.6 mm diam., the stigma truncate,
minutely lobed. Pistillate inflorescence as in staminate but (3.5-)5.5-9-5 cm
long, the branches glomerulate. Pistillate flowers as in staminate but red,
except white on corolla lobe apices; calyx obconic, 1.3—1-7 mm long, the
tube 0.6—0.8 mm long, the lobes very widely ovate to suborbicular, 0.7—
106 SiDA 18(1)
1.2 mm long, 0.8—1.2 mm wide, apically broadly rounded, corolla 2.3—2.7
mm long, rhc cube ().6-{).8 mm long, the lobes suborbicular, 1.5—1.7 mm
long, 1.5—1.8 mm wide, the margin irregtilar, hyaline, staminodial tube
1-1.3 mm long, the apically free portions of the filaments 0.4-0.5 mm long,
the antherodes 0.3—0.4 mm long and wide; pistil obnapiform, 2.2—2.4 mm
long, 1.3—1.5 mm diam, the ovary 1.3—1.5 mm long, the style thick, 1.1—
1 .3 mm long, the stigma capitate, 4-lobed, the lobes recurved, the placenta
deeply cuptililorm, the ovtdes 2—3, imbedded. Fr//it unknown.
Distribution. — Known only fi'om the Department of Loreto, in Alto Amazonas
and Loreto Provinces, Peru, at 160—220 m elevation.
Ecology and amservci turn stc/tns, — Cybianthiis I'c/sqi/ezii occurs in primary terra
jirnie lowland forest, with scattered white sand areas. Label data do not al-
low determination of whether this species occurs on the laterite or on the
sands. Because it is known only from two gatherings, its conservation sta-
tus is unknown. However, its importance as a tonic in Mayna Jivaro cul-
ture may indicate it is locally common.
Etymology. — It gives me great pleasure to dedicate this species to a great
friend and colleague, Ing. Rodolfo Vasquez Martinez, Assistant Curator and
Director of the Flora of Peru Program of the Missouri Botanical Garden.
Vasquez is an indefatigable collector, a forestry engineer, dendrologist, taxonomist
and author of numerous publications on uses of Peruvian forest products,
economic plants of the Peruvian Amazon, and Flon/la of the Biological Re-
serves of hinitos. He is a taxonomic authority on Caraipa and Myristicaceae
of the Amazon Basin.
Local names and uses. — Peru: "sesa," "kurup" (Mayna Jfvaro). The sap is
extracted and a juice is drunk to "improve hunting ability."
PARATYPii: PERU. Loreto: Prov. Loreto, Pampa Hermosa and vicinity, Rio Corrientes, 1
km S of junction with Ri'o Macusan, 03° 15' S, 75° 50' W, 160 m, 3-20 Dec 1985 (pist.
fl), W. Lcins et at. 10306 (BRIT, MO).
Cybianthits vascyiezii is most closely related to C potiaei c^f the eastern Amazon
Basin (French Gtiiana and Brazil (Amapa, Bahfa), but is easily recognized
by the conspicuously rubiginous punctate-lineate branchlets, larger
pseudoverticillate, oblanceolate leaves, longer petioles and inflorescences,
ovate, apically rounded corolla lobes that are abruptly constricted basal ly,
and coarsely crenate along the margins.
26. Cybianthus cenepensis Pipoly, sp. nov. (Fig. 20). Tyi^i : PERU. Amazonas:
Ri'o Cenepa, vicinity Huampami, ca. 5 km E of (Chavez Valdi'via, 04° 30' S, 78° 30
W, 200-250 m, 1 2 Aug 1978 (stam. fl),/\, K/ijikcit 263 (iiolotyph: MO; isotypi.s: F,
MO, NY, US, USM).
Ob folia chartacea oblanceolata atl apices acummara ad bases ciineata, petiolos canaiicnatos,
inHorescentias angnste bipinnatipaniciilatas, ad C. h/icbtieiii valde atfinis sed ab ea ramnlis
alatis (nee laevibtis), pctiolis 2-2.5 (non 1 .5—1 .8) cm longis, lobis calycinis late ovatis (ncc
PiPOi.'i', Cybianthus in Ecuador and PerLi
107
2mm
Fig. 20. Cyhianth/ts cmepensis Pipoly. A. Habit, showing winged branchlers. B, Sraminace
inflorescence branch, showing secondary inflorescence branch bracts. C. Staminate flower,
showing stamens subeqtial to corolla lobes, conspicuous square staminal tube, and coarsely
crenate corolla lobes. D. Pistillate flower, showing ellipsoid pistil. E. Pistillate inflorescence
branch, showing dense spike appearing glomerulate. D. Branchlet apex, showing puberu-
lent vestiture. A drawn from holotype. B— D, drawn from Auc/uish 522. E— F, drawn from
Kujikat 306. Figure drawn by Linda Ellis.
108 SiDA 18(1)
lineari-lanccolatis), grossc crenaris (ncc inte^'errimis), lobis corollinis grosse crcnatis (nee
enccris) clenicjue antlieris ad a|-iices acLitis (nee roninclatis) atl bases eordatis (nee obtLisis)
perhieile clisringuitLir.
Treelet to 2 m tall. Branchkts prominently longitudinally ridged, the ridges
forming small, rounded wings, (2.5-3-)5-6 mm diam., sparsely rufous
puberulent, glabrescent. Leaves alternate; blades chartaceous, oblanceolate,
(1 3.7— )18.5-23(— 26.2) cm long, (4— )6-10 cm wide, apically acuminate,
basally cuneate, glabrous above, sparsely rufous puberulent below, glabres-
cent, inconspicuously pellucid punctate, the margin entire, flat; petioles
canaliculate, 2—2.3 cm long, sparsely rufous puberulent, glabrescent. Staviinate
inflorescence a lax bipinnate panicle, 11—1 8 cm long, 1—1.7 cm wide, the rachis
densely rufous papillate; secondary inflorescence bracts chartaceous, linear-
lanceolate, 4—4.5 mm long, 1—1.2 mm wide, apically subulate, densely rufous
tomentellotis, the margin irregular, entire; inflorescence branches 3-8 mm
long, the flowers densely subspicate, appearing glomerulate; floral bracts
chartaceous, linear, 1 — 1.2 mm long, 0.1— 0.2 mm wide, apically subulate,
densely glandular-papillate, the margin crenulate, glabrous; pedicels cylindrical,
0.2—0.3 mm long, densely glandular-papillate. Staminate flowers A-mtrons,
chartaceous, brownish-purple, 1.6—1.8 mm long; calyx cotyliform, 1—1.2
mm long, the ttibe 0.3—0.4 mm long, the lobes broadly ovate, 0.7— 0.8 mm
long, 0.6—0.7 mm wide, apically acute, prominently brown punctate, the
margin hyaline, coarsely crenate, glabrous; corolla subrotate, 1.2—1.3 mm
long, the tube ca. 0.1 mm long, densely glanclular-granulose within, the
lobes widely ovate, 1 — 1 .2 mm long, 0.8—1 mm wide, apically subacuminate,
glabrous and inconspicuously orange punctate without, glabrous within except
under the filaments, the margin hyaline, coarsely crenate, glabrous; stamens
subequalling the corolla lobes, 1 — 1.1 mm long, the tube conspicuous, carnose,
square, 0.1-0.2 mm long, densely glandular-granulose, the filaments 0.6—
0.7 mm long, the anthers ovate, 0.3—0.4 mm long, 0.4—0.5 mm wide, apically
acute, basally cordate, the connective inconspicuously orange punctate; pistillode
obturbinate, 0.1—0.2 mm long, 0.2—0.3 mm diam., hollow, glabrous. Pis-
tillate inflorescence diS in staminate but 12—14 cm long, 0.8—1 .0 cm wide; secondary
inflorescence bracts 3.8—4. 1 mm long, 0.8—1 mm wide; inflorescence branches
3.5—6 mm long; floral bracts 1 — 1.2 mm long, 0.1—0.2 mm wide; pedicels
0.2—0.3 mm long. Pistillate flowers as in staminate but white, 1.1 — 1.4 mm
long; calyx 0.6—0.8 mm long, the tube 0.1—0.2 mm long, the lobes 0.5—
0.6 mm long, 0.3—0.4 mm wide; corolla 0.7—0.8 mm long, the tube ca.
0.1 mm long, the lobes 0.6— 0.7 mm long, 0,5—0.6 mm wide; staminodes
much sorter than the corolla lobes, 0.3—0.4 mm long, the tube conspicu-
ous, carnose, circular, ca. 0. 1 mm long, the filaments ca. 0.1 mm long, the
anthers 0.2—0.3 mm long, 0. 1—0.2 mm wide; pistil ellipsoid, 0.5—0.6 mm
long, 0.2—0.3 mm diam., the style 4-lobed, the lobes curved distally, pla-
PiFOLY, Cybianthus in Ecuador and Peru 109
ccnta subobose, bearing 4 partially immersed ovules. Fruit globose, 6—7
mm long and in diam., prominently red punctate.
Distribution. — Known only from the Rfo Cenepa Drainage Basin, Amazonas,
Peru, 200—250 m elevation, and from one disjimct population from Napo,
Ecuador, very close to the Colombian border.
Ecology and conservation status. — Cybianthus cenepensts inhabits varzea for-
ests in the Amazon of Ecuador and Peru. Despite lack of collections, it is
known that the Napo and the Cenepa Rivers are shallower than many with
their same volume, thus giving them strong currents. Aside from the need
for much more exploration at the edge of the Amazon Basin in Ecuador and
Peru, perhaps the strong current of these rivers makes the dynamics of their
adjacent varzea forests different from others in Amazonia. At this time, the
conservation status of this species is unknown.
Etymology. — The specific epithet refers to the region of Peru from which
the type was collected, the Rio Cenepa Drainage Basin of the Alto Amazonas
Province, Amazonas Department, Peru.
Local names and uses. — Peru: "sauka" (Huambisa).
Paratypes. ECUADOR. Napo: Canton Orellana, Yasuni National Park, Maxus Rd. and
pipeline construction project, km 15. 01° 3 I' S, 76° 32' W, 250 m, 30Jun 1994 (bud), N.
Pitman 461 (BRIT, MO, QCNE); Rio Yasunf, periodically inundated forest ca. 80 km upriver
from Nuevo Rocaluerte, 225 m, 17 Sep 1 977 (ft), R. Foster 3708 (F, QCA). PERU. Amazonas:
Quebrada Sasa, Rio Cenepa, 250 m, 2 Jun 1973 (stam. fl), E. Ancuasb 522 (MO, USM);
Ri'o Cenepa, vicinity of Huampami, ca. 5 km E of Chavez Valdivia, ca. 4° 30' S, 78° 30' W,
200-250 m, 12 Aug 1978 (pist. fl), A Kupkat 306 (F, MO, USM); Quebrada Chigki Shiunk,
4° 30' S, 78° 30' W, 11 Aug 1978 (bud), £, Amtiash 1412 (MO, USM).
Cybianthus cenepensis is closely related to C. buchtieni Pax of the Mapiri
region in Bolivia. However, Cybianthus ce?iepensis is easily recognized by its
winged branchlets, petioles 2—2.5 cm long, widely ovate and coarsely crenate
calyx lobes, coarsely crenate corolla lobes, and anthers with acute apices and
cordate bases. The ellipsoid pistil is also unique within the subgenus.
27. Cybianthus nanayensis (J.F. Macbr.) G. Agostini, Acta Biol. Venez.
10:l60. 1980. Weigeltia nanayensis ].V. Macbr., Field Mus. Nat. Hist., Bot. Set.
11:33. 1 931. Type: PERU. LORETO: Lower Rfo Nanay, 4 Jun 1929 (stam. fl),L/. Williams
658 (holotype: F; fragment, G).
Weigeltia silvestris ].¥. Macbr., Candollea 6; 16. 1934. syn. nov. Cybianthus silfestris (J.F.
Macbr.) G. Agostini, Acta. Biol. Venez. 10:163. 1980. Type: PERU. Loreto: Mishuyacu,
near Iquitos, Dec 1929 (stam. fl), G. King 724 (holotype: F; isotypes: NY, US).
Cononiorpha d/ibia } .V. Macbr., Candollea 6: 17. 1934. syn. nov. Cybianthus dubius (^ .V. Macbr.)
G. Agostini, Acta Biol. Venez. 10: 158. 1980. Type: PERU. Loreto: Mishuyacu, near
Iquitos, 24-28 Sep 1929 (pist. fl, fr), A. Killili & A. Smith 29906 (holotype: F; isotypes:
NY, US) [erroneously cited as G. King 299(^6].
Conomorpha loretensis Lundell, Wrightia 6:113. 1 980. syn. nov. Cybianthus loretensis (Lundell)
Pipoly, Brittonia 33:496. 1981. Type: PERU. Loreto: Quistococha, near Iquitos, 18
Nov 1977 (stam. fl), A. Gentry 20763 (holotype: LL-TEX; isotypes: F, MO, NY).
1 10 SiDA 18(1)
S//hshr//h to shrub to ().5(— 1.5) m tall. Braiichlets 0.2—0.5 cm thick, smooth,
rufoLis-lepidote. Leai'es in loose pseudoverticels; blades chartaceous to sub-
coriaceous, elliptic, lanceolate or oblanceolate, (7— )1 l-18(-25.5) cm long,
(2.9~)3.5— 7.5(— 10) cm, apically acute to acuminate, basally acute to cu-
neate, midrib depressed above, prominently raised below, the secondary veins
7—16 pairs, glabrous above, sparsely and minutely rutous-lepidote below,
puncticulate, the margin flat, entire; petioles marginate, (0.6— )().9— 1.5 cm
long, glabrous. Inflorescences monomorphic, a simple raceme or rarely with a
second, malformed basal branch, (1 .5— )3— 9-5(— 14.5) cm long, the rachis
densely glandular-papillate; floral bracts membranaceous, linear-lanceolate,
0.2—1.2 mm long, 0. 1—0.2 mm wide, the apically long-attenuate, sparsely
glandular-papillate, the margin glandular-ciliate, entire; pedicels cylindri-
cal, (0.7— )1— 1 .5(— 2) mm long, densely glandular-papillate. Stan/inate flowers:
white to cream, 4-merous, chartaceous; calyx cotyliform, 1—1.3 mm long,
the tube ca. 0.1 mm long, the lobes widely ovate to deltate, 0.9—1.2 mm
long, 0.8—1.2 mm wide, apically acute to actnninate, medially thickened
and prominently black punctate, glabrous, the margin hyaline, irregular,
densely glandular-ciliolate; corolla rotate, 1.9—2.3 mm long, the tube 0.6—
0.8 mm long, the lobes widely ovate to ovate, 1.3—1-6 mm long, 1.2—1.4
mm wide, apically acute to rounded, subentire or apically notched, medi-
ally thickened and prominently black punctate, glabrous without, glandu-
lar-granulose within at junction of tube and lobe, the margin hyaline, ir-
regular, glabrous; stamens 2.2—2.5 mm long, exserted or rarely subequal to
corolla, the staminal tube carnose, conspicuous, 0.5—0.7 mm long, subtruncate
between the filaments, the apically free portions of the filaments terete, (1 .2—)
1.4—2 mm long, the anthers widely ovate, 0.32-0.36 mm long, apically
rotmded to obttise, basally cordulate, the connective prominently punctate
dorsally; pistillode vestigial, obclavate or tubiform, 0.7—0.8 mm long, sparsely
translucent glandular-lepidote, hollow. Pistillate flowers as in staminate but
beige to brown; calyx 2—2. 1 mm long, the tube 0.1—0.2 mm long, the lobes
1.9—2 mm long, 1.9—2 mm wide; corolla cotyliform, 1.9-2 mm long, the
tube 0.5—0.6 mm long, the lobes widely ovate, 1.2—1.4 mm long, 1.3—1.4
mm wide, apically acute to acuminate; staminodes resembling stamens, 1.2—
1.3 mm long, the staminodial tube 0.5—0.6 mm long, the filaments 0.4—
0.5 mm long, the antherodes subquadrate, ca. 0.2 mm long and wide, apically
obtuse, basally truncate; pistil ellipsoid, 1.4—1.6 mm long, the ovary 1 —
1.2 mm long, 1.1—1.2 mm diam., densely translucent glandular-lepidote,
the placenta cupuliform, ovules 2, partially immersed, the style trunctae,
0.3—0.4 mm long, the stigma punctiform. Fr//it depressed-globose, 4.5—
5.5 mm long, 5.5— 6.5(— 7) mm diam., yellow when fresh, prominently black
punctate.
PiPOi.Y, Cybianthus in Ecuador and Peru 111
Distribution. — Endemic to the tall moist forests on white sands of the
Peruvian Amazon, primarily from the Iquitos area, 100—160 m.
Ecology and conservation status. — Cybianthus nanayensis is locally common,
and thrives in gaps left by large treefalls in overmature forests, and along
the margins of forest margins and paths, where it occurs in a rather dense
herbaceous layer. However, it does not tolerate compacted soils. With de-
creasing quantitites of habitat owing to logging pressures, the species should
be considered threatened. As a gap species growing on nutrient deficient
soils, and with very attractive fruits, Cybianthus nanayensis shows great promise
as a potentially marketable horticultural plant.
Etymology. — The epithet takes its name from the river basin where it occurs,
the Nanay River.
Representative specimens examined. PERU. Loreto: Prov. Loreco, Nauta, 0-4°32' S, 73°35'
W, 160 m, 2 Jun 1984 (stam. fl), R. Vdsquez & N.Jaramillo 3073 (AMAZ, MO, NY), (fr),
R. Vasquez & N.Jaramitlo 3086 (AMAZ, MO); Prov. Maynas, Allpahuayo, IlAP Experi-
mental Station, 04° 10' S, 73° 30' W, 120 m, 20 Sep 1990 (fr),J. Pipolyetal. 12263 (AMAZ,
MO, USM), 15 Aug 1990 (fr), R. Vasquez & N.Jaramillo 14204 (AMAZ, MO, US, USM),
1 Oct 1990 (stam. fl), R. Vasquez & N.Jaramillo 14463 (AMAZ, MO); Laguna Quistococha,
1 5 km SW of Iquitos, 8 Jul 1977 (stam. fl),J. Solomon 3466 (LL-TEX, MO); Mjshiana, 30
km SW of Iquitos, Callicebus Biological Reserve, Ri'o Nanay, 4 km S of Mishana, 19 Aug
1978 (stam. fl), R. Foster 4243 (AMAZ, F), 16 Aug ] 980 (stam. fl), 4327 (F-2 sheets); Vicinity
Mishana, between Rfo Nanay and Rfo Itaya, 130 m, 29 Nov 1977 (fr), A. Gentry et al.
21033 (F, MO); Between Iquitos and Sta. Maria de Nanay, 180 m, 31 May 1978 (stam. fl),
A. Gentry et al. 22367 (AMAZ, F, MO); 03° 50' S, 73° 30' W, 25 Feb 1981 (fr), A. Gentry
et el. 31479 (AMAZ, MO), 31 Dec. 1 982 (mixed- stam. fl, fr), A. Gentry & L. Emmons 38776
(MO-2 sheets), 5 Aug 1990 (fr), R. Vasquez el al. 14161 (AMAZ, MO, USM); Rfo Nanay,
May-Jun 1929 (stam. fl), LI. Williams 637 (F); Mishuyacu, near Iquitos, 100 m, Oct-Nov
1929 (stam. fl), G. Klug 304 (F), May-Jun 1930 (stam. fl), G. Klug 1335 (F); 6 Oct. 1982
(stam. bud), R. Vasquez & N.Jaramillo 3261 (MO), 20 Jan 1985 (stam. fl), 6123 (MO);
Puerto. Almendras, 03° 48' S, 73° 25' W, 122 m, 4 Jan. 1986 (pist. fl), R. Vasquez & N.
Jaramdlo 7070 (AMAZ, MO), 30 May (stam. bud), 7393 (MO), 20 Oct. 1986 (fr), 8073
(AMAZ, MO); Roca Fuerte (Momon), Oct-Nov 1984 (stam. fl), R. Vasquez & N.Jaramillo
3241 ■ Madre de Dios: Prov. Tambopata, Tambopata Reserve, Rfo Tambopata at mouth of
Rfo D'Orbigny, 250 m, 6 Mar 1981 (fr), A. Gentry & K. Young 32023 (AMAZ, MO, NY).
Agostini (1980) had not seen the type specimen of Weigeltia silvestris and
included it in Cybianthus subgenus Comomyrsine, probably due to matching
with a herbarium misidentification of a specimen of Cybianthus kayapii (Lundell)
Pipoly as Weigeltia silvestris in NY. I previously recognized Cybianthus dubius
and C. loretensis as distinct taxa (Pipoly 1981) on the basis of quantiative
characters, and had not seen the type of C silvestris. The exserted stamens
and obclavate or tubiform pistillode of the staminate flowers, and truncate
style with punctiform stigma in the pistillate flowers are unique features
within the subgenus.
112 SiDA 18(1)
Populations corresponding to the type oiWeigeltia silvestris have sHghtly
larger leaves, but are otherwise indistinguishable from the type of Wejgeltia
nanayemh. The type oi Conomorpha di/bia is notable only for differences at-
tributable to the fact that it is a pistillate fruiting specimen. Fieldwork in
Peru has shown that populations corresponding to the type oi Conomorpha
loretensis grow in full sun, and consequently have narrower leaves with longer
petioles than those of the type of Weigeltia nanayensu .
VII. Cybianthus subgenus Grammadenia (Benth.) Pipoly, Mem. New York
Bot. Gard. 43:47. 1987. GrammackmaV>ftnx.\\., PI. Hartw. 218. 1846. Type Spi-riES:
Grannuadenui niarginata Benth. = Cyhiaiithus iiiarginati/s (Benth.) Pipoly.
Erect or pendent, evergreen, terrestrial, epiphytic or epipetric dioecious,
bisexual, dioecious or monoecious shn/bs or small trees. Roots diageotropic.
Bark mostly smooth, gray or sometimes brown, cracking transversely on
older parts. Trunks distinguishable, normally more or less terete, leptocaulous
or rarely pachycaulous basally, growth dynamics corresponding to Rauh's
Model (Halle et al. 1978). Branchlets thin to moderately thick, terete or ridged,
smooth to verruculose to verrucose, glabrous or rarely glandular-papillate
apically, glandular-papillate at first in the leaf axils then glabrescent, with-
out lenticels. Cataphylls and pseudocataphylls absent. Leaves alternate,
supervolute, exstipulate, sessile, acrodromous, apically obtuse-mucronate,
basally auriculate, subamplexicaul, minutely glandular-papillate along midrib
above, glabrescent, glabrous below, hydropotes few or absent above, nu-
merous below, the margins hyaline, membranaceous, prominently punc-
tate and punctate-lineate, entire or minutely crenulate to denticulate; peti-
oles absent. Inflorescence monomorphic, simple, axillary raceme, at times reduced
to appear dichasial; inflorescence bract broadly ovate to deltate, acute, prominently
punctate, the margins entire to erose, early caducous; rachis terete, straight
or flexuous, minutely glandular-papillate, glabrescent; floral bracts solitary,
at pedicel base, ovate to lanceolate, persistent; pedicels cylindrical to clav-
ate, minutely glandular-papillate, glabrescent, accrescent in fruit. Floivers
unisexual or bisexual, (4— )5— 6(— 7)-merous; perianth lobes imbricate or
quincuncial or rarely and aberrantly dextrorsely contorted, promiently, con-
spicuously or inconspicuously punctate and punctate-lineate; calyx cotyliform,
the lobes erect, at times reflexed in fruit, the margins irregular or regular,
entire or minutely crenulate, glabrous or glandular-ciliate; corolla rotate,
bearing a ring of glandular granules at the junction of tube and lobe, the
lobes glabrous without, glabrous or rarely glanduiar-granulose and smooth
or rugose within, the margins regular or irregular, entire or minutely crenulate;
stamens and staminodes similar, connate to form a conspicuous, membranaceous
or carnose tube adnate to the corolla tube, elobate or with minute lobes
alternating with the anthers, the anthers and antherodes basifixed, sessile
PiPOLY, Cybianthus in Ecuador and Peru 113
or on minute apically free filaments, quadrate or ovate, apically emarginate
or rounded, basally truncate, dehiscent by apical birimose pores, usually
dorsally and rarely ventrally punctate; pistil and pistillode similar, obnapiform,
ellipsoid or umbonate, the ovary terete, lobed or costate, sparsely to densely
translucent-lepidote, the placenta umbonate, (l-)2-3(-4) ovulate, the ovules
half-immersed in the basal placenta, uniseriate or biseriate, the style short,
truncate, the stigma punctiform. Fn/it drupaceous, depressed globose, obovoid,
or ellipsoid, the exocarp sometimes fleshy, the mesocarp and endocarp stony,
prominently punctate-lineate, white, lavender or purple-black at maturity,
one seeded, the testa corrugate, the embryo cylindrical, transverse.
A subgenus of 7 species, in the Lesser Antilles, in Mesoamerica from northern
Costa Rica through Panama, in the Andes from Venezuela to southern Peru,
east through the Guayana Highland and to the Serrania de Turumiquire,
Anzoategui, Venezuela. In Ecuador, two species occur. In Peru, 3 species
have been recorded, of which one, Cybianthus lineatus (Benth.) Pipoly, for-
merly thought to be a Guayana Highland endemic (Pipoly 1987) is reported
for the first time here.
KEY TO THE SPECIES OF CYBIANTHUS SUBGENUS GRAMMADENIA
1 . Branchlets verruculose- to verrucose-papillate or red glandular-papillate; leaves
coriaceous, symmetrical; inflorescence erect, flexuous; perianth coriaceous;
epipetric or terrestrial shrubs or trees ot lagunas in paramos and elfin forests
or in "jalca" or "pajonal" alpine savannas.
2. Epipetric shrub or tree to 6 m tall; trunk leptocaulous; branchlets verruculose-
to verrucose-papillate, prominently ridged, rugose when dried; leaves oblong,
elliptic or lanceolate, (l.l-)1.4-2(-2,5)cm wide; fruit ovoid, then ellip-
soid at maturity; plants of paramos or elfin forests 28. C. marginatus
2. Terrestrial shrub to 1.5 m tall; trunk pachycaulous; branchlets red glan-
dular-papillate, terete, smooth when dried; leaves oblanceolate, (0.4-)0.6-
1(-1.2) cm wide; fruit obovoid throughout development; plants of jalca
or pajonal alpine savannas 29- C. lineatus
1 . Branchlets smooth, glabrous; leaves chartaceous, asymmetrical; infiorescence
lax, straight; perianth chartaceous; epiphytic shrub or tree in cloud lorests
below subparamos or facultative epiphytes in montane and elfin forests.
30. C. magnus
3. Leaves not bearing hydropoten above, conspicuously black punctate and
punctate-lineate below; inflorescence rachis black punctate-lineate; peri-
anth whitish-green, prominently black punctate and punctate-lineate; anthers
ovate, rounded apically; branchlets 4-7 mm diam.; fruit purple-black at
maturity 30a. C. magnus subsp. magnus
3 . Leaves bearing hydropoten above, at least proximally, inconspicuously pellucid
to orange punctate; inflorescence rachis orange punctate-lineate; perianth
maroon to purple, rarely white (then pistillate), prominently orange-punctate;
anthers quadrate, emarginate apically; branchlets 2. 5-3. 5(— 4.0) mm diam.;
fruit white, then lavender at maturity 30b. C. magnus subsp. asymmetricus
114
SiDA 18(1)
Fig. 21. Piccoriaiized distribution of C. marginatm. A-K, Variation in leaf shape, marginal
venation and punctation; note prominent apical mucron, sessile leaf base typical of subge-
nus Grammadema. A-K, drawn from: A. Cuatrecasas 21803, B. Pipoiy 6954, C. Pipoiy 6539,
D. LmiuT 8038, E. L/iteyn 9032, F. Steyermark 100867, G. Mason 13730, H. Pfpoiy 6975 ,
I. Pearce 250, ]. Leh//unin 399, K. Luteyn /Oi 7 J. Figure from Pipoly, 1 987, drawn by Bobbi
Angell.
28. Cybianthus marginatus (Bench.) Pipoly, (Fig. 1 A,B, 7C,F, 21). Mem.
New York Bot. Gard. 43:60. 1987. Grammadema marginata^mxh., PI. Hartw.
218. 1846. Type: COLOMBIA. Cauca: Near Pitayo, 3,636 m, 1843 (bisex. fl), C.
Uartweg 1200 (holotype: K; isotypes: BM, E, G-BOISS, G-DEL, LD, OXF, P, W-2
sheets).
Grammadeu/a lehmaniiii Mez in Engl., Pflanzenr. IV. 236(Heft 9): 231. 1902. Type:
COLOMBIA. Toljma: Altos de Otesas, 3,300 m, 1 1 Jan 1883, (bisex. fl), F. Lehmann
2399 (ei-CTOTYPE by Pipoly 1987: G; isolectotypes: LE, US).
PiPOLY, Cybianthus in Ecuador and Peru 1 15
Grcnumackma alp'ma Mez in Engl., Pflanzenr. IV. 236(Heft 9):231. 1902. Typh: VEN-
EZUELA. Andes of Trujillo and Merida, 1,212-4,390 m, 1842 (bisex. fl),J. Linden
447 (lectotype by Pipoly 1987; P; isoleci-otypes: BM-2 sheets, BR, G, G-DEL, G-
BOISS, K, OXF, S, VEN).
Grainmadenia pastensis Mez in Engl., Pllanzenr. IV. 236(HeFt 9):232. 1902. Type: CO-
LOMBIA. Narixo: W Cordillera of Pasto, 3, 000-3, 200 m, 20 Feb 1881 (bisex. fl), F.
Lehmann 399 (holotype: G; jsotypes: BM, LE).
Grammadema mttda Mez in Engl., Pflanzenr. IV. 236(Heft 9):232. 1902. Type: PERU.
Hl;Anuc:o: Pozuzo, 2,131-2,727 m, 1863 (bisex. fl, fr), R. Pearce 250 (holotype: K).
Grammadema weberbaueri Mez, Repcrt. Spec. Nov. Rcgni Veg. 1 6:418. 1920. Type: PERU.
Cajamarca: Jaen, cordillera E of Huancabamba, E slopes, 2,400-2,500 m, Apr 1 91 2
(bisex. fl), A. Weberbauer 6121 (lectotype by Pipoly 1987: GH; isolectotype: F).
Grammmadmia hexamera Pittier, J. Wash. Acad. Sci. 21:l40. 1931. Type: VENEZUELA.
Merida: Tabay, 2,500-3,000 m, 18 Sep 1930 (bisex. fl), W. Gehriger 47 1 (holotype:
VEN; isotypes: A, F, G, NY, PH).
Grammadenia andkola Cuatrec, Revisca Acad. Colomb. Ci. Exact. 8(3 1):32 1 . 1 95 1 . Type:
COLOMBIA. Valle: Cordillera Occidental, Los Farallones, NW slope, Qucbrada Las
Nieves, below El Diamante, 2,900 m, 30 Jtil 1 946 (bisex. fl, fr),/. Ciuitrecasas 21805
(lectotype by Pipoly 1987: F; isoLE.cTcrrYPE.s: F, COL 2-sheets, U, US).
Epiphytic shrub or small tree to 6 m, the trunk leptocaulous. Branchlets
prominently ridged, 3-4(-5) mm diam., verruculose- to verrucose-papil-
iate, rugose when dried, glabrous. Leaves coriaceous, symmetrical, oblong,
elliptic or lanceolate, (3-)3.5-6(-6.5) cm long, (l.l-)1.4-2(-2.5) cm wide,
apically acute to obtuse, mucronulate, basally acute, auriculate, nitid above,
pallid below, prominently punctate and punctate-lineate, the margin en-
tire, revolute. Inflorescence erect, the rachis flexuous, slender, (1— )1.3-2(— 3.2)
cm long, densely black punctate-lineate; floral bracts widely ovate to deltate,
(().8-)1.2-1.5(-1.8) mm long, 0.8-1.4(-I.8) mm wide, apically acute, the
margin entire; pedicels (1-)1 .5-2 mm long. Flowers coriaceous, 5(-7)-merous;
calyx 1.1-2.1 mm long, the tube ca. 0.1 mm long, the lobes deltate, (1-)
1.5-2 mm long, (1.2-)1.3-1 .5(-2) mm long, apically acute, prominently
and densely punctate and punctate-lineate, the margins minutely crenu-
late, glabrous to minutely ciliolate at first apically; corolla (2-)2.2-3 mm
long, the lobes widely ovate, (l-)1.2-2 mm long, (1.2-)1.5-2(-2.5) mm
wide, obtuse to rounded, glabrous without, glandular-granulose within basally,
densely and prominently punctate medially, the margin entire; staminal tube
carnose, 0.6-0.8(-l) mm long, the anthers sessile, alternate with fleshy lobes
to 0.2 mm long, quadrate, 0.4-0.7 mm long and wide, apically rounded,
basally truncate, the connective prominently punctate dorsally; pistil obnapiform,
(0.9-)1.2-1.5 mm long and (0.8-)1.2-1.6 mm diam., the ovary (0.4-)0.6-
1 mm long, glabrous to sparingly translucen glandular-lepidote, ovules 2(-
4), uni- or biseriate, the style 0.3-0.5 mm long, epunctate. Frai/ somewhat
ovoid, then ellipsoid at maturity, 4.5-5(-6) mm long, (2.5-)3-4 mm diam.,
white, then purple-black at maturity, the punctations red-black, prominent.
Distribution. — Cybianthus marginatus is the most common species of the
116 SlDA 18(1)
subgenus, occurring throughout the Andes from Venezuela to Peru, 2,000-
3,400 m.
Ecology and anuervatum it at US. — This species occurs in large populations
on rocks above lagunas in paramos in northern Ecuador and along water-
courses in paramoid elfin forests, and "ceja" formations in the remainder of
Ecuador and Peru. As long as there are remnant paramo formations, or elfin
forests, there will be populations o^ Cyhianthus marginatm. However, in some
instances, soil compaction due to overgrazing by sheep can render the soil
uninhabitable for this species.
Etymology. — The specific epithet refers to the scarious leaf margin, made
conspicous by the coriaceous texture. In addition, this aspect is made more
conspicuous by the prominent submarginal vein of many populations.
Representative specimens examined. ECUADOR. Azuay: "Oriente" border. Paramo deJ
Castillo, crest of E cordillera on trail between SeviUa de Ore and Mendez, 2,727-3,333 m,
18 Aug 1945 (fl, fr), W. Camp E-4809 (NY, VEN); Rio Collay, Huagarancha S of El Pan,
2,650-3,290 m, 6 Jtd 1 943 (fl) J. Steyermark 53354 (NY), carchi: Peak of Cerro Golondrinas,
00° 51' 12" N, 78° 08' 2 1" W, 3070 m, 24 Jul 1994 (fr), B. Boyle et at. 3373 (BRIT, MO,
QCNE); Canton Montufar, Loma El Corazon, Bretaiia, SE of Mariscal Sucre, Ri'o Minas,
00° 35' N, 77° 42' W, 3,150 m, 22-23 Dec 1992 (fl, fr), W. Palacios & G. Tipaz 10569
(BRIT, MO, QCNE). Imbabura: Cordillera Oriental, Camp Arelan, E of Volcan Cayambe,
2,803 m, 21 Jul 1944 (fl, fr), W. Drew E-351 (MSC); Ridge just S of Rio Clavadero, along
trail to Rio San Pedro, E of Cayambe, 2,893 m, 27 Jul 1944 (fl, fr), /, Wii^gn/s 10484 (DS,
US). Loja: Saraguro-Loja, Km 12.4, turnoff toward Fierro Urco, Km 2.5-2,7, 03° 41' 05"
S, 79° 17' 20" W, 3,150-3,300 m, 7 Dec 1994 (fl), P.Jorgensen et at. 1278 (BRIT, LOJA,
MO, QCA, QCNE); Paramos de Saraguro, 10 km S of Saraguro, 3,050 m, 2 Jan 1979 (fl),
_/. Luteyn et al. 6647 (NY, QCA). Zamora-Chinchipe: border, crest of Cordillera Oriental,
2,840 m, 28 Jan 1984 (fl),J. Luteyn & E. Cotton 1 1295 (NY, QCA); W slopes of Cordillera
del Condor and NW slopes of Nudo de Sabanillas, around Tambo Cachiyacu, ca. 2 km SE
of Yangana, 2,000-3,000 m, 19 Oct 1943 (fl, fr),./. Steyermark 54800 (NY, U); S of El
Playon de San Francisco, slopes of Cerro Mirador, 3,300-3,600 m, 29 Dec 1980 (fl, fr), L.
Hotm-Nielsen et al. 29949 (AAU),J.Jarmtito et al. 3929 (AAU, QCA). PERU. Amazonas:
Prov. Luyas, Dtto. Camporredondo, Ancxo Tullanaya, Cerro Wicsocunga, 06° 05' 35" S,
78° 19' 56" W, 3,075 m, 7 Dec 1996 (pist. fl),/ & L. Campos 3121 (BRIT, MO, USM).
Cajamarca: Jaen, SW of Querocotillo, 3,150 m, Aug 1915 (bisex. fl, fr), A. Weberbai/er
7168 (F, G, GH). Cusco: La Convencion, 2,800 m, 9 Jul 1968 (bisex. fl, fr), T. Dudley
10910 (NA), 10 Jul 1968 (bisex, fl, fr), T. Dttdley 10922 (E, NA), T. Dudley I0931B (F,
NA). Huanuco: Prov. Huanuco, 45 km on rd. from Huanuco to Tingo Mari'a, trail on S
sideofCarpish Tunnel, 09° 42' S, 76° 05' W, 2,400 m, 3 Mar 1985 (bisex. fl), C. Todzia &
B. Stew 2740 (F, TEX, USM).
As was stated previously (Pipoly 1987), Cybianthus marginatus is most
closely related to C. liueat/is (Benth.) Pipoly, previously known only from
the contiguous Guayana Floristic Province, because of its (synapomorphic)
flexous inflorescences and biseriate ovules. However, Cybianthus niargtnatus
IS easily distinguished from C. liutatiis by its glabrous, ridged, verrucose-
papillate branchlets, oblong, elliptic or ovate leaves, obnapiform pistil and
PiPOLY, Cybianthus in Ecuador and Peru 117
ellipsoid fruits. The verrucose-papillate branchlets, leaf and fruit shape are
unique (autapomorphic) features within the subgenus.
Cybianthus margtnatns is the most variable species of the subgenus, con-
taining one-third of the names attributed to Grammadmta as taxonomic synonyms.
Variation in leaf size and punctation and quantitative floral variation have
led to much overdescription. A full disctission of synonymy and variation
was provided by Pipoly (1987). Cybianthus marginatus is most closely re-
lated to C. limatus by virtue of its flexuous inflorescences and biseriate ovules.
However, Cybianthus marginatus is easily distinguished from C. lineatus by
the glabrous, ridged, verruose-papillate branchlets, oblong, elliptic or ovate
leaves, obnapiform pistil and ellipsoid fruits.
29. Cybianthus lineatus (Benth.) Pipoly (Fig. 7E), Mem. New York Bot.
Gard. 43:64. 1987. Gnimmcuknhi limata Bentham, PI. Hartw. 218. 1846. Type:
VENEZUELA. ["GUYANA"}, houvar: Savannas near Roraima, 1843 (bisex. fl, fr), R.
Schonih/a-gk 647/992 (i lOLOTYPr,: K; isotypes: B, BM, G-DC, G-DEL, P, U, W-2 sheets).
Terrestrial shrub to 1 .5 m tall, the trunk pachycaulous. Branchlets terete,
(2.5-)3— 4(— 6) mm diam., densely red glandular-papillate apically. Leaves
coriaceous, symmetrical, oblanceolate,(1.2-)1.6-3(-3. 9) cm long,(0.4-)0.6-
1 (—1 .2) cm wide, apically acute, basally subauriculate, the margin flat. Inflorescence
erect, flexuous, at times reduced to a simple dichasium, 0.6— 1.0(— 2.5) cm
long, sparingly glandular-papillate; floral bracts widely ovate, 0.8—1 . 1 mm
long, 1.1—1.4 mm wide, apically acute, the margin entire, glabrous. Flow-
ers chartaceous, 5-6(-7)-merous; calyx 1.1-1.6 mm long, the tube ca. 0.1
mm long, the lobes wide-triangular to deltate, (0.8-)l-1.5 mm long, (0.7-)
1-1 .2(-l .4) mm wide, apically acute, conspicuously black punctate and punctate-
lineate, the margins entire, glandular-ciliolate; corolla (1.8— )2— 2. 5(— 3) mm
long, the lobes widely ovate, (1-)1 .3-1 .6(-2) mm long, (1-)1 .3-1.7(-2)
mm wide, obtuse to emargmate, inconspicuously punctate medially, the
margins irregular, entire; staminal tube carnose, conspicuous, (0.6— )0. 8—1
mm long, the anthers sessile, alternate with prominent fleshy lobes, quad-
rate, 0.4-0.6 mm long and wide, apically rounded, prominently black punctate
dorsally; pistil ellipsoid, (0.9-)l-l .2(-l .4) mm long, (0.8-)l-l .3 mm diam.,
the ovary 0.8—1 mm long, glabrous to translucent glandular-lepidote apically,
ovules 2-3, when more than 2, biseriate, the style (0.1-)0.2(-0.3) mm long,
glabrous. Fruit ohowo'id, (3-)4-5 mm long, (2-)2.5-3 mm diam., purple,
than black at maturity, the punctations green, prominent.
Distribution. — Formerly thought to be endemic to the Guayana High-
land, in open savannas on tepuf summits throughout Pantepuf (Mayr & Phelps
1967), at 1,400-2,850 m elevation, but now known elsewhere only from
the collection cited below.
Ecology and conservation status. — Cybianthus lineatus grows in fully exposed,
118 SiDA 18(1)
dry montane "shrub savannas" (Huber 1995) in shallow sand over sand-
stone throughout Pantepui. Its occurrence in Pajonal ("jalca") vegetation
in Peru gives reason to expect it in other places, especially in the Cordillera
del CcSndor along the Ecuadorean/Peruvian border. Unfortunately, recent
civil Luirest has prohibited collection in that area. In the majority of the
range lor the species, Cybianthus lineatus is not threatened.
Etymology. — The specific epithet refers to the numerous and prominent
punctate-lineations of the abaxial leaf surface.
Specimen examined. PERU. Pasco: Oxapampa l-'rov., ( ^rro Pajonal, 29 km from Oxapampa,
2,680 m, D. Smith & [-Mter 2'>09 (F, MO, USM).
Cybianthus lineatiis is most closely related to C marginatum (Benth.) Pipoly,
by virtue of its flexuous inflorescence and biseriate ovules (Pipoly 1987).
However, Cyb'mnthin lineatus may be easily recognized by its pachycaulous
trunk, densely red glandular-papillate branchlet apices, ellipsoid pistil and
obovoid fruits. It is the only species in the subgenus with parenchyma in-
stead of aerenchyma in the cortex, the only one with bifacial palisade layers
in the leaf, and the only one with a pachycaulous trunk. None of these
morphological peculiarities are unexpected given its drier, wind-swept habitat.
30. Cybianthus magnus (Mez) Pipoly, Mem. New York Bot. Gard. 43:55. 1987.
Facultative epiphytic shrub or tree to 7 m tall. Branchlets terete, smooth,
glabrotis. Leaves chartaceotis, asymmetrical, narrowly oblanceolate, oblan-
ceolate or narrowly obovate, (4.5-)5.2-15 cm long, (l.()-)2. 1—5.2 cm wide,
apically acute to abruptly acuminate, tapering abruptly or gradually to base,
bearing hydropotes above or not, conspicuously black punctate and punc-
tate-lineate or inconspicuously pellucid to orange punctate below, the margins
entire, flat, or subrevolute. Inflorescence lax, straight, (1.5-)2-8(-l 1.5) cm
long, slender, densely glandular-granuiosc and papillate, prominently black
punctate-lineate or conspicuously orange to brown ptinctate-lineate; floral
bracts ovate, widely ovate or deltate, (0.7— )1. 1-2.2 mm long, (().6-)l .3-2
mm wide, apically acute to acuminate, prominently black or orange punc-
tate and punctate-lineate, the margins erose and glandular-ciliate; pedicels
1.0— 2. 2(— 5.5) mm long in flower, the smaller ones accrescent to 4(— 6) mm
long in fruit. Flowers chartaceotis, 5(— 6)-merous, whitish-green or pink to
maroon; calyx lobes widely ovate to cielatate, (0.8— )1-1 .5(— 2) mm long,
(0.8—) I -2.1 mm wide, apically acute to acuminate, prominently black punctate
and punctate-lineate or orange to brown punctate, the margins erose to fimbriate
and densely glandular-ciliate; corolla (1 .7-)2-2.6(-3) mm long, the stami-
nate and bisexual maroon, the pistillate white, the lobes widely ovate, 1.1—
1.6(-2.2) mm long, (0.9-)l . L-2.6 mm wide, apically obtuse to emargin-
ate, rugose medially within, densely and prominently black punctate and
PiPOLY, Cybianthus in Ecuador and Peru 119
punctate-lineate or orange to brown punctate, the margins irregular, en-
tire; staminal tube membranaceous, (0.4— )0. 7—1 (—1.2) mm long, lobate,
the lobes 0.1-0.2 mm long, the anthers sessile, alternate with the lobes,
ovate to quadrate, (0.3-)0.4-0.6 mm long, 0.4— 0.6(— 7) mm wide, apically
rounded or emarginate, the connectives epunctate ventrally, prominently
black or orange punctate dorsally; pistil obnapiform, 1-1.2 mm long, 0.9—
1.5 mm diam., the ovary 0.6— 0.9(— 1 -3) mm long, densely translucent glandular-
lepidote, the ovules 2-4(-5), uniseriate, the style 0.3-0.5(-0.7) mm long,
glabrous. Fr//i/ obovoid, 2.5—3.5 mm long, 2-3 mm diam. when dried, pink,
then purple-black or white, then lavender at maturity, prominently black
punctate-lineate or orange punctate and punctate-lineate when dried.
Distribution. — Cybianthus ma gnus occurs in the Andes of Venezuela southward
to Peru as an epiphyte in the cloud forest zone below subparamo thickets
and as a facultative epiphyte in montane and elfin "ceja" forests, from 1 ,100—
3,500 m. It is also known from the Serranfa de Turumiquire, in the states
of Monagas, Sucre and Anzoategui, Venezuela, at 2,000-2,400 m.
Etymology. — The specific epithet refers both to the large, branchlets, often
appearing succulent, as well as the large leaf size found in some populations.
As shown by Pipoly (1987) Cybianthus magnus is most closely related to
C. parasiticus (Sw.) Pipoly from the Lesser Antilles by its chartaceous co-
rolla rugose medially within, asymmetrical leaves, and obovoid fruits, but
is easily separated from it by its erose and fimbriate calyx lobes, lobate staminal
tube and sessile anthers. Cybianthus magnus superficially resembles C. marginatus
(Benth.) Pipoly, but may be easily separated by its subsucculent smooth
stems, and obovoid fruits
Both subspecies are known from Ecuador and Peru, with one region of
apparent sympatry on the eastern slopes of the Cordillera Oriental in Ecua-
dor and adjacent northern Peru. However, they appear to be separated by
habitat. In Ecuador and Peru, subspecies magnus appears to be restricted to
closed cloud forests, while subspecies asynmietricus is found in open mon-
tane forest and elfin ("ceja") forest. The salient features of each are summa-
rized below.
30a. Cybianthus magnus (Mez) Pipoly subsp. magnus (Fig. 8E). Mem.
New York Bot. Gard. 43:56. 1987. Gnjmvuidefiui magnaMez'm'Engi.^V'Aiinz.tnr.
IV. 236(Heft 9):231. 1902. Tvph: COLOMBIA. Santander del Norte: Ocana to
Pamplona, 2,000-2,500 m, 4 Mar 1S79 (tl), W. Kalhreyer 1087 (hou)TYFh: K).
Grammadenia oxygyna Cuatrec, Revista Acad. Colomb. Ci. Exact 8:321. 1951. T'^te:
COLOMBIA. Valle di;L Caica: Cordillera Occidental, W slope, bank of Rio Digua,
left side, Piedra de Moler, 900-1 , 100 m, 20 Aug 1943 (fl, fr),/ Cuatrecasas 14947 (lcc;to-
TYPE by Pipoly (1987): F, NY Neg 12136; lsole(;totypes:COL-3 sheets, F, U, US).
Facultative epiphytic shr/^b or tree to 7 m tall, 7-15(-30) cm diam., the
120 SiDA 18(1)
canopy often bowl-shaped. Branchlets (4.0— )5. 0—7.0 mm diam. Leaves not
bearing hydropotes above, conspicuously black punctate and punctate-lineate
below. Inflorescence rachis prominently black punctate and punctate-lineate.
Flowers with perianth whitish-green, prominently black punctate and punctate-
lineate; anthers ovate, rounded apically, the connectives prominently black
punctate dorsally. Fr//it purple-black at maturity, prominently black punc-
tate-lineate when dried.
Distribution. — Subspecies magniis occurs from the Serrania de Turumiquire
(states of Anzoategui, Monagas, Sucre), and in the Andes, from Venezuela
southward through Colombia to and Ecuador to Peru.
Ecology and conservation status. — Subspecies magnus occurs as an epiphyte
in closed cloud forests, especially those below subparamo thickets. It is a
relatively rare, but widely distributed subspecies, and is increasingly en-
dangered owing to habitat destruction.
Etymology. — The epithet refers to the leaf size and stem succulence, a novelty
within the subgenus.
Specimens examined. ECUADOR. Loja: Between Nudo de Sabanillas and Rio Cachiyacu
at Tambo Cachiyacu, 3,()()()-3,5()() m, 1 7 Oct 1943 (IDJ. Stuymuark 53584 (NY). Zamora-
Chinchipe: Nangaritza Cant6n, ridge crest of Cordillera del Condor, above Pachiciitza, on
disputed Peru-Ecuador border, ()4° 06' S, 78" 35' W, 1 ,800 m, 5 Dec 1990 (stam. (1), D.
Neill & W. ?alcicm 9318 (MO, QCNE). PERU. Cajamarca: Prov. San Ignacio, path at the
border of "La Union," 2,200 m, 1 Nov 1995 (stam, H), C. D/az & A. Torres 7805 (BRIT,
MO, USM).
30b. Cybianthus magnus (Mez) Pipoly subsp. asymmetricus (Mez) Pipoly
(Fig. 8F), JVLem. New York Bot. Card. 43:57. 1987. Cr^immckuicicisyiuuietriai
Mez, Bull. Herb. Boissier ser 2, 5:246. 1905. T^'Pi;: PERU. Loiurro: Cerro de Ponasa,
1 ,300 m, Mar 1903 (fr), E. Ule 6792 (l.i^CTOTYPi.; by Pi]M)ly 1987: HBG; i.solf.ctotyprs:
F, G, K, L).
Gramiiiaden'ui niacrocarpci Lundell, Wrightia 5:292. 1976. Type: ECUADOR. Napo: 17
km W of Lumbaque, 70-73 km W of Lago Agno, 1,130 m, 4 Nov 1974 (bisex. fl,
fr), A. Gaitry 1241') (hoi.otypp.: LL-TRX; isotyph.s: MO, S).
Facultative epiphytic .t/ir///; to 2 m tall, 7 m diam., the canopy open, conical.
Brancblets 2.5— 3. 5(— 4.0) mm diam. Leaves bearing hydropotes above at least
proximally, inconspicuously pellucid to orange punctate below. Inflorescence
rachis conspicuously orange to brown punctate-lineate. flowers with calyx
greenish-pink, the staminate and bisexial with corolla maroon, the pistil-
late with corolla white and prominently orange punctate and punctate-lineate
lobes; anthers quadrate, apically emarginate, the connectives prominently
orange punctate dorsally. Fruit white, then lavender at maturity, prominently
orange punctate and punctate-lineate when dried.
Distribution. — Cybianthus magnus subsp. asymmetricus occurs as an obligate
epiphyte, growing on detritis in open montane forests and elfin forests, from
the Darien of Panama to Cusco, Peru, from 1 ,000-2,000(-2,700) m.
PiPOLY, Cybianchus in Ecuador and Peru 121
Ecology and conservation statns. — Subspecies asymmetricus grows as an obli-
gate epiphyte, growing on rocks with deep organic detritis and large, moss-
covered trees. With increasing levels of disturbance, it is being threatened.
Etymology. — The epithet refers to the asymmetric shape of the leaf blades.
Specimens examined. ECUADOR. Carchi: Canton Tulcan, Parroquia Tobar Donoso,
Reserva Indigena Awa, Centre El Baboso, 00" 53' N, 78° 25' W, 1,800 m, 17-27 Aug
1992 (fr), G. Tipaz etal. 1 741 (BRIT, MO, QCNE); Parroquia el Chical, Centro San Marcos,
01° 06' N, 78° 14' W, 900-1,100 m, 20-30 Apr 1993 (A), P- Mendez et al. 341 (BRIT,
MO); Cerro Golondrinas, valley bottom ca. 1.5 km NNE of summit, 00° 51' 52" N, 78°
08' 10" W, 2,750 m, 25 Jul 1994 (stam. fl), B. Boyle etal. 3450 (BRIT, QCNE, MO). PERU.
Amazonas: Prov. Bagua, Imaza, Nuevo Samaria (anexo de UVT), 18 Mar 1995 C. Diaz et
cil. 7385 (BRIT, HUT, MO, USM), Cusco: La Convencion, Cordillera Vilcabamba, ca. 1/2
way between Camps 2 1/2 and 3, 1,980 m, 1 Jul 1968 (bisex. fl, fr), T. Diulley 1()66S (F,
NA), 1,800 m, 24 Jul 1968 (bisex. fl, fr), T. Diulley 1 1324 (NA). Huanuco: Prov. Pachitca,
region of Pucallpa, W part of Sira Mountains and adjacent lowland, ca 24 km SE to 26 km
ESE of Puerto Inca, next to Campamento Pato Rojo, 09° 27' S, 74° 46' W, and along crest
after Campamento Peligrosol,600 m, 14 Apr 1 988 (fr), B. Wallmfer 18-144H8 (BRIT, MO,
W, WU). Pasco: Oxapampa, Cordillera Yanachaga, Cerro Pajonal, chacos, 12 km SE of
Oxapampa, 2,700-2,800 m, 7 Oct 1983 (bisex. fl), R. Foster 9013 (MO, NY, USM).
Cybianthus magnus subsp. asymmetricus is notable for its stems with angu-
lar coUenchyma in the pith, well-developed aerenchyma in the inner cor-
tex, and tangential collenchyma in the outer cortex. It is separated from
subspecies magnus by its inconspicuous or orange punctate leaves, orange
punctate-lineate inflorescence rachis and perianth, quadrate, emarginate anthers
and open montane and elfin forest habitat.
Grammadenia macrocarpa Lundell is notable only for its large, bright or-
ange fruits and smaller flowers. I examined one fruit from each of the isotypes
of G. macrocarpa and found an insect larva in each one, accounting for the
size and peculiar morphology.
VIII. Cybianthus Mart, subgenus Cybianthus. Cyhianthus sect. Encylnanthus
Miq. in Mart., Fl. Bras. 10:292. 1856. Cybiautbi/s sett. Cybumthoidei Miq. in Mart.
Fl. Bras. 10:292. 1856. Typh SPECrES. Cyhianthns pendulifiorus M-An.
Peckia Veil., Fl. Flum. 1:5 1. 1825., nom. rej.
Terrestrial dioecious shrubs or small trees. Roots positively geotropic. Bark
mostly smooth, gray or sometimes brown, cracking longitudinally on older
parts. Trunks distinguishable, normally more or less terete, leptocaulous,
growth dynamics corresponding to Rauh's Model (Halle et al . 1 978). Branchlets
thin to moderately thick, terete, trigonal, or ridged, rufous stellate or den-
droid tomentose or rarely, with rufous, subsessile covering lepidote scales.
Cataphylls and pseudocataphylls absent. Leaves alternate, or pseudoverticillate,
supervolute, exstipulate, petiolate; blades epunctate or variously black or
red punctate, the punctations at times prominent. Inflorescence a simple, lat-
eral (axillary) raceme; peduncle 1-5 mm long, the rachis straight, minutely
122 Si DA 1,S(1)
rufous glandular-papillate, glandular-granulose or lepidote, glabrescent or
persistent; floral bracts solitary, at pedicel basally, ovate to lanceolate, per-
sistent; pedicels cylindrical to clavate, or obconic, minutely glandular-papillate,
glandular-granulose or lepidote, accrescent in fruit. Flowers unisextial or bisexual,
4-merous; perianth lobes imbricate or rarely valvate, prominently, conspicuously
or inconspicuously punctate and/or punctate-lineate; calyx cotyliform, the
lobes erect, at times reflexed in fruit, the margins irregular or regular, en-
tire or minutely crenulate, glandular-ciliate; corolla rotate, to subrotate, the
tube short, glabrous or glandular-granulose, at times papillate, the lobes
glabrous without, glandular-granulose and/or glandular-papillate throughout
within, the margins regular or irregular, entire or minutely crenulate, gla-
brous, glandular-granulose or papillate along the margins; stamens and
stami nodes similar, developmentally adnate to the corolla tube to form an
inconspicuous tube, the stamens thus appearing epipetalous, apical free portion
of the filaments present or absent, when present up to 3 times longer than
the anthers, the anthers basifixed, appearing sessile or on minute apically
free filaments, quadrate or ovate, apically emarginate, rounded, acute, api-
culate or truncate, basally truncate or subcordate, dehiscent by apical pores,
the pores at times confluent, the connective epunctate or conspicuously or
prominently punctate, at times glandular-papillate; pistil and pistillode similar,
conic, ellipsoid, or rarely, obturbinate, the ovary terete, lobed or costate,
sparsely to densely translucent-lepidote, the placenta subglobose, (l-)2-
3(— 4) ovulate, the ovules half-immersed in the basal placenta, uniseriate or
biseriate, the style long-attenuate, the stigma punctiform, lobed or capi-
tate-lobed. Fr//// drupaceous, depressed globose, the exocarp sometimes fleshy,
the mesocarp and endocarp stony, prominently punctate-lineate, white, red
or purple-black at maturity, one seeded, the testa corrugate, the embryo
cylindrical, transverse.
A SLibgenLis of 50 species in tropical South America, with the largest
concentration ol species in the Amazon Basin and adjacent Guianas, Planalto
and coastal Brazil. Subgenus Cybianthia is by far the most complicated taxonomic
group within the genus and is known from rather incomplete material. Collection
of more material is hampered by the fact that the populations studied here-
tofore have revealed population densities lower than any other subgenus.
So far, 13 species are known from Ecuador and Peru, but with additional
collections, we may expect to find Cybianthus lanceolatm Pax, and/or Cybtcinthns
psychotriijoliiis (Rusby) Mez, both from nearby Bolivia, in southern Peru.
KF.Y TO SPECinS OF O'BIANTHHS SUBGENUS CYBIANTHUS
1. Branclilecs subteretL- to ant^'iilate.
2. Indorcscence spicarc- or SLibspicatc; calyx lobes inconspicuously or promi-
nently orange piincrate; petioles pulvinace basally.
PiPOi.Y, Cybianthus in Ecuador and Peru 123
3 . Leaf blades inconspicuously pellucid punccate below; calyx lobes chartaceous
or carnose, rounded or acute apically.
4. Branchlets 4—6 mm diam.; leaf blades coriaceous, nitid above; peti-
oles 2.2-3 cm loni;; inflorescence spicate, the pedicels obsolete to
0.4 mm long; calyx lobes chartaceous, deltate, the margin entire,
undulate, glabrous 31. C. incognitus
4. Branchlet 2—3.9 mm diam.; leaf blades membranaceous, dull above;
petioles 1.7-2 cm long; inflorescence stibspicate, the pedicels 0.6-
0.9 mm long; calyx lobes carnose, ovate, the margin crenulate, glandular-
ciliace 32. C. minutiflorus
3. Leaf blades perpuncticulose below; calyx lobes membranaceous,
subacuminate apically 33- C. huampamiensis
2. Inflorescence racemose; calyx lobes densely and prominently black punc-
tate or epunccate; petioles tapered, not pulvinate basally.
5. Leaves alternate; calyx lobes acute apically, the margins hyaline; an-
ther apically rounded, obtuse or acute apically, the pores not confluent.
6. Pedicels 1.9-2.2 mm long; flowers erect; calyx carnose, 0.8-1 .1 mm
long, the lobes ovate, abruptly constricted basally, densely and promi-
nently black punctate, the margin irregularly serrate, punctate-lineate,
glabrous 34. C. granulosus
6. Pedicels 2.5-3.5 mm long; flowers nodding; calyx membranaceous,
1.5-1.9 mm long, the lobes deltate to triangular, not constricted
basally, epunctate, the margin entire, epunctate, minutely glandu-
lar-ciliolate 35. C. flavovirens
5 . Leaves pseudoverticiUate; calyx lobes rounded apically, the margins opaque;
anthers truncate apically, the pores confluent 36. C. venezuelanus
1. Branchlets terete.
7. Branchlets thick, (6-)7-10 mm in diameter.
8. Leaf blades subacuminate apically; calyx membranaceous or chartaceous,
the lobes acute or acuminate.
9. Leaf blades nitid and perpuncticulose above, 12—13.5 cm wide, the
secondary veins prominently raised above and below, basally trun-
cate, auriculate; petioles 0.5-1.4 cm long; pedicels 0.3-0.5 mm long.
37. C. grandezii
9. Leaf blades sordid and epunctate above, 2.1—5 cm wide, the second-
ary veins deeply impressed above, prominently raised below, basally
cuneate; petioles 2.1-5 cm long; pedicels 0.8-1.4 mm long.
38. C. jensonii
8. Leaf blades abruptly acuminate apically; calyx coriaceous, the lobes
rounded 39- C. fosteri
7. Branchlets thin, 1.5-3.5 mm in diameter.
1 0. Leaf blades apically subacuminate to acuminate; calyx lobes acuminate
or attentuate, the margin erose, short glandular-ciliate.
1 1 . Leaf blades chartaceous to coriaceous, somewhat to very nitid
above and below, the midrib raised above, decurrenr to base
of petiole; petioles short-pulvinate basally.
12. Leaf blades elliptic, (4-)5.5-7(-10.8) cm wide, apically
long-acuminate, basally cuneate, the tertiary veins prominently
raised, inconspicuously pellucid-punctate below; petioles
canaliculate; pedicels cylindrical in fruit 40. C. resinosus
124
SiDA 18(1)
2mm
FKi. 22. Cyhhiiitbiis ntoiv^iiitiis I'lpoly. A. Habit, showing rrit^onal branchler. B. Abaxial leaf
surface, showing miniiCL- scales. C. Portion of infrLictcscencc, showing delcate calyx lobes
with entire margins. D. Branchler apex, showing dendrojd and stellate tomentum. E. Por-
tion of staminate spike, showing conspicuous staminal tube, and obcordate anthers with
SLibapical non-confluent pores. A, B, O, drawn from Gt:)it)-y et al. 2291 1. C, drawn from
B(/r/x)//r 2567 . E, drawn from holotype, by Lintia EUis.
Pipon', Cybianthus in Ecuador and Peru 125
12. Leaf blades very narrowly oblanceolate or oblong, 2— 4(— 5) cm
wide, apically and basally long-attenuate, the tertiary veins
inconspicuous, conspicuously black or red punctate and
punctate lineate below; petioles marginate; pedicels obconic
in frti i t 4 I.e. fuscus
1 1. Leaf blades chartaceous, di.ill green above and below, the mid-
rib impressed above, not decurrent on the petiole; petioles gradually
tapering to base, withotit ptilvinus 42. C. cyclopetalus
10. Leaf blades apically acute; calyx lobes obtuse, the margin crenu-
late, long ciliate 43. C. penduliflorus
31. Cybianthus incognitus Pipoly, sp. nov. (Fig. 22). Typi;: PERU. Amazonas:
Rio Santiago Valley, 03° 50' S, 77° 40' W, QuebradaCaterpiza, 2-3 km from Caterpiza
settlement, primary foresr, 200 m, tree 9 m tall, 12 Dec 1979 (stam. fl), S. Tiinqui
289 (holotype: MO; isotypes: USM, NY).
Ob folia coriacea oblanceolata desuper nitida subter pallida anthera filamenta 3-plo breviores
C. prieuro valde arete affinis sed ab ea petiolis canaliculatis (nee marginatis), ramulis adpresse
dendroideo- et stellate- (nee erecte dendroideo-) tomentellis, pedicellis 0—0.4 (non 0.8—
1.4) mm longis, lobiscalycinis inconspicuepellucido- (non manitesre aero-) punccatis, lobisw
coroUinis ovatis (nee stiborbicularis) pistillodio globoso (nee conico) denique fructu laevi
statim recognitur.
Tree to 9 m tall, at times flowering precociously {P. Barbour 2405). Branchlets
subterete to trigonal, 4—6 mm diam., appressed rufous dendroid and stel-
late tomentose. Leaves pseudoverticillate; blades coriaceous, oblanceolate,
(16— )20— 25(-31) cm long, (5— )6.5— 8.5(— 10) cm wide, apically acuminate,
basally cuneate, decurrent on the petiole, nitid above, pallid below, the midrib
slightly impressed above, prominently raised below, the secondary veins 9—
12(— 15) pairs, slightly raised above, prominently raised below, nitid above,
pallid and minutely rubiginous lepidote below, the pellucid punctations
inconspicuous, the margin entire, irregular, flat; petioles canaliculate 2.2—
3 cm long, pulvinate, sparsely pubescent at first, glabrescent. Staminate inflorescence:
an erect, dense spike, (6— )8— 9-5 cm long, the rachis green, sparsely den-
droid pubescent, glabrescent; floral bracts lanceolate, 0.5-1 mm long, apically
attenuate, sparsely pubescent, early caducous; pedicels obsolete to stoutly
cylindrical, 0—0.4 mm long, glabrate. Staminate flowers 4-merous, yellow,
chartaceous; calyx cotyliform, 0.6—0.9 nxm long, the tube 0.2 mm long,
the lobes deltate, 0.4—0.7 mm long and wide, apically acute, inconspicu-
ously pellucid punctate, the margins scarious, entire, epunctate, undulate,
glabrous; corolla subrotate, 1.4—1.6 mm long, the tube 0.2—0.3 mm long,
the lobes ovate, 1.2—1.4 mm long, 0.4—0.6 mm wide, apically acute, gla-
brous without, densely glandular-granulose within, inconspicuously pel-
lucid punctate, the margin opaque, densely glandular-granulose, entire; stamens
1—1.2 mm long, the tube ca. 0.2 mm long, the filaments terete, 0.5—0.6
mm long, slightly reflexed proximally, the anthers obcordate, 0.3—0.4 mm
long, apically apiculate, basally cordate, dehiscent by small, subapical, ovate.
126 SiDA IS(I)
non-conHuent pores, the pores extendin^^ less than 1/2 anther length, the
connective epunctate, densely and minutely rubiginous glandular-granulose
dorsally; pistillode broadly subglobose, ca. 0.2 mm long, 0.4 mm diam.,
densely yellow glandular-papillate. P istillate inflorescence as in staminate but
9—12 cm long; floral bracts lanceloate, 0.2—0.5 mm long, glabrate; pedicels
obsolete to 0.4 mm long, glabrate. P isti I Ic/te flowers as in staminate, but ca-
lyx 0.6—0.9 mm long, the tube 0.2 mm long, the lobes 0.4—0.7 mm long
and wide, the margins opaque, entire; corolla, staminodes and pistil un-
known. Fr///'/ dark purple at maturity, globose, 0.4—0.5 mm long, 0.4—0.6
mm diam., smooth, incospicuously pellucid punctate.
Distribution. — Upper Rio Santiago Valley and adjacent Serranfa de Bagua,
Amazonas, 200—2,000 m, and Maynas Province, Loreto, Peru, along the
Rfo Napo, at 120 m elevation.
Ecology and conservation status. — Cyhianthi/s incognitas is mostly a ridgetop
species in the cloud forests of Amazonas Department, occurring infrequently
near the forest margin. The Upper Rio Santiago Valley and adjacent Serranfa
de Bagua are known for their endemic species (Pipoly 1992b). However,
one surprising collection was noted at 120 m elevation, from Caserio de
Urcumiraho, in Maynas Province of Loreto Department, where vegetation
normally associated with much higher altitudes occurs on the tops of un-
dulating hills. Much more fieldwork will be required to better understand
forest dynamics at the western limits of the Peruvian Amazon with the foothills
of the Andes.
Etymology. — The specific epithet refers to the fact that the plant was
misidentified even to family for nearly twenty years, and was finally identified
only when a flowering specimen was matched with the other fruiting specimens.
The densely spicate infructescences with numerous fruits were heretofore
unknown in the genus.
Paratypi;s. PERU. Amazonas: Prov. Bagua, 12 km E of La Peca, cloud forest, 1700 m,
20 Jun 1978 (fr), P. Barbour 2405 (AMAZ, F, MO, USM), 29 Jun 1978 (fr), P. Barhotir
2367 (AMAZ, BRIT, F, MO, NY, US); Ca. 12-18 km E of La Peca m Serranfa de Bagua,
cloud forest, 1 ,800-1 ,950 m, 14 Jun 1978 (fr), A. Gentry et al. 22839 (F, MO, USM), A.
Gentry et al. 2291 1 (F, MO, USM). Loreto: Rio Napo near entrance to Isla Inayuga, 20 Sep
1972 (fr), T. Croat 20328 (AMAZ, MO, USM); Caserio de Urcumirafio, Rfo Napo, 2 hours
along trail from village to forest, 120 m, 8 Oct 1979 (stam. fl), C. Diaz & N. Jara>nillo
1474 (AMAZ, BRIT, MO, USM).
Cybianthus incognitus is appears to be most closely related to Cybianthus
prieurii A. DC. of the Guianas, Venezuela and Brazil, because of the oblan-
ceolate, highly nitid coriaceous leaf blades and the filaments three times
longer than the anthers. However, Cybianthus incognitus is separated from
C prieurii by its canaliculate petioles, dendroid and stellate tomentose branchlets,
sessile to subsessile flowers (spicate inflorescences), inconspicuously pellu-
PiPOLY, Cybianrhus in Ecuador and Peru 127
cid-punctate calyx lobes, ovate corolla lobes, globose pistillode and smooth
fruits. Cybianthus incognitus is unique within the subgenus by virtue of its
densely spicate inflorescences, dendroid and stellate tomentum of the branchlets
and inconspicuosly punctate calyx lobes.
32. Cybianthus minutiflorus Mez, Repert. Spec. Nov. Regni Veg. 3:102.
1 906. Type: PERU. Loreto: near Rioja, W of Moyobamba, 800-900 m, 8 Sep 1904
(pist. fl, fr), A. Weberbauer 4699 (iiolotype: B -destr.; fragment, F; lectotype, here
designated: F). Because the fragment at F contains floral and leaf material, and leave
no doubt as to the identity of the species, in the absence of other duplicates, it is
most appropriate to select this "clastotype" (a fragment taken with permission) as
the lectotype.
Tree to 3 m tall. Branchlets angulate, 4.5—6 mm diam., densely rufous
stellate-tomentose, glabrescent. Leaves alternate; blades membranaceous, widely
(rarely narrowly) oblanceolate, (l4-)19-28(-40) cm long, (4-^)9-1 2(-l 5)
cm wide, apically acuminate, the acumen 1—1.5 cm long, basally cuneate,
midrib slightly raised above, prominently raised and densely rufous tomentulose
below, the secondary veins (9-) 12— 21 pairs, slightly raised above, promi-
nently raised and sparsely rufous tomentulose below, smooth and glabrous
above at maturity, sparsely rufous puberulent below, conspicuously pellu-
cid punctate, the margin entire, flat; petioles canaliculate, 1.7—2 cm long,
somewhat pulvinate, glabrous above, rufous tomentulose below, glabres-
cent. Staminate inflorescence a simple, erect raceme, 4.5—9 cm long, the ra-
chis densely rufous stellate-tomentose; floral bracts linear-lanceolate, 1 — 1.2
mm long, 0.2—0.3 mm wide, apically attenuate, densely tomentose above
and below, the margin entire; pedicels cylindrical, 0.6—0.9 mm long, densely
tomentose, persistent. Staminate flowers pale yellow; calyx carnose, cotyliform,
0.6—0.8 mm long, the tube ca. 0.1 mm long, the lobes widely triangular,
0.5—0.7 mm long, 0.7—0.9 rnm wide, apically acute, densely and promi-
nently orange punctate, sparsely rufous puberulent, the margin irregular,
opaque, somewhat crenulate, minutely glandular-ciliolate; corolla carnose,
subrotate, 1.2—1.4 mm long, the tube 0.3—0.4 mm long, the lobes widely
triangular, 0.9—1.1 mm long, 1.1 — 1.2 mm wide, apically obtuse, densely
and prominently orange punctate medially without, densely glandular-granulose
medially and above anther within, the margin opaque, glandular-granulose,
entire; stamens apparently sessile at corolla tube apically, the anthers widely
obcordate, 0.4—0.5 mm long, 0.7—0.9 rnm wide, apically acute, basally cordate,
the thecae moderately yellow glandular-granulose, the connective promi-
nently red punctate; pistillode conic, 0.3—0.4 mm long, 0.1-0.2 mm diam.,
the stigma glandular-papillate. Pistillate inflorescence as m staminate but (4—)
8—13 cm long; floral bracts 0.6—1 mm long, 0.1-0.2 mm wide, apically
attenuate, densely tomentose above and below, the margin entire; pedicels
0.6—0.8 mm long, accrescent in fruit to 1.8 mm long. Pistillate flowers as in
128 SiDA 18(1)
staminate but green; calyx 0.8-1 mm long, the tube ca. 0.1 mm long, the
lobes 0.7-0.9 mm long, 0.9-1 mm wide; corolla 1-1.2 mm long, the tube
0.2-0.3 mm long, the lobes 0.8-1.0 mm long, 1-1.1 mm wide; staminodes
as in stamens but antherodes obcordate, 0.2-0.3 mm long, 0.2—0.3 mm
wide; pistil cylindrical, 0.6-1 mm long, 0.2—0.3 mm diam., the ovary an-
gular, 0.3-0.4 mm long, the style 0.3-0.6 mm long, the stigma subcapitate,
4-lobed, the placenta cotyliform, ovules 2, naked. Fr//// depressed-globose,
2.5-3.5 mm long, 4.5-5.5 mm diam., inconspicuously pellucid punctate,
the exocarp thin. Bisexual inflorescence in staminate but 2-3(-5) cm long,
the rachis moderately tomentose; floral bracts, 0.6-0.9 mm long, 0.1-0.2
mm wide; pedicels 0.6-0.7 mm long. Bisexual flowers green; calyx 0.7-8
mm long, the tube ca. 0.1 mm long, the lobes 0.6-0.7 mm long and wide;
corolla 1.3-1.4 mm long, the tube 0.2-0.3 mm long, the lobes 1.1-1.2
mm long, 1-1.1 mm wide; stamens identical to pistillate staminodes, but
thecae full of pollen; pistil almost indistinguishable from pistillate flower
except the stigma subcapitate, 3-4-lobed, ovule 1, naked. Fruit {horn bi-
sexual flower) unknown.
Distribution. — Endemic to the eastern slopes of the Andes and adjacent
Amazonian Hylaea of Peru and adjacent Bolivia, 100-1200 m.
Ecology and conservation status. — Cybianthus minutiflorus occurs in primary
tall wet forest and premontane forest, on well drained white sands, known
as varillal in Peru. These pockets of sandstone often alternate with rolling
lateritic hills in the foothills of the eastern Andean slopes and adjacent Amazonia.
The lowland forests where Cybianthus minutiflorus is occurs are also notable
for their numerous lianas. Cybianthus minutiflorus is a rare species and should
be considered threatened.
Etymology. — The specific epithet refers to the extremely small flowers,
some of the smallest in the subgenus.
Local names and uses. — Peru: "taku kaspi" (Maynajivaro); leaves are boiled
in water and the decoction drunk to treat stomach ache.
Specimens examined. PERU. Amazonas: Prov. Bagua, Dtto. Imaza, Cerros de Putuim,
05° 03' 20" S. 78° 20' 23" W, 350 m, 1 5 Jun 1996 (scam, fl), R. Vasquez tt al. 21187 (AMAZ,
MO). Prov. Condorcanqui, Dtto. El Cenepa, NE region of Maranon Drainage Basin, Ri'o
Cencpa, Comunidad Tutmo, 04° 33' S, 78° 10' W, 750 m, 22 Nov 1993 (fr), R. Vaujuezet
al. 18520 (BRIT, HUT, JMO, USM). Cusco: Quipicanchi Prov., Camanti, Manirf, along
Rio Maniri and along the trail to Quebrada Garrote, 13° 71' S, 70° 45' W, 720 m, 8 Sep
1990 (bud), At, 71)/iatia922 (CUZ, MO, USM). Huanuco: Rio LluUa Pichis watershed,
Cerros del Sira, 1,290 m, 17 Jul 1969 (fr),/ Wolfe 12346 (F, NA, US). Loreto: Prov. Loreto,
Pampa Hermosa and vicinity, Rio Corrientes, 1 km S of junction with Rfo Macusari, 03°
15' S, 75° 50' W, 160 m, 3-20 Dec 19S5 (fr), W. Lewis et al. 10312 (BRIT, MO); Prov.
Maynas, Dtto. Las Amazonas, Explornapo Camp, near Sucusari, along Rio Napo, 03° 20'
S, 72° 55' W, 100-140 m, 3 Mar 1991 (ster.),J, Pipoly et al. 14174 (MO, UNAP); Dtto.
Iquitos, Allpahtiayo (IIAP), Permanent inventory, 04° 10' S, 73° 30' W, 150 m, 25 Mar
1992 (ster.), R. Vasquez et al. IS 1 63 (BRIT, MO, UNAP). Pasco: Oxapampa, Rd. in con-
struction between Oxapampa and Villa Rica, km 7, 10° 37' S, 75° 20' W, 2,100 m, 4 Jan
PiPOLY, Cybianthus in Ecuador and Peru
129
2mm
Fig. 23- Cybianthus huampamiensis Pipoly. A. Habit, showing pulvinate petioles. B. Abaxial,
densely perpuncticulose leaf surface. C. Portion of staminate inflorescence, showing subspicate
habit, and calyx erose-dentate and glandular-ciliate. D. Staminate flower, showing abruptly
constricted corolla lobe base. E. Fruit and calyx. F. Branchlet apex. G, Pistillate flower,
showing constricted corolla lobe base. A— B, drawn from Kayap 783 ■ C~D, drawn from Kayap
982. E-G, drawn from Kayap 933- Figure drawn by Linda Ellis.
130 Sum 18(1)
1984 (bisex. fl), R. Foster et al. 787 7 (F, MO, USM); Oxapampa, Pichi's Valley, San Marias
Ridge, 1 0—1 2 km SW of Puerto Bermudez, above Sta. Rosa de Chivis, trail to Loma Linda,
10° 20' S, 75° 00' W, 1,000 m, 29 Sep 1982 (fr), R. Foster 8624 (R MO, USM), (stam. tl),
R. Foster H981 (BRIT, MO, F, NY, US, USM). San Martin: Chazuta, Rio Huallaga, 260 m,
Mar 1935 (stam. fl), G. King 3981 (F, MO, S, US); Prov. Mariscal Caceres, Dtco. Tocache
Nuevo, Isla de Pucunchu, right bank of Rfo Huallaga, 3 Apr 1971 (lr),_/. Schunke 4779 (F,
MO, NY, US); W of bridge, 700-800 m, 16 Dec 1971 (fr),/ Schunke' 37 37 (F, NY, MO,
US). Ucayali: Prov. Coronel Portillo, Plantacion Margarita, near Loreto border, 1,500-
1,600 m, 14 Aug 1946 (fr), R. Ferreyra 1040 (US, USM). BOLIVIA. Pando: Prov. Madre
de Dios, Mobil Oil Site, 12° 10' S, 67° 15' W, 170 m, 20-25 Aug 1992 (fr), T. K/I/een
4449 (BRIT, LPS, MO).
Cyhianthus minutiflorus is most closely related to C. granulosus Pipoly by
its densely rufous furfuraceous-lepidote branchlets, erect, carnose, densely
and prominently black punctate perianth, and sessile anthers. However, the
flat petioles, subacuminate leaf apices, short pedicels, opaque, crenulate and
epunctate margins of the calyx lobes, and emarginate anthers easily distin-
guish Cyhianthus minutiflorus.
33. Cyhianthus huampamiensis Pipoly, sp. nov. (Fig. 23). Type: PERU. Ama7.ona.s:
Quebrada chigkan entsa, Rfo Cenepa, 300 m, 9 Jun 1973 (stam. fl), F. Ancuash 388
(holotype: MO; isotypps: NY, USM).
Propter folia elliptica lanceolata vel oblanceolata, longipetiolata equilaterale vel inequilarerale
secus margines irregulares, rhachides inflorescentiares graciles, flores erectes deminutosque,
necnon Irutos minores, ad as]->ecru primo intuito C. resinoso arete similans sed ab ea laminis
membranaceis (non renuiter coriaceis), utrinque sordidis (nee nitidis) subter manii:este
prominenteque atro-perpuncticulosis (nee epunctatis), petiolis (1.5— )2— 2,5 (non 0.5—1 .4)
cm longis, lobis calycinis translticentibus (non opacis) acuminatis (nee rotundatis), lobis
coroliinis extus glandulari-granulosis (non glabris) acutis vel rotundatis (nee obtusis vel
emarginatis) denique fructibus luteis (non atris) permanifeste distinguitur.
Tree to 3(— 6) m tall. Branchlets subterete to angulate, (3-5— )4— 5 mm diam.,
densely rufous tomentose, glabrescent. Leaves psetidoverticiUate; blades
membranaceous, elliptic lanceolate or rarely, oblanceolate, (12— )I6— 25(—
31) cm long, (5.2— )7— 9(— 12)cm wide, apically long-acuminate, basally acute,
not decurrent on the petiole, dull green above, pallid below, midrib im-
pressed above, prominently raised below, the secondary veins 12—18 pairs,
prominently raised below, glabrous above rufous puberulent below along
the veins, prominently and densely perpuncticulose below, the margin en-
tire, irregular, flat; petiole slightly canaliculate distally or flat, 2—2.5 mm
long, thick and pulvinate, densely rufous puberulent at first, glabrescent.
Stamina te inflorescence a lax, simple, subspicate raceme, (5—) 10— (18) cm long,
sparsely rufous puberulent; floral bracts linear-lanceolate, 1—1.3 mm long,
apically attenuate, densely rufous publerulent abaxially, the margin erose,
persistent; pedicels cylindrical 0.8— 0.9 nim long, densely rufous pubescent,
glabrescent. Staminate flowers yellowish to orange; calyx membranaceous,
cotyliform, translucent, 1 .2—1.4 mm long, the tube ca. 0.2 mm long, the
PiPOLY, Cybianthus in Ecuador and Peru 131
lobes ovate, 1—1.2 mm long, 0.6—0.8 mm wide, apically subacuminate, densely
and prominently orange punctate, glabrous, the margin hyaline, prominently
erose-dentate, sparsely glandular-ciliate; corolla chartaceous, subrotate, 2.0—
2.4 mm long, the tube 0.4—0.5 mm long, the lobes ovate, 1.6—1.9 mm long,
1.2—1.4 mm wide, apically acute to rounded, sparsely glandular-granulose
without and densely so throughout within, densely and prominently or-
ange punctate, flat, the margin scarious, erose-denticulate and glandular-
granulose; anthers apparently sessile at junction of corolla tube and lobe,
very widely ovate, 0.4—0.5 mm long, 0.6—0.8 mm wide, apically obtuse to
rounded, dehiscent by apical confluent pores extending ca. 2/3 length of
anther, the connective epunctate, densely rubiginous glandular-papillate dorsally;
pistillode conic, ca. 0.2—0.3 mm long and diam., hollow, glandular-papil-
late. Pistillate inflorescence 3.?. in staminate but (5— )7— 9(— 14) cm long, densely
rufous puberulent at first, glabrescent; pedicel 0.4—0.5 mm long. Pistillate
flowers as in staminate but yellowish to orange; calyx 1—1.2 mm long, the
tube ca. 0.2 mm long, the lobes 0.8-1 mm long, 0.4-0.5 mm wide; corolla
1.8-2.2 mm long, the tube 0.3-0.4 mm long, the lobes 1.5-1.7 mm long,
1.1-1.4 mm wide; staminodes as in stamens but antherodes 0.3—0.4 mm
long, 0.5—0.6 mm wide, apically rounded to acute; pistil conic, ca. 1.3 mm
long, the ovary 0.5—0.6 mm long 0.6-0.8 mm diam., translucent glandu-
lar lepidote, the style short, to 0.2 mm long, the stigma 4-lobate, the lobes
distally curved, glandular-papillate. Fr///V yellow, subglobose, 4—6 mm long,
5—8 mm diam. inconspicuously pellucid punctate.
Distribution. — Endemic to the Rfo Maranon, Rio Cenepa and Rfo Santiago
drainage basins in the northwest corner of the Department of Amazonas,
Peru, 200-550(-l,850)m.
Ecology and conservation status. — Cybianthus huampamiensis occurs in the foothills
of the premontane and lowland wet forest of the most underexplored area
of the Peruvian Hylaea/Andean interface. Given that the region is a border
area, and therefore, a priority for development, this species should be con-
sidered.
Etymology. — The epithet describes the place where the many of the col-
lections were made, the Haumpami area of the Rio Cenepa Drainage Basin.
Paratyprs. PERU. Amazonas: Prov. Bagua; ca. 12-18 trail km E of La Peca, Serranfa
de Bagua, 1,800-1,950 m, 14 Jun 1978 (fr), A. Gentry et at. 22859 (F, MO, USM); Ri'o
Cenepa, Quebrada tujushik entsa, 330 m, 18 Apr 1973 (fr), E. Ana/ash 274 (MO, USM);
along Rio Cenepa, 350 m, 3 May 1973 (fr), H. Amuash 303 (MO, USM); Rfo Cenepa, Quebrada
Idayua entsa, 400 m, 16 May 1 973 (fr), /;. Amuash 392 (BRIT, F, LL-TEX, MO, US, USM);
Rfo Cenepa, Quebrada Wampusik entsa, "3 Aug 1974 (stam. fl), £. Ami/ash 731 (MO-2
sheets, USM); Rfo Cenepa, Quebrada Cikan Inci, 250-330 m, 1 Jan 1973 (stam. fl), B.
Bertni 779 (MO, VSM); 10 km N of Quebrada Huampami, 200-250 m, 24 Jul 1974 (stam.
fl), B. Berlin 1760 (MO, USM); Quebrada Chigkui Shmuki Cenepa, 250 m, 1 1 Apr 1973
(fr), R. Kayap 618 (MO, USM), 23 May 1973 (stam. fl), R. Kayap 783 (MO, NY, USM);
132
SiDA l<S(l)
Fk;. 24. Cyhui}itl)iis {i^ViDndoiiis Fipoly. A. Habic. B. Branchlet apex. (]. Abaxial leaf SLirhxce,
showint; miiuicely riitous Icpitlocc indLimentum. D. Staminate flower. E. Staminate corolla.
F. Sraminace tal\x. A-f], drawn from holotypie. D— F, drawn from \' Hinishikiit 1221 .
Figure drawn by Linda Fllis.
PiPOLY, Cybianthus in Ecuador and Peru 133
Quebrada Etseketai, Rfo Cenepa, 250 m, 31 May 1973 (fr), R. Kayap 856 (MO, USM);
Quebrada Wampushik entsa, 330 m, 13 Jun 1973 (pist. fl, fr), R. Kayap 933 (F, LL-TEX,
MO, NY, USM); Huampami, Rio Cenepa, 200 m, 15 Jun 1973 (stam. fl), R. Kayap 982
(LL-TEX, MO, NY, USM); Rio Cenepa, vicinity Huampami, ca. 5 km E of Chavez Valdivia,
04° 30' S, 78° 30' W, 200-250 m, 31 Jul. 1978 (fr), A. Kujikat 30 (LL-TEX, MO, NY,
USM), 12 Aug. 1978 (fr), A. Kujikat 291 (MO, NY, USM), 15 Aug. 1978 (fr), A. Kujikat
395 (MO, USM); Mouth of Rio Santiago, without date (stam. fl), G. Tessmann 4439 (NY);
Rio Santiago Valley, 03° 50' S, 77° 40' W, Quebrada Catetpiza, 2-3 km from Caterpiza,
200 m, 28 Dec 1979 (stam. fl), S. Tunqui 488 (MO, NY, USM); Dtto. Imaza, Quebrada
Kuzu, 1 hour walk from Comunidad de Cunchim, 370 m, 21 Jul 1994 (fr), C. Diaz et al.
6930 (BRIT, HUT, MO, USM); Rfo Marafion Drainage Basin, Comunidad de Yamayakat,
Rio Maranon, 04° 55' S, 78° 19' W, 600 m, 28 Jan 1995 (fr), £. Rodriguez 283 (BRIT,
HUT, MO, USM), Quebrada Kusu-Chapi, 04° 55' S, 78° 19' W, 550 m, Feb 1995 (stam.
fl), R. Vdsquez et al. 20045 (BRIT, HUT, MO), 320 m, 1 1 Mar 1996 (stam. fl), N.Jarami/h
etal. 1351 (BRIT, HUT, MO).
Cybianthus huampamiemis may at once be distinguished from all other species
of the subgenus by its translucent calyx, externally glandular-granulose corolla
and yellow fruit. This species is one of many endemic taxa known from this
most underexplored area at the junction of the eastern Andean slopes with
the Amazon Basin in northern Peru.
34. Cybianthus granulosus Pipoly, sp. nov. (Fig. 24). Type: PERU. Amazonas:
Rio Santiago Valley, 03° 50' S, 77° 40' W, Quebrada Caterpiza, 2-3 km from Caterpiza
settlement, primary forest, 200 m, treelet 2 m tall, fls. brownish-green, 28 Nov 1979
(stam. fl), S. Tunqui 161 (hoi.otype: MO; isotype: NY).
Quoad folia alterna chartacea ad apices acuminata ad bases cuneataque, ramulos dense
rufo-furfuraceo-lepidotos, flores erectos, carnosos, dense manifesteque atro maculatos, antheras
sessiles, ad C. minutiftoro valde affinis sed ab ea petiolis canaliculatis (non planis) laminis
abrupte largo- (non sub-) acuminatis, pedicellis 1 .5-2.5 (nee 0.5-1) mm longis, lobis calycinis
secus margines hyalims (non opacis) serratis (nee crenulatis) punctato-lineatis (nee epunctatis)
lobis corollinis secus marginis erosis (nee integerrimis) antheris acutis (nee ematginatis)
facile cognoscitur.
Treelet to 3 m tall. Branchlets angulate, 2-4 mm diam., densely rufous
furfuraceous lepidote. Leaves alternate; blades chartaceous, elliptic to nar-
rowly oblanceolate, (9-)l 1-15(-18) cm long, (3-)4-5(-6) cm wide, apically
abruptly acuminate, basally cuneate, densely rufous lepidote (appearing granulose)
above and below at first, glabrate above, somewhat persistent below, mid-
rib impressed above, prominently raised below, the secondary veins 8-12
pairs, impressed above, prominently raised below, inconspicuously pellu-
cid punctate, the margin entire; petioles canaliculate, 1 .5-1 .8 cm long, glabrescent
above, densely lepidote below. Staminate inflorescence a simple raceme, (4-)
6_9(_10)cm long, the rachis and pedicels densely rufous furfuraceous lepi-
dote; floral bracts carnose, linear lanceolate, 0.8-1.2 mm long, apically long-
attenuate, the margin entire, densely lepidote; pedicels cylindrical, 1.9-
2.2 mm long. Staminate flowers erect, 4-merous, carnose, subrotate, brownish-green;
134 SiDA 18(1)
calyx 0.8-1.1 mm long, the tube 0.1-0.2 mm long, the lobes ovate, 0.7-
0.9 mm long, 0.8-0.8 mm wide, apically acute, abruptly constricted ba-
sally, densely and prominently black punctate, moderately rufous lepidote,
glabrescent, the margin hyaline, conspicuously black punctate-lineate, ir-
regularly serrate, glabrous; corolla subrotate, 1.6-1.8 mm long, the tube
0.3-0.4 mm long, square, glabrous, the lobes suborbicular, 1.3-1.4 mm
long and wide, emargmate apically, abruptly constricted basally, densely
and prominently black punctate, sparsely rufous lepidote without, glabrescent,
densely glandular-granulose throughout within, the margin erose; anthers
sessile at apex of corolla tube, thus appearing epipetalous, the tube 0.3-0.4
mm long, glabrous, the anthers widely cuadrate, 0.3-0.4 mm long, 0.6-
0.7 mm wide, apically acute, basally truncate, dehiscent by large subapi-
cal, ovate pores, the pores not confluent, extending ca. 3/4 anther length,
the connectives densely and prominently red or black punctate; pistillode
conic, 0.2-0.3 mm long, sparsely glandular-lepidote. Pistillate inflomcence
unknown. Fruit unknown.
Distribution. — Endemic to the upper Rio Santiago Valley, in the Depart-
ment of Amazonas, Peru, 180-200 m.
Ecology and conservation status. — Cybianthus granulosus inhabits wet premontane
forests above the Rfo Santiago valley, which together with the Rio Cenepa,
comprise a region now known to be host to a number of endemic species.
Given that it is most likely endemic, and not at all well-known, its conser-
vation status cannot be determined at this time.
Etymology. — The specific epithet refers to the densely rufous lepidote
tomentum, which appears granulose when examined superficially.
Par/viypes. PERU. Amazonas: Prov. Bagua, Dtto. Imaza, Comuniclad Aguaruna Putuim,
Anexo Yamayakat, Zonas Alcas de Putuim, "Campou," 700 m, 18 Jan 1996 (Fr), C. Dun et
al. 7649A (BRIT, HUT, MO, USM); Rio Santiago, 3 km from Caterpiza, 180 m, 1 2 Nov
1 979 (scam, fl), V. H/iashikat 1221 (MO, USM); Valle del Rio Santiago, 65 km N of Pinglo,
Quebrada Caterpiza, 2-3 km from Caterpiza, 200 m, 19 Sep 1977 (bud), V. H/mshikat677
(MO, USM), 28 Nov 1979 (scam, fl), V. Huashikat 1422 (MO, USM).
Cybianthus granulosus is most closely related to C. minutifiorus Mez by virtue
of Its alternate, chartaceous leaf blades with acuminate apices and cuneate
bases, densely rufous furfuraceotis-lepdidote branchles, flowers with carnose
texture, densely and prominently black punctate, and sessile anthers. However,
Cybianthus granulosus is easily separated from C. minuttjlorus by the abruptly
long-acuminate leaf apices, canaliculate petioles, the hyaline, serrate, and
punctate-lineate calyx margins, the erose corolla margins, acute anthers, and
pedicels 1.5-2.5 mm long.
35. Cybianthus flavovirens Pipoly, sp. nov. (Fig. 25). Typi;: PERU. San Martin:
Prov. Mariscal Caceres, Dtto. Tocache Nuevo, Palo Blanco near Fundo de Manuel
PiPOLY, Cybianthus in Ecuador and Peru
135
Fic. 25. Cybianthus jlcivovirem Pipoly. A. Habit, sliowing anguiate branchlecs and gradually
capering petioles. B. Abaxial leaf surface, showing sparse, rufous stellate tomentum. C.
Section of inflorescence, showing nodding flowers, widely deltate calyx lobes and corolla
with lobes constricted basally, prominent veins, and pusticulate surface. D. Branchlet, showing
rufous stellate tomentum. A-D, drawn from holotype, by Linda Ellis.
136 Sum 18(1)
Aranjo, 700-800 m, 1 Mar 1979 (stam. fl)J, Sihimke 10883 (iiolotypi;: MO; isotypes;
AMAZ, BRIT, F, TEX, US, USM).
Propter ramulos angulatos, peciolos graciare angustatos, inflorcscentiam racemosam, flores
nutanres, ancheras sessilia, C. imezi/elano valde arete affinis, sed ab ea lobis calycinis membranaceis
(non carnosis) acucis (nee rotundatis) seciis margmis hyalinis (nee opaeis), antheris obtusis
vel subacutis (non truncaris), porisque separatis (nee eonfluentibus), denique lobis eorollinis
membranaceis (non carnosis) pustieulatis (nee planis) triineatis vel emarginatisque (nee aeutisque)
praeclare distat.
Tree to 4 m tall. Branchlets lightly angulate, 2.5-3.5 mm diam., densely
rufous stellate-tomentose. Leaves alternate; blades chartaceous, elliptic, (12. 5-)
19-.30(-32) cm long, (4-)6-10 cm wide, apically subacuminate to acumi-
nate, the acumen 0.5-2 cm long, basally attenuate, dectirrent on the distal
end of the petiole, midrib somewhat elevated but canaliculate above, not
decurrent on petiole, prominently raised and densely rufous tomenose be-
low, the secondary veins 10-16 pairs, dull green above, pallid green below,
pellucid punctate above and below, sparsely rufous stellate-puberulent be-
low, the margin entire, opaque, regular, flat; petioles semiterete (1.6-)2-
2.5 cm, flat above, tapered, slightly thickend basally, not ptilvinate, gla-
brous above, sparsely rufous pubescent below, glabrescent. Staminate infloresa')ice
a lax raceme (7.5-)13-19(-40) cm long, the rachis and pedicels moderately
rufous tomentellous, glabrescent; floral bracts linear-lanceolate, 1 . 1-1.5 mm
long, 0.1-0.2 mm wide, apically attenuate, densely rufous tomentellous
below, glabrescent, the margin entire; pedicels cylindrical, 2.5-3.5 mm long
at anthesis, erect in bud, nodding in anthesis. Staminate flowers 4-merous,
membranaceous, nodding, bright yellow-green; calyx cotyliform, 1.5-1.9
mm long, the tube 0.6-0.8 mm long, the lobes widely deltate to triangu-
lar, 1.1-1.3 mm long and wide, apically acute, epunctate, medially thick-
ened, sparsely rufous puberulent, glabrescent, the margin hyaline, epunctate,
entire, minutely glandular-ciliolate; corolla subrotate, 2.8-3 mm long, the
tube 0.5-0.8 mm long, glabrous without, densely glandular-granulose within,
the lobes suborbicular to oblate, 1.5-2.2 mm long, 2.2-2.6 mm wide, apically
truncate to slightly emarginate, contracted basally, translucent, the three
veins conspicuous, glabrous without, prominently pusticulate and sparsely
glandular-granulose above but densely so toward base within, the margin
opaque, sparsely glandular-granulose, entire; stamens apparently sessile at
junction of corolla lobes and tube, the anthers sessile, very widely ovate-
obcordate, 0.4-0.6 mm long, 0.5-0.8 mm wide, apically obtuse to sub-
acute, basally cordate, the pores widely ovate, extending 1/2-3/4 anther
length, separate (not confluent), the connective epunctate, minutely red glandular-
papillate; pistiUode costate, subglobose, 0.8-1 mm long, 0.6-0.7 mm wide,
hollow, the stigma capitate, densely yellow glandtilar-papilalte. Pistillate
inflorescence: unknown. Fruit unknown.
PiPOLY, Cybianthus in Ecuador and Peru 137
Distribution. — Endemic to the junction of the westernmost Amazon Ba-
sin (Hylaea) with the foothills of the Peruvian Andes, from 100—800 m elevation.
Ecology and conservation status. — Cylnanthus flavovirens occurs in lowland
moist forests on terra firme. These forests are drier than sites with the same
general physiognomy farther to the north in Amazonas and Loreto. The few
sporadic collections may be the result of underexploration rather than rar-
ity, but owing to increasing pressure to cut forests for farming, this species
should be considered threatened.
Etymology. — The epithet refers to the bright yellow-green corolla, a unique
feature in the subgenus.
Paratypfs. PERU. Huanuco: Villa Isabel, Rfo Cuchara, 20 Sep 1961 (stam. fl),J. Schunke
5667 (F, K, iVIO, US, USiVI). Loreto: Mishuyacu, near Iquitos, 100 m, Oct-Nov 1929 (scam,
fl), G. Ktug 285 (F, US), (scam, fl), C. Kli/i^ 367 (F, US). Pasco: Prov. Oxapampa, Palcazu
Valley, Cabeza de Mono, 5-6 km W oflscosaci'n, 10° 12' S, 75° I4' W, 14-15 Apr 1983
(scam, fl), D. Smith 3 709 (MO, US, USM); Prov. Pasco, Palcazu Valley, Selva General, Cerro
de Pasco, Proyccto Especial Pichis-Palcazu, IND, 09° 50' S, 68° OO'W, 300-600 m, 13
Occ 1987 (stam. fl), G. Hartshorn i^t cil. 2996 (BRIT, MO, USM).
Cybianthus flavovirens is a member of a complex of taxa related to C. venezuelanus
Mez, as evidenced by the angulate branches, gradually tapered petioles, nodding
flowers, and sessile anthers. However, C, flavovirens is easily recognized by
its membranaceous perianth, acute calyx lobes with opaque margins, pusticulate,
bright yellow-green corolla lobes with truncate or emarginate apices, and
obtuse or subacute anthers with separate pores. This species has been con-
fused with Cybianthus cyclopetalus. However, Cybianthus flavovirens may eas-
ily be separated from that species by its angulate branchlets, racemose
inflorescences, large, yellow flowers, and deltate to triangular calyx lobes.
36. Cybianthus venezuelanus Mez in Engl., Pflanzenr. IV. 236(Heft 9):22T
1902. Type: VENEZUELA. Lara: Barquisimeco, San Felipe, 600 m,Jun 1846 (scam,
fl), N. Funck & L. SchliDi 678 (lhctotypf., here designaced: G-BOIS; isolectotypes:
BM, BR,G,LD, P W).
?eckia purpurea Rusby, Bull. New York Boc. Card. 4:405. 1907. Type: BOLIVIA. Wich-
our localicy daca, (scam, fl), A.M. Bang 2048 (holotype: NY).
Cybianthus egensis Mez in Engl., Pflanzenr. IV. 236(FIefc 9):222. 1902. syn. nov. Type:
BRAZIL. Amazonas: Near Ega [Teffe], Sep 1831 (scam, fl), E. Poeppig 2567 (holo-
type: W, F Neg. 31997; isotype: W) .
Cybianthus hrownii Gleason, Bull. Torrey Boc. Club 53:293- 1926. syn nov. Type: GUYANA
[BRITISH GUIANA]. Tumacumari, 18Jun-8 Jul 1921 (scam, fl), H. Gleason 159
(holotype: NY; lsoi-ype,: K).
Tree to 5 m tall. Branchlets angulate, (2.5— )3.5— 5 mm diam., densely ru-
fous tomentose. Leaves pseudoverticillate; blades chartaceous, oblanceolate
to elliptic, (l()-)17-27(-34)cm long, (3-)6-9(-ll) cm wide, apically acute
to acuminate, basally acute, slightly decurrent on the petiole, midrib slightly
138 SiDAlSd)
raised above, prominently raised below, the secondary veins 10—13 pairs,
rufous puberulent and smooth above, glabrescent, sparsely rufous puberu-
lent below, concentrated along the midrib and secondary veins, inconspicuously
pellucid punctate, the margin flat, entire; petioles slightly canaliculate, (15—)
20— 25(— 30) mm long, tapered, densely and minutely stellate rufous stel-
late puberulent. Staminate inflorescence: an erect, simple raceme, (7.5— )1 0.5—
18(— 23) cm long, sparsely rufous stellate puberulent; peduncle 1—3 cm long;
floral bracts coriaceous, lanceolate, 0.5-0.7 mm long, 0.2-0.3 mm wide,
apically subulate, densely rufous stellate puberulent, the margin entire; pedicels
cylindrical, 2.1—2.7 mm long, sparsely rufous stellate puberulent, glabres-
cent. Staminate floicevs 4-merous, carnose, nodding, green; calyx cotyliform,
0.9—1.1 mm long, the tube ca. 0. 1 mm long, the lobes widely ovate, O.cS—
1 mm long, 0.9—1 mm wide, apically rounded, densely rufous stellate pu-
berulent, sparsely and prominently black punctate, the margin opaque, coarsely
serrulate, densely glandular-ciliolate; corolla carnose, subrotate, 1 .6—1.8 mm
long, the tube quadrate, 0.5—0.6 mm long, glabrous, the lobes widely tri-
angular, 1.1—1.3 mm long, 1.5—1.7 mm wide, reflexed in anthesis, apically
actite, dorsally recurved, prominently and densely black punctate and gla-
brous without, densely glandular granulose throughout within, the mar-
gin slightly revolute, densely glandular-granulose; staminal tube inconspicuous,
hyaline, membranous, 0.5—0.6 mm long, aclnate to corolla tube, anthers
apparently sessile, cuadrate, 0.3-0.4 mm long, 0.5—0.6 mm wide, apically
truncate, basally trtmcate, leaning proximally at anthesis, the connective
prominently punctate dorsally, rufotis glandular-papillate apically; pistillode
obsolete or conical, 0.4—0.5 mm long, 0. 1—0.2 mm wide, hollow, glabrous.
Pistillate inflorescence as in staminate but (4.5— )8.5-l ()(-l 3) cm long; peduncle
1—2.5 cm long; floral bracts 0.5—0.7 mm long, 0.2—0.3 mm wide; pedicels
0.9—1.2 mm long. Pistillate flowers as in staminate but calyx 0.8—1.2 mm
long, the ttibe ca. 0.2 mm long, the lobes 0.6—0.8 mm long, 0.9— 1-1 mm
wide; corolla 1.2—1 .4 mm long, the ttibe 0.4-0.5 mm long, the lobes 0.7—
0.9 mm long, 1.1 — 1.2 mm wide; staminodial as in staminal tube, 0.4—0.5
mm long, adnate to corolla tube, the antherodes 0.2—0.3 mm long, 0.4—
0.5 mm wide; pistil obturbinate, 0.5—0.6 mm long, 0.3—0.4 mm wide, glabrotis,
the ovtiles 2—3, partially imbedded in the placenta. Fruit globose, 5— 7(— 9)
mm diam., black at maturity, the exocarp thick, juicy.
Distribution. — In moist forests rimming the Amazon Basin, from Guyana
through Veneztiela to the Andes of Colombia sotithward to Bolivia, 100—
1 ,00{)(-2,000) m. It is also known from the Choco Floristic Province of Colombia,
and may be expected in that corresponding region of Ecuador. The Boliv-
ian specimens cited below represent new distribution records.
Ecology and conservation status. — Cyhianthus venezuelanus is common in lowland
and premontane moist forests, and occasionally in premontane pluvial for-
PiPOLY, Cybianthus in Ecuador and Peru 139
ests. Ir is locally common and appears to do well in light gaps and forest
margins, thus, it should not be considered threatened.
Etymology. — The epithet refers to the the country from which the type
was collected, Venezuela.
Representative specimens examined. ECUADOR. Zamora-Chinchipe: Nangaritza Canton,
Pachicutza, Camino al Hito, Cordillera del Condor, 04" 07' S, 78° 37' W, 1,000-1,100 m,
19 Oct 1991 (fr), W. Palachs et al. 8313 (BRIT, MO, QCNE). PERU. Huanuco: Prov.
Leoncio Prado, Dtto. Rupa Rupa, Ynti, Rd. to Ri'o Rondos, 750 m, 24 Mar 1972 (stam.
fl),./. Sc/y/inke 3308 (AMAZ, F, G, MO, NY, US, USM). Loreto: Prov Maynas, Mishuyacu,
near Iquitos, 100 m, Oct-Nov 1929 (stam. fl), G. Ki//g 94 (F, NY, US); Peria Negra, near
Iquitos, 100 m, 20 Oct 1979 (bud), R Ayaia2I02 (AMAZ, MO, USM); QuebradaOrejon,
Purma, 29 Oct 1980 (fr), F. Aya/a et al. 2814 (AMAZ, MO, US, USM); Pumayacu, be-
tween Balsapuerto and Moyobamba, 600-1 ,200 m, Aug-Sep 1 933 (stam. fl), G. Kh/s^ 3188
(F, G, MO, NY, S, US). Madre de Dios: Prov. Tambopata, Tambopata Wildlife Reserve,
30 km S of Puerto Maldonado, 12° 15' S, 69° 17' W, 260 m, 14 Nov 1984 (stam. fl), H.
Young et al. 1 79 (MO, US, USM); Tambopata Tourist Camp at junction of Rfos Tambopata
and La Torre, 12° 49' S, 69° 43' W, 280 m, 22 Jul 1985 (ster.), A. Gentry et al. 51083
(CUZ, MO, USM), 27 Jul 1985 (ster.), A. Genttyetal 5 1400 (CUZ, MO, US, USM). BOLIVIA.
Santa Cruz: Prov. Florida, Parque Nacional Amboro, Sta. Rosa de Lima, 5-8 km N of
Cerca a La Cumbre, on path to La Playa, 1 7° 49.5' S, 64° 16' W, 2,000-2,100 m, 3-5 May
1993 (fr), /. Vargas et al. 2248 (BRIT, MO, USZ)
The perianth of the lectotype o( Cybianthus venezuelanus , with dense and
prominent black punctations, the truncate sessile anthers, large oblanceolate
chartaceoLis leaves, and short pedicels match the collections cited here. Cybianthus
venezuelanus , as treated here, is a polymorphic ochlospecies with many re-
gional variants. The type q{ Cybianthus brownii Gleason varies only in its
shorter pedicels and longer corolla lobes. Likewise, the type o{ Cybianthus
egensis Mez, in bud, has correspondingly shorter pedicels and corollas. Cybianthus
venezuelanus is easily recognized by the angulate branchlets, pseudoverticillate
leaves, rounded calyx lobes with opaque margins, and truncate anthers with
confluent pores. Its distribution, which rims the Amazon Basin, is unusual
in the Myrsinaceae but is common in other families, such as the Piperaceae
(R. Callejas, pers. comm.).
37. Cybianthus grandezii Pipoly, sp. nov. (Fig. 26). Typh: PERU. Lorf.to:
Prov. Maynas, Quebrada Yanayacu, entering from Aucayo, 25 Aug 1990 (stam. fl),
C. Grdndez. S. Vdsquez & M. F lores 1824 (holotype: MO; isotypics: AMAZ, US).
Quoad folia magna chartacea nervos secundarios tertiarioscjue laminares praeclare utrinque
conspicLia ad bases gradatim descrescentiaque necnon petiolos pulvinatos C. jensoni valde
affinis sed ab ea laminis ad bases obtusis auriculatisque (non cuneatis) manifeste desu|"ier
perpuncticulosis (nee epunctatis) desuper nitidus (nee sordidis) nerviis secundariis 18-24
(non 24-30) -jugis, 12-13.5 (non 6.5-9) cm latis, petiolis 0.5-1.4 (non 2. 1-5) cm longis,
pedicellis 0.3-0.5 (non 0.8-1.4) mm longis, bracteis florinis pedicellis 3 (non 6-7)-plo
longiores statim cognoscitur.
Shrub to 2 m tall. Branchlets terete, ca. 10 mm diam., rufous-lepidote.
140
SiDA l.Sd)
Fk ;. 2fi. Cyhianthi/s graiulezn Pjpoly. A. Habit, showing basally cruncace and somewhat auricukite
leaf bases. B. Adaxial, prominently perpuncticuiose surface. C. Abaxial, minutely rutous
lepidote surface. D. Portion ot inflorescence. E. Pistillate calyx, showing pustulate, black
punctate lobes with hyaline, erose-serrulate margins. F. Pistillate cotolla, showing pustu-
late abaxial surface. G. Branchlet apex. A— Ci, drawn from holotype by Linda Ellis.
Pipoi Y, Cybianthiis in Ecuador and Peru l4l
Leaves ahernate; blades chartaceous, oblanceolate, 34—36 cm long, 12—13-2
cm wide, apically subacuminate, basally truncate and appearing auriculate,
nitid above, pallid below, glabrous, and prominently black perpuncticulose
above, sparsely and minutely rufous-lepidote below, the midrib raised and
canaliculate above, prominently raised, black punctate-lineate and densely
rufous-lepidote below, the secondary vems 18-24 pairs, loop-connected
submarginally, the margin entire; petioles canaliculate, (0.5-)l-1.4 mm
long, 0.4—0.6 mm diam., pulvinate, densely lepidote. Staminate inflorescence
unknown. Pistillate inflorescence an erect, straight simple raceme, (8.5— )1 1—
14 cm long; peduncle 1-3 cm long; the rachis green, densely red glandu-
lar-papillate, black punctate-lineate; floral bracts membranaceous, linear-
lanceolate, 1.8-2 mm long, densely rufous lepidote; pedicels obconic, 0.5-1
mm long, densely glandular-papillate. Pistillate flowers chartaceous, creamish-
white; calyx cotyliform, 1.8-2 mm long, the tube ca. 0.5 mm long, the
lobes widely ovate, 1.3-1.5 mm long, 1.1-1.2 mm wide, apically acute,
densely and prominently black punctate, prominently translucent pustu-
late, the margin hyaline, irregular, erose-serrulate, epunctate, glabrous; corolla
subrotate, 2.2-2.4 mm long, the tube ca. 0.5 mm long, the lobes very widely
ovate 1.7-1.9 mm long and wide, apically rounded, densely and promi-
nently black punctate and translucent pustulate without, glandular-granulose
throughout and prominently black punctate within, the margin hyaline,
epunctate, minutely erose- crenulate, glabrous; staminodes 1.1-1.2 mm long,
staminal tube conspicuous, carnose, 0.4-0.5 mm long, elobate, densely glandular-
papillate, the filaments terete, proximally curved, 0.2-0.3 mm long, the
sterile anthers ovate, 0.5-0.6 mm long, and wide, the apically apiculate,
dehiscent by terminal confluent pores ca. 3/4 length, the connective prominently
black punctate; pistil obturbinate, 1.5-1.6 mm long, 0.5-0.6 mm diam.,
densely and prominently black punctate, translucent glandular-lepidote,
the ovary 0.5-0.6 mm long, the style 0.8-0.9 mm long, the stigma bi-
lobed, the lobes to 0.1 mm long, distally curved, the placenta subglobose,
with 2-4 ovules partially embedded. Fruit globose, 8-10 mm long and in
diam., the exocarp thick, juicy, purple-black at maturity.
Distribution. — Known only from Maynas Province, Dept. of Loreto, Peru,
at up to 125 m elevation; presumably endemic.
Ecology and conservation status. — Cybianthiis grandezii is restricted to pri-
mary lowland most upland terra firme forest over red lateritic clays. Label
data indicate it is rare, and given the valuable timber present in that forest
type, this species should be considered threatened.
Etymology. — It is indeed a pleasure to dedicate this species to Biol. Cesar Grandez,
professor of biology at the Universidad Nacional de la Amazonia Peruana
(UNAP), Iquitos, and authority on the systematics of Peruvian Flacourtiaceae.
Cesar is an ardent field worker, an excellent teacher and herbarium curator.
142 SiDA 18(1)
Para'iypf,: PERU. Loreto: Prov. Maynas, Drto. Fernando Lores, Panguana, 125 m, 6
Aug 1 991 (fr), S. AlcDc/i/n-I & M. R/madn 31219 (MO, IBE).
Cyhianthus grandezii is closely related to C. jensonii Pipoly, but is easily
recognized by its subsessile leaves with truncate bases. The perpuncticulose
and nitid adaxial leaf surface, and secondary veins raised prominently above
and below are also distinctive.
38. Cybianthus jensonii Pipoly, sp. nov. (Fig. 27). Typi;: PERU. Loreto: Prov.
Alto Amazonas: Andoa.s, Rio Pastaza near Ecuadorean border, 02° 48' S, 76° 28' W,
210 m, 14 Aug 1980 (fr), A. Gentry. R. Vasqi/ez & N. Jaramillo 29700 (holotyph:
MO; isoTYPns: AMAZ, NY, USM).
Quoad lolia magna charracea nervos laminare.s secundarios tertianosque praeclare utrinque
conspicLia ad bases gradacim aqdescrescentia i^eciolosque pulvinatos C. grandezi valde affinis sed
ab ea laminis ad bases cuneatis (nee obtusis auricularisque) epunccatis (non manifeste desuper
perpuncticulosis) desuper sordidis (non nitidis) nerviis secundariis 24-.^0 (non 18-24)-
JLigis, 6.5-9 (nee 12-1 3.5) cm latis, pctiolis 2.1-5 (non 0.5-1 .4) cm longis, pedicellis 0.8-1.4
(non 0..^-().5) mm longis bracteis florinis pedicellis 6-7 (non 3)-lilo longiores statim cognoscitur.
Treelet to 4 m tall. Branchlets terete, 7-9 mm diam., lenticellate, the bark
vertically ridged, densely rufous tomentose. Leai'es alternate; blades mem-
branaceous, oblanceolate, .33^9 cm long, 6.5—9 cm wide, apically subacuminate,
basally long-attenuate, dull green above and below, the veins 24-30 pairs,
buUate above, prominently raised below, the tertiary areoles prominently
raised below; glabrate above, moderately rubiginous furfuraceous lepidote
and sparsely pellucid punctate below, the margin entire, decurrent, gradu-
ally tapering to the petiole; petioles marginate, 2.1-5 cm long, to 0.5 cm
thick, glabrous, abruptly pulvinate, the pulvinus 1.3-2 cm above petiole
base. Staminate inflorescence: a simple, lax, axillary raceme 6.5—8 cm in bud;
the rachis, bracts and pedicels densely rufous tomentose; floral bracts mem-
branaceous, linear-lanceolate, 3^ mm long, 0.3-0.4 mm wide, apically subulate,
the margin entire; pedicels cylindrical, 0.8-1 .4 mm long, glabrescent. Staminate
flowers 4-merous; calyx membranaceous, cotyliform, 1.4-1.6 mm long, the
tube 0.2-0.4 mm long, the lobes widely ovate, 1-1.3 mm long, 1.2-1.5
mm wide, apically acuminate, densely and prominently black punctate, sparsely
rufous pubescent, the margin hyaline, flat, erose, epunctate; corolla (in bud)
subrotate, membranaceous, to 1.2-1.5 mm long, the lobes ovate, apically
obtuse, densely and prominently black punctate, glabrous without, sparsely
glandular-granulose within, the margin hyaline, flat, erose, epunctate, gla-
brous; stamens appearing epipetalous, the anthers sessile at the junction of
corolla tube and lobe, deltoid, ca. 0.7 mm long and wide, dehiscent by subapical
pores, the connectives red punctate medially; pistiUode, conic, hollow, ca.
0.5 mm long and 0.2 mm diam. Pistillate inflorescence as in staminate but 3-
6(-7.2) cm long; floral bracts 1-1.6 mm long, 0.2-0.4 mm wide, apically
long-attenuate; fruiting pedicels incrassate, 0.7-0.9 mm long, to 1.5 mm
PiPOi.Y, Cybianthus in Ecuador and Peru
143
Fic. 27. Cybtanthiis jemunit Pipoly. A. Habit, showing long-attenuate leat bases. B. Abaxial
leaf surface with prominently defined areoles. C. Portion of staminate inflorescence in bud.
D. Open staminate bud. E. Pedicel, calyx and fruit. A-D, drawn from Gentry et al. 33708.
E, drawn from Croat 19485. Figure drawn by Linda Ellis.
144 SiDA 18(1)
diam. Pistillate flowers as in staminate, calyx 1.4-1.6 mm long, the tube
0.2-0.4 mm long, the lobes 1-1.3 mm long, 1.2-1.5 mm wide; corolla,
staminodes and pistil unknown. Fr//// depressed-globose, orange, 5—6 mm
long, 6-7 mm diam., smooth, prommently pellucid punctate.
Distribution. — Endemic to lateritic slopes above riparian areas in the
Department of Loreto, Provinces of Alto Amazonas and Maynas, in the northern
Amazon Basin of Peru, 130-210 m.
Ecology and conservation status. — This species occurs in primary lowland
tropical varzea forest margin. It is surely a rare species, given the recent
massive collection effort at the sites during the Florula of the Biological Re-
serves of Iquitos project (Vasquez 1997), during which the species was not
relocated at the Explorama Inn (Indiana) site. Given its rarity, this species
should be considered threatened.
Etymology. — I dedicate this species to Peter Jenson, President of Explorama
Tours, conservationist and principal promoter of ecotourism in the Peru-
vian Amazon. Much of our knowledge regarding the biology of tropical
ecosystems in the Peruvian Amazon would not have been discovered were
it not for Peter and his associates' enthusiastic support, generosity and hos-
pitality at the company's biological reserves (Explorama Inn, Explorama Lodge,
and Explornapo Camp) which now serve as long-term ecological study sites.
Explorama Tours' properties, with their combination of research, public education
and collaboration with local communities and tourism, serves as the most
successful model tor tropical ecotourism known.
Parat^'pes. PERU. Loreto: Prov. Maynas, Explorama Inn, 2 km W of Indiana on Rio
Amazonas, 03° 30' S, 73° 02' W, 130 m, 12 Feb 1987 (scam, bud), A. Gentry et ^i I. 55708
(AMAZ, MO); Varadero de Mazan from Rio Amazonas to Rfo Napo, 22 Aug 1972 (fr), T.
Cnuit 19485 (AMAZ, MO, NY, USM).
Cybianthus jensonii is most closely related to C. grandezii Pipoly, but eas-
ily recognized by its long-attenuate leaf bases, epunctate, pallid abaxial leaf
surfaces, the secondary veins 24—30 pairs, narrower leaves with much longer
petioles, subobsolete pedicels 0.8—1.4 mm long, and longer floral bracts.
39. Cybianthus fosteri Pipoly, sp. nov. (Fig. 28). Type: PERU. Madre de Digs:
Prov. Manu, Atalaya, vicinity Hacienda Amazonia, 2-3 km W of village, across Rio
Alto Madre, 12'^^ 55' S, 71° 12' W, forested ridge, 600-900 m, 7 Dec 1983 (stam.
fl), R. Foster & T. Wachter 7254 (iiolotype: MO; isotypes: F, NY, USM).
Ob folia pseudoverticillata magna charcacea longipetiolata abrupte acuminataque, inflorescentia
longiracemosa, flores nutantes, coriaceosc]ue, antheras sessiles manifesce necnon dorso punctatas,
C. venez/zelano valde affinis sed ab ea ramulis teretes (non angulacis), foliis pscudoverticillatis
(non alternis), laminis denseque manifeste atro-punctatis et omnino prominens (non parceqe
plane subter atro lineato-punctatis) petiolatis canaliculatis (non marginatis ad bases abrupte
crassis (nee gracilis) lobis corollinis interius pustulatis (non planis), antheris ad apices rocundatis
(non truncatis) poris separatis (non conHuentibus) praeclare distat.
PiPOLY, Cybiantlius in Ecuador and Peru
145
Fig. 28. Cyhianthus fosten Pipoly. A. Habit, showing terete branchlets and pseudoverticillate
phyllotaxy. B. Portion of staminate inflorescence, showing pustulate corolla and apicalUy
rounded anthers with separate (not birimose) pores. C. Abaxial leaf surface detail, showing
punctations and sparse puberulence. D. Branchlet apex, showing tomentum. A-D, drawn
from holotype, by Linda Ellis.
146 SiDA 18(1)
Tree to 5 m tall. Branchlets terete, (6-)7-9 mm diam., densely terrugineotis
tomentose. Leaves pseudoverticiUate; blades chartaceous, oblanceolate, (26-)
28—34 cm long, 5.5-9(-H)) cm wide, apically abruptly acuminate, basally
long attenuate, midrib somewliat elevated above, prominently raised be-
low, the secondary veins 11 — 18 pairs, dull green above, pallid below, prominently
black punctate above and below, sparsely rufous puberulent below, the margin
entire, regular, flat; petioles canaliculate, 1 .6—2 cm, pulvinate, ferrugineous
tomentose, glabrescent. Stanihiate ijiftorescence a lax raceme 8—12 cm long,
moderately rufous lepidote, glabrescent; floral bracts lanceolate, 2—3 mm
long, 0.8—1 . 1 mm wide, apically acute, densely rufous lepidote; pedicels cylin-
drical, 3.5—5 mm long at anthesis erect in bud, nodding in anthesis, densely
rufous lepidote. Stcoiiinate flowers 4-merous, coriaceous, nodding, pale green;
calyx cotyliform, 1.2—1.3 mm long, the tube 0.5—0.6 mm long, the lobes
widely ovate, 0.7— 0.8 mm long, 1.1 — 1 .2 mm wide, apically rounded, densely
and prominently black punctate, sparsely rufous lepidote, glabrescent, the
margin stramineous, opaque, epunctate, erose-fimbriate, glabrotis; corolla
subrotate, 2-2.5 mm long, the tube 0.5—0.6 mm long, glabrous, the lobes
widely ovate, 2—2.1 mm long, 1.4—1.6 mm wide, apically acute, densely
and proniinently black punctate, sparsely rtilous lepidote without, glabrescent,
prominently pustulate and densely glandular-granulose throughout within,
the margin stramineous, erose, glandular-granulose; stamens apparently sessile
at junction of corolla lobes and tube, the anthers sessile, very widely ovate,
0.5—0.6 mm long, 0.8—0.9 mm wide, apically rounded, basally truncate,
the pores widely ovate, extending 1/2—3/4 anther length, separate (not conl^tient),
the connective prominently red ptmctate; pistiUode subglobose, 0.7-0.8
mm long, 0.3—0.4 mm wide, hollow, densely glandular-lepidote. Pistillate
inflorescence as in staminate but erect, 3—5 cm long; floral bracts 2—3 mm
long, 0.8—1.1 mm wide; pedicels 2.5—4 mm long in fruit. Pistillate flowers
as in staminate but calyx 0.8—1 mm long, the tube 0.2—0.3 mm long, the
lobes 0.5—0.7 mm long, 0.9—1.1 mm wide; corolla, staminodes and pistil
unknown. Fr/zit globose, 6—8 mm long and diam., red at maturity.
Distribution. — Known only from the type locality, presimiably endemic.
Ecology and conservation status. — Cyhianthi/s fosteri is endemic to one area
of the Manti Biosphere Reserve and National Park, one of the largest in
South America. It is a ridgetop species in lowland topical most forest. Given
the extension of the Reserve and the species narrow range, it should not be
considered threatened at this time.
Etymology. — This species is dedicated to Robin Foster, of the Smithsonian
Tropical Research Institute and a research associate of the Field iVIuseum of
Natural Flistory. Robin has served as one of the co-founders of the Rapid
Assessment Protocol, and is an expert on the reproductive biology oiTacbigali
(including Sclerolohiitin) of the Fabaceae.
PiPOi,Y, Cybianchus in Ecuador and Peru 147
Paratyfe: PERU. Madre de Dios: Prov. Manii, Aralaya, vicinity Hacienda Amazonia,
2-3 km W of village, across Rfo Alto Madre, 12° 55' S, 71° 12' W, 6()()-90() m, 7 Dec
1 983 (fr), R. Foster^G T. Wachter 7242 (BRIT, F, MO, NY, USM).
Cybianth//s fosteri is closely related to Cybianthus venez/ielanus , but is eas-
ily recognized by its terete branchlets, pseudoverticillate leaves prominently
and densely black punctate above and below, canaliculate petioles, pustu-
late corolla lobes, and rounded anthers with separate (not birimose) pores.
The pustulate corolla lobes are unique within the genus.
40. Cybianthus resinosus Mez in Engl., Pflanzenr. IV. 236(Heft 9):219.
1902. Typh: PERU. Loreto: Prov. Maynas, near Yurimaguas, without date (fr), E.
Poejipig 2428 (holotype: W; isotypc: P).
Tree to 1 5 m tall. Branchlets 2.5—3.5 mm diam., densely ferrugineous dendroid-
tomentose at first, glabrescent. Leaves pseudoverticillate; blades thinly co-
riaceous, elliptic, (ll-)15-21(-26) cm long, (4-)5.5~7(-l().8) cm wide,
apicaliy long-acuminate, the acumen 0.8— 1.5(— 2) cm long, basally cuneate,
decurrent on the petiole, midrib prominently elevated above and below,
decurrent to petiole base, the secondary veins (9-)l 1-16(-18) pairs, nitid
and glabrous above and below, inconspicuously pellucid-punctate (not vis-
ible when dried), the margin entire, irregular, flat, entire; petioles canaliculate,
l-2(-3) cm long, tapered, densely ferrugineous dendroid-tomentose, gla-
brescent. Staminate inflorescence a lax, simple raceme, 1 1-1 5 cm long, sparsely
rufous stellate puberulent; peduncle 0.9-l(-l-8) cm long; floral bracts
chartaceous, linear, 1—1.2 mm long, 0.2—0.3 mm wide, apicaliy subulate,
densely rufous stellate puberulent above and below, the margin entire, early
caducous; pedicels cylindrical, 2.1-1.7 mm long, sparsely puberulent, gla-
brescent. Staminate flowers 4-merous, membranaceous, nodding, greyish-brown;
calyx cotyliform, 0.9-1 mm long, the tube 0.1-0.2 mm long, the lobes
widely ovate, 0.6-0.8 mm long, 0.9—1 -2 mm wide, apicaliy acuminate, sparsely
rufous stellate puberulent, densely and prominently black punctate, the margin
hyaline, erose, short glandular-ciliate; corolla subrotate, translucent, 1.6-
1 .8 mm long, the tube cylindrical, 0.3—0.5 mm long, the lobes very widely
ovate, 1.2-1.5 mm long, 1.5-1.8 mm wide, flat, apicaliy obtuse, densely
and prominently black punctate and glabrous without, densely glandular-
granulose and pusticulate within, the margin irregular, glandular-granulose,
entire; stamens 0.7—0.9 mm long, the tube inconspicuous, membranaceous,
0.3-0.5 mm long, sessile, the anthers cuadrate, 0.2-0.3 mm long, 0.5-0.6
mm wide, apicaliy truncate, translucent, glabrous, the connective promi-
nently red punctate dorsally; pistillode cylindrical, 0.3-0.5 mm long, 0.2-
0.3 mm diam., hollow, densely punctate, glabrous, the stigma 3-lobed. Pistillate
inflorescence -AS in staminate but (l-)1.5-5 cm long; peduncle (0.3-)O.5-l(-
1.5) cm long; floral bracts 0.6-1 mm long, 0. 1-0.2 mm wide; pedicels slightly
148 SiDAl8(l)
obconical, 0.7—1. 1 mm long, erect in fruit. Pistillate flowers as in staminate
but 1-1.2 mm long, the tube 0.2— 0.3 mm long, the lobes 0.8-0.9 mm
long, 1-1.2 mm wide; corolla 1.3-1.5 mm long, the tube 0.3—0.5 mm long,
the lobes 0.8—1 mm long, 1.1—1.5 mm wide; stammodes as in stamens but
0.4-0.6 mm long, the tube 0.3-0.5 mm long, the antherodes ca. 0.1 mm
long, 0.2—0.3 mm wide; pistil obturbinate, 0.4—0.6 mm long, and in di-
ameter, the style very short, the stigma 3-lobed, the placenta cotyliform,
bearing 2 apically exposed ovules. Fr/iit globose, 5—7 mm diam. at matu-
rity, exocarp black, juicy, edible at maturity. Bisexual inflorescence ^s in pis-
tillate but a lax, simple raceme, or rarely a poorly formed panicle, 5-8(— 1 0)
cm long; pedtmcle 0.5-1 cm long; floral bracts 1-1.2 mm long, 0.2-0.3
mm wide; pedicels 1.2—1.5 mm long. Bisexual flowers as in pistillate, but
stamens as in staminate, 0.5-0.8 mm long, the tube 0.2-0.3 mm long, the
anthers ca. 0.3—0.5 mm long, 0.4—0.5 mm wide; pistil as in pistillate, conical,
0.4-0.6 mm long, and in diam. Bisexual jriiit globose, A~G mm diam. at
maturity, exocarp reddish-black, thin.
Distribution. — Once thought to be endemic to the Iquitos area of Loreto,
Peru, Cybianthus resinosus is now known (reported for the first time here),
from the Choco of Colombia, Amazonian Ecuador, Venezuela, with one disjunct
population in French Guiana {Oldeman 5272) growing at 1 00-200(-l ,300)
m elevation.
Ecology and conservation status. — Cybianthus resinosus is restricted to pri-
mary non-inundated forests on white sand (varillal). While it is locally abundant,
it should be considered threatened due to incre-asing habitat loss. The Ecuadorean
and Venezuelan populations are unusual because they occur in premontane
pluvial forest and wet forest on lower tepui talus slopes (on sandstone) re-
spectively, each containing numerous lowland elements. It may be expected
in the Rio Cenepa-Rio Santiago Drainage Basins, of Amazonas, Peru, an
area known to show the same environments with numerous pockets of sandstone.
Etymology. — The specific epithet refers to the highly nitid adaxial leaf surface,
giving it a laccjuercd, resinous appearance.
Specimens examined. COLOMBIA. Valle del Cauca: Bajo C;alima Concession, ca. 2")
km NW of Buenaventura, 9 km NW of San Isidro intersection on "Canalete," near gate,
5-45° slopes, 03° 59' N, 77" 08' W, 50 m, 13 Jul 1988 (ster.), D. Fciher-Langendoen &J.
Hurtado 1 757 (CUVC, MO). VENEZUELA. Territorio Federal Amazonas: Dept. Atabapo,
base of cliff and forest below slope of Cerro Huachamacari, 03" 39' N, 65° 43' W, 1 ,000-
1,300 m, 5 Mar 1985 (stam. 11), R. Liesiier 18302 (BRIT, MO, VEN). ECUADOR. Napo:
Canton El Chaco, Rfo CiranadiUo, Campamento de INECEL, "Codo Alto," 00" 08' S, 77"
28' W, 1,300 m, 13-15 Sep 1990 (fr), W. Pa/acm 5389 (MO, QCNE). PERU. Huanuco:
Prov. Pachitea, region of Pucailpa, W part of Sira Motintains and adjacent lowland, ca. 24
km SE to 26 km ESE of Puerro Inca, from Campamento Sira, 09" 28' S, 74° 17' W, SE to
valley of Rfo Negro, 750 m, 29 May 1988 (fr), B. Wal/mfcr 14-29388 (BRIT, MO, W,
WU). Loreto: Prov. Maynas, Iquitos, May 1925 (stam. fi), G. Tessiiu/nii 3143 (NY); Casen'a
PiPOLY, Cybianthus in Ecuador and Peru 149
Mishana, 30 km SW of Iquitos, Callicebus Biological Reserve, 4 km S of Mishana, 19 Aug
1980 (stam. fl), R. Foster 4404 (F-2 sheets, NY); Mishana, l6 Aug. 1978 (pist. fl, h),J.
Ramtrez 17 (AMAZ, MO); Mishana, Rfo Nanay, 03° 50' S, 73° 30' W, 140 m, 16 Aug
1978 (fr),./. Ramiyez 132 (AMAZ, MO); Mishana, along Rio Nanay, 03° 55' S, 73° 35' W,
150 m, 20 Jan 1985 (fr), R. Va'squez & N. Jaramillo 6126 (AMAZ, MO, NY); AUpahuayo,
Estacion IIAP, 04° 10' S, 73° 30' W, 150 m, 5 Jun 1985 (bud), R. Va'squez et al. 6551
(AMAZ, BRIT, MO, NY), 29 May 1990 (fr) R. Vdscjuez et al. 13764 (AMAZ, MO, USM),
16 Aug 1990 (pist. fl, fr), R. VascjM'z & H. Jaramillo 14224 (AMAZ, BRIT, F, MO, NY,
TEX, US, USM), 4 Dec 1990 (fr), R. Va'squez & N. Jaramillo 15237 (AMAZ, BRIT, E, MO,
US, USM), 150-180 m, 29 May 1991 (ster.), R. Va'squez & N. Jaramillo 166H1 (AMAZ,
BRIT, MO, USM), 150 m, 23 Mar 1992 (ster.), R. Vasquez et al. 17996 (AMAZ, BRIT,
MO, USM); AUpahuayo, ca. 26 km along Iquitos-Nauta Rd., 130 m, 25 Aug 1988 (fr), H.
van cler Werff 10273 (AMAZ, MO); Mishuyacu, near Iquitos, 100 m, Sep 24-28 1929 (fr),
E. Ktllip & A. Smith 29873 (E, US), May-Jun 1930 (bisex. fl), G. Klug 1384 (F, NY, US);
Quistococha, 200 m, 27 May 1978 (pist. fl, fr), A. Gentry & N. Jaramillo 22314 (AMAZ,
MO); Altura de Pina Negra, SW of Iquitos, ca. 3-4 km past Quistococha, 200 m, 19 Nov
1975 (fr); Caseri'o de Urcumirafio, Rfo Napo, path from settlement to tall foresr, 120 m, 8
Oct 1979 (fr), C. Diaz ON. Jaramillo i486 (MO, NY); Pena Blanca, on Rfo Itaya, 110 m,
19 Sep 1929 (fr), E. Ktllip & A. Smith 29(>72 (E, US); Between Yurimaguas and Balsapuerto
(lower Rfo Huallaga basin), 1 35-1 50 m, 26-31 Aug 1929(fr)£. Ktllip & A. Smith 28110
(F, NY, US); Prov. Requena, Dtto. Sapuena, Jenaro Herrera, Rfo Ucayali, 04° 55' S, 73°
40' W, 160 m, 16 Aug 1994 (stam. fl), R. Ortiz et al. 101 (AMAZ, BRIT, MO); without
locality, except "in Peruvia subandina, without date (fr), E. Poeppig s.n. (L).
Cybianthus resinosus is most closely related to C. penduliflorns Mart., but is
easily separated from it by the inconspicuously punctate leaves, longer pedicels,
flat corolla lobes and calyx lobes with acuminate apices and erose, short-
ciliate margins. The adaxial prominently raised midrib decurrent to the petiole
base is unique within the subgenus. The fruit oi Cybianthus resinosus is also
smaller, black and has a thick exocarp, and it inhabits terra firme forests on
white sand whereas C. penduliflorus is an igapo species.
41. Cybianthus fuscus Mart., Flora 259- 1841. Typr: BRAZIL. MatoGrosso:
"Prope rivum Cochim in Cujaba," May (pist. fl), P. da Silva Manso s. n. (i iolotyfi;: M)
Shrub or small tree to .3 m tall. Branchlets terete, 2-.3 mm diam., densely
dendroid and stellate rufous glandular-tomentose, tardily glabrescent. Leaves
pseudoverticiUate; blades chartaceous to coriaceous, very narrowly oblan-
ceolate or very narrowly oblong, (13-)l6-25(-30) cm long, 2-4(-6) cm
wide, apically long acuminate-attenuate, the attenuated portion 1-2 cm
long, terminating in a minute rounded tip, basally long acuminate-attenu-
ate, the attenuated portion 1.5-2 cm long, giving the petioles appearance
of being longer, fully decurrent on petiole to pulvinus; midrib prominently
elevated above, decurrent to petiole base, the secondary veins 12-25 pairs,
somewhat to deeply impressed, the leaf appearing subbullate to buUate above,
prominently raised and loop-connected below, somewhat nitid and glabrous
above, pallid, rufous papillate and conspicuously black punctate and punc-
tate-lineate below, the margin entire, flat, glabrous; petioles somewhat marginate,
1^'* SiDA 18(1)
5-l()(-12) mm lon^ij;, with a basal pulvinus. Staminate inflorescence: a lax raceme
(2.5-)5-8(-19) cm long; peduncle 0.8-1 .5 cm long; floral bracts chartaceotis,
linear, 1 .2-1.5 mm long, 0.3 mm wide, apically subidare, densely and promi-
nently rufous papillate; pedicels 3.5-5 mm long, densely papillate . Stami-
nate flowers chartaceous, 4-merous; calyx cotyliform, 0.8-l(-l .8) mm long,
the tube ca. 0.2 mm long, the lobes very broadly ovate or linear-lanceolate,
0.5-0.8(-l .6) mm long, 0.6-0.8 mm wide, apically acute to acuminate to
attenuate, medially thickened, densely and prominently red and black punctate
medially, with a few scattered rufous papillae, the margin scarious, highly
erose, densely glandular-ciliate; corolla subrotate, 2-2.3 mm long, the tube
0.6-0.8 mm long, the lobes suborbicular, unequally divided, 1 .4-1.6 mm
long, 1.3-1.7 mm wide, apically broadly rounded, densely and prominently
orange punctate without, densely glandular-granulose throughout within,
the margin often revolute at maturity, irregular, entire, glandular-granulose;
stamens ca. 1 mm long, the filaments developmentally fused to the corolla
tube for their entire length (the stamens appearing epipetalous), 0.6-0.8
mm long, the anthers very widely ovate, 0.4-0.5 mm long and wide, the
apically acute, basally cordate, apically dehiscent by termmal pores, confluent
at anthesis, ventrally sparsely rufous papillate basally, dorsally densely ru-
fous papillate and sparsely but prominently orange punctate; pistillode absent
or highly reduced, ca. 0.5 mm long. Pistillate inflorescence as in staminate
but 3-10(-l4) cm long; peduncle 6-10 mm long; pedicels 1.5-4 mm long.
Pistillate flowers as m staminate bur 0.7-0.9 mm long, the tube ca. 0. 1 mm
long, the lobes oblate, 0.6-0.8 mm long, 1-1.2 mm wide; corolla as in staminate
but 1.4-1.8 mm long, the tube ca. 0.5 mm long, the lobes ovate to subor-
bicular, 1.1-1.7 mm long, 1.0-1.2 mm wide, the staminodes resembling
stamens but with antherodes 0.3-0.4 mm long and wide; pistil ellipsoid,
1-1 .2 mm long, 0.6-0.8 mm diam., the stigma capitate, 3-4-lobed, densely
translucent glandular-lepidote, ovules 2-3, immersed in the placenta. Fruit
globose, 5-7 mm long and diam. at maturity, densely and prominently punctate,
with a few persistent translucent Icpidote scales.
Distribution. — As here recognized, Cybianthus fnsciis occurs from the Guianas,
to Venezuela, Colombia, Ecuador, Peru, Bolivia, and their corresponding
frontiers with Brazil. Cybianthus fliisciis rims the Amazon Basin, from 100-
200 m elevation. As stated earlier, this is an infrequent distribution, as in
Cybianthus Venezuela nus.
Ecology and conservation status. — Cybianthus fuscus occtirs in primary forest
on terra firnie, especially on steep slopes, near water courses. It is a locally
infrequent element of the understory but does not appear to be threatened
at this time.
Etymology. — The epithet refers to the color of the tomentum of the branchlets,
pedicels and calyx.
PiPOLY, Cybianthus in Ecuador and Peru 151
Represenrarive specimens examined. ECUADOR. Napo: Small area of non-inundated
forest, ca. 60 km upriver from Nuevo Rocafuerte, 1 3 Sep 1977 (fr), R. Foster 361H (F, USM).
PERU. Loreto: Prov. Alto Amazonas, Capahuari Sur (Petroleum Camp), 02° 51' S, 76°
20' W, 200 m, 25 Mar 1982 (fr), R. Va'squez et at. 3065 (AMAZ, MO, US); Prov. Maynas,
Rio Yavari, Petropolis, 3 km from Rio Amazonas, 8 Sep 1976 (fr),/. ReviIL/ 1302 (AMAZ,
BRIT, MO); 15 km from roadside along Rd. between UNAP Agricultural Experiment area
and Escuela Forescal Vivero, 9 Feb 1968 (pist. fl), D. Simpson &J. Schutike 647 (F, USM);
Puerto Almendras, 03° 45' S, 73° 25' W, 122 m, 7 Dec 1982 (pist. fl), R. Vasquez & N.
Jaramillo 35! I (AMAZ, MO, NY).
Cybianthus fusa/s is a widely defined, infrequent, but widely distributed
species, and is most variable with regard to leaf size and inflorescence stat-
ure. Populations in Ecuador and Peru are almost identical to specimens known
from the Guianas, Bolivia, and Brazil in the northeast portion of Amazonas
state, near the border of Territorio do Roraima. Populations matching the
type have leaves much smaller than the Ecuadorean and Peruvian popula-
tions do, and are more like those of the SE Amazon Basin. While Cuiaba is
located at the northern extreme of the Pantanal Region, it is not entirely
clear where the exact type locality was. If the type locality in what was Cuiaba
Province, was north of the Chapada dos Parecis, then it would be at the
headwaters of the Rio Juruena or Rio Teles Pires, both of which dump into
the Rio Tapajos, then to the Rio Amazonas. If the locality was west of Cuiaba,
toward the Bolivian border, streams there form part of the headwaters of
the Rio Mamase, a branch of the Rio Madeiras, which empties into the Amazonas
near Manaus. In either case, the type locality would be at the vety edge of
the Amazon Basin sensu strkto and it would not be surprising to see the species
in other parts ot the Basin. Therefore, while populations from the type lo-
cality are slightly smaller in stature, the leaves are more chartaceous, and
the inflorescences shorter, there is good evidence to show that they are part
of a large polymorphic ochlospecies complex, of which the populations in
Ecuador and Peru represent a commonly encountered morphotype. The same
pattern of variation seen in this species is seen in many Piperaceae (R. Callejas,
pers. comm.).
Cybianthus fuscus appears to be closely related to C. a/neifolius Mart, (in-
cluding C. indecorus Mez), a vicariant species from SE Brazil. The unique
indument, pedicels obconic in fruit, and striking leaves with very long and
attenuate apices and bases allow for easy recognition oi Cybianthus fuscus.
42. Cybianthus cyclopetalus Mez, Bull. Herb. Boissier, Ser. 2, 5:5.33- 1905.
Type: BRAZIL. Ama/.onas: near Jurua, Miry, Sep 1903 (stam. fl), E. Vie 5840 (uo-
lotype: B -destr.; leciotype, here designated: FFBG; isolectotypes: G, K, MG).
Shrub to 1.5 m tall. Branchlets terete, 1.5-2.5 mm diam., densely rufous
stellate tomentulose, the tomentum appressed. Lmves loosely pseudoverticillate;
blades chartaceous, elliptic to oblanceolate, (9.5-)l 2.5-l6(-20.5) cm long.
'^- SlDA 18(1)
3-5-5(-7) cm wide, apiailly subacuminare to acuminate, basally cuneate,
decurrcnt thtoughout petiole length, midtib depressed above, prominently
raised below, decurrent to base of petiole, the secondary veins 10-15 pairs,
dull and glabrous above, dull and sparsely rufous puberulent below, con-
centrated along the midrib and the secondary veins, prominently peipuncticulose
and black punctate-lineate, the margin slightly revolute upon drying, ir-
regular, entire; petioles marginate and canaliculate, (1-) 1 .5-2(-3) cm long,
tapered, sparsely stellate rufous puberulent, glabrescent, Staiiiniate infhmcence
an erect, simple raceme, {4-)9-l 3 cm long, sparsely rufous stellate puberulent;
peduncle (().6-)().8-l (-1 .3) cm long; floral bracts coriaceous, linear-lanceolate,
1.3-1.3 mm long, 0.2-0.3 mm wide, apically attentuate to a rounded tip,
glabrous above, densely and minutely rufous stellate tomentulose below,
the margin glabrous, entire; pedicels cylindrical, 3.7-5 mm long, sparsely
rufous stellate tomentulose, glabrescent. Stumhhite flowers 4-merous, coria-
ceous, nodding, green; calyx 0.9-1. 1 mm long, the tube 0.2-0.3 mm long,
the lobes linear-lanceolate, 0.7-0.8 mm long, 0.2-0.3 mm wide basally,
apically long-attenuate, densely and prominently red and black punctate,
sparsely rufous stellate puberulent, glabrescent, the margin irregular, erose,
minutely ciliolate; corolla subrotate, 1 .5-1 .8 mm long, the tube 0.6-0.7
mm long, the lobes very widely ovate, 0.9-1 . 1 mm long, 1 .3-1 .7 mm wide,
apically emarginate, densely and prominently red and black punctate, gla-
brous without, densely glandular-granulose througout within, the margin
irregular, entire, flat, densely glandular-granulose; staminal tube inconspicuous,
adnate to corolla throughout, 0.6-0.7 mm long, the filaments short, thick,
0.1-0.2 mm long, glabrous, the anthers widely ovate, 0.5-0.6 mm long,
0.4-0.5 mm wide, apically and basally trunctae, the connective epunctate,
glabrous; pistillode subglobose, 0.3-0.4 mm long, 0.3-0.4 mm diam., hollow,
densely translucent glandular-lepidote, the stigma obsolete. Pistillate inflorescence
as in staminate but 1-6.5 cm long; peduncle 1-1.5 cm long; floral bracts
0.9-1 . 1 mm long, 0. 1-0.2 mm wide; pedicels 1.5-2.5 mm long. Pistillate
flowers as in staminate but calyx 0.8-1 mm long, the tube 0.1-0.2 mm long,
the lobes, 0.7-0.8 mm long, 0. 1-0.2 mm wide basally; corolla, staminodes
and pistil unknown. Fr///V globose, 0.3-0.5 mm long and wide, green, exo-
carp thin, red punctate.
Distribution. — Cybianthiis cyclopetalns is restricted to the western Amazon
Basin of Brazil and southeastern Peru, to 290 m elevation.
Ecology and conservation status. — This species is retricted to varzea or "tahuampa"
habitats, subject to inundation. Rapid development along the rivers in the
Amazon Basin changes its flow and may effect these populations. There-
fore, it should be considered threatened.
Etymology. — The specific epithet refers to the very widely ovate petals of
the species.
PiPOLY, Cybianchus in Ecuador and Peru 153
Represenracive specimens examined. PERU. Madre de Dios: Prov. Tambopara, Tambopaca
Wildlife Reserve, 30 km S of Puerto Maldonado, 12° 15' S, 69° 17' W, 260 m, 9 Nov
1984 (stam. fl), H. Yoiiug etal. 146 (MO, US); Tambopata Reserve, at mouth of Ri'o Orbigny,
12° 50' S, 69° 17' W, 250 m, 6 Mar 1981 (fr), A. Gentry & K. Young 32028 (MO, USM);
Along trail Irom large laguna at end of Swamp Trail, Explorer's Inn, near confluence of Rfo
Tambopata and Rio La Torre, Reserva Tambopata, 12° 50' S, 69° 20' W, 39 km SW of
Puerto Maldonado, 14 Oct 1985 (stam. fl), S. Smith et al. 738 (US), (stam. fl), D. Bell 101
(US); Explorer's Inn, Permanent Plots, Tambopata Reserve, 12° 50' S, 69° 17' W, 290 m,
1 8 Sep I 994 (stam. fl), R. Vchqiiez et cl. 19132 (AiMAZ, BRIT, CUZ, MO) .
Cybianthus cyclopetalus is most closely related to C. restnosus Mez, but can
immediately be separated from it by the thinner branchlets, dull, chartaceous
leaves, and the unique linear-lanceolate calyx lobes with long attenuate apices.
43.CybianthuspenduliflorusMart.,Nov. Gen. Sp. PI. 3:87. 1831 [1829}.
Cylmitithm pencliilniiis A. DC, Trans. Linn. Soc. London, Bot. 17:104. 1834 [orth.
var.}. Cyhicinthin pendiflorin A. DC, Prodr. 8:1 17. 1844 [orth. var.]. Tyfi;: BRAZIL.
Amazonas: Prov. Rio Negro, near Ega, 170 m, without date, (stam. fl), C. Marti//.f
s.n. (uolotype: M).
Cyhicinthiis riiacrophyllus Miq. in Mart., El. Bras. 10:292. 1856. PI. 36. syn. nov. Peckla
nuicrophylla (Miq. in Mart.) Kuntze, Revis. Gen. PI. 2:402. 1891. T^Ti:: BRAZIL. Amazonas:
Near Ega, without date, (pist. fl), E. Poepprg 2709 (i.uctotype, here designated: W).
Shrub or small tree to 4 m tall. Branchlets terete, 2.5-3.5 mm diam., stel-
late rufuous tomentose, glabrescent. Leaves alternate; blades chartaceous,
elliptic to oblanceolate, (8-)l().5-19(-28.4) cm long, apically acute, ba-
sally broadly acute, slightly decurrent on the petiole, midrib raised above
and below, the secondary veins (10-)12-15(-23) pairs, nitid above, pallid
below, glabrous, densely black punctate, the margin flat, entire or bearing
a few rough serrulations; petioles semiterete, (0.5-)0.8-1.2(-1.5) cm long,
tapered, glabrous. Staminte inflorescence a lax, simple raceme (8-)l()-l4(-2())
cm long, densely rufous puberulent; peduncle (().5-)l-1.2(-1.5) cm long;
floral bracts membranaceous, linear-lanceolate, 1-1.2 mm long, 0. 1-0.2 mm
wide, apically attenuate, densely rufous puberulent, the margin entire; pedicels
cylindrical, l-2(-2.5) mm long, densely rufous puberulent, glabrescent.
S laminate flowers 4-merous, erect, membranaceous green; calyx cotyliform,
0.8-1.1 mm long, the tube 0.1-0.3 mm long, the lobes ovate, 0.4-0.6 mm
long, 0.6-0.9 mm wide, apically obtuse, densely and prominently black
punctate, the margin crenulate, long glandular-ciliate; corolla subrotate,
1.2-1.5 mm long, the tube 0.3-0.4 mm long, the lobes very widely ovate,
0.9-1.2 mm long, 1.2-1.5 mm wide, apically obtuse to rounded, densely
and prominently black punctate, glabrous without, densely glandular-granulose
throughout within, the margin involute, densely glandular-granulose, en-
tire; stamens 0.7-0.9 mm long, the tube completely adnate to corolla tube,
0.3-0.4 mm long, the anthers ovate, 0.4-0.5 mm long, 0.3-0.4 mm wide,
apically acute to obtuse, basally cordate, the connective prominently black
154 SiDA 18(1)
punctate clorsally; pistillode obsolete. Pistillate in florescence as in staminate
but (2.5-)4-8(-ll) cm long; peduncle (().3-)().5-l cm long; fioral bracts
1-1.2 mm long, 0.2-0.3 mm wide; pedicels 0.2-0.5 mm long. PistilUite
flowers as in stammate but calyx 1.2-1.5 mm long, the tube 0.3-0.4 mm
long, the lobes widely ovate, 0.9-1.2 mm long, L.2-1 .5 mm wide, apically
rounded, corolla and staminodes unknown; pistil conical, 1 .2-1 .3 mm long,
0.9-1.1 mm wide, the stigma 4-lobed, the placenta cotyliform, ovules 4,
naked. Fr/fit 6-10 mm long and in diam., the exocarp thin, densely black
punctate. Bisexual inflorescence: as in staminate but 6-15 cm long; peduncle
0.8-1.2 cm long; floral bracts 1-1.2 mm long, 0.1-0.2 mm wide; pedicels
1-1.2 mm long. Bisexual (lowers as in staminate but calyx 0.7-1 mm long,
the tube 0. 1-0.2 mm long, the lobes 0.6-0.9 nnm long, 0.5-0.8 mm wide;
corolla 1.2-1.5 mm long, the tube 0.3-0.4 mm long, the lobes very widely
ovate, 0.9-1.2 mm long, 1.2-1 .5 mm wide, the margin involute, densely
glandular-granulose, entire; stamens 0.7-0.9 mm long, the tube completely
adnate to corolla tube, 0.3-0.4 mm long, the anthers ovate, 0.3-0.4 mm
long, 0.3-0.4 mm wide, apically acute to obtuse, basally cordate, the con-
nective prominently black punctate dorsally; pistil conical, 0.9-1.1 mm
long, 0.7-0.9 mm diam., the style 4-lobed, the placenta cotyliform, ovules
3, naked. Bisexual fruit globose, 4-6 mm long and in diam., the exocarp
thin, pellucid punctate.
Distribution. — Cyhianthus pendidflorus is known from Brazil, Peru and Bolivia,
100-200 m. The species is not known from Ecuador, but may be expected
anywhere the habitat is appropriate within the Ecuadorean Amazon.
Ecology and conservation status. — Cybmnthuspenduliflorus is endemic to igapo
habitats, and withstands flooding. As a small shrub, it grows on riverbanks
and on hummocks, behind Triplans (Polygonaceae) and other shoreline plants.
At this time, it is not considered threatened.
Etymology. — The specific epithet refers to the lax habit ot the inflorescence,
pendent in the field.
RL-pre.sentativc specimens cxiimiiit'd. PERU. I.oreto: Prov. Maynas, Quistococha, 100
m, 1 Feb 1979 (fr), F- Maid 1623 (AMAZ, MO), Quisrococha, 00° 45' S, 73° IV W, 122
m, 27 Alii,' 19«7 (scam. H), R. Vauj/zez & N. Jaramillo 9461 (AMAZ, MO, USM); Lower
Rio Momon, tributary of RTo Nanay, near Iquitos, 8 Dec 1979 (bud), A.Juius & C. Ddvidsim
9717 (AMAZ, CAS, MO), Near Momoncillo, 1 6 Nov 1976 (stam. i\)J. Rev/t/a 1826 (AMAZ,
F, MO); Dtto. Iquitos, caseri'o near Nina Rumi, on Rfo Nanay, 23 Feb 1976 (fr),J, Rei'illa
1S7 (AMAZ, F, MO, USM); Vicinity of Iquitos, 10 Sep- 12 Oct 1976 (bud),/ Revii/a
1442 (AMAZ, BRIT, MO, USM); Morona Cocha, near Iquitos, 100 m, l4 Dec 1962 (fr),
_/, Sdwuk' 6268 (AMAZ, F, M(3), UC:LA, US, USM); Puerto Almendras, 03" 48' S, 73° 2V
W, 122 m, 17 Aug 1983 (bud), /^. Vasquez & N. Jc/nm/tto 4285 (AMAZ, MO, NY, USM);
Naiica, Quebrada Saragoza, 04° 29' S, 73° 35' W, 1 50 m, 10 Jan 1 988 (fr), R. Va'u/urz & N.
Janimillo 10339 (AMAZ, MO, US, USM); Iquitos and vicmity, 1 I Oct 1929 (bud), U.
Willhiim 3(-)76 (F). BOLIVIA. Santa Cruz: Velasco Prov., Campamcnto El Refugio, along
Rio Paragtia, SF of the house, M° 46' 09" S, 61° 02' 1 I " W, 240 m, 1 1 Oct 1994 (H bud),
PiPOLY, Cybianrhus in Ecuador and Peru 155
R. Giiillm & G. Scilvatierni 2290 (BRIT, MO, USZ); Campamento La Toledo, 1 ,()()() m E of
the house, 14° 42' S, 61 ° 09' W, I6()m,21 Oct 1994(sram. fl), /?, G//!!le>i&R. Chore2439
(BRIT, MO, USZ), 1 km W of camp, on canoe route to Campamento Toledo, 14° 45' 51"
S, 61° 02' 22" W, 30 Jan 1995 (fr), R. Guillen et al. 3114 (BRIT, MO, USZ).
ACKNOWLEIXiMF.NTS
Support for my studies on the Flora of Peru was provided by a generous
grant from the Andrew W. Mellon Foundation (during my tenure at the
Missouri Botanical Garden), which is gratefully acknowleged. I would also
like to thank the John D. and Catherine T. MacArthur Foundation for their
support of fieldwork in Peru as part of the Documentation of Neotropical
Plant Diversity Project, which I directed from 1990-1994. Thanks are also
due to Michael Dillon (F) and the late Al Gentry (MO) for their collabora-
tion with the Flora of Peru project, to Enrique Forero (COL) and Warren D.
Stevens (MO) whose advice was invaluable. Many thanks to Jon Ricketson
(MO), then project coordinator and now colleague in Myrsinaceae system-
atics and authority on the systematics of Parathesis. For their warm hospi-
tality and collaboration, whether in the field in the herbarium, I express
my sincere gratitude to Rodolfo Vasquez (MO), Camilo Dfaz (MO), Nestor
Jaramillo (MO), Cesar Grandez (UNAP), Rosa Ortiz de Gentry (MO), Martin
Timana (TEX), Abundio Sagastegui (HUT), Jose Mostacero (HUT), Joaquina
Alban (USM). My volunteer at MO, Catherine Mayo, assisted in the move-
ment of specimens and recording of determinations. Studies on Cyhianthus
subgenera Microconomorpha. Conomorpha. Triadophora and Cornomyrsine were
completed during my tenure as a Postdoctoral Fellow in the Department
of Botany, National Museum of Natural History, Smithsonian Institution.
My continued status as a Smithsonian Research Associate has permitted me
to continue studies in the group, particularly in their strong Peruvian and
Colombian collections. I thank Linda Ellis, Peggy Duke, Juan Pinzon, and
Linny Heagy for the skillfull illustrations of the new taxa described herein.
I am most grateful to the directors and curators of herbaria cited, especially
F, G, GB, K, MO, NY, S, and US, for their patience and work in arranging
for loans and various transfers of material. Finally, to Jon Ricketson (MO),
and Ted Barkley (KSC) for their painstaking review of the manuscript and
helpful comments, and to Barney Lipscomb for his unselfish, time-consuming,
and meticulous work in editing, I owe my sincerest gratitude.
REFERENCES
ACiOsriNi, G. 1 970. Notes on Myrsinaceae I. The generic assignment o': Conomorpha sodiroana
Mez, Ardisici cnnbigua Mart, and related species. Phytologia 20:401-403.
1 972. A revision ot the genus Cyhianthus subgenus Conomorpha (Myrsinaceae).
Ph. D. Thesis. The City University of New York, New York.
1980. Una nueva clasificacion del genero Cyhianthus (Myrsinaceae). Acta
Biol. Venez. 10:129-185.
156 SiDA 17(4)
CuLLEN, J. 1978. A |irfliminary SLirvcy ol pryxis (vernation) in tlic Angiospernis. Notes
Royal Bot. Ciard. F.dinburgh 37:l6U2l1.
Faun, A. 1979. Secretory tissues in plants. Academic Press. New York.
Gessnf.r, F. & G . Volz. 1951. Die kutictiia der hytlropoten von Nyz/ip/jae^/. Planta ,t9: 1 7 1-174.
Gross, J. 1972a. Die Uiltblatter der Nymphaeaceen. Ber. Detitsch. Boc. Ges. 45:454-458.
1972b. Die Haustoren der Nymphaeaceen. Ber. Dcutsch. Bot. Ges. 45:459-466.
Halle, F., R.A. Oi.di.man, and P.B. Tomlin.son. 1978. Tropica! trees and forests. An archi-
tectural analysis. Springer- Verlag. New York.
HiCKLY, L. 1984. A revised classification of the architecture ol dicotyletlonous leaves. In:
C.R. Meccalte and F. C^halk, eds. Anatomy ol the Dictyledons. Vol. 1 . Systematic Anatomy
of Feaf and Stem, With a Brief History ol the Stibject. Clarendon Press. Oxh)rd, FJ.K.
Pp. 25-.^9.
HUBER, O. 1995. Vegetation. In: J. Steyermark, P. Berry, and B. Hoist, eds. Flota ol the
Venezuelan Guayana. Vol. 1 . Introdtiction. Missotiri Botanical Garden antl limber Press.
St. Louis. Pp. 97-160.
FiNDLi-Y, J. 1 848. Illustrated dictionary ot botanical terms. Fxcerpt from illustrated dictio-
nary of botanical terms by John Lindley. [Reprint, With an Introduction by Alice Eastwood,
Stanlord LIniversity, School of Earth Sciences, 1964}.
LirnxiH, U. 1964. Mikroautoradiographische untersuchungen iibcrdie lunktion derhydropoten
von Nymphaea. Protopiasma 59:157—162.
and G. KrahI'. I 972. Die Ultrastruktur der jVyw/?/wtv/-hydro|i()ten in zuzm-
menhang mic ihi-cT. Firnktion als salztransporierende Driisen. C]\tobiologie 1:121-131-
MacBrioi., J.F. 1931. Spermatophytes, mostly Peruvian, III. Publ. Field Mus. Nat. Hist.,
Boc. Sen 11 (1): 1-^6.
Maguirl, B. 1979. Guayana, region of the Roraima Sandstone Formation. In: K. Farsen
and F.B. Holm-Nielsen, eds. Tropical Botany. Academic Press. Lontlon. Pp. 223-238.
Mayr, E. and W.H. Pi ili.i'S. 1967. The origin of the bird fauna of the Sotith American highlands.
Bull. Amer. iVlus. Nat. Hist. 136:273-327.
Mayr, F. 1915. Hydropoten an vvasser-und-sumpf-pflanzen. Beih. Bot. Centraibi. 32:278—371.
Metc;alfe, C.R. 1 98 t. Some basic types of cells and tissues. In: C. R. Metcalfe and F. Chalk,
eds. Anatomy of the dictyledons. Vol. 1 . Systematic anatomy ol leaf and stem, With a
Brief History of the Subject. Clarendon Press. Oxford, U. K. Pp. 54-62.
Ml./., C. 1902. Myrsinaceae. In: A. Engler, ed. Das Pflanzenreich 9 (IV, 236): 1-437.
Pi poly, J. 1981 . Contributions toward a monograph of Cyhicnilh/zs (Myrsinaceae): 1. Subge-
nus Iteoida and the identity of Conoiiiorpha lorcleiisi.s. Brittonia 33:491-493.
. I 983a. Clontributions toward a monograph oiCybianthin (Myrsinaceae): III.
A revision of stibgentis Lcixiflon/s. Brittonia 35:61—80.
1983b. Cj)ntributions toward a monograph of Cj/;M«//)//'i (Myrsinaceae): IV.
Notes on stibgenera Stitpjui and Alicroajno)iiorphc/. Wrightia 7:235—244.
. 1 987. A systeniatic revision of the genus Cybianthm subgenus Granimadenia
(Myrsinaceae). Mem. New York Bot. Gard. 43:1-76.
1 988. (^ontribuciones para una monografia del genero Cybiiinthiis (Myrsinaceae)
V. Dos especies ntievas de la Guayana venezolana. Ernstia 50:32—40.
1991. New specfies of Cyhiauthus subgenus Conomorplni from Amazonian
Peru. Candoliea 46:4 1-45.
I 992a. The genus Cybianthiis subgenus Comimorphci (Myrsinaceae) in Guayana.
Ann. Missouri Bot. Gard. 79:908-957.
. 1992b. Notes on the genus Myrsnie (Myrsinaceae) in Peru. Novon 2:392—407.
1993. Notes on Cybuaith/is subgenus Cyb'unithm (Myrsinaceae) in southeastern
Brazil. Novon 3:4 59-462.
PiPOLY, Cybianthiis in Ecuador and Peru
157
1994. Notes on the genus Cyhicrnthi/s subgenus Cybianthin (Myrsinaceae) in
Colombian Amazonia. Sida 16:333—339.
,. 1 996. Noras sobre el genero Cyhianthin (Myrsinaceae) en Colombia y Panama.
Caldasia. l8(3):285-294.
and Y.B. Harvey. 1995. Myrsinaceae. In: B. Stannard, ed. Flora of the Pico
das Almas, Chapada Diamantina, Bahia, Brazil. Royal Boranic Gardens, Kew, U.K. Pp.
487-491.
Prancb, G. 1982. Forest refuges: Evidence from woody angiosperms. In: G. Prance, ed.
Biological Diversification in the Tropics. Columbia University Press, New York. Pp.
137-156.
Sprucf., R. 1880. Notes of a botanist on the Amazon & the Andes. MacMillian & Co., Ltd.,
London, 1908. 2 vols.
Steyermark, J. 1981. Erroneous citation of Venezuelan localities. Taxon 30:816-817.
Theobald, W.L.,J. Krahulik, and R. RtnxiNS. 1984. Trichome description and classification.
In: C.R. Metcalfe and L. Chalk, eds. Anatomy of the Dictyledons. Vol. 1. Systematic
Anatomy of Leaf and Stem, With a Brief History of the Subject. Clarendon Press. Ox-
ford, U.K. Pp. 40-53.
VAsQLii:/., R. 1997. Florula de las Reservas Biologicos de Iquitos, Peru. Monogr. Syst. Bot.
Missouri Bot. Gard. 63-
White, F. 1962. Geographic variation and speciation in Africa with particular reference to
Diospyros. In: D. Nichols, ed. Taxonomy and geography. The Systematics Association
Publ. No. 4, London. Pp. 71-103.
Wiley, E. 1978. The evolutionary species concept reconsidered. Syst. Zool. 27:17—26.
.. 1981. Phylogenetics: The theory and practice of phylogenetic systematics.
J Wiley & Sons, New York.
NUMERICAL LIST OF CYBIANTHUS TAXA
1. Cybianthus pastensis 24.
2. C. t^igantophyllus 25.
3. C. occigranatensis 26.
4. C. spichigeri 27.
5. C. lepidotLis 28.
6. C. laetus 29.
7. C. peruvianas 50.
8. C. tomperuvianus 30a.
9. C. guyanensis subsp. pseudoicacoreus 30b.
10. C. cimanae 31.
11. C. CLiarrecasasii 32.
1 2. C. nestorii 33.
1 3. C. spicatus 34.
14. C tulvopulverulentus subsp. magnoliifolius 35.
15. C. verticilloides 36.
I 6. C. LToarii 37.
17. C. humilis 38.
18. C. sprucei 39-
19. C. simplex 40.
20. C. kayapii ■• 41.
21. C. anrhLiriophyllus 42.
22. C, schlimii 43.
23. C. poeppigii
C. pseudolongitolius
C. vasquczii
C. cenepensis
C. nanayensis
C. marginatus
C. lineatus
C. magnus
C. magnus subsp. magnus
C. magnus subsp. asymmetricus
C incognitos
C. mmutifiorus
C. huampamiensis
C. granulosus
C. fjavovirens
C. vcnezuelanus
C. grandezii
C. jensonii
C. foscen
C. rcsinosus
C. tuscus
C. cyclopetalus
C. penduliflorus
158 SiDA 18(1)
LIST OI' liXSlCCATAi;
Figures in parentheses refer to numbers from the numerical Hst of taxa.
Collecrion numbers in boldface type indicate type specimens.
Acnsca-Solis, M. 5544 (20); At;Lnlar, M^ & D. Casrro 623 (7); 655 (7); ,S()5 (9); Albert clc Escobar, L.
L't al. 3744 (16); Alvarez, A. L-t al. 381 ( 1 ); 4 1 2 ( I ); 490 (1); 1 330 ( 1 ); Anciiash, E. 93 (8); Itl (8);
21 1 (20); 220 (S); 274 (33); 392 (33); 522 (26); 580 (2); 588 (33); 731 (33); 1405 (20); 1412 (26);
1437 (2); Antlrc, E. s.n. (22); 1151 (22): 3819 (17); 45 51 (1); Arias, L. et al. 134 (22); Arroyo, L. &
K, Kciil l6-i (S); Arroyo. L. ct al. 510 (9); 67 'i (8); Asplund E. 9396 (20); 10209 (2i); Aspluiid, E.
10302 (23); 12197 (23); 18717 (2])\ Aulesna, C. & M. 1313 (3); Aulcstia, M. & J. Andi 925 (9);
Ayala, 1-. 1623 (43): 2102 (36); Ayala, I", cc al. 2814 ( t6).
Balsk-v, H. 4295 (I); Balslev, 11. & -Santos Dt-a 2850 (15); Bang, M. 2048 (36); Barbour, P. 2405
(31); Barbour, P. 2567 (31); Bell, D. 101 (42); Berlin, B. i93 (20); 1760(33); 779(33); Betancur.J.
et al.4857 (19); Bohlin, J. et al. 1493 (18); Boyle, B. 3 173 (19); 3599 (19); Boyle, B. &J. Brailtord
1878 (I 9); Boyle, B. et al. 337 i (28); 3450 (30b); Bradford, J. et al. 55 (1 ); Brandbyge, J. 42095 (1);
Brandby.ue, J. & Barlord, A. 42506 ( 1 ); Bravo, E. & R Gdmez 19 (2 I ); Brenes, A. 20530 (22); Buclitein,
O. 175.^ (8); 1758 (5); Bunting, G. & I.. Licht 775 (22): Burger, W. & R. Stoize 5803 (22).
Calderon, C. et al. 2852 (22); Gillejas, R. et al. 2737 (3); Camp, W. E-4809 (28); Campos, J. & L.
3121 (28);Campos,J.etal. 3161 (6); Castaneda, R. 2873 ( IS); 4746 (22); Castro, J. 27 (7);Cer()n,C.
2494 ( I 8); 2585 (23); 7 1 84 (17); 7409 ( I 8); Cerdn C. & M. Faetos 764 1 (20); 76>48 (18); Cerdn, C. &
N. Gallo 5063(9);Chat(')n, I. &(i. Herrera I 720 (22); Cogollo, A. & C. Estrada 296 (3); Cogollo, A.
etal. 7529(3);Collenete, B. 1 1 3 (8); Cornejo, X. & C. Bonifaz 2979 (19); 4339 (19); Croat, T. 7115
(18); 17 167 (22); 19485 (38); 20528 (U); 21 194 (8); ^7785 (23); 49361 (20); 59009 (16); 72394
(19); 73352 (I H); Croat, T. & A. Cogollo 52257 (18); Cuatrecasas, J. 14053 (18); 14138(18); 14918 (3);
14947 (30a); 15474 {22}\ 15658 (3); 15743 (18); 16272 (20); 16272 (18); 17312 (22); 17625 (18);
19892 (18); 21721 (15); 21805 (28); 21981 (15); 22131 (3); 23583 (I I); 23734 (3); 23855 122).
Daly, D. etal. 5926(1 1 ); de Nevers, (i. & H. Herrera 6642 (22); Di'az, C. 566 (22); 7265 (20); 1474
(31); l486(4());Dfaz,<;:. &J. Campos 37 11 (l);Di'az, C. & A. Torres 7805 (30a); Dfaz, C. etal. 3330
(1); 4448 (1); 6930 (3.^); 6993 (7); 7007 (2); 71 70 D; ^252 (5); 7585 (30b); 7649A (34); Dillon, M,
eral. 1 2 1 9 (8); Dodge, C:, & V. Cioerger 9283 (22); Dodson, C. 2821 (7); Dodson, C. er al. 7597 (19);
9101 (19); 151 15 (3); Domfnguez, F. 147 (20); Dressier, R. 6036(20); Drew, W. E-351 (28); Dudley,
T. 10668 (30b); 10690 (6); 10803 (6); 10910 (28); 10922 (28); 10931B (28); 11324 (30b); 1 1915
(1); 13124(13); 13513 (1).
Encarnacitin, R 864 (7); 26200 (1); Espinal, S. 2109 (22).
Faber-Eangendoen, D. & J. Htirrado 1757 (40); I'erreyra, R. 1()40 (32); E'lgueroa, A. 875 ( I 5); b'oLsom,
J. 4544 (18); Fonnegra, R. et al. 3076 (2i); I'orero, E. & L. Wriglcy 7035 (8); 1325 (22); Forero, E.
et al. 7350 (22); Eosberg, F R. 1 9969 (ID; 2()li8 (22); 22376 (II); Fosberg, R R. & Giler, M. 23 1 19
(I); Foster, R. i618 (W); 3708 (26); 4226 (4); 4243 (27); 4327 (27); 4404 (40); 7817 (32); 8624
(32); 898 1 (32); 9013 (30b); Foster, R. & T'. Watcher 7242 (39); 7254 (39); Foster, R. & B. d'Achille
10013 (23); Foster, R. 8962 (8); 10860 (1); 10993 (9); Franco, R et al. 1059 (22); Franco, R et al.
3204 (17), 15423 (23); Funck, N. & L. Schlim 678 (36).
Ciarci'a, I: &J, Echavarna 259- A (20); Garcia, E &J. Ecliavarria 259- A ( 1 7); Gehriger, W. 471 (28);
Gentry, A. 124 19 (30b); 20763 (27); 80096 (4); Cientry, A. & J. Aronson 25044 (2); Gentry, A. & L.
E.mmons i8776 (27); Gentry, A. & E. Forero 73 17 (18); Gentry, A. & N. Jaramillo 22 1 3 1 (40); Gen-
try, A. & A.Juncosa4l702 (22); Gentry, A. & S.Mori 13665 (23); 13709(23); 1 4023 (23); Genrr>
A. & G. Shupp 26565 (19); Gentry, A. & J. Solomon 44407 (5); 44668 (5); C3entry, A. & K. Voting
32025 (27); 2028 (42); Gentry, A. ct al. 16842 (3); 16867 (3); 21033 (27); 22367 (27); 22859 (33)
22859 (tI); 2291 1 (31); 27952 (2); 29700 (38); 31479 (27); t5062 (20); 35188 (19); 39040 (7)
40453 (18); -il88() i2i)\ 42183 (9); ■I22i7 (20); 51083 (36); 51400 (36); 53167 (3); 53326 (22)
53717" (18); 5>S81 (11); 5 3960 (11); 5 5053 (19); 55708 (38); 72129 (9); 72499 (19); 74630 (10)
80179 (4); 56531 (4); 53551 (3); Gleason, H. 159 (36); Grandez, C. et al. 1824 (37); 4487 (9)
4711 (7); 5321 (12); 5370(12); 5642 (12); Gravum. M. & G. Sharz ^170(22); Grubb, R etal. 744
PiPOLY, Cybianthus in Ecuador and Peru 159
(6); 1073 (1); 1633 (20); Guidino, E. 137 (7); C.uillcn, R. & G. Salvatierra 2290 (43); Guillen, R. &
R. Chore 2459 (43); Guillen, R. et al. 31 14 (4 3); Gutierrez, G. 1 186 (23).
Haenke, T. 98 (1); Hammel, B. 17150 (19); Harling, G. 25313 (1); 25334 (1); Harling, G. & L.
Andersson 1725 5 (23); 18778 (19); 21594 (1); 23540(1); 23842 (1); 24097 (23); 9764 (23); Harcman,
R. 2401 (23); 12461 (23); Hartshorn, G. 2996 (35); Hartweg, C. 1200 (28); Haught, O. 1502(22);
4660 (22); 5479 (22); Hernandez, J. & S. Hoyos 483 (22); Holm-Nielsen, L. 681 8 (1); Holm-Nielsen,
L. eraL 3965 ( 1 ); 29949 (28); 19295 (1 8); Hoover, W. 1 194 (22); Hoover, W. et al. 2456 (1 9); 2809
(19); 2815 (22); 3358 (19); Huashikat, V. 356(20); 507 (20); 581 (20); 677 (34); 1221 (34); 1248
(23); 1422 (34); Humboldt, A. von & A. Bonpland 1096 (13); Hurtado, K & D. Neill 235 (3);
Hurtado, F. & A. Alvarado 503 (20).
Idrobo, J. et al. 9671 (17); Imthurn, E. B/9 (14).
Jaramillo,J. 321 (8); 436(7); 1351 (33); 3929(28); 8298 (19); 8501 (9); 8522 (7); 30984 (23);Jaramillo,
J. & E. Grijalva 1 2988 (3); Jelski, C. von 11(1); 360 (8); Jones, A. &C. Davidson 9717 (43);j0rgensen,
P. etal. 1278(28); 1297 (1); Juncosa, A. 1255(22); 1769(22).
Kalbreyer, W. 1087 (30a); 1 534 (17); Karsten, H. s.n. (22); Kayap, R. 558 (8); 618 (33); 723 (20);
728 (22); 783 (33); 856 (33); 982 (33); 993 (33); Kegel, H. 244 (14); 2401 (19); 2601 (19); KiUeen,
T. 4449 (32); 7528 (14); 7530 (14); Killip, E. 35372 (23); 35675 (22); KiUip, E. & H. Garci'a 33802
(15); 33886 (1); 33898 (3); Killip, E. & A. Smith 24011 (8); 26073 (8); 26168 (8); 26286 (8); 26464
(8); 26548 (8); 26563 (8); 27005 (9); 281 10 (40); 29871 (9); 29873 (40); 29906 (27); 33279 (18);
Kirkbride, J. & J. Duke 610 (22); Klug, G. 94 (36); 285 (35); 304 (27); 367 (35); 724 (27); 1384
(40); 1412 (9); Klug, G. 2108 (18); 2565 (9); 2691 (20); 3165 (2); 3188(36); 3981 (32); Knapp, S.
8290 (7); Knapp, S. & J. Kress 4282 (22); Knapp, S. et al. 8514 (13); 85 1 7 (13); Kohn, E. 131 1 (18);
Krukoff, B. 1377 (22); 1388 (22); 7240 (13); 7290 (22); 7663 (20); 7663 {22)\ 10930 (8); 10987
(5); Kujikat, A. 50 (33); 265 (26); 306 (26); 395 (33).
Lawrence, A. 370 (22); 530 (22); Lehmann, F. s.n. (17); 599 (28); 2399 (28); 3027 (15); 5143 (1)
6202 (1); 651 (18); Lewis, W. etal, 10180(18); 10306(25); 10312 (32); 10340 (21); 10475 (21)
12853(21); 14051 (21); 14390(21); Liesner,R. etal. 18302(40); 15093 (22); Limach, C. 140(20):
Linden,;. 447 (28); Little, E. 7481 (1 1 ); 8532 (3); Lleras, E. et al. P17315 (22); Lowrie, S. et al. 52
(22); Lozano, G. 3366 (11); 4133 ( 1 1); Lugo, H. 3633 (23); Luteyn, J. & Cotton, E. 1 101 1 (1); 1 1 295
(28); 11414 (6); Luteyn, J. er al. 6647 (28).
Macbride, J. 5677 (8); Maddison, M. & L. Besse 7201 (22); Maddison, M. et al. 3204 ( 1 8); Malme, G.
2048 (8); 3483 (8); Malme, G. 3483 (8); Manso da Silva, P. s.n. (41); Marmillod, R. 4-R-90 (4); R-
137 (4); Martins, C. von s.n. (43) s.n.- 1 820 (9); s.n.-1826 (13); Mathews, A. s.n. (7); 1561 (6);
McDaniel, S. 10942 (7); McDaniel, S. & M. Rimachi 31219 (37); McPherson, G. et al. 13212 (1);
13397 (I); Monsalve, M. 790 (18); 797 (18); 1124 (18); Morawetz, W. & B. Wallnofer 12-27188
(13); 13-11888(8); 13-30888(8); 14-31188 (4); 22-19188 (7); Mori, S. 7015 (22); Mori, S. & A.
Bolten 7292 (l);Mutis,J. 449(22); 2919(22); 3907 (22); 5102 A (22).
Neill, D. 7494 (9); 9602 (10); 9813 (18); Neill, D. & W. Palacios 9318 (3()a); 9556 (10); 9615 (7);
Neill, D. etal. 10303 (9).
Orti'z, R., et aL 74 (9); 98 (7); 101 (40).
Palacios, W. 1466 (23); 4471 (18); 5394 (3); 5589 (40); 5815 (3); 5950 (17); 6040 (3); 6176 (3)
6187 (3); 6218 (3); 6734 (22); 10680 (18); 11407 (8); Palacios, W. & D. Neill 1584 (21); Palacios
W. & G. Tipaz 10569 (28); Palacios, W. et al. 1 040 (9); 1050 (9); 7761 (7); 8313 (36); 8346 (10)
8407 (10); 8483 (22); 8486 (7); 9269 (22); Pavon, J. s.n. (1); s.n. (1); Pearce, R. 250 (28); Pennell
E 5147 (3); Pinkus, A. 181 (13); Pipoly, J., A. Cogollo et al. 17159(3); 17182 (3); 21026 (19)
21046(19); 21051 (19); 21055 (19); 21 109 (19); 21 131 (19); 21144(22); 21148(19); 21 149(22)
21 173 (19); 21240 (17); 21241 (20); 21 249 (19); 21296 (19); 21326(19); 21328(19); 21417(19)
21 169 (19)' 21471 (19); 21520 (19); 21524 (19); 21598 (19). Pipoly, J., & A. CogUo 17322 (I)
17376(1), 17505(1); 17523(1); 17534(1); 17881 (1); Pipoly, J. & J. MuriUo 15483(20); Pipoly, J
et al. 5307 (3); 12110(7); 12112 (7); 12263(27); 12383 (20); 12490 (20); 12492 (9); 12497 (2)
12541 (20); 12706 (9); 13028 (2); 13284 (2); 13423 (2); 13426 (2); 13931 (2); 14174 (32); 14994
160 Si DA 18(1)
(7); 14997(7); 15896(20); 16075(20); 17183(23); 171,s6(n); 17253(3); 17281 (3); 17360(23);
17361 (17); 17406(22); 17542(3); 17871 (3); 17979 (3); Pirani, j. 1 326 (8); Pitman. N. 688 ( 19);
993 (19); 1161 (19); Poeppig, H. s.n. ( iO); 2428 (40); 2567 (36); 2709 (43); Prance, G. ct al. 8388
(22);P12555 (20); 19075 (8).
Quelal, (.. et al. 764 (22).
Rami'rcz,;. 17(iO); 1 t2 ( U)); RL-strcpo, C 72 i; Rfvilla.j. 187( i.^); 1302(41); 1826 (43); Riedel, L.
959(8); Rimachi, M. 72^2 (7); Robert, A. 322 (8); Rodn'^tiez, E. 283 (33); Rodrfi^uez, R. & J. Surubi'
669 (14); Rubio, D. et al. 1335 (20); 1262 (22); 2252 (1); Rudas, A. & A. Prieto 3147 (20); Rucda.
R. &J. Ruiz 597 (9); Rui'z, J. & j. Pavon 5/36 (8).
Schlim, L, 686 {22)\ Schomburgk, R. 647/992 (29); 885 ( 1 3); 1 002 ( 1 3); Schultt-s, R. E. & G. Black
8427 (20); Schultze-Rhonhof", H. 2983 (19); Schunkc, J. 10 19 (8); 3068 (22); 4779 (32); 5308
(36); 5667 (35); 5737 (32); 5881 (8); 6268 (43); 7106 (23); 10883 (35); 10895 (8); Shcpard, J. 323
(22); Silverstonc-Sopkin, P, et al. 287 1 (20), Simpson, D. & J. Scluinke 647 (41 ); 784 (9); Smith, D.
3709 (35); 3808 (24); 6782 (10); Smith, D. & R. Foster 2509 (29); Smith, S. et al. 738 (42); Sneidern,
K. von A6l2bis( 18); 919(22); 1615 (18); Sodiro, A.; 100/2(17); 100/12(19); 100/14(20); 100/14
(17); Soejarto, D. 3205 (22); 4090 (23); Solomon, J. 3466 (27); 15940 (1); 17081 (19); 17091 (19);
Spichigcr, R. & F. Encarnacmn 1224 (4); 1027 (4); Spichiger, R. ct al. 1973 (4); Spruce, R. 1040
( 1 3); 1946 ( 1 3); 5 1 75 ( 1 ); 6l43 (19); 6 1 44 (18); Stem, B. 372 1 ( I 1 ); Stein, B. & C. Todzia 2292 (6);
Stein, B. & L. McDade ^284 (3); Steyermark, J. 53345 (28); 53550 (3); 53584 (30a); 53642 (1);
53897 (1); 54552 (11); 54800(28); 58625 (5); Stork, H. & Horton, X. 10134(1).
Tate, G. H. H. 741 (5); 927 (5); Terry, M. & R. 1490 (18); Tessman, G. 5145 (40); 3525 (7); 3650
(9);4493(33);Tmiami,M. 1047 (lO); 922 (32); Tipaz, G. et al. 260 (22); 1706(20); 1709(19); 1741
(30b); 1 886 (19); 1950 (22); Todzia, C. & B. Stein 27410 (28); Toro, R. 356 (22); Torres, J. H. 1054
(11); Trail, J. 508 (13); Triana, J. 4 (22); 2562(3); 2585 (I); 2589 (3); 7594 (22); Tunqui, S. 161
(34); 289 (31); 188 (33); 549 (22); Ule, E. 5160 (23); 58)0 (42); 6792 (30b),
Ule, E. 5840 (42); 6792 (30b); 8722 (14); Uribe, L. s.n. (6).
Vargas, I. et al. 2248 (36); Vasquez, R. 8905 (14); Vasquez, R. & J. Cnollo 1801 (4); Vasquez, R. &
M. Flores 1824 (37); Vasquez, R. & N. Jaramiilo 1993 (25); 2703 (9); 3261 (27); 3511 (41); 4285
(43); 5060 (7); 5075 (27); 5086 (27); 524 1 (27); 5867 (7); 6122 ( I 2); 6126 (40); 6 1 37 (4); 6325 (20);
7070 (27); 7570 (7); 7587 (7); 7593 (27); 9461 (43); 10339 (4.^); 11100 (20); 14092 (20); 142(M
(27); 14224(40); 14465(27); 14545(9); 15016(9); 15237(40); 16681 (40); Vasquez, R. et al. 3065
(41); 5911 (4); 6551 (40); 6588 (4); 750 H'); 12108(2); 13764(40); 14161 (27); 14335(7); 17996
(40); 18163(32); 18395(20); 18520(32); 191 32 (42); 20045 (33).
Wallnolcr, B. 111-1588(13); 11-16888(1); 14-29588(10); 1 8- 1 1488 (30b); 112-13688(13); 12-
17788 (8); Warscewicz, s.n. (22); Weberbauer, A. 435 i (6); 4668 (7); 4699 (32); 6099 (1); 6121
(28); 7168 (28); Webster, G. 28702 (19); Webster, G. & R Delprete 27594 (19); Webster, G. & B.
Castro 28769 (19); Webster, G. et al. 27795 (19); 28710 (19); 28796 (19); Werff, H. van der, et al.
10273(40); 13074(11); 12430(18); 13280 (7); Werff, H. van der & 1. Cabrera 1 5786 ( 1 5); Wiggins,
1. 10484 (28); Williams, LI. 657 (27); 658 (27); 1 182 (2); 3676 (43); Wolfe, J. 12346 (32); Wolfe, J.
12295 (1 i); Wurdack, J, 2324 (20).
Yotmg, H. et al. 1 16 (42); Yotmg, K. ct al. 179 (36).
Zak,V. Ii5()(19); 1545 (19);Zarucchi,J. etal.7201 (3)
NEW FLOWERING PLANTS FROM SOUTHERN
NEW IRELAND, PAPUA NEW GUINEA
W. TAKEUCHI
Botanical Research Institute of Texas
do Papua New Guinea Forest Research Institute
R 0. Box 314. Lae. 4lh PAPUA NEW GUINEA
JOHN J. PIPOLYIII
Botanical Research Institute of Texas
309 Pecan Street
Fort Worth. TX 76102-4060 U.S.A.
jpipoly @ brit. org: clusia@latinmail. com
ABSTRACT
Botanical exploration of the Hans Meyer Range in southern New Ireland revealed the
existence of two previously unknown taxa in the families Rubiaceae and Corsiaceae. The
novelties, Psychotria osiana Takeuchi & Pipoly, (Rubiaceae) and Corsia piirpurata var. iviakabui
Takeuchi & Pipoly (Corsiaceae) are described and illustrated. Taxonomic and ecological
notes are also provided for the new taxa.
ABSTRACT (MELANESIAN TOK PISIN)
Wanpela wok bus long salt bilong botani ibin kamap long ol lain maunten bilong Hans
Meyer, Niu Hand Province, long yia 1994. Dispela wokbus o stadi ibin kamapim tupela
niupela samting. Wanpela em sotpela diwai bilong famili Rubiaisi. Namba tu em wanpela
liklik gras nating bilong famili Korsiasi. Tupela igat nem olsem: Psychotria osiana na Corsia
piirpurata var. wiakabui. Dispela stori i tok klia na soim tu sampela piksa bilong tupela.
INTRODUCTION
Papua New Guinea (PNG) is one of only four countries on earth projected
as retaining most of its original forest cover by the end of this century (Suzuki
1993). Unfortunately, it is also among the least known floristic areas within
the Malesian region (Conn 1994; Johns 1995). In the recent multiagency
Conservation Needs Assessment (CNA), 17 localities of primary conserva-
tion significance were identified for Papuasian forest environments (Beehler
1993). Southern New Ireland was one of the enumerated sites considered
vital to biodiversity preservation and in critical need of current informa-
tion. As a direct consequence of the CNA evaluation, a biological assess-
ment expedition was organized by Conservation International and the Papua
New Guinea Department of Environment and Conservation (DEC) in January-
February 1994. A multidisciplinary compilation of results from that sur-
vey has been prepared as a Rapid Assessment Protocol (RAP) Working Paper
SiDA 18(1): 161-168. 1998
162 SiDA 18(1)
by Conservation International (Beehler, in press). Orchid specimens from the
Hans Meyer trip were previously reviewed in a separate speciaUst account
(Howcroft 1994). Two new taxa discerned during general examination of
the expedition's botanical gatherings are described here.
RIJBIACEAE
Psychotria osiana Takeuchi Sc Pipoly, sp. nov. (Fig. 1). Typf; PAPUA NEW
(iUINHA. NiAV Ireland: Hans Meyer Range, pond next to 'Lake Camp,' ()4° 27.205'
S, 152" 56.489' E, 1,175 m, 29 Jan 1994 (tl, fr), W. Takei/chi &J. Wiakabu 9650
(iiOLOTVPF: LAE; isotyphs: A, BRIT, K, L).
Quoad scipLila elongata calyptrataque, stipula ab laminis juveniliLis rumpentes, P. leLiiinides
valde arete affinis, sed ab ea inttorescentiis trichotomiis reduciseiue (non monoromiLs eiongacLsque),
floribus verticellatis (non alternatis) praeditis, lamini.s secLis costis iurluraceo-lepidotis (non
glabris) denique petalis desuper papiUosis (non glabris) statim cognoscitur.
Shrub or small tree to 7 m height, 8 cm dbh. Stem straight, isodiametric,
basal swell absent; outer bark brown, smooth, slash and sapwood stramineous
to pale yellow. Branchlets terete, 2-3 mm diam., pale brown, slightly com-
pressed at the summit, laxly pilose at apical nodes otherwise glabrescent.
Stipules sheathing, calyptrate, caducous, translucent whitish-green, glabrous,
to 5.5 cm long and 6 mm wide. Leaves opposite, usually conferred; blades
coriaceous, elliptic, 10-13 cm long, 3—4 cm wide, apically acuminate, ba-
sally attenuate, the margin entire, adaxially nitid and glabrous, abaxially
light green and furfuraceous-tomentose along the midrib, the midrib prominulous
on borh sides, more elevated beneath; the secondary veins 10-13 oblique
lateral pairs, excurrently arcuate and usually closing submarginally, the tertiary
reticulum lax, irregular, bifacially raised; petioles adaxially plane, 10-20
mm long, glabrous. Inflorescence a sessile, terminal, pinnate panicle, ruptur-
ing through rhe stipule, trichotomous (with three principal branches at base),
the branches with flowers racemose, the primary rachis to 2 cm long, greenish,
somewhat angulate, laxly pilose at nodes, internodes subglabrous or pu-
berulent; floral bracts inconspicuous, not persisting; pedicels 1—3 rnm long.
Flowers opposed or in verricels, 5-merous, entirely glabrous, apparently iso-
morphic; calyx broadly cupuliform, .5 mm long, 1—2 mm wide, limb den-
ticulate; corolla tubular, 3 nim long and obtusely cylindrical in bud, the
lobes chartaceous, corniculate, elliptic to ovate, 1 .5 mm long, 1 mm wide,
spreading or reflexing at anthesis, adaxially papillose; stamens erect, epipetalous;
filaments 1 .5 mm long, adnate to rhe corolline sinuses or slightly below,
the anthers basifixed, oblongoid, 0.7 mm long, 0.2 mm wide, apically and
basally emarginate; ovary umbonate, channelled, bilocular, the style bifid,
slightly exserted, the stigmatic lobes spreading and tuberculate. Drupelet
subglobose, 5—8 mm diam., epicarp glabrous; pyrenes 2, not dorsally ridged,
approximately planoconvex, flat on the commissural face, often with one
pyrene aborted or reduced; endosperm ruminate.
Takf.uchi and Pipoly, New taxa from Papua New Guinea
163
Fig. 1 . Psychotria osiana Takeuchi & Pipoly. A.Branchlet. B. Developing inflorescence emerging
through stipule. C. Inllorescence; showing main rachis and parr of the second rachis in
back. Third axis deleted for clarity. D. Flower at anthesis; petals reflexing. E. Partially dissected
flower. A— D drawn trom holotype.
164 Sum 18(1)
Distribution aiicl ecoloii^y. — Known only from mossy montane forest in the
Hans Meyer Range of southern New Ireland. Occurring as a serai element
in gap phase regeneration among mature growth premontane forest stands,
and along forest margins.
Etymology. — It is a great pleasure to dedicate this species to Dr. Osia Gideon,
Deputy Director of the Papua New Guinea Forest Research Institute and a
specialist in Papuasian Rubiaceae and Zingiberaceae, for liis relentless ef-
forts to document the rich, largely autochthonous, flora of New Guinea.
PakatypI': PAPUA NEW GUINEA. New Ireland: Hans Meyer Range, pond next to
'Lake Camp,' 04° 27.205' S, 152° 56.489' E, 1,175 m, 27 Jan 1994 (fl, fr), W. I'akeiichi &
J. Wuikabu 9560 (A, BRIT, K, L, LAE).
Papuasian Psychotria were first revised by Valeton ( 1 927) and the nonclimbing
representatives more recently treated by Sohmer (1988). Infrageneric rela-
tionships are still poorly understood due to the taxonomic difficulty of the
genus and the absence of regionally-based studies. However a number of
informal species groups have been recognized by Sohmer from macroscopic
features such as stipule form and inflorescence structure.
Psychotria osiarui is characterized by a contracted trichotomous inflores-
cence (i.e., with 3 rachises branched at the base), flowers opposed or verticelled
along the main axes, leaf blades with abaxially furfuraceous-tomentose midribs,
completely glabrous flowers, and petals adaxially papillose. The corolline
papillae are apparent in fresh or rehydrated material, but less so i>i sicco.
A total of 17 species of Papuasian Psychotria have sheathing stipules of
the sort present in Psychotiria osiana. Developing inflorescences and young
leaves emerge by rupturing through the stipule, which subsequently disin-
tegrates or falls away intact. The combination of calyptrate (sheathing) stipules
and a trichotomous inflorescence occurs in only two species: P. leleanoides
Sohmer and P. lorentzii Valeton. Psychotria osiana is clearly related to P. leleanoides,
(a species from New Britain and the Solomon Islands), but is distinguish-
able from it by the leaf indumentum and the entirely glabrous flowers with
papillate corolla. Judging from elevational distributions, Psychotria osiana
is the montane sister species to P. leleanoides, the latter being primarily a
lowland taxon from elevations below 200 m (Sohmer 1988).
CORSIACEAE
Corsia purpurata L.O. Williams var. wiakabui, Takeuchi & Pipoly, var.
nov. (Fig. 2). Type: PAPUA NEW GUINEA. New Ireland: Hans Meyer Range,
pond nexr to 'Lake Camp,' 04° 27.205' S, 152° 56.489' E, 1,175 m, 28Jan 1994 (fl,
fr), W. Takeuchi &J. Wiakabi/ 96! 1 (holotype: LAE, in spirit).
Ad Corsia piirpuratam var. purpuratam accedens sed ab ea labello acuminate nee ad basem
extendens praeclare di.stat.
Terrestrial herb to 13 cm height, sciophytic, erect, all parts glabrous and
Takeuciii and PiPOLY, New taxa from Papua New Guinea
165
5 mm
Fig. 2. Carsia puypiiratci L.O. Williams var. iviakabui Takeuchi & Pipoly var. nov. A. I labit,
dorsal view. B. Habit, lateral view. C. Frontal aspect, petals and lateral sepals. D. Median
sepal. E. Petal. F. Basal callosity, from side. A-F drawn from holotype.
166 SiiMlS(l)
dull reddish-brown. Stenis rerete, weakly sulcate, the nodes distant. Leases
4-5, spirally arranged, acroscopic, base sheathing; lamina linear-acuminate,
conduplicate or with margins broadly revolute, chartaceous, 9—17 mm long,
4—6 mm wide, to 6-nerval, sometimes apiculate; venation visible as dark
lines, not raised on either side. Flowers solitary, terminal on the unbranched
stem, bisexual; median sepal flabellate, symmetrical, 16-1 9 mm long, 20-
23 mm wide, nodding at anthesis, unguiculate, bluntly acuminate at the
apex; the basal callosity linear, narrowly raised by 4 mm above the plane of
the 'labelkim,' summit corniculate, base abruptly truncate at the claw; median
sepal 1 2-veined, the veins paralleiodromous, diverging mainly from the proximal
half of the sepal, once or several times furcate before the margin; petals and
lateral sepals isomorphic, basiscopic and incurved, costate, lanceolate, 4 mm
long, 3 mm wide, acumen 1.5 mm long; stamens 6, in 2 whorls opposite
the perianth segments; filaments resembling the style, 0.6-0.7 mm long;
the anther cells 2, oblongoid, 1 .2 mm long; style simple, cylindrical, ca. 1
mm long. Pr/nt not seen.
Distribution and ecology. — Known only from the type locality in the Hans
Meyer Range on New Ireland. Corsia pi/rpunitci var. wicikahiii occurs infre-
quently in mature-growth, primary premontane forest, on its floor, cov-
ered by leaf duff. This new variety was encountered as a population flush-
ing after recent rains and thereafter evanescent; all individuals were seen in
comparable maturational states.
Etyiiiology. — The new variety is dedicated to Joseph Wiakabu; botanist,
explorer-collector, and colleague from the Papua New Guinea National
Herbarium.
Paratvpi:: PAPUA NEW CUIINEA. Ni:\v Iri.i.am): Huns Meyer Range, slopes above
river valley on the ascent from Mandih lake, ()4^ 26' S, 1 52^^ 59' E, 7^0 m, 10 Au.^ 1975,
M.J.S. Scnuh et cil. 2091 (K, EAE).
Corsia was revised by van Royen { 1 972) in a monograph based on 45 exsiccatae
numbers, an average of less than 2 collections per species. The limited number
of specimens available for study is due to population rarity, the inconspicu-
ous habit of the plants, an ephemeral [4ienology, and the preference for sheltered
microsites in forest where ambient light conditions favor concealment. Van
Royen (ibid.) commented that encounters with Corsia are often of a fortti-
itous nature; occasioned by understory sun flecks falling on the plants in a
certain way by chance. There is no doubt that the genus is rarely found by
collectors. The intervening years since the initial revision has seen little increase
in availability of specimens, nor is the situation likely to improve dramati-
cally in the future. The difficulty of botanizing purposefully for Corsia dis-
courages the sort of field-based studies necessary for understanding the
morphological variation between populations.
Takeuchi and PiPOLY, New taxa from Papua New Guinea 1 67
According to van Royen's (1972) conspectus, Corsia is a saprophytic ge-
nus with 25 species distributed through New Guinea, the Bismarck Ar-
chipelago, the Solomon Islands, and Australia. Most of the species are nar-
rowly defined endemics of montane forest habitats, and appear to be restricted
to specific mountains. On the basis of the venation and basal callosity of
the median sepal, the new taxon is related to C. purpurata var. purpurata,
but is distinguishable by the flabellate 'labellum' with acuminate apex and
the base truncate rather than extended.
The type locality for var. waikabui is floristically depauperate in com-
parison to similar habitats from mainland New Guinea, and is dispropor-
tionately composed of epiphytic or nonendemic species (Takeuchi and Wiakabu,
in press). Depauperate levels of biological diversity in this montane envi-
ronment was also reported by zoological specialists participating in the New
Ireland survey (Beehler in press, passim). The expedition's collective find-
ings are consistent with a supposition that the cloudy uplands of southern
New Ireland are of geologically recent origin. In view of the multidisciplinary
results from the recent survey, it is unlikely that future exploration of the
Hans Meyer Range will yield significant numbers of additional novelties.
ACKNOWLEDGMENTS
The Rapid Assessment Protocol survey of New Ireland was funded by
Conservation International and the Biodiversity Support Program. Expe-
dition leader Dr. Bruce Beehler and the Papua New Guinea Department of
Environment and Conservation performed key coordinating roles. The Liz
Claiborne and Art Ortenberg Foundation and the John D. and Catherine
T. MacArthur Foundation have provided ongoing financial support to W.
Takeuchi for studies partially based at the Papua New Guinea National
Herbarium, at the PNG Forest Research Institute, m Lae, in support of his
principal work at the Crater Mountain Wildlife Management Area.
Neville Howcroft prepared the illustration o{ Corsia purpurata var. wiakabui
and Talk Iwagu the illustration of Psychotria osiana, which we gratefully
acknowledge.
REFERENCES
Bi;eiim;r, B.M. (ecL). 1993. Papua New Guinea conservation needs assessment, vol. 2. Cor-
porate Press, Inc., Landover, Maryland.
Bi-EHLr,R, B. (ed.). In press. Rapid Assessment Program Working Papers, No. 1 0. A biodiversity
assessment of southern New Ireland, Paptia New Guinea. Conservatif)n Internatiorial.
Washmgton, DC.
Conn, B.J. 1994. Documentation of the flora ot New Guinea. In C.-I Peng and C.H. Chou,
eds. Biodiversity and terrestrial ecosystems. Institute Bot., Acad. Sinica Monogr. 14: 123— 1 56.
Howc;roft, N.H.S. 1 994. Orchid collections from New Ireland. Orchid Res. Bull 3. Papua
New Guinea Forest Research Institute.
168 SiDA 18(1)
Johns, R.J, 1995. Malcsia-an introduction. Bot. Mag. 12:'52-62.
RoYEN, P. VAN. 1972. SertLilum Papiiannm 1 7. Corsiaccae of New Guinea and surroLindint,'
areas. Webbia 27:223-255.
SoilMF.R, S.H. 19H(S. The nonclimbing species ot the genus Psychntric/ (Rubiaceae) in New
Guinea anci the Bismarck Archipelago. Bishop Mus. Bull. Bot. 1:1—3.^9.
Suzuki, l^. 1993. Time for a change. Allen and Unwin, St. Leonards.
Takeuciii, W. and J. Wiakabu. In press. A transect-based floristic reconnaissance of sourh-
ern New Ireland. In B. Beehler, ed. A biodiversity assessment of southern New Ireland,
Papua Nfew Gtnnea. Rapid Assessment Program Working Papers 10, Conservation In-
ternational, Washmgton, D.G.
Valhton, T. 1927. Die Rtibiaceae von Papuasien. 11. /.weiter Teil: Cofleoidcae. Bot. Jahrb.
61:32-163.
THREE NEW SPECIES OF CRATAEGUS (ROSACEAE)
FROM WESTERN NORTH AMERICA:
C. OKENNONII, C. OKANAGANENSIS
AND C. PHIPPSII
J.B. PHIPPS
The University of Western Ontario
Department of Plant Sciences
London, Ontario, CANADA N6A 5B7
RJ. O'KENNON
Botanical Research Institute of Texas
309 Pecan Street
Fort Worth, TX 76102-4060, U.S.A.
ABSTRACT
Three new species of Crataegus, C. okennonii J.B. Phipps (from series Douglasianae), C.
okanagane)isis J.B. Phipps & O'Kennnon (from the newly described series Pi/rpureofri/cti J.B.
Phipps & O'Kennon), and C. phippsii O'Kennon (near series Molles) are described from western
North America. All are striking, widespread taxa with a substantial number of records.
Key Worus: Taxonomy, Crataegus, Rosaceae, new species
RESUMEN
Tres nuevas especies de Crataegus, C. okennonii J.B. Phipps (de la serie Dauglasianae), C.
okanaganensis J .B . Phipps & O'Kennnon (de la nueva serie descrita Purpureofructi J.B. Phipps
& O'Kennon), y C. phippsii O'Kennon (proximo a la serie Molles) se describen del oeste de
Norte America. Todos son taxa sorprendentes y de amplia difusion con un sustancial numero
de ciras.
Recent fieldwork with R.J. O'Kennon of Fort Worth, Texas in the American
states west of the Rocky Mountains and in British Columbia for the first
author's revision of the red-fruited Crataegi (Phipps 1998) of this region
has serendipitously uncovered three new species of hawthorn. The first, Crataegus
okennonii, a new purplish-fruited taxon of ser. Douglasianae (Rehd. ex Sarg.)
Rehd., striking and obvious in the field, is also usually easily resolved in
the herbarium. Crataegus okanaganensis, rarely collected before our studies,
is also a striking new taxon with brilliant red (late August) to deep purple
(late September) fruit. It is accorded a new series, Pupureofructi, together
with the long-described and long-overlooked C. williamsii Eggl. The third
new species, C. phippsii, though first collected as far back as 1908, remained
otherwise unnoticed until our fieldwork. It is a very distinct purple-fruited
species related to series Molles.
SiuA 18(1): 169-191. 1998
170 SiDA 18(1)
One of the reasons that these new Crataegus species west of the Rocky
Mountains have been overlooked has been the fliiltire of nearly all collectors
to take note of fruit color (Phipps 1998). Each of the following species is
quite distinct in this respect and we therefore take the opportunity to present
color plates of the three species illustrating this point. Hawthorns are con-
sidered to have mostly black or red fruit. The discovery of these three ex-
tends the number of those which are burgundy to deep purple at some stage
in their development. Another feature emerging from this work are new
centers of diversity for western North American Crataegus appearing m the
northern Okanagan of British Columbia and northwest Montana.
A further reason that these taxa have been overlooked is that in the Pa-
cific Northwest region (to nw Montana) and British Columbia where they
occur there has been a belief that generally, only one red-fruited native spe-
cies, C. 'Columbiana Howell,' one black-fruited native taxon, C. douglam Lindl.
(inch C. suksdorfii (Sarg.) Kruschke) and one introduced red-fruited taxon
(C wrwog}'Wi:7jacq.) occurred. This belief has persisted (Phipps 1998) in spite
of the widespread presence in this region of C. niacracantha Lodd. ex Loud.
In view of this lack of taxonomic inattention, including the disappearance
from the floristic literature of the distinctive C. icillianis'n, it is perhaps not
surprising that earlier botanists did not recognize potential new taxa in this
region. Consequently, we take this opportunity of reviewing the main western
North American species (west of the Rocky Mountains) in a numerical taxonomic
analysis. This analysis uses one synthetic OTU per taxon, each described
from typical material over 38 morphological characters (Table 1) of the kind
commonly used in Crataegus taxonomy. The eleven native species generally
recognized in the west are all present, together with C. monogyna, a useful
outgroup, and five species from east of the Rocky Mountains (C. mollis (Torr.
& A. Gray) Scheele, C. suhmolUs Sarg., C. brachyaeantha Sarg. & Engelm., C.
schuettei Ashe and C punctata Jacq.) to give structure and scale. Also the
three new taxa described in this paper are included. A dendrogram was prepared
by NT-SYS using the options taxonomic distance and unweighted pair-group
arithmetic averaging. The results (Fig. 1) place C. inonogyiia and C. punctata
outside all the other species. Among results of general interest, varieties
(C chrysocarpa Ashe var. chrysocarpa and C. c. var. piperi (Britton) Kruschke)
cluster at about the 0.6 level while distinct species cluster at the 0.8 or greater
level, e.g. C. schuettei 'dnd C. wnotoniana Eggl. (series Tenuifoliae), C. suksdorfii
and C. do//glasii {'iQUi^s Douglasianae) and C. erythropoda Ashe and C. rivularis
Nutt. (series Cerrones). Other pertinent results are discussed under the new
taxa described in this paper,
TAXON DESCRIPTIONS
1. Crataegus okennonii J.B. Phipps, sp. nov. (Fig. 2). 'Ywv. CANADA.
Piiipps AND O'Kennon, New species of Crataegus from Western North America 171
TAiii.n 1. The 38 characters scored for tlie numerical caxonomic analysis.
PLoit. gctiercil
1 . Plant habit
2. Bark type
3. Branch arrangement
'r barns and I'uigs
4. Thorns: indeterminate present?
5. Thorns: length
6. Thorns: curvatLire
7. Thorns, color: browns
(S. Thorns, color: gray
9. Twigs, I yr old, color; browns
10. Twigs, I yr old, color; gray
Lamina: length
Lamina: length/breadth
Lamina: position ot widest part
Lamina: venation, number
Lamina: veins to sinuses
Lamina: lobe number
Lamina: lobe shape
Lamina: max. sinus depth (LII)
Lamina: abaxial pubescence, young
Petiole: glands
Leaf teeth: glands
Lecij
11.
12.
13.
1-1,
15.
16.
17.
18.
19.
20.
21.
bijloreKena
22. Flower: number
23. Pedicel: pubescence
24. Anthesis time
Floivtr
25. Diameter
26. Calyx lobes: margins
27. Stamen: no.
28. Anther: color
29. Style: numbei
Fntit
30. Fruit: color, month before lull ripeness
3L Pfuit: color at full ripeness
32. Fruit: shape, 1
33. Fruit: shape, 2
34. Fruit: pubescence
35. Fruit: calyx orientation
36. Fruit: length (height)
37. Pyrenes: lateral faces
Autumnal joltage
38. Color
British Columbia: Okanagan Valley, E side of 97A 0.5 mi N of intersection with
97, scrubby slopes with long cleared area, 50°2rN, 119°15'W, 19 Aug \99A,Phipps,
J.B. & (rKennon, R.J. 6998 (hoi.otype: UWO; isotypes: BRIT, CAN, TRT, US).
Frutices grandes vel arbores parvae, plerumque trunco uno, ad 10 m alti, spinosi; ra-
muli unius anni nirentes atro-cascanei, ramuli veteriores brunneo-cinerei; spinae plerumque
< 2.0 cm longae, crassae, plus minusve recurvatae, in juventute brunnae nitenter. Folia
decidua;petioli plerumque 1—2 cm longi; laminae 4— 6 cm longae, ellipticae vel lare-ellipticae,
interdum aliquantum anguste ovatae vel rhombeae, paribus 4 venarum sccondariarum longiorum
vel aliquando non-lobatae vel vix-lobatae vel vadosirer 2-3(-4) lobatae per latus, marginibus
serratis, ± coriaceae, in mattiritate tenuiter appresso-pubescentes supra, subter ± glabrae
venis exceptis, colore autumnah prunino ad carmineum vel pallide carmineum. Inflorcscenciae
12-20 florarae, bracteolis parvis caducis glandulo-marginatis, glabrae vel raro pubescentes.
Flores 15—20 mm diam.; hypanthium glabrum extrinsecus; lobi calycis 3 mm longi, ob-
scure gland uloso-marginati; petala ± circulaha; stamina 10—12, antheris pallide roseis; carpelli
et styla 3—4. Fructus ampulliformo-orbiculares, ca 10 mm diam., triste-purpurei vel nigri
subinde in maturitate, in mense Augusto rubro-vinosi vel spadiceo-castanei, reliquiis calycis
parvis; pyrenae 3—4, vadositer sulcatae dorsaliter, lateribus cum foveis vel interdum planis.
Large bashes or small trees, usually single-trunked, up to 10 m tall, thorny;
one year old twigs deep shining mahogany, older branches gray/brown; thorns
usually < 2.0 cm long, stout, slightly recurved, shiny brown when young.
Leaves deciduous; petioles usually 1—2 cm long; blades elliptic to broad-
172
N
ew Crataegus
species
of western North America
Sum 18(1)
UPGMA of taxonomic aisiance
19 taxa
38 characters
.J.ulUli
chr piped — — '
-J
1 ' '
4D
8C
1 20
1 60
200
Fig. 1 . Dendrogram of western North American Crcitctegi/s taxa sliowing relation.shi|TS of
species describee! in this paper. Newly described taxa in capitals. See text lor more infor-
mation.
elliptic, sometimes somewhat narrow ovate or rhombic, 4—6 cm long, with
four pairs of longer secondary veins, barely lobed to shallowly 2-3(— 4) lobed,
the margins also serrate, somewhat coriaceous, at maturity finely appressed-
pubescent above, below ± glabrous except on the veins; autumnal color plum
to crimson or light crimson. Inflorescences 1 2—20 flowered, with small caducous
gland-margined bracteoles, glabrous to rarely shortly pubescent. Flowers
15—20 mm diam.; the hypanthium externally glabrous; calyx lobes 3 mm
long, obscurely gland-margined; petals ± circular; stamens 1 0—12, anthers
pale pink; carpels and styles 3—4. Fruit ± amptflliform-orbicular, ca. 10 mm
diam., dull-purple or occasionally black at full maturity, red-burgundy to
chestnut or reddish-brown in August, calyx remnants small; nutlets 3-4,
shallowly grooved dorsally, sides variably eroded or sometimes plane.
Distribution. — Cratciegiis okennouii is fairly common in southern interior
Britisli Columbia and the eastern half of northern Washington, ranging to
Idaho and nw IMontana (Fig. 3), usually along streams or other sources of
water. It is a tall distinctive plant, usually with a straight trunk, easily rec-
ognized in the field both in flower and in frtiit. This new species is most
closely related to C. Joiiglasii , sens, str., from which it differs in fruit shape
Phipps and O'Kpnnon, New species of Crataegus from Western North America 173
Fic;. 2. Line drawing ot C. okeinio/aiJ.B. Phipps. Inflorescence, flowers and parts from Phipps
6959 (UWO); mfructescence, fruit and parts from Phipps 6991, 6995 aiuKYKennon (UWO).
Scale bars = 1 cm.
\1^
SiDA 18(1)
Fig. 3. Distriburion map of C, okef///oi/!/ ) .\i. Fhij^ps.
and color, flower size, color of marure leaves (often yellowish-green), habit,
thorn length, leal shape and autumnal color of foliage.
Crataegus okemiomi is one of a small grotip of 'hhick-frnited' (fruit ± black
at full mattirity) species comprising series D()nglasianae(Kt\\d. ex Sarg.) Rehd.
which I restrict to C. douglasii Lindl., its immediate relative C sz/ksclorfii
and the new species described here. A greatly widened concept of C douit^lasn
which includes C. saligna Greene and C. riviilarh is advocated by Welsh
(1982) and Holmgren (1997) but in my opinion this cannot stand up due
to many significant differences in leaf-shape, venation type, bark and thorn
type. Moreover intermediates do not exist between saHi^na-rmdaris'^ind douglasii-
okennonii-suksdorjii . Diffictdties for the broad concept are also emphasized
by the fact that C. saligna (called C. doiiglasi! Linth. van diichesnensis Welsh
by Holmgren) has 20 stamens and cream anthers, contrary to the Intermountain
Flora. I am (JBP, 199X) therefore placing C saligna and C. rividar'is in a
different grouping.
Crataegus okennonii is a clearly defined segregate of C. douglasii differing
by a considerable number of correlated characters of which thorn length
(the shortest in series Douglasianae) and flower size (kirgest in the series) are
the most unambiguous. The following couplet separates C. okennonii from
the restricted interpretation of typical C. douglasii 'as, found in western Canada
and the adjacent United States.
Phipps and O'Kennon, New species of Crataegus from Western North America 175
1. Usually tree-like with a straight trunk, 5-10 m tall; young twigs deep glossy
purple-brown; thorns usually 1.5—2 cm long; flowers ca. 15—20 mm cliam.;
fruit crimson lake or chestnut-colored in late August, becoming deep plum-
black at maturity, not bloomy, broadly ampulliform; autumnal foliage usu-
ally crimson, plum or pale shades of same; styles and nutlets usually 3-
C. okennonii
1 . Usually btishy, though often large, not conspicuously single or straight-trtmked,
3-8 m tall; thorns often longer, to 2.5(-3) cm long; young twigs mid to
deep glossy brown, only occasionally as dark as C okennonii; flowers often
12-15 mm in diam.; fruit variably colored in late August though never chestnut,
shape ellipsoid or suborbicular; autumnal foliage color often bronzy, not
consistently ± crimson or paler; styles and nutlets 3—5 C. douglasii
The differences between C. okennonii and a somewhat restricted C. douglasii
are not by any means large but recognition at the specific level is also sup-
ported by the numerical analysis (Fig. 1), the lack of intermediates among
good quality specimens both in the field and herbarium and the consequent
immediacy of recognition. A more elaborate comparison with C. douglasii
in which detailed leaf shape comparisons will be used may be expected consequent
on the publication of R. Dotterer and T.A. Dickinson's detailed biosystem-
atic studies of C. okennonii and C. douglasii.
It is possible that C. okennonii is the same as C. douglasii f. hadia Sarg.
(1907), a taxon in which the fruit is described as "chestnut", certainly not
a common color in ripening fruit of section Douglasii. However, some of
the unripe fruit of C. okennonii seen by us certainly was the color of reddish-
brown, highly polished, shoe leather. Piper, the original collector of forma
hadia, variously described the fruit as brown, dark shining brown, chest-
nut, etc. Forma badia is a taxon systematically disregarded in the floristic
literature. Both color changes during the ripening of western North American
Crataegus fruit and the final color of the autumnal foliage have been poorly
understood (Phipps 1997) as taxonomic characters so we wish to record them
here. Crataegus douglasii fruit is shown in Figures 4d and 4e. The latter,
photographed in August, illustrates the black, pendant, ellipsoid fruit while
the other is another British Columbia specimen showing plenty of bloom.
Figures 4a-4c illustrate C. okennonii, plate 4a showing the late August "chestnut"
color while Figure 4b is a later version of the same color. Figure 4c shows
the final, near ripe color, in late September, close to that of C douglasii. The
foliage is frequently a distinctive bright pale crimson color in late Septem-
ber as is shown in Figure 4f. Variation in leaf form of C okennonii tends to
fall into two classes, both illustrated in Figure 2, there being a broader,
somewhat rhombo-deltoid, distinctly lobed form (Fig. 4a) and a more el-
liptic or narrow-ovate, shallowly lobed shape (Fig. 4b). Crataegus okennonii
also bears conspicuous, expanding, caducous, coral-red bud scales.
I (J.B.P) am pleased to name this distinctive and handsome new species
176
Si DA 18(1)
Fjg. 4. Cratiicgin okmiion'ii compared with C. dniigliisii: a) C okenntDiii showing remarkable
chestnut-coloured fruit, Kclowna, BC, 18 Aug 1995; b) C, okennonii, Form with pale foli-
age, note ampulliform fruit, northern Okanagan, BC, 20 Aug 1995; c) C. okennonii with
some triut close to the final purple-black color, northern Okanagan, BC, 19 Aug 1995; d)
C. doiigLnii showing full ripe color and a high bloom, northern Okanagan, BC, 19 Aug
1995; e) full ripe color ot C. donglasii with ellipsoidal fruit, northern Okanagan, BC, 19
Aug 1995; f) C. okennonii showing typical cerise color of full ripe foliage; JBP in foreground;
Palmer Lake, Washington State, 28 Sep 1993.
after Robert (Bob) J. O'Kennon whose companionship and critical abiHties
as a field botanist helped to make this discovery possible. It is mtriginng
that so widely distributed a taxon, of which we are able to cite 36 speci-
mens, has remained undetected for so long.
Additional specimens examined; CANADA. BRITISH COLUMBIA: Northern Okanagan,
Spallumcheen Municipality, Powerhouse Rd., M side, near Stardel Drive, alt. ca. 1350 ft,
50°27'N, 1 19°09'W, bush, 1.5 m tall, ± erect; fruit deep plum/chestnut, ellipsoid-turbi-
nate, 19 Aug 1995, ././3, Phtpln -"164 (UBC, UWO); Okanagan Valley, Oyama, ca. 0.5 mi
E ofE end of spit, alt. ca. 1320 ft, 50n)7'N, 119°2rW, hedgeline on clay soil, bush 5 m
Phiprs ani:i O'Khnnon, New species of Crataegus from Western North America 177
tall, frtiit plum-purple, turbmate, 17 Aug 1 995 J, C. Ph!pps7158 (UWO); Castlegar, Selkirk
College grounds, towards bottom of slope beyond beehives, scrubby area, 49° 1 9'N, 1 1 7°38'W,
alt 950 ft, tree 6 m tall; stamens 5, anthers pale pink, buds forced til 10 May, then 10
stamens seen, 06 May 1994 J. B. Phipjn 6928 (BRIT, CAN, DAO, TRT, UBC, UWO, US);
Okanagan Valley, Westside Rd., ca. 9 mi W of jet. with 97 and ± due opposite Vernon,
wooded edges on west of road, 50°1 5'N, 1 19°27'W, ca. 1200 ft, tree, 5 m tall, fruit small-
ish, bright red-burgundy, 19 Aug 1994, Phipps.J.B. & O'Kennon, R.J. 6990 (CAN, TRT,
UBC, UWO); Northern Okanagan, Hwy 97, E side, between Westside Rd. and Silver Creek
Rd., alt. 1800 ft, equals JBP 6969, beautiful 7 m tree, 50°2.3'N, 119"17'W, convex light
yellow-green foliage, fruit light red, turning to burgundy, roundish, fatter at base, glossy,
19 Aug 1994, PhtppsJ.B. & O'Kennon. R.J. 6993 (BRIT, UWO); Salmon Arm, large haw-
thorn pasture just N of railroad and W of central business section, alt. ca. 1900 ft, 50°43'N,
119°16'W; bush, 4.5 m tall, fruir burgundy, locally frequent, (equals JBP 6959); 20 Aug
1994, Phipps.J.B. & O'Kennon. RJ. 7027 (CAN, TRT, UBC, UWO); Okanagan Valley,
Westside Rd., ca. 9 mi W of jet. with 97 and ± due opposite Vernon on E side of road,
fenceline 50° 1 5'N, 1 19°27'W; 7 m multitrunked tree, fruit burgundy, alt. ca. 1200 ft, 19
Aug 1994, PhippsJ.B. & O'Kennon. RJ. 6991 (BRIT, CAN, TRT, UWO); Okanagan Val-
ley, NNE of Vernon at jet, 91197 A, scrubby hillside E side of road, 26 Sep 1993,7.6. Phipps
andRJ. O'Kennon 6821 (BRIT, UWO). Pass Creek Rd., N of Castlegar, 1.3 mi along, mesic
grassy slopes, alt. ca. 1800 ft, 49°19'N, 1 17°40'W, bush, 4 m tall, fruit reddish-burgundy
(somewhat 'chestnut'-ROK), 23 Aug 1994, PhippsJ.B. & O'Kennon. RJ. 7036 (TRT, UWO);
Castlegar, Selkirk College, path behind beehives towards river edge, open woodland-scrub,
alt. 950 ft, 49°I9'N, 117°38'W, tree, 20 ft tall, reddish-purple foliage, dark purple, ±
round fruit, 20 stamens, 26 Sep 1993, PhippsJ.B. & O'Kennon. RJ. 6824 (UWO); Slocan
Valley, S of Slocan, above swamp along small road off Hwy. 6 & just N of Perry Siding
across river, 49°40'N, 1 17°30'W, hedges and brushy slopes, alt. 1775 ft, bush 6 m tall,
fruit red-burgundy, 22 Aug 1994, PhippsJ.B. & O'Kennon, RJ. 7047 (UWO); NNW of
Vernon on W side Rd about 2.5-3 km SW of intersection of 97 & 97A, alt. 450 ft, 50°20'N,
119°20'W, back of field on west side of road, bush 5 m tall, 24 Sep 1993, PhippsJ.B. &
O'Kennon. RJ. 6785 (UWO); Okanagan Valley, Hwy. #97, ca. 2 mi S of Silver Creek Rd.
exit, (200 m S of 'road narrows' sign), (100 m S of JBP 6968), 50°23'N, 119°17'W, alt
1800 ft, in cut-over forest, young tree 7 m tall, 11 May 1994 J. B. Phipps 6969 (BRIT,
TRT, UWO); Okanagan Valley, ca. 8 km NNE of center of Vernon, Pleasant Valley Rd.,
behind Vowle's residence, hedge at S boundary of Burke's PYO, 50°18'N, 119°l4'W, alt.
380 m, dense, broad, mature hedge, dominant species, 3-7 m bush, 10 pink anthers, thorns
long for species, 2 May 1994 J. B. Phipps 6879 (UWO); Salmon Arm, large hawthorn pas-
ture, just N of railroad and W of central business center; 50°43'N, 119°16'W, alt. 1900
ft, bush, 5 m tall, 10 stamens, pink anthers, 10 May 1994, J. B. Phipps 6939 (TRT, UBC,
UWO, V); Okanagan Valley, Newport Beach campsite on W side Rd., approx. due NW of
Vernon, end of parking lot, S end of camp, 50°20'N, 119°22'W, below trees, alt, 1140 ft,
tree 5 m tall, erect habit, burgundy fruit, short thorns, 19 Aug 1994, Phipps. J. B. & O'Kennon,
RJ. 6988 (BRIT, DAO, UBC, UWO, V); NNE of Vernon, S of jet. of 97 and 97 A just N
of Baker Hogg Rd., bottom of hill, 50°21'N, 119°15'W, alt. 350 m, 4 m bush, purplish
fruit, 23 Sep 1993, PhippsJ.B. & O'Kennon, RJ. 6821 (UWO); Okanagan Valley, E side of
97, half mi N of intersection with 97; scrubby slopes with long cleared area, 50°21'N,
1 19°15'W, most abundant species here, bush 6 m tall, red fruit; 19 Aug 1994, Phipps. J. B.
& O'Kennon. R.J. 6999 (UWO); Castlegar, grassy floodplain ENE of Selkirk College, dense
hawthorn thickets on banks of old oxbow, alt. 950 m, 49°19'N,1 17°38'W, tree, 9 m tall,
foliage elliptical, slightly lobed, purple, fruit globose, dark purple, ? = 20 stamens, 27 Sep
1993, Phipps. J. B. & O'Kennon. RJ. 6827 (CAN, ID, MONTU, TRT, UBC, UWO, US);
178 SiDA 18(1)
Norrhcrn Okana^'an, ca. 1 mi S ol Enderby, gravel driveway to NE starting at Indian Om-
etery, alt. 1 300 ft, 5()"3 I'N, 11 9°08'W, bush, 4 m tall, fruit burgundy, only one here, 20
Aug 1994, Phipps.J.B. &iyKeinmi. R.J. 7006 (UWO).
U.S.A. IDAHO. Idaho Co.: US 12, ca. 10 road mi E of Kooskia, bank at back of field,
N side of road, with large hawthorns, 46°07'N, 1 ^"50'W, alt. 1 550 ft, tree, 9 m tall,
fruit burgundy, 26 Aug 1994, Phipps.J.B. & (rKenrioii. R.J. 7089 (UWO). MONTANA.
Flathead Co.: Eew mi E of Columbia Falls, at dead end of Mt. Creek Rd. (off Berne l^d.),
on fenceline opposite Shoal 330 sign, 48°22'N, 1 14°08'W, alt. 3000 ft, bush, 2 m tall, 08
May 1994,y.Z3. Phipp.^ 693 > (UWO). Lake Co.: Route 211, ca. 4 mi S of Ronan, nr. MP
44, at farm entrance, driveway throtigh pasture, near scenic turnoiu , alt. 3030 ft, 47°28'N,
114°06'W, grove of hawthorns on bank S side of driveway, 20 ft apple-like tree, reddish
foliage, black orbicular fruit, 28 Sep 1993, PhippsJ.B. & O'Kenmm. RJ. 6841 (UWO);
Rte. 93, ca. 4 mi S of Ronan, near Milepost 44, near scenic turnout, one of group on S side
of farm tirive, 47°28'N, 114"06'W, alt. 3030 ft, tree 5 m tall, quite large buds forced 'til
1 1 May, stamens 10, anthers pink, (equals JBP 684 1 ), 08 May 1 994J.B. Phipps 6940 (BRIT,
iMONTU, TRT, UWO, US). Sanders Co.: Route 200, 30 mi W of 93 on N side of road
above Flathead R., in group of hawthorns, alt. 2840 ft, 47"I9'N, 1 14°43'W tree, 1 1 m
tall, ± orbicular, black fruit, 29 Sep \99:\ Pb/pp.^. J. B. OiYKeinion. RJ. 6« 58 (BRIT, CAN,
DAO, MO, UWO, V); Route 200, 50.6 km W of jet. with US 93, group of hawthorns
above Flathead R., 47"19'N, 1 14°43'W, alt. 2840 ft, tree 9 m tall, stamens 10, anthers
pale pink,(=JBP & RO'K 6955), 09 May 1 994,./. B. Phtpps 6955 (UWO); Route 200, 25
mi W of 93 on N side of road above Flathead R., in long hawthorn hedge, alt. 2845 ft,
47°19'N, 1 14°38'W, btish 6 m tall with spheroidal burgundy fruit, 29 Sep 1993, Phipps,
J.B. & O'KeHrwn. R.J. 6857 (UWO); Route 200, exactly 42.6 km W of jet. with US 93,
long hawthorn hedge, above Flathead River, 47°19'N, 1 14'38'W, alt. 2840 ft, (equals JBP
& RO'K 6857), 09 May 1994, J.B, P/j/>/v 6933 (BRIT, MO, MONTU, UWO). WASH-
INGTON. Ferry Co.: 1 Iwy. 2 1 , ca. 3 km SW of Danville, alt. ca. 500 m, 18 = 59'N, 1 1 8°3 1 'W,
marshy roadside with trees and scrub, tree, 7 m tall, fruit burgLindy, (C. cloniilcisii very common
here), 23 Aug 1995, ./.B. Phipp.s 7174 (UWO). Okanogan Co.: Chopaka Rd., 3 km di-
rectly NW of Palmer Lake, open thicket, low rocky slopes, edge of floodplain, alt. 340 m,
48°56'N, 1 19°4] 'W, tree, 5.5 m tall, fruit plum-red to burgtmdy, 28 Aug 1994, Ph/pps,
J.B. &0'Kennon. R.J. 7105 (UBC, UWO, WS); Okanogan Valley, W side road about 7 km
N of Tonasket, huge hawthorn stand along riverside, ca. 975 fr, 48°44'N, 1 19°25'W; bush,
6 m tall, crimson foliage, no fruit, 3 Sep 1 993, Ph/pps. J.B. & O'Kef/i/ou. RJ. 6865 (UWO);
Sinlahekin Valley, ca. 5 mi S of Loomis, in depression near roadside, alt. 450 m, 48°45'N,
119°37'W, bush, 4 m tall, black fruit, 28 Aug 1994, Phipps.J.B. & O'Kenmm. RJ. 7] 09
(UWO); ca. 6 km NE of Omak, dense hedges in valley-bottom among farm lands, alt. 250
m, 48°27'N, 119"28'W, dense bush, 4 m tall, fruit red, changing Irom orange, 28 Aug
1994, Phipps.J.B. & (YKmrinn. RJ. 7100 (UWO). Whitman Co.: US 195 along Spring
Flat Creek few mi S of Colfax at MP 35.3, alt. ca. 775 m, 46"51'N, 1 17°2rW, tree, 6 m
tall, fruit plum-red (some apiiroaching chestnut), group of same, 28 Aug 1994, Phipps.
J.B. &(yKe)imii. RJ. 7099 (UWO); 66 Staley Rd., 1 km NE of Chambers, alt. 775 m, dry
roadside, young tree in front yard of Harold Bough, 46''39'N, 117"! 1 'W, bush, 4 m tall,
fruit shrivelled, 27 Aug 1994, Phipps.J.B. & (rKennon. RJ. 7096 (UWO).
2. Crataegus okanaganensis J.B. Phipps & O'Kennon, sp. nov. (Fig. 5).
Typi;: CANADA, British Columbia: Rough grass with hawthorns on Hwy, 97 near
entrance to Kelowna airport, alt. 1800 ft, equals JBP 6907, bush, 4 m tall, frtiit,
slightly dull and deep red, ± ellipsoid, 17 Aug 1994, Phipps.J.B. & (YKcnmni. RJ.
6974 (iiolotype: UWO; isotypes: CAN, TRT, UBC, US).
Phipps and O'Kennon, New species of Crataegus from Western North America 179
Fig. 5. Lme drawing of C. okanaga?iensnJ.B. Phipps and O'Kennon. Inflorescence and flower
parts from Phipps 6929 (UWO); infruccescence and fruit parts from Phipps & O'Kennon 7002
and 6975 (UWO); and sterile shoot from Phipps 6894a (UWO). Scale bars = 1 cm.
180 SiDA 18(1)
Frutex, 3—6 m altiis, ramLili LiniLis anni mt-diocriter brunnci vel arrobrLinnei, in JLivencuce
sparsim pLibescentes; spinae valiclae, 2—4 cm lon^ac, in jLivcnCLitc nitcntcr atrobrunneae,
rccurvatae. Folia dccickia, petiolata; pctioli 1—2 cm loni^'i, sparsim pubescences, eglandulares
vel cum glanckilis nigribus in jiiventLite (Oeston); laminae 35—6 cm longae, ovatae vel
ovaro-rhombeae, alicjuanclo late ellipticae vel angnsre ovatae, lobis 3—4 ± acutis, marginibus
SLibtiliter dentatis, venis 4-5 per latiis, alicjuantmn impressis stipra, in juventtite dense
appresso-pilosis, pilis attntis m matiintate, siibter pLibescentes praecipne m venis, coriaceae,
super nitentes praesertim in juventute, crescentes rubrae, aeneae in autumno. Inflorescentia
panicula convexa, 10—20 florata; ramuli varie villosi, glandulo-maculati, bracteolis linearibus
caducis glandulo-marginatis. Flotes 12—13 mm diam., conspicue ctipLililormes in |LiventLice;
hypantliiLim externe villosum solo basale vel glabrum omnino; lobi calycis angnste trian-
gulares, 3 — i mm longi, marginibus glandularibus, abaxiale cum pilis paticis; petala ± circularia,
alba; stamina 10, antheris albis vel raro pallide roseis; carpelli et styli 2— 3(— 4). Fructiis 8
mm diam., plcrumcpic urceolattis, glabcr, coccincLis in juventute sed in matuntatem coloris
vini vel atropurpuretis (raro tere niger), si in conditione bona ctim lobis calycis longis in-
signe erectopatentibus (seel aliquanto brevioribtis vel reflexis); pyrenae 2—3, dorsaliter stilcatae,
latenbus ± planis, paulo pinguibus vel patilo erosis.
Bush, 3—6 m tall; 1 year old rwigs mid to dark brown; young shoots thinly
pubescent; thorns stout, 2—4 cm long, shiny deep brown when young, re-
curved. Foliage deciduous, petiolate; petioles 1—2 cm long, thinly pubes-
cent, eglandular or with small black glands when young (as at Creston, B.C.);
blades 3.5—6 cm long, ovate to ovate-rombic, or less commonly broad el-
liptical to narrow-ovate, with 3^ sharp lobes per side, margins finely double
toothed, 4—5 veins per side, somewhat impressed above, densely appressed
hairy when young above, this abrading with age, pubescent principally on
the veins beneath, somewhat coriaceous, glossy above especially when young,
expanding leaves reddish, the fall colour bronze. Inflorescence a domed panicle,
10—20 flowered, the branches variably villous, gland-dotted, with cadu-
cous linear, gland-margined bracteoles. Flowers 12—15 mm diam, notably
cup-shaped when young; hypanthium externally villous only at biise or completely
glabrous; calyx lobes narrow triangular, 3—4 mm long, the margins glan-
dular, abaxially with a few hairs; petals ± circular, white; stamens 10, an-
thers ivory or occasionally very pale pink; carpels and styles 2— 3{— 4). Fruit
8 mm diam., generally flask-shaped, glabrous, red when young but later
burgundy to deep purple (occasionally almost black), when in good condi-
tion with strikingly long erecto-patent calyx lobes but these sometimes shorter
or reflexed; nutlets 2—3, dorsally sulcate, their sides flat, a little plump, or
shallowly eroded.
Distribution. — Widely distributed and common from the Okanagan val-
ley of southern British Columbia and northern Washington to northern Idaho
(Fig. 6). This is an extraordinarily handsome hawthorn in late August and
early September.
Crataegus okanaganensis in similar to C. willianisii of northwest IMontana as
illustrated by the numerical analysis (Fig. 1) but with more glossy foliage
Phipps and O'Kennon, New species of Crataegus from Western North America 181
Fig. 6. Distribution of C okariaf^anensis }.^. Piiipps and O'Kennon.
of a much more coriaceous texture and the color usually yellowish-green in
mid-season, the blades a little broader and the lobes usually less sharp, thorns
generally stouter, different anther color, and mature fruits deep purple (Fig.
9d) instead of sometimes blood or deep red; the young fruit is a brilliant
red (Fig. 9e) at the stage when C. williamsii may be pale to deeper dull red.
The color change from brilliant red (late August) to a deep dull purple (late
September) is nearly unique among Crataegus species. In northern Idaho at
2300 ft, C, okanaganemh flowered in 1 994 a good week before C. williamsii at
the same altitude in northwest Montana. Otherwise identical material with
five stamens is occasionally found. Crataegus okanaganensis was repeatedly
collected in the 198()s in Idaho by Fred Johnson though without recognition.
The two taxa may of ser. Purpureofructi be separated as follows:
1. Fresh anthers white or cream; inflorescence branches thinly to moderately
hairy; hypanthium usually glabrous; leaves coriaceous (Idaho and west).
C. okanaganensi s
1. Fresh anthers bright pink; inflorescence branches densely hairy; lower part
of hypanthium densely hairy; leaves thin (nw Montana) C. williamsii
Additional specimens examined: CANADA. BRITISH COLUMBIA: Okanagan Val-
ley, Westside Rd. NW of Vernon, ca. 2 km S of intersection with 97, 1450 ft, hedges on
both sides of road, 03 May 1994, Phipps. J. B. 6900 (UWO); ca. 1.2 mi E of Enderby on
road to Mabel Lake between 200 m & 25 m W of "Autobody collision & repairs" sign, alt.
1275 ft, hedgerow, bush, 3 m tall, fruit burgundy, 20 Aug 1994, Phipps, J. B. & O'Kennon,
R.J. 7016 (UWO); Rough grass with hawthorns on Fiwy. 97 near entrance to Kelowna
182 SiDA 18(1)
airport, S side of Airport Dr., alt. ISOO ft, btish 6 m tall, fruit red, slightly deep and dull,
± ellipsoid, 17 Aug 1994, P/npps. J. B. &0-Kc,nw,i. R.J. 697 5 (UBC, UWO); ca. 8 km NNE
ol Vernon center at Burke's Pick-your-own on Pleasant Valley Rd., bottom of roadside,
4()7 m; 5 m bush, dull burgundy, ellipsoid fruit, 24 Sep 199.% Phlpps. J.B. & O'KtiPwn,
R.J. 6777 (BRIT, MONTU, UWO, WS); Shuswap Valley, just E of Enderby, across bridge,
S side ol rd. on mound by roadside depression, alt. ca. 1 250 ft, bush, 5 m tall, 10 stamens,
in bud, 05 May 1994, Phtpps.J.B. 6912 (UWO); Okanagan Valley, ca. 9 km N of center of
Vernon off E side of 97, just S of jet. with 97A & just N of Baker Hogg Road, alt. .t50 m,
hawthorn thickets to N side of long cleared ride, bush, 5 m tall, 10 white anthers, 03 May
1994, Phipps.J.B. 6H94 (UWO); Okanagan Valley, ca. 9 km N of center of Vernon off E
side of 97, just S of )ct. with 97A & just N of Baker Hogg Road, alt. 350 m, hawthorn
(mainly) thickets to N side of long cleared ride, bush, 5 m tall, 10 white anthers, 03 May
1994, Phipps.J.B. 6894a (UWO); Shuswap Valley, ca. 2.8 km E of Enderby on Kingfisher
Rd., on hedgerow S side of road opposite house wirh wooden fence, N side, alt. I 280 ft,
btish, 4 m tall, possibly = JBP & RO'K 6812, foliage somewhat reddish, stamens 10. cream
anthers, 05 May 1994, Phipps.J.B. 6921 (BRIT, UWO); ca. 1.2 mi E of Enderby on road
to Mabel Lake between 200 m & 25 m W of "Autobody collision & repairs" sign, alt. 1275
ft, hedgerow, bush, 5 m tall, fruit reddish burgundy, 20 Aug 1994, Phipps.J.B. & (YKeinimi.
R.J. 701 7 (UBC, UWO); Salmon Arm, large hawthorn pasture just N of railroad and W of
central business section, alt. ca. 1900 ft, bush, 3-5 m tall, opposite building with blue
strip, flowers very cup-shaped, anthers 10, very pale pink, 1 I May 1994, Phipps.J.B. 6966
(BRIT, CAN, UWO); \l side of 97 A 0.5 mi N of inter.section with 97, scrubby slopes with
long cleated area, bush, 5 m tall, scarlet fruit, 19 Aug 1994, Phipps.J.B. &0'Kenno)i. RJ.
-^000 (UWO); Okanagan Valley, about 8 km NNE of center of Vernon, Pleasant Valley
Rd., behind Vowle's residence, hedge at S boundary of Burke's PYO, alt. ca. 380 m, dense,
broad mature hedge, biish, 3 m tall, nearest top of hedgerow, 10 anthers, ivory, 02 May
1994, Phipps.J.B. 68H2 (DAO, UBC, UWO); Castlega'r, near Selkirk College,' left diand
slope towards bottom of Rosedale Rd., alt. 970 ft, scrubby area, bush 2.5 m tall, prob.
=JBP & RO'K 6826, stamens 10, anthers white, 06 May 1994, Phipps.J.B. 6929 (BRIT,
TRT, UBC, UWO); Okanagan Valley, Westside Rd. NW of Vernon, ca. 2 km S of inter-
section with 97, 1450 ft, hedges on both sides of road, 03 May 1994, Phipps.J.B. 6901
(UWO); Okanagan Valley, west side of hwy. at entrance to Kelowna airport, alt. 1800 ft,
open scrub, bush, 4 m tall, stamens 1 0, anthers ivory to palest pink, 05 May 1994, Phipps,
J.B. 6907 (BRIT, TRT, UBC, UWO); West side of hwy. 21, just N of, and in view of,
Canada Customs post, roadside/forest edge, 2200 ft, tall, 5 m bush, 27 Sep \99oJ.B. Phipps,
J.B. & 0'Ke>i>!on. R.J. 6H32 (ID, UWO); NNW of Vernon on Wesrside Rd., ca. 2.5 km SW
ol intersection ot 97 and 97A, hedges along roadside, back of field, W side of rd., bush, 5
m tall with purple shrivelled fruit, 24 Sep 1993, Phipps.J.B. & O'Kennon. R.J. 6788 (C;AN,
TRT, UBC, UWO); E side of hwy. 21, ca. 250 m S of Creston Ferry Rd. to Hood Rd., this
species very common here, 1970 ft, dense bush, 3.5 m tall, ellipsoid, burgundy fruit, 27
Sep I 993, Phipps.J.B. & O'Ke/i/ioii. RJ. 6831 (ID, MO, LJWO); E side of hwy. 2 I , imme-
diately S of jet. with B(; 3, north of Oeston, alt. 1850 ft, scrubby bank, 2.5 m tall bush,
fruit shrivelled, black, 27 Sep 1993. Phipps.J.B. & O'Kennon. R.J. 683('>(UWO); Castlegar,
edge of Campus Rd. off Frank Beinder Rd., grassy slopes with scattered hawthorns, bush,
3 m, Ivs sharply lobed, abundant btirgundy IrLiit, 27 Sep 1993; Phipps.J.B. & O'Kennon.
R.J. 6826 (BRIT, UBC, UWO, US); Larry Calder property, E of 97 about 9 km NNE of
Vernon, scrubby partly cleared hillside, 475 m, bush, 4 m, reddish foliage, fruit a good
purple, 26 Sep 1993, Phipps.J.B. & O'Kennon. RJ. 687 9 (UWO); Kalamalka Prov. Park, S
of Vernon, within 200 m of yellow gate, at parking lot, damjiish depression, ca. 1375 ft,
PuiPPS A\D O'Kennon, New species of Crataegus from Western North America 183
common, multistemmecl bush, fruit oblong-orbicular, burgundy, 24 Sep \9^y,Phipps.J.B.
6 O'Ktminii. R.J. 6798 (BRIT, UWO); NNW of Vernon on Westside Rd., ca. 2.5-3 km
SW of intersection -of 97 and 97A, hedges along roadside, back of field, 5 m bush with
purple fruit, 24 Sep 1993, Phipps.J.B. & O'Kennon. R.J. 679/ (UWO); E side of hwy. 21,
immediately S of jet. with BC3, north of Creston, scrubby banks, 1850 h, 2.5 m bush,
leaves green turning purple, ellipsoid, burgundy fruit, 27 Sep 1993, Phipps.J.B. & O'Kennon,
R.J. 6828 (UWO, V); N of Creston, E side of Hwy. 21, grassy banks with hawthorns on
top, just S of junction with Hwy. 3, alt. 1850 ft, bush, 2.5 m tall, stamens 10, anthers
white, equals JBP & RO'K 6828, 07 May 1994, PhippsJ.B. 6930 (UWO); Salmon Arm,
large hawthorn pasture just N of railroad and west of central business section, alt. ca. 1900
ft, bush, 3.5 m tall, fruit bright red, equals JBP 6966, 20 Aug 1994, Phipp.s.J.B. &0'Kenuon.
R.J. 7024 (BRIT, TRT, UWO); E side of 97A 0.5 mi N of intersection with 97, scrubby
slopes with long cleared area, young plant on south side of site, scarlet Iruit, 19 Aug 1994,
Phipps.J.B. & O'Kennon. RJ. 7001 (UWO); E side of 97A 0.5 mi N of intersection with
97, scrubby slopes with long cleared area, young plant on south end of site, scarlet fruit,
19 Aug 1994, Phipps.J.B. & O'Kennon. R.J. 7002 (UWO); Hwy. 6 ca. 6 road mi W of
Fauquier, west of Arrow Lake in Inookalin Valley, along fenceline in farmland, alfalfa fields,
alt. 475 m, bush, 4 m tall, fruit bright red, 22 Aug 1994, Phipps.J.B. & O'Kemion. R.J.
7045a (UBC, UWO); Okanagan Valley NNE of Vernon, Pleasant Valley Rd., hedge be-
hind Vowle's residence, E end boundary fence with Burke's P-Y-O, alt. 380 m, bush 3 m
tall, fruit large, plump, purple-red, hairy, with stout erect calyx lobes, 19 Aug 1994, Phipps,
J.B. & O'Kennon. R.J. 6996 (UBC, UWO); Slocan Valley, S of Slocan, above swamp cross-
ing river along small road off Hwy. 6 & just N of Perry Siding, hedges and brushy slopes,
alt. 1775 ft, bush, 5 m tall, fruit bright red, 22 Aug 1994, Phipps.J.B. & O'Kennon. R.J.
7048 (UWO); Shuswap Valley, just E of Enderby, across bridge, E side of road, 200 m N of
gate, alt. ca. 1250 ft, level open field, growing along roadside across from cattle feeder, 4
m tall, fresh anthers cream, 1 5 JVIay 1 995, Dowotwk. L.S. & D. 2741 (UWO); Shuswap Valley,
just E of Enderby, across bridge, E side of road, alt. ca. 1250 ft, level open field, growing
along roadside across from cattle feeder, 2.5 m tall, fresh anthers cream, 15 May 1995,
Donovan. L.S. & D. 2744 (UWO); Okanagan Valley, Kelowna, service road at W side of
airport, alt. ca. 1250 ft, fenceline, bush 3 m tall; fruit brilliant red, ellipsoid, 17 Aug 1995,
Phipps.J.B. 7155 (UWO); Okanagan Valley, Oyama, ca. 0.5 mi E of E end of spit, alt. ca.
1320 ft, hedgeline on clay soil, bush, 4 m tall, fruit ellipsoid, red, 17 Aug 1995, Phipps,
J.B. 7157 (UWO); Okanagan Valley, Oyama, ca. 0.5 mi E of E end of spit, alt. ca. 1320 ft,
hedgeline on clay soil, bush, 4 m tall, fruit ellipsoid, 17 Aug \995, Phipps. J.B. 7^9 (UWO).
U.S.A. IDAHO. Bonner Co.: Eowcr Priest River, river bank, alt. 660 m., scarlet fruit,
07 Aug 1897, Leiber}^.John B. 285^ (OSC); Pend Oreille R. across from Oldtown, N of
bridge. Section 24,T56N, R6W, scrub just above high water mark, dominated by C. coliimb'iana,
2000 ft, 07 Aug \9'66 Johnson, pyed 86126 (IDF); Eeclede, Riley Creek, 8 mi E of Priest R.
at Pend Oreille R. (Albenai Falls Reservoir) T56N, R3W sec. 30, riparian fringe along res-
ervoir climax forest, gentle slope, common, 12 Aug \9^'8, Johnson, P. 88079 (IDF); Pend
Oreille R. across from Oldtown, N of bridge, T56N R6W S24, 2000 ft, shrub fringe just
above high water mark, abundant, 07 Aug \986, Johnson. Fred 861 26 (ID (#93728); 6 mi
E of Priest R, along Albenai Falls Reservoir in T56N, R4W sec. 34&35, riparian fringe,
1900 ft, common, 12 Aug 1988, Johnson. P. 88082 (IDF); Section 28,T56N, R3W along
Pend Oreille R. reservoir, roadside fringe of trees/shrubs by hayfield, many thornless trees,
height to 15 ft, 07 Atig 1986, Johnson. P. 86124 (ID, IDF). Boundary Co.: US 95, about
5 km S of jet. with Idaho 1 & S of turnout on right below tall conifers, alt. 2250 ft, on E
plowed field backed by steep rocky, treed slope, dense bush, 3 m tall, stamens 10, anthers
184 Sum 18(1)
ivory, OH May L99 i, Plu/>/>s. J.B. 69U (ID, IJWO); US 95, ca. 5 km S of jet. with Idaho 1
& slightly S ol turnout on ri,i^ht below tall conifers, alt. 2250 ft, on E plowed field backed
by steep rocky, treed slope, dense bush, 3 m tall, stamens 10, 07 May 1994, Phipps. J.B.
6933 (ID, UWO, WS); US 95 2 mi S of jet. with Idaho 1 , W side of road, roadside, 21 Sep
199.^, Phipps. J.B. & (yKt^inin)i. R.J. 6S34 (UWO); 2.6 mi S of jct. US 95 & Hwy. 1, in
T64N, R 1 E sec.2H (SE of C;o|->eland), roadside shrub frmge, 2200 ft, gentle slojx', 18 ft tall
X 22 (t wide, 5 main stems 3-5, possible escape, no farms near, 12 Aug \99S, Johnson. F.
88072 (IDF). Kootenai Co.: floodplain of Spokane R. at Coeur d'Alene, 1/4 mi W of US
95, grassy meadow with scattered trees, no slope, open, 2000 ft, tare with C. cloz/^hisii, 10
jun 19cS2,>/mTO;/. P.D. (IDE (#8604 IS)); Spokane River, 1/4 mi E of Washington border,
2100 ft, among granitic rocks and sand, shrubs to 3 m, 01 Aug \9't\G, Johnson, F. 8644 (ID
(#93729)); Coeur d'Alene, 20 Jun 1930, Ri/st.J.H. (ID), 20 Jun 19t0; floodplam of Spo-
kane R. at Coeur d'Alene, 1/4 mi W of US 95, grassy meadow with scattered tree/shrubs,
elev. 2000 ft, 10 Jun \9^2, Johnson. Fiecl{\r) (#93730)); lloodplain of Spokane R., at Coeur
d'Alene, 1/-I mi W of LJS 95, grassy meadow with scattered trees & shrubs, no slope, open,
2000 ft, 16 Jun [9^, Johnson. ED. (IDF). WASHING'EON. Okanogan Co.: Westside
Rd. ca. 7 km N of Tonasket, 975 ft, along riverside, 7 m bush, smaller leaves, burgundy
fruit, 30 Sep 1993, Phipps. J.B. & (VKcnnon. RJ. 6868 (UWO); Westside Rd. about 4 km
N of Tonasket, alt. 97 5 ft, bush 5 m tall, riverside hawthorn thicket with a few poplars,
fruit + erect, lake-red, broad-elliptic, 23 Aug 1 995, PA//;/;.i-,./.B. 7/7.5 (UWO, WS). Spo-
kane Co.: Spokane, damp places, Siiksdorf. W.N. 919 (WS), Aug 1889. Whitman Co.:
Pullman. P/>w. C.V. (CAS), May 1902; Westend; 1800 ft, spring 1976, May. R. 91 (RMV).
ENGLAND: cult. Surrey, Kew, Royal Botanic Gardens; Kew #197-67, 19728; 22 May
1985, Phipps. J.B. 5651.
The discovery of C. ()kcinaganeiLsi.s and its clear relationship to C. wiUianisi'i
merit the creation of a new series, as follows:
Series Purpureofructi J.B. Phipps &0'Kennon, ser. nov. Fvpu: C. okmaganensis
J. 11 Phipps and O'Kennon.
Frutices, plerumc]tie grandes, vel arbores parvae; spinae mediocres in longittidine (2—4
cmlongae). Folia plerumque ± coriacea(|uaeterC. inllicnnsii), laminae plerumqiie + rhombeae,
(vel eliptico-oblongae vel rhombo-ovatae), plerumque lobis 3-4 brevibus patentibus pier
latLis. Inflore-scentia 10-25 florara. Flores mediocres (10-15 mm)diam; stamina 5-10; carpelli
et styli 2-1. Eructus coccineus, plerumcjue dare coccineus in Augusto exeunte, in maturitate
Septembri exeunte fierens .sanguineus vel atropurpureus, orbicularis vel ampulliformis; pyrenis
3—4, lateribus erosis vel |Tlanis.
Bushes, ustially large, to small trees; thorns medium in length (2-4 cm).
Leaves generally ± coriaceous (except C. willtamsii), blades usually ± rhom-
bic (to elliptic-oblong or rhomb-ovate), usually with 3-4 short outward-
pointing lobes per side. Inflorescences 10-25 flowered; flowers not large
(10-15 mm diam.); stamens 5-10; carpels and sryles 2-4. Fruit red, usu-
ally brilliant red in late August, becoming deep blood-red to dark purple
at maturity, orbicular to flask shaped; pyrenes 3-4, sides eroded or plane.
Distrihiituni. — A distinctive series with two known species and perhaps
a couple of others found from the interior Pacific Northwest and southern
British Columbia to northwestern Montana. Crataegus wiliamsii was origi-
Phipps and O'Kennon, New species of Crataegus from Western North America 185
nally placed in series Rotundifoltae by Eggleston but the ellipsoid fruit, long
calyx lobes in fruit, dark-colored twigs and somewhat erose nutlets suggest
a different affinity. Our numerical analysis (Fig. 1) indicates no relation-
ship whatever to C. chrysocarpa in ser. Rotundifoliae.
3. Crataegus phippsii O'Kennon, sp. nov. (Fig. 7). Type: CANADA. British
Collmbia: Okanagan Valley, Pleasant Valley Rd., ca. 8 km NNE of Vernon center,
bush, 4 m tall, fruit red, 24 Sep 1993, Phipps, J. B. and O'Kennon, R.J. 6780 (holo-
type: UWO; isotyphs: BRIT, CAN, TRT, US, WS).
Arbor vel frutex magnus, 5-6(-7) m alta, trunco uno dominante sed aliquanto cum surculis;
ramuli unius anni atropurpureiZ-brunnei, veteres atrocinenerei; in juventute dense et breviter
pubescens; spinae aliquantum sparsae, atrobrunneae vel nigrae, leviter recurvatae, aliquantum
tenues, 2—4 cm. longae. Folia decidua, petiolata; petioli 1.5—2.0 cm longi, pubescentes;
laminae 4-8 cm longae (sed ad 10 cm vel plus in blastis elongarionis), late ellipticae vel
ovatae vel deltoideo-ovatae, lobis fere 3 ± vadositer obtusis vel late angulatis per latus,
marginibus breve dentatis; ca. 4 (—5) venis secondariis per latus, supra breviter appresso-
pubescentes subter breviter pubescentes praecipue in venis, aliquanrum coriaceae postea.
Inflorescentiapaniculadepressa, 6—12 florata; ramuli dense breviter-tomentosi, bracteolati;
bracteoli lineares glandulo-marginati. Flores 1 5—22 mm diam.; hypanthium externe breviter
tomentosum; lobi calycis late triangulares, ca. 5 mm longi, abaxiale pubescentes praecipue
proximale, marginibus glandulo-laciniatis; petala ± circularia; stamina 10, antheris roseis
pallide; styli carpellique 2—4, stigmatis capitatis. Fructus 12 mm diam., breve-pubescentes,
maturentes ab coccineo ad pruneo-purpureum vel nigrum, ± orbiculares vel late urceolati,
lobis calycis reflexis; pyrenae 3, vadositer sulcatae dorsale, lateribus planis.
Tree or large bush, 5— 6(— 7) m tall, with main trunk, but sometimes suckering;
1 year old twigs dark purple-brown, older dark gray, young shoots densely
short pubescent; thorns somewhat sparse, deep brown to black, slightly recurved,
somewhat thin, 2-4 cm long. Foliage deciduous, petiolate; petioles 1.5-
2.0 cm long, pubescent; blades 4—8 cm long (but on shoots of elongation
to 10 cm or more), broadly elliptic to ovate or broad deltoid-ovate, with
about 3 ± shallow rounded to broadly angled lobes per side, the margins
short-dentate; about 4—5 secondary veins per side, shortly appressed hairy
above, shortly hairy especially along the veins below, somewhat coriaceous
later. Inflorescence a 6— I2-flowered flattened panicle, the branches densely
short-tomentose, bracteolate, the bracteoles linear, gland-margined. Flow-
ers 1 5-22 mm diam; hypanthium externally densely tomentose; calyx lobes
broad-triangular, ca. 5 mm long, pubescent abaxially, especially below, the
margins glandular— laciniate; petals ± circular; stamens 10, anthers pale pink;
inside of hypanthial cup mostly glabrous, bristly at center; carpels and styles
2—4, the stigmas capitate. Fruit 12 mm diam., short-hairy, ripening through
shades of red to plum-purple to black in colour, ± orbicular to broadly flask-
shaped, pubescent, with reflexed calyx lobes; nutlets 3, shallowly dorsally
ridged, lateral walls ± plane.
Distribution. — This species is not so common as the other two described
186
SlDA IS(1)
Fig. 7. Line drawing ot C. phippsii O'Kennon. Inflorescence flowers and parts from Phipps
6891 (UWO); mfructescence, fruit and parts from Phipps 6780 and O'Kennon (UWO); two
smaller isolated leaves from Phipps 6780 aud O'Kennon (UWO); magnified leaf (bottom left)
from Phipps 6891 (UWO). Scale bars = 1 cm.
Phipps and O'Kennon, New species of Crataegus from Western North America 187
in this paper but is nevertheless widespread and has been found at fourteen
focahties ranging from the Okanagan VaUey of southern British Columbia
and northern Washington to the Flathead Valley of Montana (Fig. 8). We
have never found it numerous at any site.
Crataegus phippsii is one of the most distinctive species of hawthorn to
have been found in North America in recent decades and does not appear
to be closely similar to any other known in the west, a feature emphasized
in Figure 1 . It is a large hawthorn, reminiscent of C, ?nollis, the latter a common
species of the interior of the continent, in foliage, habit and pubescence while
in its red (Fig. 9a) changing to purple (Fig. 9b) to black fruit it is similar to
a number of mostly previously undescribed species of the Pacific North-
west and British Columbia. The foliage colours attractively in autumn (Fig.
9b, 90. The mature bark sometimes flakes off in longitudinal strips, an unusual
feature in hawthorns (Fig. 9c).
Crataegus phippsii was first collected by B.T. Butler (nos. 321-325) in
1908 and these specimens were cited by Eggleston (1909) in his protologue
for C. williamsii. The many differences from C. williamsii, however, show
that C. phippsii in not in this affinity. That Eggleston thought it somewhat
distinct is shown by his annotations on the sheets of "var. ovata," a name,
however, never published. Beyond this brief glimpse of notoriety, the present
species, very striking to the present authors, remained totally overlooked
until our collections of recent years.
The large, broad leaves with rounded lobes and hairiness of all parts sug-
gests an affinity with series Molles. Superficially, it most nearly resembles
the local species C. noelensis Sarg. of Missouri, a taxon which, however, has
not been seen for many years. A combination of stamen number, anther color,
foliage size and shape, and fruit color are the main characters distinguish-
ing the species in the Molles group as shown in the following key. For con-
venience, I am including C. greggiana in this key although it is now placed
in set. Greggianae.
KEY TO C. PHIPPSII AND MEMBERS OF SERIES MOLLES
1. Foliage small, on short shoots mainly 4 cm long.
2. Flowers 1 — 1.5 cm diam.; stamens usually 1 C. greggiana
2. Flowers ca. 2 cm diam.; stamens 20 C. lanuginosa
1. Foliage larger, on short shoots generally 5-8 cm long.
3. Anthers pink.
4. Stamens 5-10.
5. Fully ripe fruit orange-red; leaves ± unlobed C. noelensis
5. Fully ripe fruit deep purple to black; leaves evidently lobed C. phippsii
4. Stamens 20.
6. Fruit red C. cexana
6. Fruit yellow C. viburnifolia, C. kellogii
3. Anthers white to cream.
188
SiDA 18(1)
Fic. 8. Distribution map of C. phippsii O'Kennon.
7. Stamens 20 C. mollis
7. Stamens 10 C. submoUis, sens. lat.
Additional Specimens examined: CANADA. BRITISH COLUMBIA: Okanagan Val-
ley, 8 km NNE of center of Vernon, Pleasant Valley Rd., behind Vowle's residence, hedge
at S boundary of Burke's PYO, dense, broad mature hedgeline, bush 4 m, commencing
flowering, 7-8 pale pink anthers, 02 May 1994, Phipps.J.B 6878 (BRIT, CAN, TRT, UBC,
UWO, US); Okanagan Valley, 8 km NNE of center of Vernon, Pleasant Valley Rd., be-
hind Vowle's residence, hedge at S boundary of Burke's PYC), dense, broad mature hedgeline,
bush 4 m, foliage glossy, pedicels hairy, 10 pink anthers, 02 May I 994, Phipps.J.B. 6880
(CAN, TRT, UBC, UWO); Okanagan Valley, rough, scrubby slope above dirt road NW-
bound into O'Keefe Ranch, opposite L. & A. Cross Rd. off 97A, alt. 1290 ft a.s.I., stamens
10, anthers light pink, equals JBP & RO'K 6803, 03 May 1994, Phipps.J.B. 6891 (UBC,
UWO, WS); ca. 1 km N of Vernon cenrer, on dirt road N-bound into O'Keefe Ranch off
Hwy. 97 A at L.& A. Cross Rd., dry rocky hillside, suckering clump of trees to 5 m, 25 ft
across, bark coarse, plated, peeling, iruit purple, 25 Sep 1993, Phipps.J.B. & O'Kennon.
R.J. 68VJJ (CAN, UWO); Pleasant Valley Rd. ca. 8 km NNE of Vernon Center, southern
boundary hedge of Burke's PYO behind Vowles' residence, alt. ca. 380 m, dense, broad
mature hedgeline o'^ Cratciegiis, bush 4 m tall, fruit deep red, 24 Sep 1993, Phipps, J.B. &
O'Kennon. R.J. 6780 (BRIT, CAN, MO. UWO); ca. 5 km NNE of Enderby, on E-W rd.
between Shuswap R. and Hwy. 9^ A, long hetlgerows, alt. ca. 1375 ft, bush, 5 m tall, truit
red, 21 Aug 1994, Phipps.J.B. & O'Kennon. R.J. 7038 (UWO); ca. 3 km NE of Enderby on
road to Sicamous (Hwy. 97 A), bottom of scrubby slope on east side, alt. ca. 1400 ft, clump
of 5 m bLishes, reddish fruit, 2 1 Aug 1994, Phipps.J.B. & OKainon. RJ. 7037 (BAO, UWO);
ca. 5 km ENE of Armstrong on E-W aligned rd. jusr ENE ot Armstrong airport, hedgerows,
S side of road, alt. 1450 ft, bush 5 m tall, 21 Aug \994J'hipps.J.B. & O'Kennon. RJ. 704!
Phipps and O'KiiNNON, New species of Crataegus from Western North America 189
Fig. 9. Various pictures of C. phippsii O'Kennon and C okanagai/ensis J.h. Phipps & O'Kennon:
a) typical reddish late August fruit color of C. phipps//, northern Okanagan, BC, 19 Aug
1995; b) typical later September fruit color of C. phippsii with fall h)liage color beginning,
northern Okanagan, BC, 24 Sep 1993; c) trunks of C. ph/ppsii showing shredding bark,
shores of Palmer Lake, Washington, late Aug. 1 994; d) full ripe fruit color of C okanaganensis,
Westside Rd., nr. Vernon, BC, 24 Sep 1 99.3; e) late August fruit color of C okanaganensis,
nr. Kelowna, BC, 17 Aug 1995; 1) R.J. O'Kennon holding fruiting branch, Palmer Lake,
Okanagan Co., WA, 01 Oct 1993; note fall foliage.
190 SiDAhSd)
(UWO); ca. 6 km N of Armstrong, branch of Deep Creek, thickecs, primarily C. doKghtsii,
in narrow flood-plain, alt. 1450 ft, tree 4 m tall, Fruit red, 21 Aug 1994, Phipps.J.B. &
O'Kennon. R.J. 7034^ (UWO); ca. 3 km NW of Armstrong, N-S road parallel to Schubert
Rd. and to the W of it, E side, alone by telephone pole, alt. ca. 1400 ft a.s.l., bush 3.5 m
tall, dark green leaves, dull reddish fruit, 2 1 Aug 1994, Phipps.J.B. & O'Kennon, R.J. 7033
(BRIT, UBC, UWO).
U.S.A. MONTANA. Flathead Co.: about 0.5 km along Tower Rd. off US 93, nearSW
corner of Flathead Lake, hawthorn thicket along trail, rear of small field, alt. 2950 ft, equals
JBP 7078, bush 7 m tall, overtopping all others, at popcorn, anthers pink, 20 May 1997,
Phipps.J.B. & O'Kemimi. RJ. 7 591 (MONTU, UWO). Lake Co.: Hwy. 93, mipost 63.1,
Tower Rd., (major pablillo), 2.0 mi. NW of Poison Bridge, 8-( 1 1-12)-14 flowers, 10 pink
anthers, leaves hairy, shaggy, in green bud stage (forced), 18 May 1 995, O'Kennon, R.J. 13 1 56
(UWO); Flathead Lake, SW border, 29 Aug 1908, Buthr. B.T. 322 (NY); Rocky Point
Road NW of Poison at 1.7 mi N of hwy 93 and 0.3 mi NF of Jim's Road, alt. 2920 ft,
extensive thickets near lakeshore behind new housing, bush, 5 m tall, Iruit orbicular, red-
dening, 25 Aug 1994, PhtppsJ.B. &0'Kenmin. RJ. 7080 (BRIT, ID, MO, MONTU, UWO);
Flathead Lake, SW border, 19 Aug 1908, Bu/hr. B.T. 323 (NY); Flathead Lane, off Tower
Rd., olThwy 93 few mi NW of Poison, thickets along creek back of field, alt. 3050 ft, tree
5 m tall, fruit partly deep oxblood, partly paler, 25 Aug 1994; Phipps.J.B. & O'Kennon,
RJ. 7078 (DAO, MONTU, UWO, US); Flathead Lake, SW border, 19 Aug 1908, Butler.
B.T. 321 (NY). WASHINGTON: Okanogan Co.: Palmer Lake, N side., open hawthorn
thicket at upper level of pebble beach at picnic spot/cam|\sife, just E of Cdiopata Lodge,
small tree, 6 m tall, mostly finished flowering, stamens 10, anthers pale pink in bud,=JBP
6 RO'K 6874, 04 May 1994, Phipps.J.B. 6904 (ID, rVIO, UWO, WS); W of US 97 on
crossroad at EUisford, in floodplain to S side of road, bush, 7 m tall, anthers 1 0, pale pink,
equals JBP & RO'K 6876, ()4 May 1 994, Ph/pps.J.B. 6906 (US, UWO, WS); Palmer Lake,
N side, open hawthorn thicket at upper beach level on campsite just E of Chopata Lodge,
alt. 1 150 ft, fine tree, 7 m tall, burgundy fruit; 30 Sep 1993, Ph/pps.J.B. & O'Kennon. RJ.
6874 (DAO, ID, MO, UWO, WS; E shores of Palmer Lake, 0.7 mi S of campsite at N end,
thickets, alt. 355 m, bush, 4.5 m tall, deep plum-red fruit, 28 Aug 1994, Phipps.J.B. &
O'Kennon. RJ. 7/0^' (BRIT, ID, UWO, WS); Okanogan Valley, cross-road at EUisford, alt.
950 ft, bush 7 m tall in field in floodplain, S side of road, fruit blackish, 01 Oct 1993,
Phipps.J.B. & O'Kennon. RJ. 6876 (UWO, WS).
ACKNOWLFIXIMENTS
We wish to thank the National Research Council of Canada for supporting
this research, Susan Laurie-Bourque of Hull, Quebec for the line illustra-
tions and Antony Littlewood of the Department of Classical Studies, Tlie
University of Western Ontario, for checking the Latin diagnoses. Peter Wood
of Selkirk College, Castlegar is thanked for field assistance. J. B. Phipps thanks
R.J. O'Kennon for supporting the color printing.
REFERnNC;ES
Eggleston, W.W. 1909. New North American Cnitaegi. Bull. Torrey. Bot. (4ub 36:639-
642.
Hoi.MGRi'N, N.H. 1997. Rosaceae. In: A. Oonciuist, et al., etis. Intermountain Flora, vol.
3A. New York Botanical Garden, Bronx.
PiiiFPS, J.B. 199x. Taxonomic and nomenclatural problems in black and burgLmdy-truitetl
Cviitciegns (Rosaceae) sp)ecies: C erythropodci , C. rivnLiyis and C sciligna. Sida, submitfeil.
Phipps and O'Kknnon, New species of Crataegus from Western North America 191
Phipps, J.B. 199H. Introduction to the red-truited hawthorns {Crataegus, Rosaceae) of western
North America. Canad. J. Bot. 76(ui press).
Sakcent, C.S. 1907. The black-lruited hawthorns of western Nortli America. Bot. Gaz.
44:64-66.
Welsh, S.L. 1987. Rosaceae. In: A Utah Flora. Great Basin Naturalist Mem. 9:520-543.
192 SiDA 18(1)
BOOK REVIEW
Annr HurcHiNcs, and Ai.ak Haxton Scott, Gillian Lfavls, and Anthony
Cunningham. 1996. Zulu Medicinal Plants: An Inventory. (ISBN
0-86980-893-1, pbk; ISBN 0-86980-923-7, hbk). University of Na-
tal Press, Private Ba^^ XOI, Scottsville 3209, Republic of South Af-
rica, in association with University of Zululand and National Botani-
cal Institute. Exclusive distributor: hiternational Specialized Book Services,
Inc., 5804 N.E. Hassalo Street, Portland, OR 97213-3644, U.S.A.
(503) 287-3093; FAX (503) 280-8832. $110.00. 450 pp.
The sLibtitk- Lindersratcs the amount ot intormation in tliis tightly organized compendium
of information about plant u.sed by Zulu traditional healers. The authors have chosen to
arrange their entries in phyh)genecic order to facilitate comparisons between related plants.
As this publication deals exclusively with plants of southern Africa, the classification fol-
lows that used in the National MerbariLim. Cieiuis numbers reflect the standards tised in
herbaria arranged in the Englerian system with additional standards for ferns and grasses.
These are referenced in the introduction.
The entry lor each species includes geogra|-ihical distribution, a short ticscription, local
names in English, Alrikaans and Zuhi followetl by documented use by Zidu healers and
then those referring to use by other grtjtips in sotithern Africa. Physiological effects, Chemical
constituents and biological i:)ro|ierties are added where information is available. These gaps
will tease and stimulate further research condticred, it is hoped, under the ethical and legal
codes iirotecting inieljecttial propert)' rights.
There is a wealth of information not only on the medicinal uses of these plants, but by
the social and cultural uses too. These range from love potions to charms against lightning
and incense to invoke the goodwill of ancestors. However, the autiiors do warn readers nor
to experiment!
For those who do not own a copy ol Watt and Breyer-Brandwijk's 1962 edition the Me-
dtctthil and Poisonous plants of Southern and Eastern Afriea this book is a must. For those for-
ttinate to own a copy this will be a vakiable supplement, for at least a third of the 1032
species mentioned here are new.
With the worldwide renewal of interest in natural piroducts this book has a huge poten-
tial readership. In each foreword, we arc- reminded of the value, and ephemeral nature, of
traditional knowledge. 'Fhis book makes it less likel\' we will lose the traditional ethnobo-
tanical knowledge of the Ztilu nMion.- 1- ion a Norris
SiDA ISd): 192. 199S
TAXONOMIC CLARIFICATION OF ATRIPLEX
NUTTALLII (CHENOPODIACEAE)
AND ITS NEAR RELATIVES
HOWARD C. STUTZ
Department of Botany and Range Science
Brigham Young University
Provo. Ur 84602, U.S.A.
STEWART C. SANDERSON
USD A Forest Service Shrub Sciences Laboratory
Provo. UT 84601. U.S.A.
ABSTRACT
In 1874 Sereno Watson described Atriplex niittallii as a new species, citing his number
981 collecrion, made in 1868 from northern Nevada, as a representative. His collection
consisted ot three plants each collected from a separate population all mounted on one sheet.
One is A. canescem^ two are the new species, A. nuttall'ii. Failure to recognize the original
cohection and collection site of A. niittallii has resulted in assignment of the name A. niittallii
to several different A /'r//' /ex species in western North America. However, by using cjuanti-
tative as well as qualitative differences, A. nuttciUii is readily distinguished from its near
relatives A. cuneata, A. falcata, A. ^ardnen, and A. tridmtata. The principal distinguishing
features include plant size and habit, leaf dimensions, fruiting-bract size and shape, fruit-
ing bract appendages, chromosome number, saj^onin content, flavonol content, dates of
antliesis, and geograpiiic distribution.
RF.SUMI-N
En 187-1 Sereno Watson describio Atriplex niittallii como nueva especie, citando su recoleccion
niimero 98 1 , hecha en el norte del estado de Nevada, E.E.U.U., como representativa. Consiste
esta recoleccion en tres plantas, de distintos lugares pero colocadas todas en un mismo pliego
de herbario. Una de ellas es A. canescens y dos son de la nueva especie, A. nuttalli. El no
darse cuenta los botanicos de la colecta original y su lugar de origen ha tenido como consecuencia
la aplicacion del nombre de Atriplex niittallii a varias otras especies de Atriplex del oeste de
Norte America. Sin embargo, usando tanto caracterfsticas cuantitativas como cualitativas,
se distingue facilmente A. niittallii de las especies alines, A. ciineata, A. falcata, A. ganlneri,
y A. tridentata. Las caracterfsticas de mayor utilidad son la altura de la planta, las cHmensiones
de la hoja, la forma y tamafio de kis bracteas frucn'feras, sus apendices, el numero de cromosomas,
contenido de saponinas, contenido de flavonoicles, fecha de antesis, y la distribucion geognifica.
INTRODUCTION
In 1 874, Sereno Warson described A /r//?/f.v nuttallii as new (Wacson 1 874)
and listed his collection number 981, obtained in 1868 in northern Ne-
vada, as a representative of the species. This collection consists of three plants
SiDA 18(1): 19.0-212. 1998
194 Sii)AlS(l)
collected at different times in separate locations in northern Nevada, all
mounted on one herbarium sheet (GH!) (Fig. 1), and each labeled by Watson
as Obione carmcens Moq. The plant on the right is A. canescens (Pursh) Nutt.
collected in June, 1868, in Unionville Valley, Pershing County, Nevada.
The other two are A. nuttallii. The middle plant, which appears to have
been broken and folded back, was collected in September, 1868, in Thou-
sand Springs Valley, Elko County, Nevada. The plant on the left was col-
lected in July 1868 in Reese Valley, Lander County, Nevada. Currently there
are still extensive populations of these species in the valleys where Watson
collected them.
As shown in Figure 1, several botanists have recognized differences be-
tween these three plants and have provided annotations for them. In 195 1 ,
Grant D. Brown annotated the plant on the right as Atnplex canescens (Pursh)
Nutt. In 1962, C.A. Hanson annotated the plants on the left and right as
Atriplex bonnevillensh Hanson and the central one as Atriplex falcata (Jones)
Standley. In 1972, I.J. Bassett and C.W. Crompton annotated the plant on
the right as Atriplex canescens, the central one as A. mittallii -vat. falcata, and
designated on the annotation label, the left specimen as the lectotype of
Atriplex nuttallii S. Watson. Later, McNeil et al. (1983) designated as the
lectotype of A. nuttallii, a specimen collected in Saskatchewan, Canada, by
Bourgeau {s.n.), in 1857, and included by Watson among 10 other collec-
tors of A. nuttallii, in addition to Watson 981 . However, as directed in the
International Code of Botanical Nomenclature, Article 9-9 (Greuter et al.
1 994), — " when the material designated as type is found to belong to more
than one taxon, a lectotype — may be designated," and, as described in
Article 7.5 of the 1988 edition of the code (Greuter et al. 1988), "A lecto-
type is a specimen or illustration selected from the original material to serve
as a nomenclatural type when no holotype was designated at the time of
publication — . When two or more specimens have been designated as types
by the author ... the lectotype must be chosen from among them." Since
Watson listed in the protologue, his number 981 as a representative of the
new species, he ostensibly considered it typical and, since this collection
consists of three specimens, one of them must be chosen as the lectotype.
Consequently, the lectotype of Atriplex nuttallii is the specimen on the left
(GH) as annotated by Bassett and Crompton in 1972. The designation by
McNeil et al. of a specimen collected in Saskatchewan, Canada as the lecto-
type of A. nuttallii, was therefore unnecessary. The illustration in Bassett et
al. ( 1 983) of the Saskatchewan plants, is representative of a diploid A, gardneri
(Moquin-Tandon) Hall & Clements which is common throughout south-
ern Alberta, southern Saskatchewan and northern Montana but very differ-
ent from A. nuttallii of northern Nevada. In his protologue, Watson de-
Stutz and Sanderson, Atriplex nuttallii and irs near relatives
195
iSt
S -'■
r ■ - -y Y'
/^>
Fk,. 1. Atrip/ex ni/thiHii. The leccotypc o\' Atriplex nintnllii (Walsou 98/, upper left corner)
collected in Reese Valley, Lander County, Nevada, July IH^iH- The central element is A.
riiittallii collected in Thousand Springs Valley, Elko County, Nevada, September, 1868.
The specimen on the right is Atripkxaimsmis collected in Unionville Valley, Pershing County,
Nevada, June 1S6H.
196 Si DA 18(1)
scribes A. yiuttallu -^"^ being 1—2 feet high. This is the stature of the A. nuttalln
plants in the populations he collected in northern Nevada (Table 1) but not
A. gcirdnert plants which are mostly 6-1 2 inches tall, nor A. cciuescens plants
which are seldom less than 30 mches in height. The Saskatchewan plant
illustrated in Bassett et al. (1983) appears to be ca 12 inches in height.
In the botany treatment of the King expedition (Watson 1871), Torrey
provided identification for Watson's collections and Watson provided the
descriptions (footnote, page 287). Referring to Watson 98 1 , Torrey identi-
fied it as Obionecanescens Moq. and Watson applied that name to each of the
three plants. Watson's description of the collection specifically referred to
his number 981 and indicated that "most of the specimens have nearly or
quite wingless fruit. Others have the bracts considerably dilated, though
still less than is frequently the case (982)," indicating that he recognized it
as being anomalous, quite unlike other Ohuma (Atriplex) caiiescens plants. As
shown in Figure 2, there is considerable variation in the fruiting-bract char-
acteristics of plants in the Reese Valley population, with none being truly
4-winged.
Since Watson, following Torrey 's identification, labelled each of the three
specimens {Watson 981), Ohione canescens, he apparently assumed the varia-
tion he saw in the populations in northern Nevada to be representative of
the variation displayed by Atriplex shrubs throughout western United States.
This is confirmed in his later description of A . n/tttallii (Watson 1 874) wherein
he lists Atriplex canescens Nuttall, not of James, Ohione canescens Moq., and
Atriplex Gordoni Hook, as synonyms and gives its distribution as "from the
Saskatchewan to Colorado and Northern Nevada."
This broad interpretation by Watson appears to have been the initiation
of a series of misconceptions concerning the geographic distribution of A.
nuttallii that we interpret to be confined to northern Nevada and north-
western Utah (Fig. 3). Standley (1916) listed the type locality of A. nuttallii
as "on the denuded hills of the Missouri River, about 15 miles below the
confluence of the White River, South Dakota." Since this is the location
given by Nuttall (1818) as the type locality (){ Atnplex canescens (^\xi'i\\) Nutt.,
Standley clearly confused the two. Furthermore, this locality for Atriplex
canescens {Calligonium canescens Pursh) invites correction because its origin
given by Pursh (1814) is "in the plains of the Missouri River, near the Big
Bend" which is several kilometers upstream from the confluence of the White
River.
Standley (1916) also mistakenly listed the distribution of A. nuttallii as
"alkaline plains and hillsides, Manitoba and Saskatchewan to northern Utah,
Colorado and western Nebnuska" without mentioning Reese Valley and Thousand
Springs Valley in northern Nevada.
Stutz and Sandhrson, Atriplex nutcallii and its near relatives
197
Fig. 2. Fruitin^'-bracts of six randomly selected plants oi Atriplex nuttallii from a popula-
tion in Reese Valley, ca 1 km west of Battle Mountam, Lander Co., Nevada. Each row across
represents one individual. Bar = 15 mm.
198
SlDA 18(1)
Tic;. 3. DismbLirion ol known populations of /\/r//;/(-.v «////■/////.
Hall and Clements (1923) likewise failed to include the type locality of
A. niittciliii in their description of the distribution of /\ . uiittallu as "Saskatchewan
and South Dakota to western Nebraska, New Mexico, Arizona, northwest-
ern California, eastern Washington, and Alberta."
Recently, Basset et al. (1983) cited the distribution of /i. uiittalliicV^ "from
the Peace River District of Alberta to Manitoba. In the United States specimens
have been found as far south as Colorado." Clearly this does not refer to the
A. nuttaliii described by Watson from Nevada.
The failure to identify the original collection site of /\. unttalli'i in all
references has caused considerable misunderstanding of its identity and has
resulted in the assignment of this binomial to most of the suffrutescent shrub
species o{Atriplex in western North America. Ulbricht (1934) listed A. nuttallu
S. Wats., A. bnxijolict Rydb., A. ciinecitci A. Nelson, A. eremicola Osterh., A.
falcata Standley, A. i^ordomi Hook., A. nmniexkaua Standley, A. ohlanceolata
Kydh., A. pabular is A. Nelson and A . tridentata Kuntze as synonyms oWbione
gardneri Moq. {Atripkx gardnen [Moq.] Standley). Hall and Clements (1 923)
treat A . buxifolia, A . cuneata, A . eremicola, A. falcata, A . gardneri, A . pabiilaris
and A. tridentata as subspecies of nuttaliii.
Standley (1916) listed eight species as close relatives of A. nuttaliii (Se-
ries Nuttallianae): A . buxifolia, A . cuneata, A .falcata, A . gardneri, A. neomexicana,
A. oblanceolata, A. pringlei Standley, and A. tridentata. As suggested by Hall
Stutz and Sandhrson, Atriplex nuttallii and its near relatives 199
and Clements (1923), A. pringki appears to be an insignificant variant of
A. acanthocarpa (Torr.) S. Wats and A. ohlanceolata is non-distinguishable
from A. gardneri. A. neomextcana is a common reoccurring hybrid between
A. cuneata and A. conferufolia (Torr. & Frem.) S. Wats. (Hanson 1962) and
A. bnxifolia is synonymous with A. gardneri. The remaining four species, A.
cuneata, A. falcata, A. gardneri, and A. tndentata, were treated by Hall and
Clements (1923) as subspecies of A. nuttallii. Since they and A. nuttallii are
clearly distinct phenotypically (Pope 1976, Tables 1, 2, Fig. 4) and geo-
graphically (Figs. 3, 5-8) we consider them best treated as separate species
as proposed by Standley (1916).
Because Watson listed A. gordoni (A. gardneri) as a synonym of A . nuttallii,
Hanson (1962) argued that the epithet nuttallii was superfluous and hence
illegitimate and that A . gardneri was the correct name for Watson's A. nuttallii.
This view was later accepted by Pope (1976) Stutz (1 978), Stutz et al. (1979),
and Welsh (1984). However, as noted by McNeil et al. (1983), the listing
of A. gordoni (A. gardneri) as a synonym of A. nuttallii was accompanied by
a question mark and is therefore not a legitimate synonym and, according
to the Code, Article 55.2, note 1 , (Greuter et al. 1994) the epithet nuttallii
is therefore not nomenclaturally superfluous.
As discussed by McNeil et al. ( 1 983) the other synonyms of A . nuttallii
listed by Watson: {Atriplex canescens Nuttall, and Ohione canescens Moq.), are
also not legitimate synonyms.
MATI- RIALS AND MF.THODS
To better define A. nuttallii and its near relatives, herbarium specimens
of the suffrutescent Atriplex species of western North America were exam-
ined in several herbaria (BRY, CAS, GH, MO, NY, PH, RM, RSA, US, UC)
and specimens were collected and studied from the original collection sites
of each of the species included in the study (except A. gardneri for which
the exact location is not known). In addition, morphological measurements,
saponin production, flavonoid content, and cytological studies were made
of plants in several populations of each species. The morphological mea-
surements included fruiting-bract characteristics, plant height and width,
and leaf length and width. Chromosome counts were obtained from plants
in several populations of each species (Figs. 5-8). In some cases the counts
were obtained from root tips squashed in aceto-carmine stain, but most were
from aceto-carmine squashes of pollen-mother-cells derived from anthers
fixed in 5% acetic acid and stored under refrigeration. Saponin content was
determined by hemolysis of red blood-cells as described by Sanderson et al.
(1987). Flavonoid content was determined by paper chromatography as described
by Sanderson and Stutz (1984).
Tahu; 1 . Plant and leaf characcensncs oi Atnl^lcx ni<t!uliu and its near relatives. Meastirements were obtained from 20 plants in each population. Diploid A. nnieata
IS not included because ot extensive between-population variation. N = number of populations. Data = mean with coefficient of variation in parentheses. Values
in eacli column with the same letter are not significantly different (p<.()5).
Plant
Leaf
'^^^"" f^' Heit,dit (cm) Width (cm)' Volume (dm ~)'' Len<_'rh(mm) Width (mm)
A.withillniGx) 9 32.6(0.41)'^ 47.4(0..39)'^ 101 .5(0.94)'^ 24. 1(0. r^)"^ 4.0(0.21)'* 6.0(0.22)'^
A.a/iieah!(AyO 5\ 12.8(0..t])'^ 5 1.2(0. .3 .3)'^ 41.3(0.88)'^'* 20.9(0.22)'^ 8.3(0.28)'^ 2.6(0.19)'^^
A.l],lcjta{2^) 12 12.9(0.44)" 24.6(0.,^ 1 )'* 9.4(0.^4)" 23.7(0,58)'^ 3.6(0.41)" 6.5(0.23)'^
A.i^arclnendy.) 6 9.8(0.52)" 48.9(0.11)'^" 25. 2(0. 76)'^" 19.9(0.09)'^ 3.8(0.21)" 5.4(0. 1 1 )'''f^
A.i^m-J,m;{Ax) 8 10.9(0.59)" 49.8(0.44)-^ 42.3(1.45)'^" 19.4(0. 19)'"' 5.3(0.28)" 3.8(0.18)"™
A.tnJentataiGx) 1^ 23.9(0,38)'^ 41.8(0.46)^" 63.8(1 .25)'-^" 28.7(0.53)'^ 4.1(0.33)" 6.79(0.28)'^
' Width of A. tridentcitd does not include root sprouts.
^ Volume = height X (width)-
c
>
00
Table 2. Fruiring-bracc characteristics oi Atripkx niittaU'ii and its near relatives. Measurements were made on 20 Fruits from several randomly selected plants in
each sampled population. Diploid A. ctineata is not included because of extensive between-population variation. N = number of populations. Data = mean with
coefficient ol variation in parentheses. Values in each column with the same letter are not significantly different (p<.05).
Taxon
A. niittciUii{Gx)
Fruitint,'-bract
N Length (mm) Width (mm)
1/w Volume (mm")''
Apex'^
No. of
Terminal Teeth
10
4.5(0.10)'"
.^.0(0.07)
BCtJ
1.6(0.14)'^" 39.0(0.15)'^
2.4(0.07)'^ 3.2(0.24)R
A.cuneata{Ax) 15 6.1(0.16)'^
A.falcata(2-K) 12 4.6(0. 19)^^
A.gardneri(,2x) 1 3.1(0.14)"=
A.gardnertiA-y:.) 10 4.2(0.21)''^
A. tridentataiGx) 13 4.6(0.12)^
5.0(0. 22)^^ 1.3(0.14)^ 165.3(0.59)'' 1.8(0.19)*^ 3.6(0.24)^^
2.6(0.14)1^ 1.8(0. 10)'^ 32.0(0.48)" 3. 0(0.00)''^ l.CKO.OO)^-
2.5(0.21)^^ 1.3(0.17)"^ 19.7(0.45)^* 1.7(0.19)"= 4.64(0.46)"
3.0(0.31)''^° 1.5(0.12)'3c 45.1(1.12)13 1.5(0.05)^' 3.8(0.12)"
3.9(0.16)"^: 1.2(0.15)^ 72.6(0.39)" 2.6(0.15)" 8.5(0.45)'^
'^ Volume = length X (width)'.
^ Prominence of central apical tooth was scored to 3 with central tooth absent = 0, small = 1, conspicuous = 2, prominent = 3.
■^ Number of lateral appendages was scored to 5 with no appendages = 0, numerous appendages = 5.
CO
H
C
H
N
>
a
>
•z.
D
m
Appendages'- "5
2.0(0.14)"
3.9(0. 17)'^
2.0(0.27)1"'
1.4(0.19)"
1.9(0.37)"
2.0(0.49)"
X
C
CI-
re
<
rn
O
202
Si DA 18(1)
Fic;. 4. I'riiiting bracts ui Atr/p/cx niittallii and its near relatives. A, A. HHttaUii. B, A. trulviitcita.
C, A. ciDiadtd. D, A. garciiien. E, A. falaita. Although there is considerable variation m the
frtiiting bracts of plants within and between populations, tliose shown are hiirly represen-
tative of each species. Collection sites h)r those shown are: A. unttcilUi, 1 km W ol Battle
Mountain, Lander Co.: Nevada; A. tridisiitiitci. Rush Valley, Tooele Co.: Utah; /\. mneata,
Navajo Mine lease site, ca 1 km SW of Farmington, San Juan Co.: New Mexico; A . y^ardneri,
ca 10 km S oFBridger, Uinta Co.: Wyoming; A. falcutct, ca 5 km N of Ontario, Malheur
Co.: Oregon. Bar = 15 mm.
Stlitz and Sandhrson, Atriplex nuttallii and its near relatives 203
RESULTS AND DISCUSSION
Probably because they are dioecious and wind-pollinated, most of the
shrubby species oi Atriplex in North America are unusually rich genetically
and can often be best defined by quantitative, in addition to available qualitative,
attributes. This is particularly true tor /\ . nuttallii and its relatives. As shown
in Figures 3, 5-8 and Tables 1-4, although A. nuttallii and each of its near
relatives can usually be distinguished from each other by some qualitative
differences, when combined with differences in quantitative attributes and
geographic distributions, they are quite distinct.
Atriplex nuttallii S. Watson, Proc. Amer. Acad. Arts 9:116.1874. Lhcto-
TYPi;: NEVADA: Reese Valley, Jul 1868, Watim 9H1 (GUI).
Phenotypically, A . nuttallii appears to be most closely related to A . tndmtata.
Although highly variable in A. nuttallii (Fig. 2), most fruiting bracts of
both taxa have multiple, terminal, marginal teeth. Both have oblong or
oblanceolate leaves and occupy similar habitats (primarily saline bottom-
lands). Both are hexaploids (2/?= 54) (a few tetraploid populations of A. tridentata
have also been found). A. nuttallii differs from A. tridentata in being taller
statured (30-60 cm vs 10-30 cm) (Table 1), woodier, and more phenotypi-
cally variable. Much of its variation appears to be the result of hybridiza-
tion and subsequent introgression from other species, including A. tridentata
with which it is often sympatric. Plants of A. tridentata do not produce sa-
ponins whereas some plants of A. nuttallii do, some do not (Table 3). A.
tridentata usually shows aggressive root-sprouting whereas A . nuttallii is distinctly
caespitose with numerous (60—120), slender (1—3 mm in diameter), woody
stems emerging from a single woody crown, 10—50 cm in diameter. Geo-
graphically, A. tridentata is common in northern Utah, southwestern Wyo-
ming, northwestern Colorado and northeastern Nevada (Fig. 8); A. nuttallii
appears to be restricted to the alkaline valleys in northern Nevada and north-
western Utah (Fig. 3).
Atriplex cuneata A. Nelson, Bot. Gaz. 34:357.1902. Type: UTAH. Emhry
Co.: Emery, alt, 7,000 ft, 1894, M.E.Jones 3443 (holotype: US!; isotypes: MO! NY!
RSA!).
Atriplex ohlaucmlatd Rydb., Bull, Torrey Bot. Club 31:403. 1904. Type: COLORADO:
Delta, 3 Sep 1897,././/. Onvm 4071 (holotype: US!; isotype: GH!).
Populations of A. cuneata are restricted primarily to eastern Utah, west-
ern Colorado and northwestern New Mexico (Fig. 5). In many places they
constitute the dominant vegetation on thousands of acres. The common name
of A. cuneata^ "Castle- Valley clover," connotes its value as a range plant, partly
because it is both palatable and nutritious for livestock and wildlife and
partly because, were it not there, there would apparently be nothing at all,
O
Table 3. Cytology, saponin production, tlavonoid production, and geographic distribution ot Atrip/ex niittallti and its near relatives. The number of plants exam-
ined is in parenthesis.
Taxon
In
Chrom. #
Saponins
s
I'lavonol
Is
6-MeO
3-MeO
A. nultallt!
54(11)
+ ,-(11)
-121)
A . ciineata
36(126)
- (69)
+
-(43)
A . ciineata
18(87)
-(72)
-
+ (62)
A . fakata
18 (115)
+ (74)
-
+ ,-(66)
A . ii^anbien
18(183)
+ ,-(32)
-
+ (31)
A . gardnen
36(250)
+ ,-(63)
+
- (40)
A . tridetitata
54 (257)
- (Ill)
+
-(105)
Geographic Distribution
Date of
Antliesis
valleys in n NV and nw UT Jun-Jul
e UT, w CO, nw NM Apr-May
e UT, w CO, nw NM Apr-May
s ID, n UT, sw WY, n NV, s OR, nw CA May-Jun
MT, WY, AB,SK May-Jun
MT, WY, AB, SK May-Jun
n UT, n NV, s WY. s ID, nw CO )ul-Aug
D
>
00
Stutz and Sanderson, Atriplex nuttallii and its near relatives 205
Table 4. Key to Atriplex nuttallii and its near relatives [Series Nuttallianae of Standley (1916) con-
sisting of species in which the plants are low-statured (less than 6 dm), suffrutescent, perennial, dio-
ecious, shrubs. Leaves are densely furfuraceous, alternate, entire, with Kranz-type anatomy. Fruiting
bracts are 2 — 6 mm long, 2 — 5 mm broad, usually longer than broad and usually appendaged.}
1. Leaves linear, length more than 5 times the width
2. Root-sprouting extensive — A. tridentata
2. Root-spfouting none or minimal
3. Stems 50 or more from a woody crown, each 30 — 80 cm long — A. nuttallii
3. Stems smgle ot few, each less than 30 cm long — A. falcata
1. Leaves ovate to oblong, length less than 5 times the width
2. Fruiting btacts globose, 3—5 mm long with numerous flattened appendages — A. cuneata
2. Fruiting bracts elliptical to ovoid, 1—3 mm long with few appendages — A. gardneri
of forage value. This is conspicuous in many places where populations of A.
cuneata terminate abruptly against clay slopes that are completely devoid of
vegetation. It is also evident in sites where populations of A. cuneata have
experienced large-scale "die-back" and remain empty until repopulated by
new A. cuneata plants.
Atriplex cuneata is mostly tetraploid but throughout its range there are
several morphologically distinct diploid populations (Fig. 5). The variation
present in disjunct populations of tetraploid A. cuneata is probably the result
of introgression from these diploids, or in some cases, tetraploid A. cuneata
may have originated polyphyletically from different diploid ancestors.
Atriplex cuneata plants are mostly caespitose and erect with no evidence
of root-sprouting or layering. Their fruiting-bracts are usually much larger
than the fruiting bracts of A. gardneri (Table 2) and are covered with nu-
merous lateral appendages (Table 2, Fig. 4). They are usually free of saponins,
with exceptions only in populations in which introgressive hybridization
from other species is suspected. Tetraploid A. cuneata plants test positively
for the presence of 6-methoxy flavonols and negatively for the presence of
3-methoxy flavonals. In contrast, diploid A. cuneata plants test negatively
for the presence of 6-methoxy flavonols and positively for the presence of
3-methoxy flavonols (Table 3).
Atriplex falcata (M.E. Jones) Standley, N. Amer. Fl. 21:68. 1916. Type:
IDAHO. Washington Co.: Weiser, alt. 2,000 ft, 7 Jul 1899, M.E. Jones s.n. {uow-
type: RSA!; isotype: UC!).
Populations oi A. falcata are sporadically common in southern Oregon,
northeastern California, northern Nevada, northern Utah, southwestern
Wyoming and southern Idaho (Fig. 6). It is mostly diploid (2;z=18), but
occasional tetraploid populations occur (Pope 1976).
Phenotypically, A. falcata is most easily recognized by its small stature
(ca. 13 cm tall X 25 cm broad), small, linear leaves (ca. 24 mm long x 4
mm wide), and small fruiting bracts (ca. 4.5 mm long X 2.5 mm wide),
206
SiDA 18(1)
otm0
•0
•
A. cuneata 2x
• A. cuneata 4x
-!i
COLORADO
NEW MEXICO
Fk;, 5. (icoyraphic distribution ot chromosome counts of plants of diploid (2w=18) and
tetraploid (2w = 36) Atripkx cuneata.
usually without appendages, and terminating in a distinct, acute, central
apex (Tables 1,2, Fig. 4).
The leaves of A.falcata produce abundant saponins (Table 3), a trait that
can be useful in field identification either by blowing bubbles in a water
emulsion of leaves or by tasting a leaf as it is chewed. When saponins are
present, bubbles form readily in a leaf-emulsion and the leaves have a dis-
tinctly bitter taste. Both of these tests are positive iov A.falcata leaves, negative
for leaves of A. trulcutata plants with which A. falcata plants are sometimes
confused. A. falcata plants are also readily distingtushed from A. trkkntata
plants by their rooting liabit: A. triclaitata plants are vigorous root-sprouters,
whereas A. Jalcata plants are caespitose with a single prominent taproot.
Also, A . falcata plants produce flowers and fruits early in the spring (May-
June) whereas A. tridentata plants flower and set fruit mostly in mid to late
summer (July— August) (Table 3).
Atriplex gardneri (Moq.) Standley, N. Amer. Fl. 2 1 -.GG. 1916. Typi - fSrare:-'}.
La Platte River [date?], Gonhn 250 (hoi.otypi;: GH!, fragments).
O/j/om- liantnen Moc]. in DC, Prod, l.rl 11. 1819.
Populations of A. gardneri (2x, 4x) dominate thousands of acres of clay
slopes and swales throughout much of southern and eastern Wyoming, central
and eastern Montana, western North and South Dakota, southern Alberta,
Stutz and SAiNDDRSON, Atriplex nuttallii and its near relatives
207
^?'
•
•
• •
•
•
% Ob
• A. falcata 2x
1 i A. falcata 4x
•
• •
•
• ••
•
Fic;. 6. Geographic distribution of chromosome counts of plants of diploid (2«=18) and
tetraploid (2w = 36) Atriplex falcata.
and southern Saskatchewan (Fig. 7). Most populations are tetraploid but
diploid ones are common. In southern Alberta, southern Saskatchewan and
north-central Montana, diploid plants can usually be distinguished from
tetraploid plants by their smaller stature, smaller, thinner leaves, finer-tex-
tured, less woody stems and smaller fruiting bracts. However, throughout
most of the saline deserts of Wyoming and southern Montana, diploid and
tetraploid plants are phenotypically very similar although both show con-
siderable phenotypic variation both within and between populations. Chro-
mosome counts of plants in populations bordering Interstate Highway 80
in southern Wyoming, showed a preponderance of tetraploids west of the
Continental Divide and a preponderance of diploids to the east (Fig. 9)-
However, no conspicuous ecological differences were evident between the
sites occupied by tetraploids and diploids, nor was it possible to consis-
tently distinguish diploids from tetraploids, phenotypically. This was true
for plants in natural populations as well as garden-grown specimens. They
are, however, clearly distinguishable by flavonoid differences: tetraploids
H)H
SiDA 18(1)
o
o
9
(D O \
• O
o i
\ o \
\ C> i
I QO O gD; O
' O'"^ o
*••
••
o
1 \
o
'\ o
o
oo
o
o
A. gard neri 2x
A. gardneri 4x
^^ 5
C3
&
Fic. 7. Geographic distribution oi chromosome cotints of plants oi' diploid {2>i=- 18) and
tetraploid (2«= •>6) Atr/p/ex giirdiieri.
always produce 6-mechoxy flavonols, diploids do not; diploids produce 3-
methoxy flavonols, cetraploids do not (Table 3). Although both diploids
and tetraploids sporadically produce saponins (Table 3), diploids are most
olten heavy producers, tetraploids usually produce little or none.
Atriplex gardneri plants are short-sratured, mostly 4-10 cm in height,
(Table 1), but are often more than 80 cm in diameter. The plants are cae-
spitose, arising from a single deep taproot but often show considerable lay-
ering. Their fruiting-bracts are usually smaller (ca. 3^ mm) and have fewer
lateral appendages than those of most near relatives (Table 2, Fig. 4).
Stutz and Sanderson, Atriplex nuttallii and its near relatives
209
Fig. 8. Geographic distribution of chromosome counts of plants of tetraploid (2« = 36) and
hexaploid (2«=54) Atriplex tridentata.
Atriplex tridentata Kuntze, Revis. Gen. PI. 2:546. 1891- T^t^e: UTAH: Corinne,
1874, D.E.O. Kiintze 3084 (holotype: NY!).
Two chromosome races of Atriplex tridentata have been found: tetraploid
(2« = 36) and hexaploid (2/2=54). Hexaploids are, by far, the most common,
occurring throughout much of western United States and in most of the
valley bottoms in Utah and Nevada that were recently occupied by Pleis-
tocene lakes (Fig. 8). Only three major tetraploid populations are known:
one in Juab County, Utah, one in Eureka County, Nevada, and one in Lin-
coln County, southwestern Wyoming (Fig. 8). Although tetraploid and hexaploid
A. tridentata are to some extent, morphologically distinct, the differences
do not appear to be sufficient to warrant designation as separate taxa.
Atriplex tridentata differs from other Atriplex species in several features,
including linear to oblong, furfuraceous leaves, fruiting bracts with con-
spicuous terminal, marginal teeth (Fig. 4), late-flowering habit (July-Au-
gust), and vigorous root-sprouting. A. tridentata plants are mostly herba-
ceous above ground with a few slender stems growing erect from woody
underground crowns and roots.
The extent of root-sprouting in A. tridentata plants is often made con-
spicuous by its dioecious flowering habit. By noting its flowers, the extent
of a single male or single female plant may be easily determined. In some
cases individual plants have been found to occupy more than 200 m^. Sometimes
210
SiDA I.S(l)
10
15
20
25
30
VUL
35
40
45
I—
45
50
55
60
Lilllc
Anicrica
75
90
JL
135
95
100
105
T
Rock
Springs
110 115
JL
jlj
140
145
150
155
_,_EL
„J«1_
la E]
EiirsicsitTi rj iiiprsi M [I , _ij IL
180
185
190
195
200
r
205
!
Wamsutler
ja tpjEiaj^iMjp
210
215
220
225
Rawlini;
j~in r:i m
225
230
235
240
245
250
255
260
265
270
Fic;. 9. Distribution ol diploid and tetraploid populations o{ Atripkx gardneri alongside
Interstate highway HO in southern Wyoming from the Utah border (mile 0) to near Elk
Ivlountain Wyoming. (Chromosome counts were made on plants at each mile marker, when
avaikible. Stippled bars = diploids, black bars = tetraploids.
a single plant forms a complete carpet in which there are no other plants; at
other times the root-sprouting carries a plant around and between plants of
other species.
Possibly because of its low saponin content (lable 3), its root-sprouting
habit and predominantly herbaceous tissues, A. trtdentata is often regarded
favorably by ranchers as forage for livestock (personal reports).
Stutz and Sanderson, Atriplex nuttallii and its near relatives 211
ACKNOWLEDGMENTS
The authors thank Dr. Richard W. Spellenberg and Dr. Ronald L. Hartman
for numerous helpful suggestions, Broken Hill Proprietory Minerals and
Brigham Young University for financial assistance and the curators of the
following herbaria for loans of specimens and access to their collections:
BRY, CAS, GH, MO, NY, PH, RM, RSA, UC and US.
REFERENCES
Bassftt, I.J., C.W. Crompton, J. Mcneiu., and P.M. Taschereau. 198.1 The genus Atriplex
(Chenopodiaceae) in Canada. Monograph No. 31. Communications Branch, Agriculture
Canada, Ottawa.
Greuter, W., H.M. Burdet, W.G. Chaloner, V. Demonlin, R. Grolle, D.L. Hawksworth,
D.H. NiCHOL.s()N, P.C. Silva, F.A. Stafleu, E.G. Vo.ss, and J. McNeil. 1988. Interna-
tional code of botanical nomenclature. Adopted by the fourteenth International Botani-
cal (Congress, Berlin, July-August 1987. Regnum Veg. 118.
, F.R. Barrie, H.M. Burdet, W.G. Chaloner, V. Demoi'lin, D.L. Hawksworth,
P.M. JORGENSEN, D.H. NicoLSEN, P.C. SiLVA, P. Trehane, and J. McNeill. 1994. Inter-
national code of botanical nomenclatute (Tokyo Code). Adopted by the fifteenth Inter-
national Botanical Congress, Yokohama, August-September 199.3. Regnum Veg. 131.
Hall, H.M. and RE. Clements. 1923. The phylogcnetic method in taxonomy. Carnegie
Institution of Washington Publication No. 326.
Hanson, C.A. 1 962. Perennial Atriplex of Utah and the northern deserts. Thesis. Brigham
Young University, Provo, Utah.
McNeil, J., I.J. Ba.ssett, C.W. Crompton, and P.M. . 1983. Taxonomic and nomenclatural
notes on Atriplex L. (Chenopodiaceae). Taxon 32: 549-556.
Nuttall, T 1818. The genera of North American plants. Vol. 1. Philadelphia. D. Heartt.
Pope, C.L. 1976. A phylogenetic study of the suffrutescent shrubs in the genus Atriplex.
Ph.D. Dissertation, Brigham Young University, Provo, UT.
PuRSii, RT. 1814. Florae Americae Septentnonalis. Vol. II. London. White, Cocchrane and
Company.
Sanderson, S.C, and H.C. Stutz. 1984. Flavonoid aglycones of diploid and polyploid Atriplex
amfertifolia. In: A.R. Tiedemann, E.D. McArthur, H.C. Stutz, R. Stevens, and K.L.Johnson,
K.L. (compilers). Proc. symposium on the biology oi Atriplex and related chenopods.
Provo, UT.
, R.L. Pendleton, E.D. Mcarthur, and K.T. Harper. 1987. Saponin effect
on small mammal forage preference in a planting o{ Atriplex canescens. In: F.J. Provenza,
J.T. Flinders and E.D. McArthur (compilers), Proc. Symposium on Plant-Herbivore
Interactions. USDA Forest Service General Technical Report INT-222, 74-77. Ogden,
UT.
Standley, P.C. 1916. Chenopodiales. North American Flora 21(1). New York. New York
Botanical Garden.
Stutz, H.C. 1978. Explosive evolution of perennial Atriplex in western America. In: K.T.
Harper and J.I. Reveal, organizers. Intermountain biogeography: A symposium. Great
Basin Naturalist Mem.. 2:l6l-l68.
, C.L. Pope, and S.C. Sanderson. 1979. Evolutionary sttidies ok Atriplex: Adaptive
products from the natural hybrid SN A.tridentata _Al^ A. canesceiis. Amcr.J. Bot. 66:1181-
1193.
212 SioA 18(1)
Ul.BRlCH, E. 1934. Chenopodiaceae. In: A. Engler and K. PrantI, etls. Die Naturlichen
PflanzenEimilien. Ed. 2. l6C:379-5Hi. Engelmann, Leipzig.
Wa'ison, S. 1871. Botany. In: C. King. Report of geological exploration of the fortieth
parallel. Vokime V. U.S. Government Printing Pre.ss, Washington, D.C.
1 874. A revision of the Nortli American Chenopodiaceae. Proc. Amer. Acad.
Arts. 9:82-126.
Whi.sh, S.L. 1984. Utah flora: Chenopodiaceae. Great Basin Naturahst 44:183-209.
FLAVONOIDS IN STROPHOSTYLES SPECIES
AND THE RELATED GENUS DOLICHOPSIS
(PHASEOLINAE, FABACEAE):
DISTRIBUTION AND PHYLOGENETIC
SIGNIFICANCE
JUAN PABLO PELOTTO and MARIA A. DEL PERO MARTINEZ
Centro de Estudios Farmacoioguvs y Botdnkos
Consejo Nacional de Investigaciones Ckntifkas y Tecnkas
Serrano 663, Buenos Aires, 14 14. ARGENTINA
ABSTRACT
Strophostyles Elliot: is the only genus within the Phaseolinae with a center of distribu-
tion in the United States. It comprises three species, namely S. hdvula (L.) Elliott, 5. umbellata
(Willd.) Britton, and 5, leiosperma (Torrey & A. Gray) Piper, and it is considered as allied to
Dolichopsis Hassler, a monotypic genus endemic to South America. This study analyses the
leaf flavonoid content from the three Ssrophostyles species and Dolichopus [laraguariensis Hassler
with the aim of examining the phylogenetic relationships among taxa. We isolated 38 gly-
cosides based on kaempferol, quercetin and isorhamnetin. All Strophostyles species were characterized
by the presence of isorhamnetin glycosides. However, S. leiosperma showed a distinctive
profile while S. hdvula and S. umbellata clustered together. In contrast, D, paraguariensis
lacked isorhamnetin-based compounds. A cladistic analysis of flavonoid plus morphologi-
cal data supported Strophostyles monophyly and showed 5". leiosperma as the sister taxon of
the clade S. helvula-S. umbellata.
RESUMEN
Strophostyles Elliott es el linico genero de las Phaseolinae con un centro de distnbucion
en los Estados Unidos. Comprende tres especies: S. helvula (L.) Elliott, S. umbellata (Willd.)
Britton y S. leiosperma (Torrey & A. Gray) Piper, y se considera afin a Dolichopsis Hassler, un
genero monotfpico endemico de Sudamerica. Este estudio analiza el contenido de Oavonoides
foliates en las tres especies de Strophostyles y en Dolichopsis paraguariensis Hassler con el objeto
de examinar las relaciones filogeneticas entre estos taxa. Se aislaron 38 glicosidos de kaempferol,
quercetina e isoramnetina. Todas las especies de Strophostyles se caractenzaron por la presencia
de glicosidos de isoramnetina. Sin embargo, S. leiosperma mostro un perfil distintivo mientras
que S. helvula and S. umbellata se agruparon juntas. Por el contrario, D. paraguariensis no
sintetizo compuestos basados en la isoramnetina. Un analisis cladistico conjunto de los datos
de flavonoides y caracteres morfologicos apoyo la monofilia del genero y mostro a^'. leiosperma
como el taxon hermano del clado S. helvula-S. umbellata.
INTRODUCTION
Strophostyles EUiorr is the only genus within the Phaseolinae with a cen-
ter of distribution in the United States. Its current taxonomic treatment
follows the original generic concept (Elliott 1822), but it was previously
SiDA 18(1): 213-222. 1998
214 SiOA 1<S(1)
associated with unrelated species and reduced to a section o^Phaseolns (de
CandoUe 1825; Bentham 1837, 1865) until it was restored as a separate
genus including three species (Britton & Brown 1897; Piper 1926). A set
of characters precludes merging Strophostyles with Phaseoh/s, namely erect
style (not coiled), lack of hooked hairs, pedicels shorter than the calyx, and
nodes of the inflorescence somewhat swollen (Marechal et al. 1 978). More-
over, Strophostyles plants can be recognized by their nearly asymmetric flowers
arranged in subumbellate inflorescences, bracts and bracteoles persisting
through seed maturation, cylindrical seeds (often pubescent), and linear pods.
Strophostyles has also been considered as allied to Dolichopsis Hassler (Marechal
et al. 1978; Lackey 1983). Nevertheless, both genera are easily distinguishable
by many characters and their quite distinct geographical disrributions. The
monotypic genus Dolichopsis^ resembles Strophostyles mainly in floral mor-
phology (purplish corolla, keel with a not curved to somewhat curved beak,
style slightly thickened distally, and stigma terminal oblique) and general
appearance, but it is unlike the latter in having symmetric flowers clustered
in elongate pseudoracemes and the unique fruit traits such as oblong, very
flat pods with oblong seeds implanted through a very long funicle and with the
hilum perpendictilar to the placenta. The geographic range oi Dolichopsis is Paraguay
and Argentina, in South America, whWt Strophostyles occurs throughout eastern
USA, eastern Canada up to south of Quebec and extreme northeastern Mexico.
Phytochemical data on Strophostyles species are scanty and include the absence
of both leuco-anthocyanins (Baudet 1978) and canavanine (Lackey 1977),
and a recent report of flavonoids (Williams et al. 1995). In this study, we
expanded on the survey of foliar flavonoids by considering a larger number
of samples of the three Strophostyles species, i.e. S. helviila, S. mnbellata and S.
leiospermci , and we added Dolichopsis paraguariensis for comparison.
MATERIALS AND MliTHODS
We analyzed the constitutive flavonoids present in the leaves of herbarium
specimens belonging to the three Strophostyles species and Dolichopsis paraguariensis .
Samples (100-200 mg) were powdered and extracted under reflux with 80%
methanol (x 3). Concentrated methanoiic extracts were two-dimensionally
chromatographed on paper (BAW/ 1 5% acetic acid). Compounds were identified
by standard methods (Mabry et al. 1970; Markham 1982). These incltided
complete and controlled (3 min.) acid hydrolysis, enzymatic hydrolysis (B-
glucosidase), co-chromatography with authentic markers and UV-Vis spec-
^Doltchupm was a genus with two species, D. paraguarienm and D, monticola (Lackey 1983,
Lewis 1991), but recently Delgado Salinas & Lewis (1997) created the new genus Oryx'n
where they placed D. monttadci. Tiierefore, D. paniy^/uinensis has became the tinique repre-
sentative of the genus.
Pelotto and Martinez, Flavonoids in Scrophoscyles and Dolichopsis species 215
troscopy. Glucosides were separated from their galactosidic analogues by
TLC in the appropriate system according to Budzianowski (1991).
Plant material. — Specimens were provided by the Institute de Botanica
Darwinion Herbarium (SI), San Isidro, and the Centro de Estudios Farmacologicos
y Botanicos Herbarium (BACP), Buenos Aires.
Strophostyles helvula (L.) Elliott
U.S.A. Arkansas. Jefferson Co.: Arkansas river bottoms, 220 ft, 17 Sep 1937, Demaree
16245 (SI). Illinois. Mc Donough Co.: Argykle Lake, near Colchester, 2 Aug \95H, Jones
22335 (SI). Iowa. Dickinson Co.: N shore ofSpirit Lake, sand (older beach), 5 Aug 1913,
Si>/ek 14... (number illegible) (SI). Mississippi. Harrison Co.: near the coast on sand, 6
Jan \9'5UDewaree30675 (SI); Ship Island, P.O. Biloxi, in stabilized sand, moist, long trailing,
Demaree 31059 (SI). Virginia. Prince George Co.: rich alluvial thicket back of sand-beach
of James River, Jordan Point, SE Virginia, 16 Sep 1938, Feriialil & Long 9353 (SI). Un-
hwuH loadity.movd of the Western Reserve] 15 Atig 1897, G.B. Asduroft. Berea 0. s.ii. (SI).
Strophostyles leiosperma (Torrey & A. Gray) Piper
U.S.A. Oklahoma: 5 mi NW of Breckenridge, 25 Jul 1941, Gephardt 747 (SI). Woods
Co.: in waste place, hard soil, near Alva, 24 Sep \9U, Stevens 2824 (SI). Texas. Smith Co.:
Amigo, neglected sandy field, 10-17 Aug 194'5, Moore. Jr. 995 (SI). Morris Co.: Aug 1891,
Carleton 420 (SI).
Strophostyles umbellata (Willd.) Bntton
U.S.A. Virginia. Greensville Co.: dry pine and oak woods, about 1 mi N of Skipper's,
l4-\5 ]n[ 1938, Fernalct & Long 8737 iSl).
Dolichopsis paraguariensis Hassler
ARGENTINA. Entre Rios: Depto. La Paz, R 1 26, desvi'o a Ombties, borde camino, 3 1
Jan 1 98 1 , fl. azul-violaceo, Troncoso de Bi/rkart & Baagalupo 3096 (SI). PARAGUAY. Depto.
Pte. Hayes: EstanciaLomaPyta, 23° 40'S, 59° 35 'W, 2 Apr 1974, enredadera casi rastrera,
fl violaceas, crece en pastizal, n.v. 'kekleichetas,' Arenas 544 (BACP). Depto. Boqueron:
Mision Santa Rosa, 21° 45'S, 61 ° 35 'W, Feb 1981, enredadera, fl. violaceas, crece en pajonal,
n.v, 'ceihlowey'. Arenas 1726 (BACP).
Data analysis. — A cluster analysis was performed on flavonoid data of
15 herbarium specimens. Similarity matrix was measured using Jaccard's
coefficient and a dendrogram was constructed applying the unweigthed pair-
group method of arithmetic averages (UPGMA). All calculations were done
using NT-SYS program (Rohlf 1993).
A cladistic analysis of the flavonoid data plus a set of morphological characters
was carried out according to the maximum parsimony principle. Outgroup
criterion was used for character polarization. Vigna adenantha was chosen as
the external group which flavonoid data were obtained following the methods
above mentioned (Pelotto, unpublished manuscript). For flavonoids, char-
acter states that occurred in the outgroup were scored as and those in the
ingroup (D. paraguariensis and the three Strophostyles species) were scored as
1 (see Appendix, Tables A and B). Morphological data were gathered from
the literature and included some multistate characters that were treated as
non-additive (see Appendix, Tables A and C). Cladograms were calculated
-^^ SiDA 18(1)
using the implicit enunieration routine (i.e.*) of the program Hennig86
(Farris 1988) with all characters equally weighted.
RESULTS ANO DISCUSSION
Chromatographic properties of the identified flavonoid glycosides and
Its distribution in Strophostyles species and D. paragiiarienm are shown in
Tables I and 2, respectively. All detected compounds were O-glycosides of
flavonols with sugars attached at positions 3 and 7 of the aglicone skeleton.
This pattern of glycosilation is very common among the Phaseolinae (Zallocchi
& Pomilio 1994; Williams et al. 1995, Pelotto unpublished manuscript).
All three Strophostyles species produced glycosides based on the methy-
lated flavonol isorhamnetin plus kaempferol and quercetin. Notwithstanding
S. helvnla and S. nmhellata showed very similar chromatographic patterns,
while the flavonoid profile of 5. lewspmna was quite distinctive. No rhamnosides
were detected in S. kmpmna and it only shared the presence of monoglycosides
with the other two species. Based on a three-sample analysis within a fla-
vonoid survey of the Phaseolinae, Williams et. al (1995) have also reported
the occurrence of isorhamnetin glycosides in Strophostyles species and no-
ticed the same interspecific differences. In contrast with our results, Will-
iams and co-workers isolated fewer compounds and did not detect kaempferol
glycosides from leaves, although they did from stems and/or flowers.
In turn, Dolichopsisparaguariensis samples were characterized by the presence
of kaempferol and quercetin glycosides, lacking isorhamnetin. Noticeably,
Paraguayan samples contained only kaempferol glycosides while Argentinean
one had kaempferol plus quercetin glycosides. However, in a previous work
(Zallocchi et al. 1995) both kaempferol and quercetin glycosides were re-
ported from one sample of D. pcircigiiarmisis from Paraguay, but of the eight
flavonol glycosides the authors identified only rutin and kaempferol-3-O-
rutinoside were also present in our .samples. These differences may be due
to the fact that Zallocchi and co-workers analyzed a whole plant extract
and therefore their results are difficult to compare with ours.
After the cluster analysis S. helvnla and S. umhellata are closer to D. paraguariensis
than to S. lewsperma (Fig. 1). This is because S. helvula and S. umhellata have
more glycosides (based on kaempferol and quercetin) in common with D.
paraguariensis than with S. leiosperma, even though D. paraguariensis does not
produce isorhamnetin glycosides.
Cladistic analysis resulted in two most parsimonious trees (length, L -
35, consistency index, CI =91, retention index, RI =70. Fig. 2). Both cla-
dograms support Strophostyles monophyly but differ in the depicted rela-
tionships among Strophostyles species. One tree (Fig. 2. A) shows S. helvula
and S. leiosperma as being sibling species, but this hypothesis needs the par-
allel gain of the characters 19, 22 and 31 on the S. umhellata and S. helvula
branches. The other tree (Fig. 2.B) supports the clade S. umhellata-S. helvula
Pelotto and MARTfNEZ, Fkvonoicls in Strophostyles and Dolichopsis species 217
Table 1. Chromatographic characteristics of the identified compounds.
SPOT IDENTITY
1 K-3-O-glLicoside + K-3-O-gahictoside
2 K-7-O-glucoside + K-7-O-galactoside
3 K-3-O-rutinoside + K-3-O-robinobioside
4 K-3-O-diglticoside + K-3-O-digalactosidc
5 K-3,7-0-ch^ducoside
6 K-3-0-rLitinoside-7-0-glucoside +
K-3-0-robinobioside-7-0-glucoside
7 K-3,7-0-tnglucoside
8 K-3,7-0-triglycoside (glu + rha + gal):|:
9 Q-3-O-glucoside + Q-3-O-galactoside
10 Q-7-O-glucoside + Q-7-O-galactoside
1 1 Q-3-O-rutinoside + Q-3-0-r()binobioside
12 Q-3-O-digliicoside + Q-3-O-digalactoside
13 Q-3,7-0-diglucoside
14 Q-3-0-rutinoside-7-0-glucoside +
Q-3-0-robinobioside-7-0-giLicoside
15 Q-3,7-0-triglucoside
16 Q-3,7-0-triglycoside (gki + rha + gal)
17 IR-3-O-glucoside + IR-3-O-gaiactoside
18 IR-7-O-glucoside + IR-7-O-gahictoside
19 IR-3-O-riitinoside + IR-3-O-robinobioside
20 IR-3-O-diglucoside + IR-3-O-digalactoside
21 IR-3,7-0-digIucoside
22 lR-3-0-rutinoside-7-0-glucoside +
IR-3-0-robmobioside-7-0-glucoside
23 IR-3,7-0-triglucoside
'DP: deep purple, Y: yellow, YO; yellow-orange
^glu: glucose, rha: rhamnose, gal: galactose
and requires three reversions (characters 1, 9 and 17). This scenario is pref-
erable to that portrayed on Fig. 2. A since a mutation lost is a more prob-
able event than the homoplastic acquisition of isorhamnetm glycosides. Even
more, if we suppose reversal of characters 1, 9 and 17 on the S. mnbellata
branch as being a consequence of samplmg error (undersampling), the cla-
dogram becomes shorter with only 32 steps (CI= 100, RI= 100) and the
unique solution of a similar analysis. Thus, we consider the tree depicted
on Figure 2.B a more plausible ingroup phylogeny.
Flavonoid evolution shows methylation of the flavonol skeleton as an advanced
character shared by all Strophostyles species and the absence of rhamnosides
in S. leiosperma as an (aut)apomorphic loss.
Morphological traits are congruent with flavonoid data. Subumbellate
inflorescence (character 24), persistent bracts and bracteoles (character 26),
linear, terete pods (character 27) and seed pubescence (character 28) ^\vp-
^otxStrophostyles monophyly, and are correlated with isorhamnetin monoglycoside
production (characters 17 and 18). Strophostyles helvula strongly resembles
COLOUR"
Rf(x 100)
UV + NH3
BAW
15% aa
DP Y
69
45
Y Y
45
15
DP Y
52
54
DP Y
33.5
61.5
DP Y
29
70
DP Y
24
75
DP Y
3
82
DP Y
4
80
DP Y
56
39
Y YO
27
9
DP Y
40
51
DP Y
26
52
DP Y
23
63
DP Y
15
70
DP Y
3
78
DP Yo
5
79
DP Y
56
42
Y Y
38
10
DP Y
40
54
DP Y
29
58
DP Y
25
68
DP Y
19
74
DP Y
3
80
00
Table 2. Glycoside disrriburion in tlie analyzed samples. Compounds are numbered according to Table 1 . K; kaempterol glycosides; Q: cjuercetin glycosides; IR:
isorhamnetin glycosides; -r: present; -: absent.
K Q IR
10 11 \2 \i 14 15 16 r 18 19 20 21 22 2t
S. helvitla
Jones 22335 - + + -- + --- + + -- + --- + + -- + -
Simek 14... ++ + -- + -- + + + - - + - + + + + - - + -
Aschroft & Berea s.n. ++ + -- + -- + + + -- + - + + + + -- + -
Demaree 3067 5 ++ + -- + -- + + + - - + - + + + + - - +
Demaree 1624 5 -+--- + _-- + --- + ^ + - + --- + -
Demaree 31059 ++ + -- + -- + + + -- + - + + + + -- + -
Fernald & Long 935 3 - + + ._ + --- + + -- + - + - + + -- + -
S. umbellata
Fernald & Long 87 37 - + + -- + --- + + -- + - + - + + -- + -
S. leioiperma
Moore Jr. 995 ++-- + --- + + -- + --- + + --^--
Carleton 420 +__ + + - + - + -- + + - + --i--- + ^- +
Gephardt 747 +-- + + - + - + -- + + - + - + -- + + - +
Stevens 2824 +-- + + - + - + -- + + - + - + -- + + - +
D. paraguariemii
Arenas 1726 _ + + __ + __-_____---------
Arenas 544 __^ + _. + -......-._. ------
Troncoso & Bacigalupo 3096 - + + -- + - + - + + -- + - + ------- (y^
— — >
00
Pelotto and Martinez, Flavonoids in Strophostyles and Dolichopsis species
JACCARD'S COEFFICIENT
219
0.00
0.25
0.50
0.75
1.00
— HEL-22335
_jHEL-9353
'UMB-8737
HEL-14...
_ HEL-sn
HEL-30675
HEL-31059
— HEL- 16245
— DOL-3096
_|DOL-1726
'DOL-544
— LEI-995
I LEl-747
-HLEI-2824
IlEI-42()
Fk;. 1. Dendrogram of the Strophostylts and Dalichopsii specimens constructed from a simi-
larity matrix (Jaccard's coefficient) using tiie UPGMA method. Cophenetic correlation co-
efficient, r = 0.969
S. umbellata, except for its more fobed leaflets and larger pods and seeds.
Strophostyles leiosperma is rather different from the other two species because
of the smaller flowers (character 25) arranged in more pauciflorous inflo-
rescences and its seeds glabrous and shining at maturity (character 29).
Similarly, both S. helvnla and S. umbellata are more widespread and northerly
distributed, with the first species reaching Canada, while S. leiosperma has a
more limited distribution ranging from south of the United States to the
extreme northeastern of Mexico (Britton & Brown 1897, Marechal et al.
1978). Species divergence at chemical and morphological level also correlates
with their ecological features; S. helvula and S. umbellata mostly grow in
more mesic sites, while S. leiosperma is adapted to live into more xeric habitats.
Regarding the evolution of the growth form, overlapping this character
onto our preferred topology suggests that annual growth would have evolved
independently in both S. leiosperma and .S". helvula, whereas perennation would
be the plesiomorphic state shared by D. paraguariensis and S. umbellata.
In summary, universal occurrence of isorhamnetin-based compounds in
Strophostyles species is a good chemical character in defining generic monophyly
while individual glycosides are useful characters to trace species evolution. With-
in the Phaseolinae isorhamnetin glycosides have sporadically been recorded
in four Phaseolus species (Pelotto, unpublished manuscript) and some Vigna
and Macroptilium species (Zallocchi & Pomilio 1994; Williams et al. 1995).
This fact suggests that flavonol methylation has appeared several times in
the tribe, making it a valuable phylogenetic marker at the infrageneric level.
Beyond this contribution, flavonoid data from the related genera Oxyrhynchus
and Oryxis are wanting for a complete view of this little group of American
species around Vigna.
220
A-
SlDA 18(1)
t
/ 3-7
-11-15
20,21,23
25(2),29
^ 18,24,26,27(2},28
^ 2,10,25(1)
B-
7,]8.24.26,27(2),28
13
Fic. 2. The two most parsimonious rrees (L ^ 35) ^'eneratecl using the data matrix (see
Appendix, Table A) and Vigthi iiJenuntha as outgroup. Characters are majDped on the trees
as Follows: solid bar - non-homoplasiotis apomorphy, clear bar = homoplasious apomorphy,
and cross = reversal. Numbers on the right of the character symbols stand for character
numbers (and character state).
Pelottc) and Martinez, Flavonoids in Strophostyles and Dolichopsis species
221
ACKNOWLEDGMENTS
We thank the curators of the Institute de Botanica Darwinion Herbarium
(SI) and the Centro de Estudios Farmacologicos y Botanicos Herbarium (B ACP)
for providing the plant material, and CONICET for financial support. We
are also grateful to two anonymous reviewers for suggestions that improved
the earlier manuscript.
APPENDIXES
Table A. Data matrix for the cladistic analysis including botii flavonoid (characters 1-15, 17-23,
codified according to Table B) and morphological (characters 24-31, codified according to Table C)
data sets and using S/igrui ddencintha as outgroup.
Taxon
1
6
character number
8 9 1(1 11 12 13 li 15 n 18 19 20 21 22 2i 24 25 26 r 28 29 30 31
y'lgna admanlki (1 >
D. para^^uarmiili 1 1 1 1) 1 1 :' 1
S. helviila 110 1 10 1 110 10 1 I 12 10 1
5. iimheUata 10 10 110 10 1112 10 1
.V. lampnma 1 1 1 1 I 1 I 1 I 1 1 1 1 1 1 1 1 I I 1 2 1 2 I 1
Table B. Flavonoids. Character numbers are as in Table 1. Character 16 was not included because no
hypothesis about homology can be made on a partially identified compound.
Character states
1. 0= absent, 1= present
2. 0= absent, 1== present
3. 0= present, 1= absent
4. 0= absent, 1 = present
5. 0= absent, 1= present
6. 0= present, 1= absent
7. 0= absent, 1= present
8. 0= absent, 1= present
9. 0= absent, 1= present
10. 0= absent, 1= present
11. 0= present, 1= absent
12. 0= absent, 1= present
13. 0= absent, 1= present
14. 0= present, 1= absent
15. 0= absent, 1= presenr
17. 0= absent, 1= present
18. 0= absent, 1= present
19- 0= absent, 1= present
20. 0= absent, 1= present
21. 0= absent, 1= present
22. 0= absent, 1 = present
23. 0= absent, 1 = present
Table C. Morphological characters, states and polarities.
character
24. inflorescence
25. flower size
26. bract and bracteole
27. pods
28. seed coat
29. seed pubescence
30. hilum
31. leaflets
0= pseudoracemose; 1= subumbellate
0= great (> 20 mm); 1= medium (7-15 mm); 2= small (< 7 mm)
0= persisting no longer anthesis; 1= persisting through seed maturation
0= linear, compressed; 1= oblong, very flat, with false cellulosic septa; 2 =
linear, cylindrical
0= smooth; 1= pubescent
?= inapplicable; 0= persisting in mature seeds; 1 = absent from mature
seeds
0= parallel to the placenta; 1= perpendicular to the placenta
0= entire; 1 = lobed to somewhat lobed at base
222 Sum 18(1)
RVA-VMV.NCV.S
Baudet, J.C, 1 978. Prodrome d'une classiiicarion gcnerique dcs Papilionaceae — Phaseoleae.
Bull. Jard. Bot, Belg. 48:183-220.
Bp.n riiAM, G. 1 837. Commentationes de legLiminosanim generibiis. J.P Sollinger, Vienna.
Bentliam, G. 1865. Leguminosae. In: Bencham, G& Hooker, J. D. Genera Plantarum 1:434-
600.
Brittok, N.L. and A. Brown. 1897. An illustrated flora of the northern United States,
Canada and the British Possessions. Vol. II. Charles Scribner's Sons. New York.
Bi:iv,iAN()\X'SKi, J. 1991 . Separation of flavonoid glucosides from their galactosidic analogues
by thm-layer chromatography. J. Chromatography 540:469-474.
DE CandollI', a. p. 1825. Prodomus systematics naturalis regni vegetabilis Vol. 2. Paris,
Strasbourg, London.
Del(;aix) Saunas, A. and G.P Lewis. 1 997. Oryxis, a new genus in tribe Phaseoleae (Leguminosae:
Papilionoideae) from Brazil. Kew Bull. 52:221-225.
Ellio-it, S. 1822. A sketch of the botany of South Carolma and Georgia. 2:229.
Farris, J.S. I 988. Hennig86 version 1.5. Program and software documentation. Published
by the author, Port Jellerson Station, New York.
Lackey, J .A. 1 977. A revised classification ol the tribe Phaseoleae (Leguminosae-Papilionoideae)
and its relation to canavanine distribution. J. Linn. Soc, Bot. 74:163-178.
Lackia, J. A. i 983. A review of generic concepts in American Phitseolinae (Fabaceae, Faboideae).
Iselya 2:21-64.
Lewis, G.P. 1991. A new combination in Dolichopsis (Leguminosae: Papilionoideae). Kew
Bull. 46:35 L
JVIabrv, T.J., K.R. Makkham, and M.B. Thomas. 1970. The systematic identification of
flavonoids. Springer Verlag, New York.
Mareceial, R., J.M. Mascherpa, and F. Stainier. 1978. Etude taxonomique d'un groupe
complexe d'especes des genres Phaseot/is et Vigtui (Papilionaceae) sur la base de donnees
morphologiques et polliniques, traitees par I'analyse informatique. Boissiera 28: 1—273-
Markham, K.R. 1982. Techniques ol flavonoid identification. Biological Techniques Se-
ries. Academic Press. London.
Piper, C. V. 1 926. Studies in American Phaseolineae. Contr. U.S. Natl. Herb. 22:663-70 1 .
RoHij-, F.J. 1993. NTSYS-pc. Nimierical taxonomy and multivariate analysis system, ver-
sion 1 .8. Exeter Software, Setauket, N.Y.
WiELiAMS, C.A., J.C. Onyieagha, andJ.B. Harborni-;. 1995. Flavonoid profiles in leaves,
flowers and stems of forty-nine members of the Phaseofinae. Biochem. Syst. Ecol. 23:655-
667.
Zaei.occhi, E.M. and A.B. Pomilio. 1994. Evolution of flavonoids in the Phaseolinae. Phy-
tochemistry 37:449-453.
ZAEi.oa:En, E.M., A.B. Pomilio, and R.A. Pai,ac:ios. 1995. Estudio quimiotaxondmico de
la SLibtribu Phaseolinae (Phaseoleae-Papilionoideae-Leguminosae) — III: Flavonoides de
las especies argentinas de los generos Phciseohis y Dolichopsis. Darwiniana 33:1 35-148.
RUELLIAJIMULCENSIS (ACANTHACEAE),
A NEW SPECIES FROM THE CHIHUAHUAN
DESERT AREA, MEXICO
JOSE A. VILLARREAL Q.
Departaniento de Botdnica
Universidad Autonotna Agraria "Antonio Narro"
Buenavista, Saltillo, Coahuila 23315, MEXICO
ABSTRACT
Rutllici jhiii/lcensis Villarreal sp. nov., from the Jimulco mountains area (southwestern
Coahuila and northeast Durango), is described and illustrated. It is similar to R. ocadentalh
(Gray) Tharp and Barkley, and recognized by its telatively smallet flowers, shorter corolla
basal tube and its distribution.
RESUMEN
Ri/elliii jimulcensis Villarreal sp. nov. del area de las montaiias de Jimulco (suroeste de
Coahuila y noreste de Durango), es dcscrita e ilustrada. Es similar a R. occidmtalh (Gray)
Tharp y Barkley, y se reconoce por sus flores mas ]:iequenas, tubo basal de la corola mas
corto y su distribucion.
Key Words: Acanthaceae, Ruellia, Coahuila, Flora of Mexico.
Ruellia, a tropical and subtropical genus of about 200 species is repre-
sented in the Chihuahuan Desert Region by six species (Henrickson & Johnston,
in press). A new species is proposed as part of the study of the Flora of Coahuila.
Ruellia jimulcensis Villarreal, sp. nov. (Fig. 1). Typr,: MEXICO. Coahuila: Mpio.
Torreon, Sierra de Jimulco, mina San Jose, vereda hacia la cima, 103 1.3' W, 25 ()8'N.
Matorral con Bonetidla anomab. Agave kchitgialla. Acacia herlandien, Flourensia, Hechtia,
SpiraeayAralia, 1 800-1850 m, 10 Ago 1994,/ A. Villarreal Q_. 1181 yM. A. Carranza.
(holotype: MEXU; isotypes: ANSM, ENCB, TEX).
Ruellia occidentalh (Gray) Tharp & Barkley similis sed differt flores plus brevis, corolla
tubis plus brevis, calyx lobis longius quam corolla tubis; flores cleistogamous absens et
differt distributio.
Perennial herb from clustered fibrous roots; stems erect to ascending 30-
50 cm tall, densely glandular pubescent with straight hairs about 1 mm
long; the mternodes 4-10 cm long; leaves with petioles 6-30 mm long,
blades broadly ovate to obovate, 3-10 cm long, 2-8 cm broad, the base
obtuse to rounded, briefly decurrent along the petiole, apex obtuse to acute,
the margin entire to undulate-crisped, both surfaces viscid with abundant
glandular hairs, the dried leaves usually green-yellowish; flowers in dicha-
sia and terminal thyrsoid panicles 5-20 cm long, to 10 cm broad, strongly
SiDA 18(1): 223-226. 1998
224
SlDA 18(1)
Fici. 1. R/icllici jh}]//Iieiisi.s, a complete phmt
ViLLAREAL Q., A new species of Ruellia 225
glandular-pubescent; peduncles ascending; calyx lobes 1 5—25 mm long, linear
attenuate, united at the very base, 1.0—1.2 mm broad at the base, unequal,
strongly glandular pubescent; corolla funnelform, bluish-purple 30—40 mm
long, the basal tube 8—12 mm long, the broadly campanulate throat 15—
20 mm long, the lobes 8—14 mm long, almost as broad, erose; cleistoga-
mous flowers absent; stamens didymanous, borne at the distal portion of
the corolla tube, free filaments 4—10 mm long; anthers 3—4 mm long; style
15—20 mm long; fruit ellipsoid 12—20 mm long, 3—4 mm broad, glandu-
lar pubescent; seeds 10—12, circular to oblate, 2.0—3.0 mm long, narrowly
winged to the apex, brownish, covered with dense apressed trichomes.
Additional specimens examined: MEXICO. Coahuria: Mpio. de Torrcon, Sierra de Jimulco,
proximiclacles ai ejido Trinidad, 25 08' N, 103 22' W, Matorral de Agave lechugiiiUa, Boiiteloua
raniosa, Caesalpinia ses5iliflora,Jatropha y Yucca, ladera rocosa, 1900—2000 m, 25 Ago 1983,
J. A. Villarreal 4387, Al.A. Carrairza y A. Rodriguez (ANSM); Sierra de Jimulco, mina San
Jose, 25" 08' N, 103" 13' W, Matorral desertico, 1800-1850m, 11 Oct 1993, M. A. Carranza
1951 yj. Noriega (ANSM); ca. 54 air km SSE of Torreon in canyon above Estacion Otto in
SW side ot Sierra de Jimulco near Mina San Jose, in limestone area with Acacia, Celtis,
Viguiera, Parthenium,Jatropha, Trixis, Fo/fq/zieria, Yucca, 25 04' N, 103 13' W, 1850 m, 12
Sep 1980,7, Henrickson & P. Bekey IH5()4 (TEX). Durango: Mpio. de Cuencame, Sierra El
Rosario, camino a la estacion de microondas Sapioris, carr. 49, 30 km al SE de Lerdo, 25
24'N, 10343' W, Matorral de Acacia crassijol/a, Viguiera stenoloha, Bursera Khlen dent all , Opuntia
imbncata y Fouquieria splendem, 1750-1800 m, 16 Ago 1991, J-A. Villarreal 6243 y M.A.
Carranza (ANSM); Estacion de microondas Sapioris, ca. 30 km SW of Gomez Palacio on
Hwy to Durango 25" 24'30" N, 103" 43' W, matorral desertico microfilo, 1400-1 500 m,
25 Mar 1973, M.C. Johnston, T.L. Weridt & F. Chiang 10409 (TEX); just SSE of Estacon
Microondas Sapioris, about 20 km NW of Estacion Chocolate, 25" 25' N, 103" 43' W, 1450-
1500 m, 14 Aug 1973, M.C.Johnston. T.L. Wendt. F Chiang &J. Henrickson 12210 (TEX);
Microondas Sapioris, along cobblestone road which departs from Hwy 40 N of Estacion
Chocolate, ca 15 air mi (25 km) W-SW of Torreon, 25" 25' N, 103" 42' W, Teconia stans.
Agave lechuguilla. Euphorbia antisyphilitica 'And d'wevsc c-Act'i, 1300 m, 30Jul 1991, AI. Mayfield,
A. Hempel & A.Jack 1093 (TEX); Mexico Hwy 40, 25 mi SW of Lerdo, 6 nov 1964, D.
Flyr 251 (TEX); Mpio. de Lerdo, 4 mi southwest ol Chocolate, route 31, growing benerh
Prosopis on clay flat in valley, 23 Jul 1 958, D.S. Correll & I.M.Johnston 20008 (TEX); ca. 4
mi SW ot Ciudad Lerdo along Hwy 40 to Zacatecas, on limestone hillside with Larrea,
Jatropha, Acacia, Opuntia, Agaveetc.,2'i" 3V N, 103 32' W, 1200 m, 2 I Sep 1978,/. Henrickson
&E. Lee 17474 (TEX).
Ruellia ji??iulcensis is found on sandy hillsides and dry canyons at eleva-
tions of 1200— 2()()() m, in xeric shrublands at the complex of mountains
near Sierra de Jimulco and Sierra del Rosario.
The new species has leaf blades broadly ovate, obtuse to rounded at the
base as R. occidentalis and often with strongly stipitate glandular trichomes
covering the stems and inflorescence. It differs in its flowers 3—4 cm long,
the basal tube of the corolla 8—12 mm long, the calyx lobes longer than the
basal tube, the cleistogamous flowers unknown and its distribution, as marked
in the key below. Tharp and Barkley (1949) gave the name R. occidentalis
226 SiDA 18(1)
WAt. ferrisae to a specimen from the mountains near Monterrey, Nuevo Leon
which is recognized by Henrickson and Johnston (in press) by this name
for the specimens described here. The type specimen from the Monterrey
area has oblong-obovate leaf blades, lacks stipitate glands on stem and leaves
and flower dimensions that better fit with R. nudiflora, a common species
in Monterrey area. The populations of the proposed species grow allopatri-
cally in an isolated area at the southwestern corner of the Chihuahuan Desert
Region while most of the related species are distributed in the eastern Si-
erra Madre Oriental (Turner 1991)-
The three species of Ruellia previously mentioned can be separated by
the following key:
i . Leaf blades ovate to elliptic-obovate, mostly less than 3 cm wide, sparsely to
moderately pubescent; lower internodes pubertilent to glabrate R. nudiflora
1. Leaf blades broadly ovate to subdeltoid, 4-7(— 9) cm wide, usually densely
pubescent; lower internodes with abundant stipitate glands 2
2. Flowers (chasmogamous)4.5-5.5{— 6.5) cm long; basal tube of the corolla
2.5—3.5 cm long, longer than the calyx lobes; cleistogamous flowers (smaller
than the chasmogamous) frecuently located in the lower nodes; se Texas,
ne Mexico (e Coah, ne N.L., Tamps, ne Vcr.) R. occiclentalis
2. Flowers 3.5—4.0 cm long; basal tube of the corolla H— 1 2 mm long, shorter
than the calyx lobes; cleistogamous flowers absent; sw Coahuila and ne
Durango R . j i mulcensis
ACKNOWLEDGMENTS
I thank Tom Wendt for the loan of specimens for revision from TEX/LL.
The illustration was prepared by Cuauhtemoc Gonzalez de Leon.
REFERENCES
Henrickson, J. and M.C.Johnston, (m press). A flora of the Chihuahuan Desert region.
{Ruellia). Los Angeles, California. 2:1 151-1 155.
Tharp, B.C. and F. Barki.i;y. 1949. Genus Rtidlui in Texas. Amer. Midland Nat. 42: 1-86.
Turner, B.L. 1991 . Texas species oiRi/ellia (Acanthaceae). Phytologia 7 1 :281-299.
UNA NUEVA ESPECIE DE AGAVE, SUBGENERO
LITTAEA (AGAVACEAE) DE GUERRERO Y
OAXACA, MEXICO
ABISAI GARCIA-MENDOZA
Jardin Botdnico, U.N. A.M.
A. P. 70-614. Del Coyoacdn
04310 Mexico. D.F., MEXICO
ESTEBAN MARTINEZ SALAS
Depto. de Botdnica, Imtituto de Biologia, U.N. A.M.
A. P. 70-233. Del. Coyoacdn
04310 Mexico, D.F., MEXICO
RESUMEN
Se describe e ilustrs. Agave gracilis de los estados de Guerrero y Oaxaca, Mexico. La especie
pertenece al grupo Striatae Baker, del subgenero Littaea (Tagliabue) Baker, y muestra si-
militudes con A. dasylirioides ^diCohi & Bouche.
ABSTRACT
Agave gracilis from Guerrero and Oaxaca, Mexico, is described and iUusrrated. The spe-
cies is a member of group Striatae Baker, subgenus Littaea (Tagliabue) Baker. It is similar
to A. dasylirioide.s JdLQohl & Bouche.
Las exploraciones botanicas recientes realizadas en los estados de Guerrero
y Oaxaca, Mexico, revelaron la existencia de una nueva especie de Agave, pertene-
ciente al subgenero Littaea (Tagliabue) Baker, grupo Striatae Baker, de la
familia Agavaceae. El grupo Striatae es endemico de Mexico y los cinco taxa
que lo conforman se caracterizan por presentar hojas estriadas, lineares, con
el margen serrulado, flores campanuladas o cilindricas con el tubo bien desarroUado
y ovario sin cuello que se proyecta hacia el interior del tubo (Gentry 1 982).
Agave gracilis Garcia-Mend. & E. Martinez, sp. nov. (Fig. 1). Tipo: MEXICO.
GuHRRniio: Municipio de Tlapa, El Salado, 8 km al N de Tlapa, camino a Huamuxtitlan,
selva baja caducifolia, 990 m, 16 Nov 1982 (fls), £. Martimz et al. 2659 (iiolotipo:
MEXIJ; isoTiPo.s: BRIT, ENCB, K, MO).
Plantae perennes, caespitosae. Folia 40-60 x 0.4—0.9 cm, linearia, striata, margine subtiliter
dcnticLilata, glauca vel viridi-glauca. Flores campanulati 2— 2.2(— 2.5) cm longi, tubo 3—4
mm longo, 5—7 mm lato; filamenta 2.7—3 cm, longa, in apice tubi inserta; ovaritim collo
carens, in tubi interiorem projeccum; capsulae globosae 9—10 x 8—9 mm.
Plantas perennes, cespitosas, con troncos rastreros de hasta 1 m de largo;
rosetas hemisfericas, compactas, 50—80 cm de diametro, 50—60 cm de alto.
SiDA 18(1): 227-230. 1998
228
SmA 18(1)
Fig. 1 . Agave gracilis, a) planca complt-ra con infloresccnt ia, b) bracteas superiores del pedunculo,
c) hoja, d) detalle del margen dc la hoja, e) flores pareatlas en la inflorescencia, f) flordisectada,
g) capSLilas, h) scmillas. Ilustrat ton basada en los especi'mcnes E. Mitrti'iiuz et uL, 2639 y A.
Gam'd-Mi'iuioZLi y L. de hi Rosa 6"^ OH.
Garci'a-Mendoza and Martinez, Una nueva especie de Agave de Mexico 229
Hojas mas de 100 por roseta, 40-70 cm de largo, 0.4-0.9 cm de ancho en
la parte media, ensanchandose en la base hasta 1-1.5 cm, lineares, estriadas,
planas, flexibles, glaucas o verde-glaucas, subcoriaceas; margen finamente
denticulado, amarillento; espina 3— 6(— 8) mm de largo, debil, de color pardo-
rojizo. Inflorescencia de 1.8-2 m de largo, erecta o ligeramente inclinada,
espiga en el cuarto superior o mitad superior del pedunculo, bracteas del
pedunculo 5-15 cm de largo, 2-4 mm de ancho en la base, disminuyendo
en tamaiio hacia el apice, lineares, pardas, debiles, sin espina o esta apenas
marcada; bracteas florales 2-3.5 cm de largo, 1-2 mm de ancho, lineares,
pardas, persistentes, mas iargas que las flores. Flores 2-2.2(-2.5) cm de largo,
campanuladas, verdes, con el apice de los tepalos rojizo-oscuros; pedicelos
1 mm de largo, alargandose a 2 mm durante la fructificacion; ovario 7—10
mm de largo, 2-4 mm de ancho, cilindrico, sin cuello, glabro o glabrescente,
penetrando ligeramente en el tubo del perianto, este de 3-4 mm de largo,
5-7 mm de ancho; tepalos 0.9-1 ■ 1 cm de largo, 2.5-3.5(-4.5) mm de ancho,
oblongos; fdamentos 2.7-3 cm de largo, insertos en el apice del tubo; anteras
7-9 mm de largo, 1 mm de ancho, centricas, rojizas. Capsulas 9-10 mm de
largo, 8-9 mm de ancho, globosas, pardo-oscuras, con el perianto, estambres
y estilo de la flor persistentes. Semillas 3-3.5 mm de largo, 2-2.5 mm de
ancho, engrosadas en el lado curvo, negras.
Paratipos; MEXICO. Guerrero: Municipio de Atlixtac, 1 km al O de Santa Isabel, 30
km al E de Chilapa, carr. a Tlapa, 25 Nov 1989 (A), J.L. Contreras 2651, 2632 (FCME);
Municipio de Tlapa, rio El Salado en su union con el rio Tlapaneco, 8 km al N de Tlapa,
carr. a Huamuxtitlan, 7 May 1997 (fr), A. Garci'a-Mendoza y L. de la Rosa 6508, 6311,
6514 (ENCB, MEXU); Municipio de Zumpango del Rio, Xocohite, Xochipala, 23 Abr
1993 (fr), A. Gomez s.n. (MEXU). Oaxaca: Distrito de Huajuapan, El Boqueron, cafion del
n'o Mixceco, 5 km al N de Tonala, 13 Ago 1993, A. Garcia-Mendozay F. Palma 581 1 (MEXU).
La especie aqui descrita se locaHza en la cuenca interinedia del rfo Balsas,
en los estados de Guerrero y Oaxaca. Seguramente su distribucion se extiende
hacia otros afluentes del rio y posiblemente mas alia, siendo quiza, la planta
senalada por Ullrich (1990) para el cerro Yucuyu en el Distrito de Tlaxiaco,
Oaxaca. Agave gracilis crece en laderas y barrancas abruptas, sobre suelos derivados
de rocas calizas, entre los 850 y 1300 m snm. Habita preferentemente en
las selvas bajas caducifolias y su ecotonia con los bosques de Quercus-Pinus.
Algunos de los generos frecuentes con los que convive son: Acacia, Agave,
Bur sera, Euphorbia, Hechtia, Ficus, F orchhammeria, Neobuxbaumia y Pachycereus.
Agave gracilis presenta similitudes morfologicas con Agave dasylirioides
Jacobi & Bouche, especie de la que se diferencia por su habito cespitoso,
roseta mas reducida en tamaho, hojas mas cortas y angostas, inflorescencia
erecta o ligeramente inclinada y flores mas pequehas; las capsulas globosas
y de tamano mas reducido contrastan con las capsulas oblongas y mas grandes
230 SiDA 18(1)
'['ahi_a 1 . ('ompuracion lIc algunas caracai'i'sricas niorfolof^icas entrc Agari' i^ycnilis y /i- cliisyHrio'ulei.
(^anicrer Agiive gnu'ilts Agave dcisylirinida
Flantas cespitosas solitarias
'IVonco hasta Im, rastrcro hasca 1.5 m, rasrrero
Diamecro de la roseca 50-80 cm 100-200 cm
'nimano de hojas 40-70 X 0.4-0.9 cm (40_)6()-l 00 x 2-.^. 8 cm
C^olor de hojas ghuicas o vcrdc-^i^diiLicas verdes o verde-i,daLicas
Iiitloresccncia 1.8-2 m, erecta o (1.5-)2-2.6 m, arqucatla
ligeramenre inclinatia
I.ongiCLid de florcs 2-2.2(-2.5) cm (2.6-)3-,t.5 cm
Tubo de la tlor 3-4 X 5-^ mm (8-)l()-15 X (8-)12-]6 mm
Tamano de Hlamcntos 2.7-3 cm 35-5 cm
Inscrcioii de filamenros apice del tubo mitad del rubo
Capsulas 9-10 X 8-9 mm, giobosa.s (I0-) 15-20 X 6-9 mm, obiongas
Semillas 3-V5 X 2-2.5 mm 3 — i X 2.5-^ mm
de A. clcLsylirioides; asi mismo, las semillas son de tamano menor en A. gra-
cilis. Una comparacion de los caracteres mas sobresalientes de ambas especies
se presenta en la Tabla 1 . Las medidas de A. dasylirioides se tomaron en parte
de Gentry (1982) y se ampliaron de acuerdo con observaciones propias. Ambas
especies habitan en la cuenca del rio Balsas, pero A. dasylirioides ocupa la
parte alta, mas htlmeda y fria, entre los 1900 y 2500 m snm, sobre laderas
con suelos derivados de rocas volcanicas, en bosques de Querms-Pin//s con
Alniis. Arbutus, Clethra, Cremnophila, Garrya, Hechtia y Salvia. La epoca de
floracion de las dos especies se presenta en los meses de octubre y noviembre.
El epfteto especifico se refiere al caracter esbelto de la planta.
ACRADF.CIMIF.NTOS
Agradecemos la revision del manuscrito a Raquel Galvan, Susan Verhoek,
Wendy Hodgson y Fernando Chiang. La descripcion latina fue hecha por el
ultimo autor. La ilustracion es aportacion de Albino Luna.
REFERENCIAS
Gentry, H.S. I9H2. A^^avcs ol Continental North America. University o( Arizona Pres.s,
Tucson.
Ui.LRiOi, B. 1990. Em neuer Standort Hir Agave chisylirionlcs ]i}i<:oh\ & Bouciie in Oaxaca.
Kakreen Sukk. A 1 (H): l64- 1 66.
A NEW SPECIES OF MANDEVILLA
(APOCYNACEAE) FROM JALISCO, MEXICO
JUSTIN WILLIAMS
Department of Botany
University of Texas
Austin. TX78713, U.S.A.
abstrac:t
In preparation for a forthcoming treatment of tlie Apocynaceae of Mexico, a routine examination
of herbarium specimens revealed the following new species, MandevUla pririgki ].¥^. Wil-
liams, sp. nov. The new species is distinct from other members of its alliance (sect. Torosae)
in having subsessile leaves and larger peduncles and flowers. In addition, Mandevilla apocynifoUa
(A. Gray) Woodson is here presented to be a synonym of Al. foliosa (Miill. Arg.) Hemsl.
RESUMF.N
En la preparacion de un proximo tratamiento tie las Apocynaceae de Mexico, un examen
de rutina de los especimenes de herbario revelo la nueva especie siguienre, Mandevilla pringlei
J.K. Williams, sp. nov. La nueva especie se distingue de otros miembros de su alianza (sect.
Torosae) por tener hojas subsesiles, y pedunculos y flores mas grandes. Ademas, Mandevilla
apocynifoUa (A. Gray) Woodson se presenta aquf como un sinonimo de M. foliosa (Miill.
Arg.) Hemsl.
Key words: Apocynaceae, Mandevilla, }'A\%c<.^, Mexico
Mandevilla pringlei J. K. Willliams Sp. nov., (Fig. 1). Type: MEXICO. Jalisco:
Ri'o Blanco, near Guadalajara, 12 Jul 1902, C.G. Pringle 1 1337 (hoeotype: US!).
Mandevilla foliosa (Miill. Arg.) Hemsl. affinis sed (oliis subsessilis petiolis 2-3 mm longis
(vice 10-18 mm in M. foliosa), pedunculis longioribus (7-10 cm vice 0.3-1.2 cm) lobis
coroUae longioribus latioribusque (12-15 X 10-14 mm vice 5-8 X 2-4 mm) plantis
suffrutescentibus (vice lignosibus ramificantibusque).
Suffrutescent herbs to 0.4 m tall, stems pubescent. Leaves 3-7 cm long,
opposite, subsessile, pubescent; petioles 2-3 mm long; blades 3-7 cm long,
1.8-2.5 cm wide at middle, ovate-elliptic, apex acute, base sub-cordate,
with 2-4 glands at apex of petiole on upper side. Inflorescence lateral, in-
determinate, racemose, with 2-10 flowers; peduncles 7-10 cm long, pu-
bescent; bracts 4-6 mm long, narrowly triangular to lanceolate, straight;
pedicels 10-17 mm long, pubescent, occasionally twisted. Sepals 5, basally
fused, 5-7 mm long, ca 0.5 mm wide, narrowly triangular to lanceolate,
straight, pubescent. Corolla salverform, yellow; tube 11-18 mm long, constricted
at mouth, lower half internally glabrous, upper half internally pubescent;
lobes 12-15 mm long, 10-14 mm wide, obovate, acuminate, occasionally
equal in length to the tube but always greater than half its length, spread-
SiDA 18(1): 231-235. 1998
232
SlDA 18(1)
UNITED STATLS NATiONAL MUSEUM
P(.ANTA-: MEXICANS.
s:Ars or ;ai.:s:o
'-— ■■» /■— ..
Fic;. 1. H()loty|ie of AL/Wfr///i//^;7;/,(,'/(7 J.K. Williams.
Williams, A new species of Mandevilla 233
ing, pubescent. Stamens 3—4 mm long; filaments ca 0.5 mm long, pubes-
cent, straight; anthers ca 3 mm long, bases sagittate with blunt lobes. Pis-
tils 7—11 mm long; style 5—7 mm long, glabrous; ovary ovoid, ca 1 mm
long, glabrous; pistil head pentagonal, 2—3 mm long. Nectaries 5, as long
as to slightly shorter than ovary. Mature follicles unknown, immature fol-
licles fused at apex, pubescent.
Additional specimens examined: MEXICO. Jalisco: Mpio. iVIazamirIa, Fraccionamiento
Los Cazos, to the S of Mazamitla, 9 Jul 1995, Machuca 7351 (TEX); Cerro Viejo, S face,
above Zapotitan de Hidalgo, a village 1 mi N of Hwy iVIEX 15, ca 25 mi due S, or 45 road
mi from Guadalajara, base of mountain in open meadow just E of trail, alt 1890 m, 27 Jun
1956, D. P. Gregory & G. Eiten 225 (P, SMU); Huejotitan, Jul 1912, Digi/et s.u. (P).
Distribution. — Mandevilla pringlei is known only from five collections
made from the Pine-Oak forest near Guadalajara, Jalisco, Mexico.
The new species honors Cyrus Guernsey Pringle (1838-1911) promi-
nent collector of the Mexican flora, and whose collections led to many a
new species in the Apocynaceae, and other families.
All specimens are with flowers and buds, Machuca 7331 is also with fruit
but the fruits are extremely immature, only 5 mm in length. The closest
relative o{ Mandevilla pringlei, M. foliosa (MiLill. Arg.) Hemsl., has an aver-
age follicle length of 10 cm.
It should be noted that according to Pringle's diary (Davis 1936, p. 199)
on the days prior to and after his collections made on July 22, 1902 he was
collecting numbers in the 8600 series. This, however, is in contrast to the
type collection of M, pringlei which is numbered 1 1357. The diary, how-
ever, does specifically state that Pringle was collecting along the Ri'o Blanco,
the type locality of AI. pringlei, on July 22, 1902. In the numerical listing
of Pringle's collections (ibid) the number 11337 is used twice. One listing
is for E. apocynifolia A. Gray (= M. foliosa see below) the second is for an
unidentified species of Asclepiadaceae. In the absence of a thorough expla-
nation of Pringle's numerical system it is reasonable to assume that the collection
number of the above type specimen is simply an error in numbering.
To date the most taxonomically thorough investigation o(I\Aanckvilla remains
Woodson's (1933) revision. Infrageneric relations are at present moderately
unclear, however, Woodson provided both subgeneric and sectional divi-
sions which remain undisputed.
I^landevilla pringlei is a member of subgenus Mandevilla (as evidenced by
the lack of glands along the midrib of the upper surface of the leaves) where
it relates to section Torosae, evidenced by its suffrutescent habit and non-
twinning stems (Woodson 1933). Mandevilla pringlei is most closely related
to M. foliosa, sharing with it an erect habit (opposed to the trailing habits of
M. karwinskii (Miill. Arg.) Hemsl. and M. torosa (Jacq.) Woodson) and hav-
234
Sum 18(1)
Fic;. 2. Comparision of flowers. A. Mandevilla foliosa (Miill. Arg.) Hemsl. (V. Fmik 2766,
TEX). B. Mandevilla prmglei].K. Williams (P. Gregory & G. Etten 225, SMU). Bhick bar
represents i cm.
ing leaves with an average length greater than 5 cm (vs. 2—5 cm of AI, mexicana
(Mlill. Arg.) Woodson). Mandevilla pringlei differs from M. foliosa in a number
of floral and habit characters that are contrasted below:
1. Branching shrubs to 1.5 m tall; leaves petiolate, petioles 10—18 mm long;
peduncles 0.3—1 -2 cm long; corolla lobes 5-8 ( 1 0) mm long, 2—4 mm wide,
up to btit not exceeding half the length ot corolla tube (Fig. 2a); stems, leaf
blades, inflorescence, and fruit glabrous or pubescent, but midrib of leaves
always pubescent; midrib hairs linear lanceolate O.f 5— 0.3 rnm long (Fig.
3a); throughout Mexico M. foliosa
1 . Sutfrutescent herbs ro 0.4 m tall; leaves subsessile, petioles 2—3 mm long;
peduncles 7—10 cm long; corolla lobes 12—15 mm long, 10—14 mm wide,
half or more the length of the corolla tube (Fig. 2b); stems, leaves, inflores-
cence and fruit pubescent; midrib hairs triangular, 0.1—0.15 mm long (Fig.
3b); Jalisco IVI. pringlei
In the most recent keys to the species oi Mandevilla (Woodson 1933, 1938),
M. pringlei keys out to M. apocynifolia (A. Gray) Woodson (BASfONYM: Echites
apocynifolia). In fact, Woodson (1933) cites the type of Al. pringlei as a specimen
of AI. apocynifolia. An examination of an isotype of Al, apocynifolia {Palmer
734; holotype: GH; isotype: NY!), however, reveals that it is a synonym of
M. foliosa {Ghiebreght s.n.\ holotype: G n.v., photo-holotype F! MO!). Al-
though the type of AI. apocynifolia is without flowers, it is evident that the
sheet represents a specimen of AI, filiosa due to its branching, petiolate leaves,
glabrous stems and fruits, and linear-lanceolate midrib hairs. Mandevilla
foliosa has both glabrous and pubescent-stemmed populations scattered
throughout its range. However, in the region of Jalisco where the type of
AI, apocynifolia was collected the populations have consistently glabrous stems
and fruits.
Williams, A new species of Mandevilla
235
m^m
Fig. 3. Scanning electron micrographs of Mandevilla midribs of leaf undersurface showing
the length and shape of midrib hairs. A. Mandevilla foliosa (Mull. Arg.) Hemsl. {R. King &.
T. Soderstrom 4632, TEX). B. Mandevilla prmglei J. K. Williams (Machuca 7351, TEX). White
bar on both photos represents 0.1 mm. Microphotographs made by theauthor using a Phillips
515 SEM (Cell Resource Center, University of Texas Austin).
ACKNOWLEDGMENTS
I would like to thank Paul Fryxell for providing the Latin diagnosis and
to him and Billie Turner for reviewing the original manuscript. Thanks is
also extended to the curators of F, MO, NY, P, SMU, TEX and US for al-
lowing me the opportunity to observe their specimens. I would also like to
credit the staff of the Cell Research Center (University of Texas at Austin)
for allowing me access to their scanning electron microscope.
REFERENCES
Davis, H.B. 1936. Life and work of Cyrus Gtiernsey Pringle. Univ. Vermont, Burlington.
Woodson, R.E., Jr. 1933. Mandevilla: In; Studies in the Apocynaceae IV. Ann. Missouri
Bot. Card. 20:645-777.
1938. Mandevilla. In: N.L. Britton et al., eds. N. Amer. Fl. 29:645-777.
236 SiDAlS(l)
BOOKS RECEIVED
Fautin, Daphni; Gail, Douglas J. Futuyma, and Frances C. Jamls, eds.
1996. Annual Review of Ecology and Systematics. Volume 27.
(ISBN 0-8243-1427-1 , hbk). Annual Reviews Inc., 4139 El Camino
Way, P.O. Box 10139, Palo Alto, CA 94303-0139. $52.00. 648 pp.
There is quite a mix ot articles reviewing' tlie fields of ecolo^^y and systematics for J 996. A
total of 20 articles are presented followed by a Subject Index, Cumulative Index of Con-
tributing Authors, and a Cumulative Index of Chapter Titles, Volumes 23-27. The con-
tents are as loUows: Early history and progress of women ecologists: Emphasis upon re-
search contriburions. Forest Canopies: Methods, hypotheses, and future directions. Extinction
by hybridization and introgression. Evolutionary significance of resource polymorphisms
in fish, amphibians, and birds. Management of the Spotted Owl: A case history in conser-
vation biology. Trouble on oiled waters: lessons from the Exxon Valdez Oil Spill. Evolu-
tionary significance ot local genetic differentiation. Rates of molecular evolution: Phylo-
genetic issues and applications. Herbivory and plant defenses in tropical forests. Mechanisms
creating community structure across a freshwater habitat gradient. Natural freezing sur-
vival in ainmals. Demograjihic and genetic models in conservation biology. Gene trees,
species trees, and systematics. Incidence and consequences of inherited environmental ef-
fects. Recruitment and the local dynamics of open marine populations. When does mor-
phology matter.-' Adaptive evolution of photoreceptors and visual pigments in vertebrates.
Microbial biodiversity: Domains and Kingdoms. The geographic range: Size, shape, boundaries,
and internal structure.
Jami:.s Stijbhendii;ck, Stephan L. Hatch, and Charles H. Butji-rfield. 1997.
North American Range Plants. Fifth edition. (ISBN 0-8032-9243-
0, pbk). University of Nebraska Press, 312 Nortii l4th Street, Lin-
coln, NE 68588-0484. (402)472-3581. $25.00. 501 pp. Illustrated.
"The 200 species in this book were selected because of their abundance, desirability, or
noxious properties. The list of plants was developed over a nearly 40-year period by coaches
of range plant identification teams and faculty from the colleges and universities with range
management programs. The formal list is now the Master Plant List for the International
Range Plant Identification Contest sponsored by the Society for Range Management (1839
York Street, Denver, CO 80206)." Each species is treated with the following information:
Tribe, species, common name, life span, origin, and season, followed by more detailed in-
formation on Inflorescence Characteristics, Vegetative Characteristics, Growth Character-
istics, Forage Value, and Habitat. An dlustration and range map are provided for each plant.
SiDA 18(1): 236. 1998
A NEW COMBINATION IN MEXICAN
MANDEVILLA (APOCYNACEAE)
JUSTIN WILLIAMS
Department of Botany
University of Texas
Austin. TX 78713, U.S.A.
ABSTRACT
In preparation tor a forthcoming treatment oi the Apocynaceae of Mexico, it was discov-
ered that Echites holosericea of Sesse & Mo^. has priority over the species Mandevilla sertnligera
Wocxlson and Al. syrinx Woodson, accordingly Mandevilla holosericea (Sesse & Mog.) J.K.
Williams comb. nov. is here proposed.
RESUMEN
En la preparacion de un proximo traramiento de las Apocynaceae de Mexico, se descubrio
que Echites holosericea Sesse & Mog. tiene prioridad sobre Mandei'illa sertidigera Woodson y
M. syrinx Woodson, como consecuencia se propone Mandevilla holosericea (Sesse & Mog.)
J.K. Williams comb. nov.
Mandevilla holosericea (Sesse & Mog.)J.K. Williams, comb. nov. Basionym:
Echites holosericea Sesse & Mog., Fl. Mex. (ed. 2) 45. 1894. Type: MEXICO. MiCHOACAN:
"Ahualulci sylvis," Oct 1787-1795, ^&v.«'<& Mogino 3073 (holotypr: MA!; isotype: F!).
Mandevilla syrinx Woodson, Ann. Missouri Bot. Gard. 19:53. 1932. Typi:: MEXICO.
Jalesco: barranca of Tequila, 8 Oct 1893, Pringle 5422 (Hc:iE(yrYPE: MO n.v.).
Mandevilla sertuligera Woodson, Ann. Missouri Bot. Gard. 19:383. 1932. Type: MEXICO.
Michoac;an: rocky hills, Coru Station, 23 Jan 1907, Pringle 1 3890 (hoeotype: US!).
Representative specimens. MEXICO. Jalisco: baranca near Guadalajara, Jun 1886, Palmer
98 (US). MEXICO: San Lucas, district Temascaltepec, 28 Nov 1933, Hvnton 5274{VSy,
San Lticas, district Temascaltepec, 26 Dec 1933, Hinton 7 176 (US); Dist. Temascaltepec,
Volcan, 1410 m, 9 Aug 1932, Hinton 1295 (US), MICHOACAN: rock-fields, Coru Sta-
tion, 15 Oct 1904, Pringle 13106 (TEX, US-2 specimens). MORELOS: lava beds near Cuernavaca,
500 ft, 23 Jun 1896, Pr/ngle 6329 (US). OAXACA: 5 km E of Tamazulapan, road to Chilapa,
district of Teposcolula, 1800 m, 7 Jun 1985, Mendoza et al. 1467 (NY).
Sesse and Mo^ino (1887, 1894) described 19 species of Echites and the
types for a majority of these names have not been located. In preparation of
a forthcoming treatment of the Apocynaceae of Mexico, the author studied
the collections of "Echites "of Sesse and Mogino during July 1995 at the Madrid
herbarium (MA), and from material borrowed from the Field Museum (F)
during December 1996 in order to locate and observe the types of their
Echites species. These observations revealed the following novelty.
The type of the species Echites holosericea Sesse and Mog. has up until now
Sida 18(1): 237-239. 1998
238
SiDA 18(1)
REAL JARDIH BOTANICO MADRID
/at.
txV
OB °~ ~i
or
ZB " ^
col
X
C'^ /^^ ^ ^* ^* t*^ J
/^J-tit^ii* f - ' ^ /£4.,^ZCc/^-<--^- ^l l*^lh^^
.1 S,.vso. .M....in<i. 0--ini.. vt .MaiJ.'i-ad.. I. .I..V
/
,/,.--'■/.
CHICAGO NATURAL
HISTORY MUSEUM
NECaii^f MO
TT"fn-T"
liilj
Ji -i ^1
Fic;. 1 . Holotypc <.->{ Echitcs holosericea Sesse & Mo^.
Williams, A new combination in Mandevilla 239
not been formally identified and presented. The typification of £, holosericea
has been made difficult because there are no specimens so labeled among
the Sesse and Mogifio collections at F or MA. Many of the posthumously-
published species names of Sesse and Mogifio are not the same ones that are
written on the specimen labels (Fuertes & Fryxell 1993). Sesse and Mogino
(1894) described the species as having opposite subcordate leaves with rusty
red-white tomentum covering all parts of the plant and white flowers with
a hairy tube arranged in a compact raceme. Only one specimen (Fig. 1) in
the Sesse and Mogino collections matches this description. This specimen
was labeled by them as ''Echites tubulosci N". The name E. tubulosa was never
validly published by them, however, the letter "N "after the name indi-
cates Sesse and Mogino considered this specimen to be a new species (Fuertes
& Fryxell, 1993). Since only one specimen matches the description oiEchites
holosericea and has a letter "N "on the label it is considered here to be the
holotype of £. holosericea.
In 1936, while the Sesse and Mogino collections were on loan to F (McVaugh
1 980), the specimen disscussed above was identified by Woodson as Mandevilla
sertuligera Woodson. This identification was later confirmed by the present
author. With the presentation of the above type it becomes evident that
the of Sesse and Mogiiio name has priority over Woodson's. A new combi-
nation is thus required and is accordingly made above.
The closest relative o'l Mandevilla holosericea is believed to be AI. syrinx
Woodson. The two species appear to have a close resemblance to one an-
other. Indeed, there are two sheets of the same collection {Pringle 13106,
US) that Woodson annotated differently, one sheet he annotated AI. sertuligera
and the other AI syrinx. Woodson (1938) keyed M. sertuligera apart from
AI syrinx by its corymbose inflorescence (vs. racemose) and sepals 4—6 mm
long (vs. 2-3 mm). Both Pringle specimens key more favorably to AI. sertuligera
(now AI. holosericea). At present I have only observed paratypes of AI, syrinx
(having been unsuccessful at locating the type at MO), I am inclined, how-
ever, to recognize the two species as undifferentiated and regard them both
as a synonym of AI holosericea.
REFERENCES
Fi]i-ini;s, J. and P.A. Fryxell. 1993. Types of names of New World Malvaceae of Lagasca,
Ortega and Sesse and jVIo^ino. Taxon 42:345—354.
JMcVauch, R. 1980. Botanical results of the Sesse and Mogifio expedition (1787-1803).
II. Tlie icones florae iVIexicanae. Contr. Univ. Michigan Herb. 14:99—140.
SESSfi, M. and J.M. MogiNO 1887. Flora Mexicana. Mexico.
1894. Flora Mexicana, ed. 2. Mexico.
Woodson, R.E., Jr. 1938. Apocynaceae. In: N.L. Britton et al., eds. North American flora.
New York Botanical Garden, New York. 29:103-192.
240 SiDA 1S(1)
BOOKS RECEIVED
Fautin, Daphne Gail, Douglas J. Futuyma, and Frances C. James, eds.
1997. Annual Review of Ecology and Systematics. Volume 28.
(ISBN {)-cS243-l427-l, hbk). Annual Reviews Int., 4139 El Camino
Way, P.O. Box 10139, Palo Alto, CA 94303-0139. $60.00. 712 pp.
Volume 28 is the lisluiI variety ot papers reviewing the tields oi ecology and systemarics.
The following list ot 25 articles reflects the diversity ol contributions in this 1997 Annual
Review. (A)ntents: Molecular population genetics ot social insects. Evolution ol eusociality
in termites. Evolutionary genetics and genetic variation of haplodiploids and x-linked genes.
Dissecting global diversity patterns: Examples from the Ordovician Radiation. A com-
parison of alternative strategies for estimating gene flow from genetic markers. The evolu-
tion of morphological diversity. Insect mouthparts: Ascertammg the paleobiology of in-
sect feeding strategies. Haldane's Rule. Echinoderm larve and phylogeny. Preserving the
information content of species: Genetic diversity, phylogeny, and conservation worth. Theoretical
and empirical examination of density-de|iendent selection. Toward an integrarion of land-
scape and lood web ecology: The dynamics of spatially stibsidized food webs. Settlement of
marine organisms in flow. Species richness of parasite assemblages: Evolution and patterns.
Hybrid origins of phmt species. Evolutionary genetics of life cycles. Phylogeny estimation
and hypothesis testing using maximum likelihood. Species ttirnover and the regulation of
trophic structure, Extinction vulnerability and selectivity: Combining ecological and pa-
leontological views. Tree-grass interactions in savannas. Plant competition tmdergrotmd,
IMale and female alternative reproductive behaviors in fishes: A new approach using inter-
sextial dynamics. The role of hybridizcuion and introgression in the diversification of ani-
mals. The ecology of interfaces: Riparian zones. Allometry for sexual size dimorphism: Pattern
and process in the coevolution of body size in males and females.
The usLial indexes follow: Subject Intlex, Cumulative Index of Contribtiting Atithors,
and a Cumulative Index of (Chapter Titles, Volimies 24-2H.
Janick, Jules, ed. 1997. Plant Breeding Reviews. Volume I 5. (ISBN 0-
471-18904-9, hbk). John Wiley & Sons, Inc., 605 Third Avenue,
New York, NY 10158-0012. $145.00. 395 pp.
This volume of Plant [breeding Reviews is eletlicatetl to Arnel R. Ilallauer, scientist, maize
breeder, .uul c]uantitative geneticist. The contents start with a dei-lication to Dr. Hallauer:
1 ) Detlit at ion: Arnel 11. Hallatier, Scientist, Maize Breeder, Quantitative geneticist. The
remaining eight chapters cover a variety ol areas in plant breeding. Contents: 2) Molecules
involved in self-incompatibility in flowering plants. 3) Genetic mosaics and plant improvement.
4) Quaiitirative trait loci: Separating, pyramiding, antl cloning. 5) Doubleil haploid breeding
in cereals. 6) Spelt: Agronomy, genetics, and breeding. 7) Cowpea breeding. 8) Recurrent
selection in soybean. 9) Gene action and plant breeding. All of this followed by a Stibject
Index, Ctmiulative Subject Index and a Cumulative Contributor Index for volumes 1-15.
Sii:)A 18(1): 240. 1998
TRACHYPOGON MAYAENSIS (POACEAE:
ANDROPOGONEAE): A NEW SPECIES
FROM BELIZE
JOSEPH K. WIPFF
Pure-Seed Testing, Inc.
P.O. Box 449
Hubbard, OR 97032, U.S.A.
joseph@turf-seed. com
STANLEY D. JONES
Herbarium (BRCH)
Botanical Research Center
R 0. Box 67 17
Bryan. TX 77805-6717. U.S.A.
sdjones@startel. net
ABSTRACT
Trachypogon mayaensis, a new species from the Maya Mountains in Belize, is described
and illustrated. A key is provided to separate it from other species in Central America.
Trachypogo?! mayaernis is readily distinguished from other Central American species by its
annual life cycle, height, leaf blade length and width, and prominent and conspicuous sheath
auricles/ligules, that can reach to 6 cm in length.
RESUMEN
Se describe y ilustra una especie nueva, Trachypogoii mayaensis., de las montaiias Maya de
Belize. Se presenta una clave para separarla de las otras especies Centro-americanas. T. mayaensis
se distingue facilmente de ortras especies Centro-americanas por su ciclo de vida anual,
altura, longitud y anchura de las laminas foliares, sus Ifgulas con aurfculas prominentes y
cnnspicuas, y que pueden Uegar hasta 6 cm de largo.
Trachypogon, a genus of approximately six species, is found in tropical and
subtropical America and Africa (Davila 1994). Two species are found in Central
America: T. spicatus (C. von Linne) K.E.O. Kuntze and T. vestitus N. Andersson.
Trachypogon spicatus [syns. = T. montufari (K. Kunth) C. Nees von Esenbeck;
Trachypogon palmeri Nash; T . plumosus (F. von Humboldt & A. Bonpland ex
C. von WiUdenow) C. Nees von Esenbeck; and T. secimdus (J. Presl) F. Lamson-
Scribner} is found from southern Texas in the United States to Argentina,
and in Africa. Trachypogon vestitus is found from Honduras to Brazil. A col-
lection o'^ Trachypogon was made in the Maya Mountains in Belize that could
not be assigned to any known species and is here described as new.
SiDA 18(1): 241-245. 1998
242 SiDA 18(1)
Trachypogon mayaensis Wipff & S.D.Jones, sp. nov. (Fig. 1). 'r> pus: BELIZE.
Cayo District: 1 .7 mi SE from the c-nrrancc gate inside Mountain F^ine Ridge (SSW
of Belmopan) in the JVIaya Mottntains, disturbed moimtain pine forest, associates:
Pitii/s, Andropo^i^on, Paniciim, Axonopiis, Sderici, D'uid'ici, and Aristula\ occasional, L9 Jan
1991, S.D.Jones 10489 & R. Oliver (holotype: MICH; isotypps: BRCH, MC), US).
Ciramen anntitun caespitostmi, culmis ( I .59— )1 .95—2.10 m altis; auriculis (3— )4— 5(— 6)
cm longis (atirictilae ligulae adnatae); tolia lamiiiis (sC— )43-''il cm longis; inflorescentia
9—30 cm longa.
Plants (1.59— )1. 95— 2.10 ni rail, robust annual, densely cespitose, erect.
heaves cauline (measurements taken from mid-culm leaves); nodes ca. II,
appressed pubescent with trichomcs to 2 mm long, rooting at lower nodes;
sheaths with ascending to appressed scattered trichomes or glabrous, when
pubescent, trichomes 1.3—5.5 mm long, becoming denser at apex of sheath;
collar glabrous; sheath auricles (3— )4— 5(— 6) cm long, erect and adnate to the
ligules, appressed pubescent or glabrous; ligules (3— )4-5(-6) cm long, mem-
branous, firm, brown, veined, appressed pubescent attenuate, adnate to the
sheath auricles; blades (30— )43— 61 cm long, 6—7.5 mm wide (measurements
taken from mid-culm leaves), flat, apically long attenuate and basally long
cuneate with the basal portion becoming involute, antrorsely scaberulous
on both surfaces, margins antrorsely scabrous. Inflorescence 9—30 cm long, a
spicate raceme or a panicle with two racemose branches; central axis of spi-
cate raceme or the racemose branches (if a panicle) short pubescent, intern-
odes 3—4 rnm long (in middle); one pedicellate spikelet terminating spi-
cate raceme or branch. Spikelets paired, one (lower spikelet) of each pair short
pedicellate, persistent, awnless, staminate, dorsally compressed; other spikelet
(upper sj:»ikelet) of pair, longer-pedicellate, deciduous, perfect-flowered, awned,
nearly terete. Florets without paleas. LOWER (SHORT PEDICELLED)
SPIKELETS: staminate, sometimes with a vestigial ovary present; the lower
spikelet either absent or rudimentary in the lowest 4—5 pairs of spikelets in
the inflorescence or racemose branch. Pedicels 1—2 mm long, pubescent. Spikelets
6.5-7.6 mm long, 1 — 1.6 mm wide, pubescent] first glumes 6.5-7.6 mm long,
1—1.6 mm wide, 9— 11 -veined, coriaceous, partially enclosing rest of spike-
let, narrowly elliptic, sparsely to densely short pubescent, keeled, the keels
strigose-ciliate, apex bifid, the teeth 0.2—0.3 mm long; second glumes 6.5—
7.6 mm long, ca. 1.4 mm wide, 3-veined, narrowly elliptic, the margins
overlapping, ciliate; Lower Floret: lenmias 5.2—5.5 mm long; O-veined, hya-
line, ciliate on margins; /w/(?i:?j absent. Upper Floret: lemmas 4.7—4.9 mm long;
3-veined, hyaline, ciliate on upper margins; paleas absenr. UPPER (LONG
PEDICELLED) SPIKELETS: perfect flowered, awned. Pedicels 2.5-3.1 mm
long, pubescent. Spikelets (including callus) 9-5—10.6 mm long, 1-1.4 mm
wide, pubescent; callus., at base of spikelet, 1.8—2.0 mm long, pubescent
with trichomes to 3.0 mm \ong\ first glumes 7.5—8.5 mm long, 1—1.4 wide,
WiPl-F AND Jones, A new species of Trachypoyon from Belize
243
Fic. 1. Trachypogon mayaensis [S.D.Jones !04i^9 & R. Oliver (BRCH)}. A. Habir (bar equals
5 cm). B. Section of culm showing the auricle/ligule (bar equals 1 cm).
244 SiDA 18(1)
9— 11-veined, coriaceous, margins involute and partially enclosing rest of
spikelet, elliptic, densely short pubescent throughout, lateral veins near apex
strigulose, apex rounded and ciliolate; second glumes 7.7-8.9 mm long, 0.9—
1.3 rnm wide, 3-veined, coriaceous, narrowly elliptic, sparsely pubescent
between veins; upper margins overlapping and ciliate, apex broadly acute
and ciliate; Lower Floret: neuter (i.e. without reproductive structures); lem-
mas 6.5—7.3 nim long, 1.1—1.2 mm wide, 2-veined, hyaline, ciliate on up-
per margins, apex truncate and c\\\?iX.t\ paleas absent. Upper Floret: perfect;
lemmas 6—6.9 tnm long, 0.7—1.0 mm wide; glabrous, the lower 1/3 is 3-
veined, hyaline and easily tearing, the upper 2/3 is 0-veined, subcoriaceous
to coriaceous and flattened, turning into a terete awn; awn 38—50.5 mm
long, twice geniculate, pubescent from base to second h^t^c^di; paleas absent.
Stamens 3, anthers ca. 3.8 mm long, 1-1.1 mm wide. Chromosome number
unknown.
Etymology. — The specific epithet refers to the Maya Mountain Range in Belize.
Phenology. — November-February.
Distribution. — Known from the Maya Mountains in the Cayo District of
Belize and from the state of Oaxaca in Mexico.
Additional specimens examined: BELIZE. Cayo District: Rio Privacion, JMountain Pine
Ridge, 26 Feb 193 1 , H.R Bartktt 1 1 785 (MICH). MEXICO. Oaxaca: District of Tuxtepe;
Chiltepec and vicinity, alt. ca. 200 m, 15 Nov 1941, G. Martinez Calderon 812 (MICH).
KEY TO THE SPECIES OE TRACHYPOGON IN CENTRAL AMERICA
(Modified from Davila 1994)
1. Plants robust annuals, 1.6—2.10 m tall; auricies/liguies (3— )4— 6 cm long;
leaf blades (30— )43— 61 cm long, 6—7.5 mm wide T. mayaensis
1. Plants perennial, less than 1.5 m tall; auricles/liguies 2(-5)cm long or less;
leal blades less than 30 cm long, 1—5 mm wide 2
2. Plants glabrous or rarely with basal sheath sparsely pubescent; inflores-
cence a spicate raceme, rarely a panicle with two racemose branches T. spicatus
2. Plants with sheaths and blades conspicuously pubescent; inflorescence a
panicle with 3 (rarely 2) racemose branches or rarely a spicate raceme
T. vestitus
The closest relative of Trachypogon mayaensis is probably T . spicatus., from
which it differs by the characters given in the above key; Trachypogon palmeri
Nash ( = 'r. spicatus) is the name given to longer liguled forms of T. spicatus
in Mexico. There is also a taxon in Brazil with long ligules, T. macroglossus
Trinius, but this taxon is perennial with very narrow, involute leaf blades.
Seeds of T. mayaensis were planted in the greenhouse, of the five plants to
germinate, only one survived the transplanting to a larger container. Once
the plant flowered, it began to branch at the lower aerial nodes (typical of
annuals), but eventually the entire plant died and there were no new tillers
produced from rootstock. This was also observed in the field. The original
WiPFF AND Jones, A new species of Trachypogon from Belize 245
material was collected in November 1994, but when we returned to the
same site in June 1995, there was no sign of this taxon, though the other
associated perennial grasses were still present.
ACKNOWLEDGMENTS
We thank Paul A. Fryxell (TEX) and Tony A. Reznicek (MICH) for their
assistance with the Latin diagnosis and review of the manuscript; Paul A.
Fryxell (TEX) for providing the Spanish translation of the abstract and Tony
A. Reznicek (iVIICH) for loaning specimens oi Trachypogon; and Guy L. Nesom
(SHST) thank for his initial review of the Latin description. We would also
like to thank Kelly Allred (NMSU) Gerrit Davidse (MO), WE. Fox, III
(formerly at TAES), Gretchen D.Jones (USDA, AWPMRU) and Paul Peterson
(US) for their review and suggestions. We would like to acknowledge the
generosity of the late Royce L. Oliver (BRIT) for making the collecting trip
to Belize possible; his generosity is greatly appreciated. We would also like
to acknowledge Ben Shaw for the illustration.
REFERENCES
DAvii.A, P. 1994. Trachypogon Nees. In. G. Davidsc, M. SousaS., and A.O. Chacer, etls. Flora
Mesoamericana, Vol. 6, Alismataceae a Cyperaccac. Universidad National Autonoma de
Mexico, Mexico City, Mexico. Pp. 3iSO-381.
246 Si DA 1H(1)
BOOKS RECEIVED
Holm, Lhroy, Ji.rry Doll, Eric Holm, Juan Pancho, and James Hbrberger.
1997. World Weeds: Natural Histories and Distribution. (ISBN
0-471-04701-5, cloth). John Wiley & Sons, Inc., 605 Third Avenue,
New York, NY 10158-0012, U.S.A. $195.00. 1 129 pp. Illustrated.
Tlie LiLitliors have jiresenced most ot rhc known biology of each species, maps oi their dis-
tribtition in more than 100 cotmtries, illustrations designed to lielp witli identiHcation,
and an index of common names, and a bibli()gra|-ihy of 3300 references to lead students
and researcliers ro further details that are contamed in the jTrmcipal papers. From the cover:
"The culmination of four decades of gloloal research, World Weeds jiresents comprehen-
sive and Lip-to-date information on over 100 weeds-addressing recent changes in such ar-
eas as cro|T tillage methods, herbicide Lise, and agricultural runoff. This monumental work,
featLiring a wealth of original data from the authors, provides extensive coverage of the
know biology of each species. Each entry contains a full botanical description, plus impor-
tant details on iiabitat recjuiremenrs and distrif^ution, seed production, ecology, physiol-
ogy, croji impact, and more." It is generously supplied with excellent line drawings.
Hodgson, Geoifrey M. 1995. Economics and Biology. (The International
Library ol Critical Writings in Economics). (ISBN 1-85898-050-X,
hbk). Edward Elgar Publishing Limited, England and Edward Elgar
Company, Old Post Road, Brookfield, VT 05036, LJ.S.A. $189-95.
606 pp.
The International Library of Critical Writings in Economics series is an essential reference
soLUce for students, researchers and lecturers in economics. Economics and Biology is number
50 in the series. There are six parts each divided into several chapters. Part 1: Biological
and Mechanical Analogies. Part II: Economics and Sociobiology. Part III: Classical Eco-
nomics aiul the Darwinian Revolution. Part IV: Alfred Marshall and Economic Biology.
Part V: Evolution, Optimization antl Rationality, j-'art Vi: Biology and Modern Econom-
ics. A name index follows.
SiDA lcS(l): 246. 1998
TAXONOMY OF THE SPOROBOLUS FLORIDANUS
COMPLEX (POACEAE: SPOROBOLINAE)
ALAN S. WEAKLEY
The Nat//re Conservancy
P.O. Box 2267
Chal)dWll, NC 27313-2267, U.S.A.
PAUL M. PETERSON
Department of Botany
National Museum of Natural History
Smithsonian Institution
Washington, DC 203(^0-0166, U.S.A.
ABSTRACT
The Sporoboliisfloridamn complex is defined to include five North American species. Keys,
descriptions, distributions, illustrations, and habitat information are provided for: S. curtissii,
S. floridaniis. S. silveanus. S. teretifolius. and a new member of the complex from North Caro-
lina, South Carolina, and eastern Georgia, Sporoholiis pinetoruvi Weakley and P. M. Peterson.
A lectotype is chosen fori', jlrnidantn. The taxonomic and ecological relationships ol these
five species, as well as the related S. juiiceus and S, beterokpis, are compared and discussed.
Spomlmliii cnrtisiii, S. floruUinii.s. S. pinetorum, S. silveanus, and S. teretifolius are all relatively
narrow endemics of various portions of the southeastern Coastal Plain. Each ot these five
species is the locally dominant or codominant grass in fire-maintained pinelands with open
canopies q{ Pirius palustris, P. serotina. and/or i^ elliottii y3.r:. elliottii. The geographic distri-
butions and many ecological requirements of the 5. curtissii. S. floridarius. S. pinetor//ni, and
S. teretifolius overlap, but they can be separated on a hydrologic gradient.
RESUMEN
El complejo Sporoholus floridanus es definido para incluir cinco especies de Norte America.
Se proporcionan claves, dcscripciones, distribucion, ilustraciones, e inlormacion del iiabitat
para: S. curtissii, S. floridanus, S. silveanus, S. teretifolius, y un nuevo miembro del complejo
de Carolina del Norte, Carolina del Sur y del este de Georgia, Sporobolus pinetorum Weakley
& P.M. Peterson. Un lectotipo es escogido pzitd^S. floridanus. Eas relaciones taxoncjmicas y
ecologicas de estas cinco esjiecies, asi como de las especies alines S. junceus j S. heterolepis,
son comparadas y discutidas, Sporobolus curtissii, S. floridanus. S. pinetorum. S. silveanus, y S.
teretifolius son todas relativamente endemicas restringidas a varias partes del sureste de la
llanura costera. C^ada una de las cinco especies, es el pasto localmente dominante o codominate
en terrenos de i^inos mantenidcjs por quemas con dosel abierto de Pinus palustris, P. serotina,
y/o P. elliottii var. elliottii. La distribucion geografica y muchos de los requerimientos ecologicos
de S. curtissii, S. floridanus, S. pinetorum, y S. teretifolius se solapan, pero pueden ser separados
en base a un gradiente hidrologico.
Sporobolus R. Br. is a worldwide genus of approximately 160 species oc-
curring in the tropics, subtropics, and warm temperate regions (Clayton &
SiDA 18(1): 247-270. 1998
248 SiDA 18(1)
Renvoize I 986). In the New World the genus is represented by approxi-
mately 45 species that generally occur on disturbed habitats, i.e., roadside
to open prairies and savannas (Peterson et al. 1995, 1997; Peterson et al.,
in press). Sporohoh/s is characterized by having spikelets with one floret, 1-
veined lemmas, fruits with free pericarps, and ligules with a line of hairs.
These characteristics also are found in two other genera, Calamovilfa (A.
Gray) Hack, and Crypsis W.T. Alton. These three genera seem to share a
common ancestor and are the only New World members included in the
subtribe Sporobolinae (Chloridoideae; Eragrostideae).
The species oiSporobolus occurring in the southeastern United States form
a heterogeneous assemblage that may be informally divided into groups based
on characteristics of the inflorescence, spikelet, plant longevity, and gen-
eral aspect. One of these groups, characterized as long-lived, perennial, clump-
forming species, with open panicle inflorescences and relatively large spikelets,
consists oiS.floriclani/s Chapman, S. airthsit (Vasey ex Beal) Small ex Scribner,
S. teretijolius Harper. S. silveanus Swallen, S. pinetorum Weakley & P.M. Peterson
(a new species described here), and, more peripherally, S. junceus (P. Beauvois)
Kunth and S. heterolepis (A. Gray) A. Gray. In the course of conducting con-
servation, taxonomic, ecological, and herbarium studies in the southeast-
ern United States, it has become apparent that this group is poorly under-
stood and has been much confused by earlier authors, current collectors,
and field workers. As a consequence, an overlooked species has remained
unnamed. We will try to clarify this situation with a detailed discussion of
the five members of what we call the S. flortdaniis complex {S. flondanus, S.
curttssit. S. teretijoln/s. S. silvean//s, and S. pinetoriini), with less detailed dis-
cussion of .S. junceus and S. heterolepis.
TAXONOMIC HISTORY
Because of the general similarities of the taxa in the Sporobolus jloruhvius
complex, there has been widespread confusion about their circumscription
and distribution, and much erroneous information can be found in earlier
systematic treatments and in the ecological literature, and numerous
herbarium specimens are misidentified. A review of previous treatments of
the complex and its closest relatives will help define problems resolved in
this paper.
The two morphologically peripheral taxa were the first to be named.
The species currently known as S. junceus was described by Beauvois in 1812
in Heleochloa, and transferred to Sporobolus by Kunth in 1 829- Sporobolus heterolepis
was first described as Vilfa heterolepis by A. Gray in 1835, and transferred
by Gray to Sporobolus in 1848.
Weakley and Peterson, Sporobolus floridanus complex 249
The first of the core members of the S. floridanus complex was named by
Chapman (I860) as S. floridanus. It is fairly well characterized in his de-
scription:
"S. floridanus, n. sp. Panicle clilhise, large; spikclets (purplish) on long hair-like stalks;
glumes acute, the lower one barely shorter than the obtuse paleae, the upper a third longer;
leaves rather rigid, flat, pungent, very rough on the edges— Low pine barrens. Middle and
West Florida. September.— Ctilm 2—4 feet high. Leaves 1—2 feet long. Panicle 1 — 1.5 feet
long" (Chapman I860, 1883, 1897).
The description, the location in Florida, and specimens collected by Chapman
at the time make the identity of ^. floridanus clear. The only species occur-
ring in Florida likely to be confused with S . floridanus would be S. curtissii,
but it is contradicated by various parts of Chapman's description, especially
the "panicle ... large," the markedly unequal glumes, the rough-edged leaves,
and the length of the blades and the panicle. No type specimen was cited,
however, and we lectotypify S. floridanus below.
The next to be named was S. curtissit, as S. floridanus var. curtissii (Beal
1896). A year later, Lamson-Scribner elevated the taxon to specific rank (Lamson-
Scribner 1897). The distribution of 5". floridanus was given as "moist pine
barrens near the coast, North Carolina to western Florida," apparently on
the basis of G. McCarthy's specimen at US (cited below) from Wilmington,
North Carolina, which is actually S. pinetorum. We know of no specimens of
S. floridanus from locations north of southern South Carolina.
Harper (1901) made a characteristically idiosyncratic addition to infor-
mation on the group with his collection of i', floridanus in Sumter Co., Georgia,
about which he commented "not definitely known outside of Florida be-
fore." He elaborated on the toughness of the leaves mentioned by Chapman
and stated "I made some tests of their strength. A leaf from the first collec-
tion (no. 547) 5.5 mm. wide, not twisted, and perfectly dry, sustained a
weight of 27 pounds without breaking."
In 1906, Harper described S. teretifolius from collections in Georgia, com-
menting that it is "a frequent and characteristic inhabitant of moist pine-
barrens in the Altamaha Grit region," and that "it is unmistakable when
seen in the field" (Harper 1906). He provides excellent and clear character
differences between S. teretifolius, S. curtissii, and S. floridanus, as well as an
excellent illustration of the highly distinctive leaf cross-section.
The first two editions of J.K. Small's flora (1903, 1913) treated S. curtissii
and S. floridanus, and gave generally accurate information about morphol-
ogy, habitat, and distribution. Sporobolus floridanus was stated as occurring
in "Georgia and northern Florida," and S. curtissii was found in "Georgia,
Florida, and Alabama." Small then added S. teretifolius (Small 1933). The
habitat of all three species was given as "moist pinelands, Coastal Plain,"
250 SiDA 18(1)
with distributions of "Fla. and Ga" for S. floridanus, "Fla. to N.C." for S.
c//rtissii, and "Cja." for ^. teretifoliiis. The extension of the distril:)ution of 5".
airtissii to North Carolina was apparently based on North (Carolina speci-
mens of S. p'Dietor/oii misidentified as S. curt'nsii. In his key, Small (1933)
distinguished the three species as follows:
Leaf-blades narrowly involute S. ttretifali/is
Leaf blades flat.
Pedicels appresscd to the branches: leaf-blades al^toiit I mm wide S. c/irtisiti
Pedicels spreading: leal-blades 2-4 mm wide S. floridanin
Although this key generally serves to distinguish the three species, it has
been, in part, responsible for continued taxonomic confusion about the three
species and a fourth, 5", ptnetornni. The blades of 5, teretijolii/s are anatomi-
cally oval, not involute; the blades of .V. cnrtissii, S. floruhiniis, and .V. phietomvi
are flat, becoming involute when dry, either during drought conditions in
the field, or as a result of drying for herbarium specimens. The narrow blades
of S. airtmii and S. pmetorum usually fold when dried in a plant press. Specimens
of .V. curtissii and S. pimtonmi often have involute blades that superficially
resemble those of 5. teretifolius. A second problem with the key involves the
width of the blade; both S. c/irtissii and S. pmetorum have blades regularly
reaching and sometimes exceeding 2 mm in width (when flat), and S. floriclaii/ts
characteristically has a leaf blade 3-10 mm wide (though the very narrow-
est blade on a plant may be as narrow as 2 mm wide). Therefore, specimens
of S. cnrtissii and the heretofore unnamed S. pmetorum have often been in-
correctly identified as S. teretifolins (by taking the first lead) or S. floridanus
(by correctly taking the second lead in the first couplet, but then incor-
rectly choosing the second lead in the second couplet by placing too much
emphasis on a blade width of ca. 2 mm).
Hitchcock (1935) treated S. fhyridanm ("low pine barrens, Georgia and
Floricfa"), S. cnrtns'i'i ("dry pine barrens. North Carolina, Georgia, and Florida"),
and S. teretifoliz/s ("moist pine barrens, Georgia"). Once again, the inclusion
of North Carohna in the distribution of .S', cnrtissii was apparently based on
miss identification of specimens of S. pmetoriini.
Blomquist's ( 1 9 IH) book on the grasses of North Carolina treated all material
of this complex in North Carolina as S. a/rtissii, but states "According to
Swallen (1941) the North Carolina plants assigned to this species may be-
long to S. floridcinns Chapm." As will be seen below, neither species has been
documented for North Carolina; instead. North Carolina is within the dis-
tribution o{S. piiietor/nii (undescribed at the time and in some ways gener-
ally intermediate in characteristics) and S. teretijolms {not collected in North
Carolina until 1 991).
Swallen (1941) added S. silveciuns to the group, based on material from
eastern Texas. Its distribution in Texas and the western (nilf Coastal Plain
Weakley and Peterson, Sporobolus floridanus complex 251
of Louisiana makes it allopatric relative to other members of the S. floridanus
complex. Swallen contrasted it with S. floridanus and S. teretifolius, yet vari-
ous characters (such as the appressed and larger spikelets) suggest a closer
relationship to S. curtissii. It also seems to show a clearer connection to 5".
heterolepis than do other members of the 5^, floridanus complex, a relation-
ship which seems especially plausible given its proximity to S. heterolepis
and its habitat preferences for barrens with prairie affinities.
Hitchcock and Chase (1950) provided the only treatment to date with
as many as four of the species in the complex: S. floridanus ("low pine bar-
rens, North Carolina to Florida"), S. curtissii ("dry pine barrens. North Carolina
to Florida"), S. teretifalius ("moist pine barrens, North Carolina and Geor-
gia"), and S. silveanus ("open woods, western Louisiana and eastern Texas").
The attribution of 5^. floridanus, S. curtissii, and S. teretifllius to North Caro-
lina are all based on misidentifications oiS. pimtorum. This is because of the
generally intermediate morphology oiS. pinetorum, and because the key was
not constructed very carefully for even the then-known members of the group.
Among the problems are that S. silveanus (which has markedly appressed
spikelets) can only be reached by following the lead for "spikelets not ap-
pressed, the branches and pedicels somewhat spreading"; S. floridanus is separated
from S. silveanus by having glumes "about equal" rather than "unequal,"
when they actually have similarly subequal first glume to second glume
ratios; S. teretifllius is separated from S. floridanus by the accurate but often
mis-interpreted "blades terete vs. blades flat or folded" character; and S.
curtissii is separated from the others by a subjective and difficult to inter-
pret couplet about pedicel length and orientation.
Radford et al. (1964), by contrast, recognized only one species {S. teretifldius)
in the complex as occurring in North Carolina and South Carolina. Unfor-
tunately, all material seen by them was actually the undescribed S. pinetorum.
Radford et al. (1968) added S. floridanus (alleged to occur in pinelands in
Lancaster County, South Carolina) and attributed S. teretifllius to "savannahs"
in thirteen counties in southern North Carolina and northern South Carolina.
Their key distinguished S. teretifllius from S. floridanus by "blades terete or
subterete" vs. "blades flat or folded." All material seen by them was actually
5, pinetorum, often with narrow folded blades so as to superficially mimic S.
teretifldius. Additionally, many of the county records are based on misidentified
specimens oiCalamovifla brevipilis (Torrey) Scribner, and likewise specimens
supposedly documenting county record distributions for Calamovifla brevipilis
are in many cases actually Sporobolus pinetorum. In truth, both Sporobolus pinetorum
and Calamovilfl brevipilis occur (or formerly occurred) in most or all counties
in southeastern North Carolina and in the adjacent counties of South Carolina.
Godfrey and Wooten's (1979) manual of southeastern United States wetland
plants treated only S. floridanus and S. curtissii, inexplicably omitting S. teretifalius
252 SioA 18(1)
(which occupies wercer habitats than S. ani/s.w/) and S. nlveanus (which oc-
curs in wetland situations in Louisiana, part of the geographic range of their
flora). Because of this, their treatment is of limited value in understanding
this group.
Brown (1993) addressed confusion between S. silveaniis, the most west-
ern species in the complex, and S. heterolepis, resulting in the deletion of ^.
heterolepis from the floras of Texas and Louisiana, and the addition of 5. silveanm
to the flora of Oklahoma.
S YSTF. M ATIC 'i' R F. ATM Ii N T
KFY TO THE SPECIFS
1. Panicle branches tlisrinctly whorled ac lower nodes S. junceus
1. Panicle branches single at lower nodes (though a Few branches may be ir-
regularly approximate in pairs or threes) 2
2. hirst ghime scaberulous, subulate above an expanded base; spikeiets gray
to nearly black; base of |ilant relatively fibrous; grain spherical; plants of
rocky barrens and prairies of physiographic provinces inland from the Coastal
Plain S. heterolepis
2. First ghime glabrous, lincar-lanceohxte to lanceolate, the base not abruptly
expainled; spikeiets purplish (fadmg tan); base of plant smooth and hard,
made up of the indurated leaf bases; grain oblong (when presenr, usually
abortive); plants primarily of pine savannas and seeps of the Coastal Plain
of North Carolina, South Carolina, Georgia, Florida, Alabama, Louisiana,
and Texas, bur exrending ftuTlier inland in southern Oklahoma and
eastern Texas 3
3. Blades terete or subterete (wiry), 0,5-1.2 mm wide, oval in cross-
section (dee|-)er than wide), sometimes irregularly channeled for por-
tions of their lengths (but lacking any flat portion), margins smooth,
distal portions often curling and twisted; pedicels witii scattered as-
cending hairs 5. S. teretifolius
3. Blades flat, O.S-IO mm wide, flat or V-shaped in cross-section (much
wider than deep), with free margins their entire length, margins scaberulous
(glabrous to scaberulous in S. ciirtisui), disral portions normally stiff
and straighr (note that the blades of the narrower-leaved species can
appear superficially wiry); pedicels without scattered ascending hairs,
either glabrous to scaLoerulous or scabrous 4
4. Lower ghime usually as long or longer than the upper glume with
lower/upper ratio averaging 0.90-1 . i 5; culms 30-80(-90) cm tall;
panicle 1 0-25 cm long; pedicels 0.5— K-H) mm long, appre.ssed, usually
shorter than the spikeler; leaves less than 30 cm long, smooth on
the margins 1. S. curtissii
4. Lower glume tisually shorter than the upper glume with a lower/tipper
ratio averaging 0.60-0.90; c ulms (30-) i5-2(){)(-250) cm tall; panicle
15-50 cm long; pedicels 1-11 mm long, spreading or appressed;
leaves mostly more rhan 30 cm long, scaberulous on the margins 5
5 . Pedicels appressed; lemmas 4.4-6.5 mm long, purple; anrhers 3.5-
5 mm long; blades 1-2.5 mm wide and bluish-green; jilants known
west of the Mississippi River 4. S. silveanus
Weakley and Peterson, Sporobolus floridanus complex 253
5 . Pedicels spreading; lemmas 3—4.3 mm long, purplish-brown; anthers
2-3.4 mm long; blades either wider than 2.5 mm, or dark green;
plants from east of the Mississippi River 6
6. Blades (2-)3-10 mm wide, bluish-green; panicles (18-)30-50
cm long, 4-15 cm wide; lower/upper glume length ratio aver-
aging 0.75-0.90; plants from southctn South Carolina, Geor-
gia, eastern Alabama, and northwestern Florida 2. S. floridanus
6. Blades L2-2(-3) mm wide, dark green; panicles 1 5-30 cm long,
2-6 cm wide; lower/upper glume length ratio averaging 0.60-
0.80; plants from eastern North Carolina, northern South Carolina,
and eastern Georgia 3. S. pinetorum
1. Sporobolus curtissii (Vasey ex Beal) Small ex Scribn., U.S.D.A. Div.
Agrostol. Bull. 7:142. 1897. (Figs. 1, j; 2). Sporobolus flonclanus wav. iurtissii
Vasey ex Beal, Grass. N. Amer. 2:290. 1896. Type: U.S.A. Florida: 1883, Curtiss
S.71. (holotype: MSC; lsotype: US-556876!).
Caespitose perennials. Culms 30-8()(-90) cm tall, erect, nodes all basal;
base diameter 1-2 mm, flattened; inrernodes glabrous. Sheaths glabrous to
appressed hairy, hairs up to 4 mm long; base shiny and endurated; margins
hyaline; summit with a tuft of hairs, hairs up to 4 mm long. Ligules 0.2-
0.6 mm long, a line of hairs. Blades 5-22(-28) cm long, 0.8-2(-2.2) mm
wide, flat to folded or involute, green, remaining green well into winter,
mostly glabrous above and below; base densely pilose on upper surface, white
to grayish hairs up to 4 mm long; margins glabrous to scaberulous. Panicles
10-25 cm long, 2-10(-13) cm wide, mostly open, contracted when imma-
ture, pyramidal to ovate; main axis glabrous to scaberulous; pulvini in axils
of primary branches glabrous or occasionally hairy; primary branches 2-
9(-10) cm long, ascending to spreading 10-80" from culm axis, not florif-
erous on lower 1/3; secondary branches mostly appressed; pedicels 0.5-4(-
8) mm long, usually shorter than spikelet, appressed, glabrous. Spikelets
3.5-6(-6.6) mm long, purphsh-brown. Glumes (2.9-)3.5-6(-6.6) mm long,
linear-lanceolate, membranous, 1 -veined, equal to subequal; ratio of lower/
upper glume length 0.90-1 . 1 5(- 1 .33); lower (2.9-)3.5-6.2 mm long, apex
acuminate; upper 3-2-6.6 mm long, apex acuminate. Lemmas 3.4—4.5 mm
long, ovate to lanceolate, membranous, 1 -veined, glabrous; apex acute. Paleas
3.4-4.5 mm long, ovate, membranous, glabrous; apex acute. Stamens 3,
anthers 1.5-2.8 mm long, yellow to purplish. Grains 1.1-1.4 mm long,
fusiform, reddish-brown.
Common name. — Curtiss' dropseed.
Distribution and habitat. — Eastern South Carolina south to central pen-
insular Florida, west to Florida Panhandle. Mesic to dry-mesic pine wood-
lands, in soils seasonally saturated at the surface or rather well-drained throughout
the year, usually under Pinus paiiistris, and sometimes also with Quercus spp.\
0-100 m. Sporobolus curtissii characteristically occurs in the following Na-
254
Si DA 1<S(1)
G
Fi(,. I. A-E. Sjmrohfil//s pn/ctoiy/iii ih-d'^.ijd on \\'t:cikU'y s.n., 31 Ali^' 1993). A. Habic. B. Lii^'iile.
C. Spikelcc. 15. Pistil and stamens. E, Blade, lateral view. F. Sporobohis teyetifoliiis {Wedkky &
Schafcile }■.!/., 16 JliI 1991). F. Blade, transverse section. G. Sporoho/us s//reai/us {Waller &
Bci/iml M 28). G. Four spikelets on a branch of the inflorescence. H, I. Sporobolin flor'nhinin
(Ciirtiis 4034}. H. .Spikelet. I. Blade, abaxial view. J. Sporohoh/s a/rtissii {Diniuiu 785 5). J.
Sjiikelet.
Wf.aki.f.y and Peti-.RSON, Sporobolus floridanus complex
255
Fig. 2. Geographic clistribLition oiSponbDliis a/rtissii {K), Sporobolus pinetnrum (B), and Sporohol//s
teret if alius (C).
tional Vegetation Classification plant associations: Vinm palustrislSerenoa repens-
Vaccinmm myrsinitesi Anstida beyrkhiana— Sporobolus curtissii Woodland; Finns
palustrhlQiiercus mcana-(2piera(S stellata/A ristula beyrkhiana-Sporobolus juncem-
Nolina georgiana Woodland; Vinus palmtrh-iPtnus elliottii var. elliottii) /Sporobolus
pmetm-um- Aster reticulatus-iSporobolus curtissii) Woodland (Weakley et al. 1998).
Flowering July to November (or less typically at other seasons if stimu-
lated by fire).
Comments. — When fertile, S. curtiss'ii is readily distinguished by the short-
pediceled, appressed spikelets, and by the large lower glume. Vegetatively,
S. curtissii is distinctive in its short leaf blades and tendency to have gla-
brotis leaf margins. Post-fire foliage tends to be narrow, stiff, and erect, while
unburned plants produce leaves that are shorter, wider, and laxer. It occu-
pies drier habitats than S. floridanus, S. teretifolins, and S. pinetorum, though
it can be found in mixed populations with all three species.
256 SiDA 18(1)
Representative specimens. FLORIDA. Alachua Co.: Gainesville, Coiiihs i.n. (GH). Baker
Co.: Okisfee Ex|>erimental I'orest, Okistee, 9 jun 1936, Wtitknis I (US). Bradford Co.:
(FLAS). Clay Co.: 7 mi NW of Midclleburg, abundant on peaty pond pme savanna, 28 Jul
1967, McDunid 9M' / (BRIT/VDB, G A). Columbia Co.: Lake City, 1 1-1 9 Jul I 'A^■^,Hash
22 1 3 (GA, GH, NCU, US). Duval Co.: moist pine barrens near Jacksonville, 6 Au^i; & 13
Nov 1 89-1, C//r//xv 3!8I (GA, GH, US). Manatee Co.: 1996, Alr/M/7/./;/ 1534 (NCU). Okaloosa
Co.: (reported in Clewell 1985). Orange Co.: Cirasmere, 23 Sep 1898, C&w/;j & Baker J 1 27
(US). Polk Co.: 1 996, McMi//a?i I 534, (NCU). Putnam Co.: E of Palatka, open field, 26
Oct 1940, S//i'i'//s 6737 (US). St. Lucie Co.: dry pme barrens, Fort Pierce, 25 June 1939,
Silve//s 4429 (DUKE). Taylor Co.: wet pine llatwoods, 1 979, GM/frey 77245 (Ci A). GEORGIA.
Berrien Co.: Ala]-)a Experimental Range, 2 mi S of Alapa, dominant (reported by Lemon
1949); Brantley Co.: moist open pinelands 1.5 mi W of Nahunta, 1953, Diniani 17036
(GH). Bryan Co.: Fort Stewart, 1996, McM/7A/// 1061. 1064. 1065 (NCU). Charlton Co.:
Camp Cornelia, 1992, Ricka- 932 (GH, US). Clinch Co.: pme barrens, Okifionokee [sic]
pocket, 8 Oct 1938, liyles 396 (DUKE). Coffee Co.: 5 mi south of Ocmulgce River, 18
May 195i, Cooley 2780 (US). Cook Co.: 2.7 mi NE of Barney on Ga. 76, 7 Nov 1993,
Sorrie 7786 (NCU). Dougherty Co.: dry sand, fossil dunes. East Albany, 30 Aug 1947,
Thorne 61 il (CiA). Echols Co.: longleaf pine-palmetto flatwoods near Ga. 94, 17 Aug
1967, Clewell 2624 (FSU). Glynn Co.: 1996, McMillan 1082. 1082a (NCU). Liberty Co.:
Fort Stewart, \996, McMillan 1068. 1074. /07_5 (NCU). LongCo.: \996, McMillan 1048.
1048a (NCLJ). Lowndes Co.: S of Melrose, 4 Sep 1902, Harjm- 1603 (GI 1, US). Tattnall
Co.: 1 996, McMillan 1081 (NCLJ). Toombs Co.: 1 3 Dec 1 995, Some & Weakley 7801 (GA).
Wayne Co.: 1 mile S ofjesup, 1 947, Duncan 7855 {GA, GH, FLAS, US). SOUTH CAROLINA.
Berkeley Co.: Francis Marion National Forest, 21 Jul 1994, Sorrie 807 5 (NCU); locally
dominant, growing in dense and obviously quite old tussocks in gummy longleaf/loblolly
flatwoods W of Tiger Corner, 30 May 1995, McMillan & K/elln/ark 10 H (NCU, USCH),
24 Jul 1996, McMillan 1 748 (CLEMS, NCU, USCH).
2. Sporobolus floridanus Chapm., Fl. South. U.S. 550. 1860. (Figs. 1, h,
i; 3). Typi:: U.S.A. FLORIDA. Franklin Co.: Apalachicola, Chapman .Lti. {L\-x:ny\Y\>\-,
here designated: NY, bar code, 00127474!; iscilhctotypr: US-998263!).
Robust caespkose perennials. Culms (40-)100-200(-250) cm tall, erect,
nodes all basal; base diameter 1.5-6 mm, flattened; internodes glabrous.
Sheaths glabrous to appressed hairy, hairs up to 5 mm long; base shiny and
endurated; margins hyaline; summit with a tuft of hairs, hairs up to 6 mm
long. Ligules 0.2-0.7 mm long, a line of hairs. Blades (l()-)25-50 cm long,
(2-)3-10 mm wide, flat to folded, pale bluish-green and yellowing in age,
mostly glabrous above and below; base often sparsely hairy, hairs up to 6
mm long; margins scaberulous. Panicles (1 8-)30-5() cm long, 4-15 cm wide,
mostly open, contracted when immature, pyramidal to ovate; main iixis scabrotis;
pulvini in axils of primary branches hairy or glabrous; primary branches 4-
15 cm long, ascending to spreading lO-yOr* from culm axis, not floriferous
on lower 1 /3; secondary branches ascending to spreading; pedicels 2-14 mm
long, usually longer than spikelet, spreading, scaberulous. Spikelets (3.7)
4-6 mm long, purplish-brown. Glumes (2.5-)2.8-5.7 mm long, linear-
lanceolate, membranous, 1-veined, subequal; ratio of lower/upper glume
length ((),60-)().75-0.90(-0.94); lower 2.5-5. 1 mm long, apex acuminate;
Weakley and Peterson, Sporobolus floridanus complex
257
Fig. 3. Geographic distribution of Sporobolus floricLniifS (A) and Sporohnlns itlvmnin (B).
Upper 3.7-5.7 mm long, apex acuminate to acute. Lemmas 3-4 mm long,
ovate to lanceolate, membranous, 1 -veined, glabrous; apex acute. Paleas 3-
4 mm long, ovate, membranous, glabrous; apex obtuse to truncate, rarely
minutely bifid. Stamens 3, anthers 2-3.1 mm long, purplish. Grains 1.7-
2 mm long, fusiform, reddish-brown.
Common name. — Florida dropseed.
Distribution and habitat. — Southeastern South Carolina south to north-
ern peninsular Florida, west to Florida Panhandle. Wet to wet-mesic pine
woodlands, in soils semi-permanently to seasonally saturated at the surface,
and even in places where water may pond for weeks, usually under Finns
elliottii var. elliottii. Pim/s palustris. or Taxodt/nn ascendens Brongn., and also
in seepage bogs, treeless swales, and depressional wetlands in pineland landscapes;
0-100 m. Sporobolus floridanus characteristically occurs in the following National
Vegetation Classification plant associations: Pinus palustris— P inns elliottii var.
elliottiilCtenium aromaticum-A ristida bey rich iana-Sporobolus floridanus Woodland ;
Pinus palustris— Pinus elliottii var. elliottiitStyrax americanus var. pulverulentusl
Sporobolus floridanus Woodland (Weakley et al. 1 998). Flowering July to November
(or less typically at other seasons if stimulated by fire).
Comments. — This is a common and conspicuous grass within its range,
258 Sum 1<S(L)
and is often a dominant to codominant component of wet pinelands. The
wide leaves with distinctly bluish cast are distinctive in this species.
Representative specimens. ALABAMA. Houston Co.: (Ciiinn, pers. tomm). FLORIDA:
Dixie Co.: Ilatwoods near Hines, 26 Aui,' 1 937, West ct uL HH-n4 (FLAS, US). Duval Co.:
moist pine barrens, near Jacksonville, Oct I 894, Ci/rtiss ]]7^ (GA, GH, NCU). Franklin
Co.: pine-turkey oak woodland [likely a wetland mclusion in or near this habitat}, 1 mile
from brid_t;e over Ochlochonee Bay, 14 Jul 1954, Fonl ]} 24 (US). Gulf C]o.: St. joe, open
pmeland, 5 Oct 1940, Silveus 649H (US). Levy Co.: Fllzen [or Ellzey], Ilatwoods, infre-
c]Lieiu, 3 Sep 1898, Coiuhs HIH (US). Liberty Co.: AiicJersun 12503 (FSU). Madison Co.:
1955, Godfrey ^4004 (DUKF, FSU, GH). Wakulla Co.: St. Marks Refu^^e, Some 7^85
(NCU). Walton Co.: de Funiak Springs, 1 898, Combs 446 (GH). GEORGIA. Baker Co.:
Jones Ecological Research Station, Ichauway Plantation, Jack Baker Woods, 17 Aug 1 995,
AlcM///i/>/. Kjellii/ark. & Dreu 1110 (USCH). Berrien Co.: Forest Experiment Station, 5
mi NE of Tilton, open pme woods, moist sandy soil, 1943, Biswell & Lemon PL-270 {¥LAS,
and reported in Lemon 1949). Brooks Co.: wet seepage area in upland pine woods, 1.4 mi
N of Morven, 6 Sep 1967, Pciirdoth & Fuircloth 4i329 (GA, NCU). Bulloch Co.: , 20 April
1994, Some 7859 (GA, NCU). Cook Co.: i mi SE of Adel, open pme forest, 27 Jun I 957,
Pohl 7636 (GH). Dooly Co.: open partially drained cypress pond, on W side of railroad,
2.7 mi SSE of Unadilla, 31 Jtil 195 i, H,/rper. H/nnphrey. & Diinani I6H39 (GA). Echols
Co.: open pine-palmetto flatwoods on the E side of the Alapaha River, 3.6 mi S of Mayday,
21 Oct 1967, fcinrloth 497H (GA, NCU). Lanier Co.: open pine woodland alongside LIS
221, 2.5 mi WSW of Lakeland, 12 Oct 1967, Pa/rc/oth 4930 (GA, NCU). Long Co.: dis-
turbed pocosin, 6.3 mi W of 301 on Ga. 26 1 , 6 Oct 1962, Bozemai/ & Radjonl 1964 (NCU).
Lowndes Co.: open longleal pine forest on level terrain 5 mi E of Valdosta, 12 Oct 1963,
Duncan 22210 (GA). Sumter Co.: moist pine barrens, 31 Aug 1900, Harper "^47 (GH).
Tattnall Co.: common in slash pine flatwoods clearing, ca, 2 mi S of Glenville, 1 I Sep
1 972, Godfrey 72107 (FSU, NCU). SOUTH CAROLINA. Jasper Co.: dominant on mesic
to wet Pi)ii/s ellioltiilP. palnstris savanna on Rains series sand}' loam with Paxville fine loamy
inclusions, 28 Jun 1995, McMillan & Kjellmark 1035 (CLEMS, NCU, USCH).
3- Sporobolus pinetorum Weakley & P.M. Peterson, sp. nov. (Figs. 1, a-
e; 2). Tyvv.: U.S.A. North Carolina. Cumberland Co.: Fort Bragg Army Base, Macridge
Danger Zone, ecotonal seepages and moist swales in uj:iland Pnii/s palustris savannas
Weakley s.n., 31 Aug 1993 (hoi.otypi:: US!; lsotypr: NCU!).
A S. fluridaui laminis foliorum 1 .2-2(-3) mm latis atrovirentibus, inflorescentisl 5-30
cm latis, culmis 45-120 cm altis, ratione longitudinus glumarum inferiorum/superiotLim
0. 60-0. 80, recedit.
Caespitose perennials. Culms (3()-)45-12()(-18()) cm tall, erect, nodes
all basal; base diameter 1-3 mm, flattened; internodes glabrous. Sheaths
glabrous to appressed hairy, hairs up to 4 mm long; base shiny and endurated;
margins hyaline; summit with a tuft of hairs, hairs up to 4 mm long. Ligules
0.2-0.6 mm long, a line of hairs. Blades 20-50 cm long, 1.2-2(-3) mm
wide, flat to folded or involute, dark green, remaining green well into win-
ter, mostly glabrous above and below; base often sparsely hairy, hairs up to
4 mm long; margins scaberulous. Panicles 15-30 cm long, 2-6 cm wide,
mostly open, contracted when immature, pyramidal to ovate; main axis
Weakley and Peterson, Sporobolus floridanus complex 259
scaberulous; pulvini in axils of primary branches hairy or glabrous; primary
branches 2—8 cm long, ascending ro spreading 0—50" from culm axis, not
floriferous on lower 1/3; secondary branches ascending to spreading; pedicels
2-22 mm long, usually longer than the spikelet, spreading, scaberulous.
Spikelets 3-5— 6.5 mm long, purplish-brown. Glumes 2.4-6.5 mm long,
linear-lanceolate, membranous, 1— veined, subequal to unequal; ratio of lower/
upper glume length (0.58-)0.60-0.80(-0.83); lower 2.4-4.5 mm long, apex
acuminate; upper (3.5-)4-6(-6.5) mm long, apex acuminate to acute. Lemmas
3.4—4.3 mm long, ovate to lanceolate, membranous, 1-veined, glabrous;
apex acute. Paleas 3.4—4.4 mm long, ovate, membranous, glabrous; apex
obtuse to truncate, often bifid with teeth up to 0.4 mm long. Stamens 3,
anthers 2.5-3.4 mm long, purplish. Grains 1.8-2.2 mm long, fusiform,
brown.
Common name. — Carolina dropseed.
Distribution and habitat. — Eastern North Carolina south to northern South
Carolina; and disjunct in eastern Georgia. Wet to moist pine woodlands, in
soils seasonally to semi-permanently saturated, usually under Pinus palustris
and Pinus serotina, also sometimes associated with Taxodium ascendens and
(in Georgia) Pinus elliottii var. dliottii. Sporobolus pinetorum characteristically
occurs in the following National Vegetation Classification plant associa-
tions: Pinus palustris-iPinus dliottii var. elliottiij/Sporobolus pinetorum-Aster
reticulatus—Sporoholus curtissii Woodland; Pinus palustns— Pinus elliottii van elliottii/
Styrax americanus var. pulverulentuslSporobolus floridanus Woodland; Pinuspalustris—
Pinus se)vtina/Sporobolus pinetorum— Aristida stricta-Eryngium integrifolium Woodland;
Pinus palustr is— Pinus serotinaiSporobolus pinetorum— Ctenium aromaticum—Eriocaulon
decangulare var. decangulare Woodland; Pinus elliottii var. elliottii— Taxodium
ascendens /Hypericum brachyphyllum / Sporobolus pinetorum— Dichanthelium scabriusculum
Woodland (Weakley et al. 1998). July to November (or less typically at
other seasons if stimulated by fire).
Comments. — This species has been overlooked until now becatise its over-
all morphology places it centrally in the complex, allowing it to be vari-
ously confounded with S. floridanus. S. curtissii, and .V. teretifolius. Despite
its "intermediate gestalt," it has a unique combination of characters that
warrant taxonomic recognition. Moreover, it occurs in mixed populations
with S. floridanus (in eastern Georgia), 5'. curtissii (in eastern Georgia), and
S. teretifolius (in southeastern North Carolina), and in these situations the
taxa are easily distinguishable and show no signs of intermediacy or hy-
bridization. Sporobolus pinetorum is locally abundant in seasonally saturated
pinelands within its range; because it does not typically flower except fol-
lowing fire, casual field observers have often overlooked its presence and
misidentified it as Aristida stricta.
260 SiDA 18(1)
Specimens examined. GEORGIA. Chatham Co.: pineland, Ogeechee Canal, 4 Aug 1 939,
Eylei 6447 (US). Liberty Co.: in ecotone oi: Piii/ts pcih/strislAristichi htyrichicoiii flatwood.s
and Taxiuiiiim cticcnckns tlrain, 1995, h\ch\'ilhin 1967 (NCU); codominanr wirli SlMmiholiis
ciirtisiii in mesic longleat pine/saw palmetto tlatwoods on Fort Stewart Military Reserva-
tion sec. A-5, 4 Sep 1995, McMillan. Kjelhiuirk, & ThoiiipMn 1 171 (USCH); Long Co.: in-
frec]Lient to rare on mesic longleaf pine savanna and pond cypress depression ecotone on
Leesfield series soil transition, I'ort Stewart Military Reservation on RSPAC in sect. D-12,
4 Sep 1995, /VlcAI///rf«, Kjel/mark, & Thow/iwi/ 1 1 70 (USCH). NORTH CAROLINA. Bladen
Co.: dry savanna 10 mi N of White Lake, 8 Oct 1 944, Blomq/tist / j6/ 7 (DUKE); flat pine
woods, 6.4 mi ESE of Ammon on road to Garland, 19 Oct 1957, Ahhs 37439 (NCIU).
Brunswick Co.: low burned savanna on W side ol road to Ec. Caswell, near junction ol
Sawdust Trail, 1938, Bloiiiquist 1044H (CtH, US); burnt longleaf pine-live oak area near
inland waterway bridge. Long Beach road, 29 Oct 1950, Boyce & Goilfrey 1602 (NCSC);
Southport-Supply, savanna, 23 Aug 1930, Bloniqiiht 97 (DUKE, PH); Southport-Supply,
savanna, 31 Aug 1931, Blomquist 667 2 (YyDKXL, US); Pireway, savanna, 4 Aug \'433,Scbiilleyt
s.n. (US); shrubby area in pine barrens, Boiling Springs Lakes subdivision, about 2 mi N ot
jet. of NC 87 & 133, off rte. 133 (N of Southport), 24 Oct 1962, Temll & Smith 3697
(NCU); plowed fire lane through a bog, Orton, 28 Sep 1941, Godfrey 10053 (NCU). Co-
lumbus Co.: cut over pine savanna to E ol Co. road, 2.6 mi straight NW of Nakissa [sic;
probably Nakina], 17 Oct 1958, Bell 13739 (NCU); Schulkens Savanna, 1.7 mi W of Old
Dock, 17 Jul 1991, Weakley, Schafale. & LeBloml s.n. (NCU, US). Cumberland Co.: Fort
Bragg Army Base, Macridge Danger Zone, ecotonal seepages and moist swales in upland
Pinus palustris savannas, 31 Aug 1993, Weakley s.n. (US, NCTJ). Duplin Co.: transition
near KenansviUe, 14 May 1925, n. c. (NCSC); burned savanna between Clinton and Seven
Springs, 7 Jul 1946, Blomquist 13936 (DUKE); field woodland border, 0.3 mile NW of
Pin Hook, 2 Aug 1957 , Ahles & Leisner 33 173 (NCU). Greene Co.: savanna 1.1 mi SE of
Jason, \95H, Radford 40402 (FLAS). HarnettCo.: l6Nov I992,.S'«me 7/ /6 (NCU). Hoke
Co.: 7 Oct [991, Sorrie 5945 (NCU); 9 Oct 1991, Some 5959 (NCU); 25 Nov 1 991, Sorrie
6101 (NCU); 28 Sep 1992, Sorrk 7007 (NCU). Jones Co.: savanna, 1 .2 mi SSW of Pleas-
ant Hill, 9 Sep 1958, Radford 39935 (NCU); savanna, 3 mi SE of Pink Hill, 9 Sep 1958,
Radford 39838 (NCU); savanna, 4.3 mi WNW of Hargetts Store, 1 8 Jul 1958, Radford
36898, (GA, GH, NCU). Lenoir Co.: pine forest, near NC 1 1 , 1 mile N of Pink Hill, 24
Oct 1957, Radford 31655 (NCU). New Hanover Co.: Wilmington, 1885, McCarthy s.n.
(US). Onslow Co.: Folkstone, 3 Aug 1939, S/lvens 4856 (US); savanna, 9.6 mi N of HoUyridge,
27 Jul 1957, Ahles & Leisner 32690 (NCU); Camp Lejeune, 23 Sep 1990, Sorr/e. Weakley.
LeBlond 5282 (GH); , 5 Sep 199 1 , Churchill 91-170 (BRIT/VDB); cut-over savanna about
10 mi NW of Holly Ridge toward Maple Hill, common, 1 6 Aug 1 967, Wilh/ir9432 (DUKE);
WsideofUS 1 7, 0,6 mi S of fire tower, Jun 1991, Weakley & Peel s.n. (\]S). Vender Co.: 30
Aug. 1991 , Some & LeBlond 5889 (GH, NCU). Richmond Co.: seep in powerline west of
NC 177, 1 2 Sep 1993, Some 7679 (NCU); Sandhills Game Land, 28 Oct \997 , Sorrie 9621
(GA, GH, NCU, US). Robeson Co.: woodland border, 2 mi N of Allenton, 2 1 Jun 1957,
Ahles & Haeslooji 29011 (NCLJ). Sampson Co.: pme savanna, 1.3 mi SE of junction US
42 1 and 70 1 on US 42 1 (SSE of Clinton), 8 Aug 1951 , Ahles &Lei.uier 33702 (NCU). Scotland
Co.: 8.3 mi SSE of Eloffman, 1 1 Oct 1959, McNeely 916 (NCU); 4 Sep 1994, Sorrie 8187
(NCU). [no Co.]: in Oriente Carolina Septentrionalis, locis paludosis [in eastern North
Carolina, boggy places], Aug \^^5, McCarthy s.n. (NCU, PH, US). SOUTH CAROLINA.
Berkeley Co.: Jun 1 998, Peet & McMillan 684 1 (NCU). Chesterfield Co.: cleared, burned-
over shrub bog m the Sandhills 1 mile W of McBee, 7 Sep 1939, Godfrey 8052 (DUKE,
GH, NCSC, PH, US); savanna, Montrose near US 52, 29 Sep 1956, Radford 18(^(-y7 (NCU);
Whakley and Peterson, Sporobolus floridanus complex 261
sandhill, 1 .3 mi SW of" Patrick near US 1 , 29 Sep 1 956, Radford 18761 (NCU); Sandhills
National Wildlife Refuge, 16 Apr 1992, Some et al. 6246 (USCH); Sandhills National
Wildlife Refuge, 1995, Pittifian s.u. (US); Hudsoma Flat, 6 Oct 1993, ^rtrr/e 7746 (NCU).
4. Sporobolus silveanusSwallen,J. Wash. Acad. Sci. 31:350. 1941. (Figs.
1, g; 3). Type: U.S.A. TEXAS. Orange Co.: 10 mi NE of Orange on U.S. 90, 30
Sep 1 940, Silvern 644 J (holoiype: US-18 1 7963!; isotwes: US-1867557!, US-2209343!).
Densely caespitose perennials. Culms 70-1 20 cm tall, erect, nodes sometimes
visible, mostly basal; base diameter 1 .5^.5 mm, rounded or flattened; internodes
glabrous. Sheaths mostly glabrous to appressed hairy, hairs up to 4 mmi long;
base shiny and endurated; margins hyaline; summit with a tuft of hairs,
hairs up to 4 mm long. Ligules 0.2-0.8 mm long, a line of hairs. Blades
15—52 cm long, 1—2.5 mm wide, flat to folded or involute, bluish-green,
glabrous above and below; margins scaberulous. Panicles 21—50 cm long,
5-12(— 15) cm wide, open and few flowered; pyramidal to ovate; main axis
scabrous; pulvini in the axils of primary branches glabrous; primary branches
6-20 cm long, ascending and loosely spreading 20-50? from culm axis,
not floriferous on lower 1/4—1/2; secondary branches appressed to loosely
spreading; pedicels 3-8(-l4) mm long, longer or shorter than spikelet, mostly
appressed, scabrous. Spikelets 4.5-7(-7.2) mm long, purplish. Glumes 3-
7 mm long, linear-lanceolate to lanceolate, membranous, 1 -veined, subequal
to unec]ual, ratio of lower/upper glume length 0.6-0.9; lower 3—4.6 mm
long, apex acuminate; upper 4-7.2 mm long, often appearing 3-veined with
lateral folds that resemble veins; apex acuminate. Lemmas 4.4-6.5 mm long,
lanceolate, membranous, 1 -veined, glabrous; margins often hyaline; apex
acuminate to acute. Paleas 4.5-6.7 mm long, lanceolate, membranous, glabrous;
apex obtuse to truncate, minutely erose. Stamens 3, anthers 3-5-5 mm long,
purplish. Grains 1 .8—2.5 mm long, obovoid, laterally compressed, light brownish.
Common name. — Silveus dropseed.
Distribution and habitat. — Western Louisiana west to eastern Texas and
north to southeastern Oklahoma. Wet to mesic pine woodlands under P/>z//j
palmtris, also in adjoining glade and barren openings, and in blackland prairies,
5-200 m. Sporobolus silveanus characteristically occurs in the following Na-
tional Vegetation Classification plant associations: Finns palmtrislSporobolus
silveanus-IWnhlenbergia capillaris -Liatris pycnostachya var. lasiophylla Wood-
land; Sporobolus silvmniis-Carex meadit Herbaceous Vegetation; Sporobolus silveanus-
Tridens strictus Herbaceous Vegetation (Weakley et al. 1998). Flowering
July to November (or less typically at other seasons if stimulated by fire).
Comments. — There has been some confusion between this species and S.
heterolepis. Allen (1992) and Thomas and Allen (1993) reported S. silveanus
from Calcasieu Parish, Louisiana, and S. heterolepis from Allen Parish and/or
Calcasieu Parish, but all specimens are actually S. silveanus. Similarly, both
262 SiDA 18(1)
S. silvea////s and S. heterolepis have been historically reported from eastern Texas
(Correll and Johnston 1970; Gould 1975; Johnston 1990; Hatch etal. 1990).
Brown (1993) determined all material Irom Texas and Louisiana to be S.
silveanus, and reported S. silveaii/is as a state record for Oklahoma, a conclu-
sion followed by Taylor and Taylor ( 1 994) and Jones et al . ( 1 997). Sporoboli/s
silveanus has a bluish color in the field, and closely resembles MNhleiiberg/a
expansa (Poir.) Trin., with which it often grows.
Representative specimens. LOUISIANA. Allen Parish: longleaf pine woods, W of Kinder,
20 Oct 1940, Bnnvn etal. 37/7 (LSLI). Calcasieu Parish: 5 mi S of Starks, 1949, Su./lleu
105 1 1 (GH). OKLAHOMA. Bryan Co.: pnurie near Durant, 2 1 Oct 1953,.Mftr j.;/. (OKLA).
TEXAS. Angehna Co.: longleaf pine uplands ol the proposed Graham Creek Wiklerness,
8.8 mi S of Zavalla on US 69 and E on FR 314, 22 Sep 1 979, N/xon & Ward 9639 (ASTC).
Brazos Co.: along highway 6, 1 2 mi S of College Station, 8 Oct 1969, Leonard 250 (TAES);
Galveston Co.: on l4th street one block W of Ave. I (FM 517), 19 Sep 1974, Waller &
Ba/aiil M2H (SBSC, TAFS, TliX). Hardin Co.: pine forest border, 6 mi SW of Kountze,
15 Oct 1964, Gould 11028 (BRIT/SMU, TAES, TEX, US). Harris Co.: Red Bluff Road,
1.5 mi W of SH 146, NW of Seabrook, 18 Sep 1974, Waller & Ba/fwl 3 1 28 (GH, TAES,
TEX, US). Jasper Co.: SE of Zavalla on US 6i, 1 .6 mi SE of the Plum Ridge Road, 29
Aug 1978, Alar/e//a & N/xon 486 (ASTC, TEX). Lamar Co.: "Tridens Prairie" 7 mi W of
Paris at intersection of highway 82 and FR 32, Coll///s .v.//., Nov 197 1 (LL, TAES, TEX).
Newton Co.: 16 mi N of Newton, 1 1 Oct 1934, Parks & Cory 10832 (TAES). Orange
Co.: open woods about 10 mi NE ol Orange, 30 Sep I940, S/lveus 6441 (US). Rains Co.:
in line sandy clay between RR and highway, 3.5 mi NW ol Point, I 2 Sep 1948, Shiiniers
7(J2.)y (BRIT/SMU). Tyler Co.: longleaf pine grassland, 6.5 mi E of Chester on route 1745
then left 3 mi to xenc Oligocenc outcrop, 19 Oct 1967, Correll 35 172 (LL). Van Zandt
Co.: sands. Wills Point, I 5 Oct 1903, Reverchoii 3484 (US).
5. Sporobolus teretifolius R.M. Harper, Bull. Torrey Bot. Club 3.3:229.
1906. (Figs. 1, f; 2). Tyim;: U.S.A. GEORGIA: Colquitt Co.: S of Moultrie, 20
Sep 1902, Harper 1642 (molotvpi;: NY; lsotvpe: US-431954!).
Caespitose perennials. Culms (20-)35-cSO(-l()0) cm tall, erect, wiry, nodes
all basal; base diameter 1—2 mm, flattened; internodes glabrous. Sheaths
glabrous to appressed hairy, hairs up to 4 mm long; base shiny and endurated;
margms hyaline; summit with a tuft of contorted hairs, hairs up to 4 mm
long. Ligules 0.2-0.4 mm long, a line of hairs. Blades (I()-)25-54 cm long,
0.5— 1 .2 mm wide, tightly involute or terete, green to yellowish-green, senescing
or turning tan in late autumn, glabrous above and below; base often sparsely
hairy, hairs up to 3 mm long. Panicles 10-26 cm long, 1—9 cm wide, mostly
open to somewhat contracted when immature, narrowly pyramidal to ovate;
main axis scabrous; jTulvini in axils of primary branches often hairy; pri-
mary branches 1-8 cm long, ascending to spreading 0-40" from culm axis,
not floriferous on lower 1/3; pedicels 3-18 mm long, longer than spikelet,
usually spreading, with scattered ascending hairs. Spikelets 4—5.6 mm long,
purplish-brown. Glumes 2—5.6 mm long, linear-lanceolate, membranous,
1 -veined, unecjual, ratio of lower/upper glume length (0.53-)0.55-0.70(-
WiiAKLEY AND Petfrson, Spofobolus floridanus complex 263
0.11); lower 2—3.8 mm long, apex acuminate; upper 4—5.6 mm long, apex
acuminate. Lemmas 5 A— 4.4 mm long, ovate, membranous, 1 -veined, gla-
brous; apex acute. Paleas 3-3-4.4 mm long, ovate, membranous, glabrous;
apex acute. Stamens 3, anthers 1 .5-2.6 mm long, purplish. Grains not seen.
Common name. — Wireleaf dropseed.
Distribution and habitat. — Southeastern North Carolina south to south-
ern Georgia, west to extreme southeastern Alabama. Wet to moist pine
woodlands, \xx\Att'Pinus serotina. Pinus elliottii var. elliottii, and Pinus palustris,
in soils; 10—150 m. Sporobolus teretifolius characteristically occurs in the fol-
lowing National Vegetation Classification plant associations: Pinus palustris—
Pinus serotina/ Magnolia virginianalSporobolus teretifolius— Carex striata
Woodland (Weakley et al. 1998) and others not yet described. Flowering
July to November (or less typically at other seasons if stimulated by fire).
Comments. — The terete leaves (well figured by Harper 1906) are distinc-
tive. Dry blades (either on dried specimens or in nature under dry field conditions)
of the other eastern species with narrow blades (5'. curtissii. S. pinetorum) can
superficially resemble those of 5'. teretifolius, and have caused confusion. Sporobolus
curtissii, however, has much shorter blades and generally occurs in drier habitats,
and the blades of S. pinetorum have distinctly rough margins (best felt by
running ones finger along the blade towards its base, or seen at 10" or greater
magnification).
Representative specimens. ALABAMA. Houston Co.: James Hughes pitcher plant bog,
1.5-2.0 mi NE of Cottonwood, burned annually, 4 Aug 1996, MacDomild 9833 (IBE).
GEORGIA. Berrien Co.: very broad powerline on N side of route 76, about 1.7 mi SW of
Nasliville, 27 Oct 1 994, Sorrks.n. (NCU). Bulloch Co.: sloping moist pine barrens, about
7 mi NW of Statesboro, 12 Sep 1954, Harper 4309 (GA). Candler Co.: about 3.25 mi W
of Matter on Stillmore Road, ecotone on E side of a N-S swampy tributary of Sams Creek,
S side of road, 22 Apr 1994, Some & Stoireli s.n. (NCU). Coffee Co.: rather dry pine-bar-
rens near Douglas, upper Eocene overlaid by Lafayette and Columbia, 22 Sep 1900, Harper
677 (NY). Colquitt Co.: moist pine barrens near Moultrie, 20 Sep 1902, Harper 1642 (G A,
NY, EJS. Cook Co.: low seepage area (pitcher plant bog) alongside GA 76, 2.7 mi NE of
Barney, 2 1 Sep 1 965 , fainioth & O'Neal 2994 (GA). Dodge Co.: reported without precise
location by Harper (1906). Dooly Co.: reported without precise location by Harper (1906).
Emanuel Co.: moist mown margin of GA 57 just NW of milepost 5, S side of road, at
edge of shrubby ecotone of the W side of Flat Creek, NW of Swainsboro, 6 May 1994,
Sorrie s.n. (NCU). Screven Co.: E side of US 301 at South Fork Ogeechee Creek (milepost
7), S of Sylvania, 25 Oct 1994, Sorrie s.n. (NCU). Thomas Co.: Greenwood Plantation,
Thomasvillc, damp peaty soil bordering a Magnolia virginiana—Nyssa hiflora drainage, area
burned June 1985, 23 Oct 1985, Gholson, Godfrey, Komarek. & Baker 11522 (GA). Tift
Co.: Vicinity of Irby P.O. (Cycloneta Station), 28 Jul 1890, Tracy s.n. (NY) {note that present
Tift Co. was part of Irwin Co. at the time of Tracy's collection}. Toombs Co.: among a
colony of pitciier plants, 3 mi N of Lyons, 28 Jul 1961, Banks s.n. (GA). Turner Co.: seep-
age slope flanking route 32, W side of Little Sand Creek, 25 Aug 1994, Sorrie s.n. (NCU).
Wheeler Co.: moist pine barrens, 2 mi W of Alamo, Plummer sandy loam, Pullen & Pliitmner
881 (GA). NORTH CAROLINA. Brunswick Co.: Camp Branch Savanna remnant, just
N of SR 1335, ca. 1.2 mi E of its junction with SR 1334, back edge of savanna, near fire
264 Si DA 18(1)
(.lirch at c'cocone CO j-iocosiny swamj-i, K\)ul 1991, Wtakk']& Schiiliile s.ii. (US). Columbus
Co.: Old Dock Savanna, south ol SR 1928, ca. 0.9 mile W of Old Dock, then south on
logging road ca. O.t mi, 15 Jul 1991, Wt'akky & Schajalc s.n. (US). SOUTH CAROLINA.
Georgetown Co.: pine savanna in Bates Hill Plantation, ca. 4.0 km SE of the US 701/SC
261 intersection, 5 Sep 1989, 'I'iii^^ii^iirt s.ii. (USCH). Horr)' Co.: along powerline, situated
between Mose Swamp anti Grass Bay, take SC 1 09 ca. 2 mi NW oi Baker Crossroads, then
W and intersect with Sancee Cooper Power right-obway, 22 Sep 1992, Pittman & Jordan
s.n. (USCH). Kershaw Co.: mesic slope, SC 28-102, 2.8 mi N of SC 12, 29 Sep 1958,
D//^<t'2979(NCU).
RELATIONSHIP.S WIT] UN SPOROBULUS
Stapf ( 1 89<S), Bor (I960), Clayton er al. (1974), and Baaijens and Veldkamp
(1991) have suggested infrageneric classifications in Sporoholiis based pri-
marily or strictly on Old World species, and a more worldwide classifica-
tion is still lacking. Based on possession of an open or contracted panicle,
caespitose perennial lifeform, non-whorled branch insertion, intravaginal
new shoot initiation, and first gltime shorter or as long as the second Pilger
(1956) erected group four in subgenus Sjiorohoh/.s. He further divided group
four into two smaller groujis a cs: B. Sporoholiis beterolepis, S. floruhnuis. S.
teretifoluis. and S. lasiophyll/is Pilg. are members of Pilger's "B" group. We
agree with Pilger's assessment that.V, heterolepis. S. floridanus, and S. teretifoPiiis
appear to be closely related. However, S. Lisiophylh/.s lias a few unique char-
acteristics, such as its strictly basal leaves with sheath bases that are densely
lanate and thickened, and its plumbeous to dark-brownish spikelets.
Small (193.3) divided Sporoholiis of the southeastern United States into
seven groups, which are not considered validly published because their rank
wiis not indicated. He placed S. jiinceiis [iis ".V. gracilis (Trin.) Merr. "], S.florickiniis,
S. ciirtissii, and S. teretifoluis in '^Grcuiles. " distinguished by perennial, bunchgrass
habit, large (more than 3 mm long), purplish to brown spikelets, and glumes
unequal, the second as long as the floret. Sporoholiis heterolepis was not known
by Small to occur in his "flora area," so it is unknown whether he would
have placed this with ''Graciles" or not.
Wc informally recognize the S. floridaniis complex, consisting of five species
(.V. ciirtissii. S. floridaniis, S. pinetoriim, S. silveaniis, and S. teretifoliiis) with
veiy similar morphological features and generally similar ecological requirements
(primarily restricted to pine savannas on the Coastal Plain). All five species
exhibit the following characteristics: caespitose perennials; basal sheaths shiny
and endurated (apparently as "fire-proofing"); panicles open at maturity,
with ascending panicle branches, pyramidal to ovate, primary branches not
floriferoLis on lower 1/3; spikelets purplish-brown to purplish; lower glume
linear-lanceolate to lanceolate, apex acuminate; grain fusiform to obovoid.
Additionally, all five species occupy fire- main rained pinelands of the southeastern
Coastal Plain, and produce culms only following removal of foliage by fire
Weakley and PetersoiN, Sporobolus floridanus complex 265
(or rarely, by mechanical disturbance) [Weakley 1998}.
The S. floridan//s complex appears to have its closest affinities with two
pairs of species: 1. S. heterolepis and S. interriiptiis Vasey, and 2. S. jiinceiis and
S. purpurascens (Swartz) Hamilton. Sporohh/s heterolepis and S. interrupt/is ap-
pear to be sibling species, S. heterolepis being widely distributed in central
North America, and extending as an uncommon disjunct into eastern North
America, and S. interrupt //s being endemic in northern Arizona. They share
several characters which distinguish them from the S. floridanus complex:
spikelets plumbeous (vs. purplish fading to tan), grains globose (vs. elon-
gate and laterally flattened), scaberulous lower glumes (vs. glabrotis), and
fibrous sheath bases (vs. shiny and indurated). Sporobolus heterolepis appears
to be particularly closely related to S. silveanus, the most western of the S.
floridanus complex, and the one most likely to have been geographically
and ecologically in contact with S. heterolepis in recent times. Ecologically,
S. heterolepis is a species of glades, prairies, and barrens, usually overmafic,
ultramafic or calcareous rock outcrops in the interior, "hard rock" physi-
ographic provinces of central North America (extending as a rare disjunct
east to and occasionally beyond the Blue Ridge Mountams), as well as on
loess and glacial tills.
Sporobolus junceus and S. purpurascens also appear to be siblings, S. junceus
being a species primarily of the southeastern United States Coastal Plain
(from southeastern Virginia west to eastern Texas), but extending inland to
adjacent provinces, while S. purpurascens is distributed in southern Texas,
Mexico, the West Indies, and into tropical America. They share several characters
which distinguish them from the S. floridanus complex: panicle branches
distinctly whorled in well-marked verticils (vs. panicle branches alternate
or sometimes irregularly paired or approximate) and spikelets smaller (3-0-
3.8 mm long vs. 3.5-7.2 mm long).
DISTRIBUTION AND ECOLOGY
Sporobolus curtissii, S. floridanus, S. pinetorum, S. silveanus, and S. teretifolius
are all relatively narrow endemics of various portions of the southeastern
Coastal Plain (Figs. 2, 3). Each of these five species is the locally dominant
or codominant grass in fire-maintained pinelands with open canopies oiFinus
palustris, P. serotina, and/or P. elliottii var, elliottii (Weakley et al. 1998). The
geographic distributions and many ecological requirements of the four more
eastern species overlap, but they can be separated based on a hydrologic gradient.
Understanding the differing but overlapping distributions and ecological
niches of the species within the group provides an important basis for the
systematic treatment.
The five species of the complex share a set of apparent adaptations to the
266 SiDA 18(1)
fire-maintained habitats in wliich they ocx-tir. Ah have the basal sheaths thickened
and cartilaginous, tightly investing the growing tips of the rhizomes, and
protecting it from damage or destruction by fire. This is a conspicuous fea-
ture of herbarium specimens (if not removed by overzealous preparation),
appearing as a stramineous, shining, thickened (almost bulbous) base of the
plant. The bases of Calamovilfa brevipiln and Calamovilfa cun'mit (Vasey) Scribn.
are very similar. Species of the S. floridanus complex re-sprout quickly fol-
lowing fire, with green leaf material protruding beyond the blackened basal
sheaths within days following fire. Like other southeastern bunchgrasses
adapted to fire-maintained pinelands, including Calamovilfa hrevipilis. C.
ii/i'tissii. Aristida stricta. A. beyrichiana. and Ctenii/ii/ aro»/atu'//>!J, all fwe Sporobol/zs
species generally produce culms only after having their leaves removed by
fire, although they will sometimes flower in response to mechanical distur-
bance. For this reason, nearly all herbarium specimens have the upper por-
tions of the basal sheaths conspicuously blackened.
General exclusion of fire from much of the pineland habitat of the five
species of the S. floridanus complex, combined with their flov/ering only in
response to fire, means that the species are often overlooked or misidentifled,
and that they are severely under- represented in herbaria despite their local
abundance. The four narrower-bladed species, S. pmetornn/, S. ciirtissit, S. teretifoli/is.
and S. silveaN//s, are often actively or passively misidentifled as other sym-
patric pineland bunchgrasses, ^uc\\ 'j&Anstidastncta, A. beyrichiana, or Muhlerihergta
expansa. Entire savannas of a hundred hectares or more dominated hyS. pinetorum
have been assumed to be "longleaf pine/wiregrass," thotigh in reality Aristida
stricta may be completely absent. For this reason, it is important that field
biologists become familiar with the vegetative characters which allow rec-
ognition of these taxa in sterile condition.
The complex reaches its greatest diversity in the Coastal Plain of C^eor-
gia, whf^tt S.floridan//s. S. airtissii. S. teretifolius, and S. pinetoriini are all found,
and co-occtir in various combinations of two and three species, along with
the related S. juucciis. Sporoboh/s silvcaiuis is the only species of the complex
that is fully allopatric, and the only one that occurs west of the Mississippi
River. In southeastern North Carolina, S. pinetor/nii and S. teretifdias occur
in wet pinelands dominated by mixtures oi Piniis palustris, Finns serotina,
and Taxodimn ascendens. They generally co-occur with other savanna bunch-
grasses, notably Cteniumaromaticuni. Calamovilfa brevipilis, M//hlenbergia expansa,
and Aristida stricta. Sporoboliis teretijolij/s is restricted to the wettest pine sa-
vannas, usually so wet as to exclude Aristida stricta, and it may be the sole
dominant, or codominant with S. pinetorum. Ctenii/m aroniaticiim, and Muhlenbergia
expansa. Slightly less wet savannas have varying mixtures of 5", pinetorum,
Aristida stricta. Muhlenbergia expansa, and Cteniiim aroma tici/ni. A similar
Weakley and Peterson, Sporobolus floridanus complex 267
composition, though often with substantial Calamovilfa hrevipilh as well, is
seen in sandhill/pocosin ecotones in the inner Coastal Plain.
A general hydrologic gradient of the five species and the sympatric S.
junceus would be (from wetter to drier) S. floridanus. S. teretifolius, S. pinetorum.
S. silveanm, S. curtissti, S. junceus. Sporobolus floridanus is restricted to habi-
tats which are saturated at least seasonally (and often semi-permanently)
and may even have shallow ponds for weeks or even months. The other spe-
cies are generally intolerant of ponds for more than a few days. Sporobolus
teretifolius occurs in habitats with semi-permanent to seasonal saturation,
whereas S. pinetorum occurs in habitats with seasonal saturation. Sporobolus
stlveanus and S. curtissii occur in a range of sites, from seasonally saturated
to sites which are rarely saturated at the surface for periods of short dura-
tion only, and may range up to rather well-drained (though not xeric) sites.
Sporobolus junceus strictly occupies well-drained sites, ranging into truly xeric,
sandhill situations, where it reaches its greatest abundance.
Sprobolus floridanus generally occupies the wettest habitats of the com-
plex. Particularly towards the edges of its distribution, it is generally found
in very wet situations, often where water stands for periods of time, often
in seepage bogs or swales, and generally associated with Pinus elliottii var.
elliottii and Taxodium ascendens. Near the center of its distribution, especially
in the eastern portion of the Florida panhandle, S. floridanus ranges into
less wet habitats, and occurs in "mcsic flatwoods." Where its distribution
overlaps with 5", pinetorum. S. curtissii, and S. teretifolius, it can occur (with
clumps side by side), but it clearly ranges ecologically into wetter sites and
avoids drier sites.
The four eastern species in the S. floridanus complex commonly co-occur
in pineland landscapes, and sometimes occur in mixed populations, with
individuals of various species intermixed. In these circumstances, the vari-
ous species are always readily distinguishable by morphological characters.
No intermediates or likely hybrids have been seen. None of these species
has been investigated cytologically.
ACKNOWLEDCiMENTS
We wish to thank Michael P. Schafale, Richard LeBlond, Bruce A. Sorrie,
Brian van Eerden, Harry LeGrand, Jr., Patrick MciVIillan, and Robert K.
Peet for discussions regarding this group and for providing additional dis-
tribution information; Susan C. Escher for preparing the illustration; Alice
Tangerini for preparing the distribution maps; Victoria E. Batista for pre-
paring the Spanish abstract; Dan H. Nicolson for help preparing the Latin
diagnosis; Dieter C. Wasshausen and Gene Rosenberg for help translating
German; the curators of the listed herbaria, and especially Jim Massey at
268 Si DA 1,S(1)
NCU for providing facilities to the first author; and David W. Hall, Stephan
L. Hatch, and Robert K. Peet for critically reviewing the manuscript.
REFER l-NCF.S
Ali.kn, cm. 1992. Gras.ses of Louisianii. Second cd. Ca)un Prainc- Mabirat Prc-serxation
Society, Eunice, Loui.siana. Pp. 272—283.
Andi.r.son, M., p. Bocrcikon, M. Br^ir, R. CJ<,'\\\'K)ri), L. Enclekinc;, D. Eai)er-Lakc;enix)i-n,
M. GAij.>(n \, D.H. CjR()SSMa\, K. Cjoodin, S. Eandaai,, K. Mi t/.i.er, K.D. Patti-r.son,
M. Pyne, M. RiiiD, L. Sneddon, and A.S. Weakeey. 1998. International cla.ssification of
ecological communities: terrestrial vegetation of the United States. Volume II: list of
vegetation types. The Nature C'onservancy, Arlington, VA.
Baaijins, (i.J. and J.E Veedkami'. 1991 . Spnriihi)lin (Gramineae) in Malesia. Blumea 35:39.3—158.
Beal, W.J. 1896. Cirasses of North America. Henry Holt and CJjmpany, New York. Pp.
282-306.
Beaevoes, l-'.B. 1812. Essai d'une nouvelle Agrostographie, ou nouveaus genres des Ciraminees,
avec figures representant les caracteres de tous les genres. Pp. 21, 147, 164.
Beomqi 1ST, H.L. 1948. The grasses ol North Carol ma. Duke University Press, Durham,
NC Pp. 91-96.
l^OK, N.L. 1960. I^oideae-Sjiorobolcae. in: The grasses of BLn"ma. Geylon, India and Paki-
stan. Pergamon Press, London. Pp. 622—636)
Brown, \..\\. 1993- The deletion of .V. heterolejiis (Poaceae) from the Texas and Louisiana
floras, and the addition of.^. s/lrL-tiiii/s to the Oklahoma flora. Phytologia 74:371-381.
Chaeman, A.W. 1S6(). Flora of the southern United States. Ivison, Phinncy, & Co., New
York, NY.
. -. 1883. Flora of the southern United States. Ivison, Blakeman, Taylor, & C^o.,
New York, NY. P 5 50.
1897. Flora of the southern United States. Cambridge Botanical Stipply C^o.,
Cambridge, MA. Pp. 597-598.
Clayton, W.D., S.M. Piiieips, and S.A. Rt:NV()izE. 1974. Sjmrohoh/s. In: R.M. PolhiU, ed.
Gramineae (part2). Flora of tropical east Africa. Crown Agents, London. Pp. 353-388.
and S.A. Renvoize. 1 986. (ienera graminum. Cirasses of the world. Her Majesty's
Stationary Office, London. Pp. 224-228.
Ceeweee. A.E 1985. Guide to the vascidar plants of the Florida panhandle. Florida State
University Press, Tallahassee. Pp. 163—161.
Corki:ee, D.S., and M.C. Johnston. 1970. Manual of the vascular plants of Texas. Texas
Reseatch I'oundation, Renner, Texas. Pp. 217—223.
GoDiREY, R.K. and J.W. Wooten. f 979. Aquatic and wetland plants of .southeastern United
States: Monocotyledons. LJmversit}- of Georgia Press, Athens. Pp. 193-197.
CtOUEd, R W. 1 975. The grasses ol Texas. Texas A&M Press, College Station. Pp. 286-3 1 1 .
Gray, A. 1835. North American Grannneae and Cyperaceae, part 2. Ann. Lyceum Nat.
Hist. New York 3:233.
1848. A manual of the botany of the Northern United States, I'dition 1.
James Monroe and Com|iany, Boston. Pji. 576.
Gro.ssman, D.IL, D. FABER-LANoiiNDoi^N, A.S. Weakeey, M. Ander.son, p. Bourgeron, R.
Craweord, K. Goodin, S. Landaal, K. Met/.eer, K.D. Patter.son, M. Pyne, M. Reid,
and L. SNi:i)n()N. 1998. International classification of ecological commLUiities: Terres-
trial vegetation of the United States. Volume 1: The national vegetation vlassification
standard. The Nature (Conservancy, Arlington, VA.
Harper, R.M. I9OI. On a collection of plants made in Georgia in the summer of f90().
Wf.aklcy and Peters(5n, Sporobolus floridanus complex 269
Bull. Torrey Boc. Club 28:454-484.
1906. Some new or otherwise noteworthy plants from the coastal plain of
Georgia. Bull. Torrey Bot. Club 33:229-245.
Hatch, S.L., K.N. Gandhi, and L.E. Brown. I 990. Checklist ol the vascular plants of Texas.
Texas Agric. Exp. Sta. MP- 165 5. Pp. 31-32.
Hit(:hcck:k, A.S. 1935. Manual of the t^rasses of the United States. U.S.D.A. Misc. Publ.
200:392-403.
, and A. Chasi-.. 1950. Manualof the grasses of the United States, second edition.
U.S.D.A. Misc. Publi. 200:413-432.
Johnston, M.C. 1990. The vascular plants of Texas. A list, up-dating the manual of the
vascular plants of Texas. Second edition. Published by the author, Austin, Texas. Pp.
10-11.
JONF.S, S.B., Jr., and N.C. Coile. 1988. The distribution of the vascular flora of Georgia.
Dept. of Botany, Univ. of Georgia, Athens. Pp. 47-48.
Jones, S.D., J.K. Wipff, and P.M. Montgomery. 1997. Vascular plants of Texas. A com-
prehensive checklist including synonymy, bibliography, and index. Univ. of Texas Press,
Austin, Texas. P. 261.
Kenth, K.S. 1829- Revision des Cjraminees. In: I'.II.A. Humboldt and A.J. A. Bonpland,
eds. Voyage aux regions ec]uinoctiales du Noveau Continent, fait 1799-1804, partie 6,
Botanique, sect. 6. P. 68.
Lamson-Sc.ribner, F. 1897. American grasses (illustrated). U.S.D.A. Bull. 7:M0-143.
Lemon, P. C. 1949. Successional responses of herbs in the longleaf-slash pine forest after
fire. Ecology 30:35-145.
Peterson, P.M., S.L. Hatch, and A.S. Weakm:y (In Press). Sporobob/s. In: M.E. Barkworth,
K.M. Capels, and L.A. Vorobik, eds. Manual of grasses for the Continental United States
and Canada. Dept. Agric. Misc. Publ.
, R.D. Webster, and J. Valdes-Reyna. 1995. Subtribal classification of the
New World Eragrostideae (Poaceae: Chloridoideae). Sida 16:529-544.
1 997. Genera of New World Eragrostideae (Poaceae: Chloridoideae). Smithsonian
Contr. Bot. 87:1-50.
PiLGER, R. 1956. Sporobolinae. In H. Melchior and E. Werdermann, eds. Gramineae II,
Unterfamilien: Micraiodeae, Eragrostideae, Oryzoideae, Olyroideae. Die Nattirlichen
Pflanzenfamilien, second ed. l4d:53— 67.
Radeord, A.E., H.E. Ahles, and C.R. Bele. 1 964. Guide to the vascular flora of the Caro-
linas. University of North Carolina Book Exchange, Chapel Hill. P. 68.
1968. Manual of the vascular flora of the Carolinas. University of North
Carolina Press, Chapel Hill. Pp. 104-106.
Schaeale, M.P, and A.S. Weakley. 1990. Classification of the natural communities of North
Carolina, third approximation. North Carolina Natural Heritage Program, Raleigh.
Small, J.K. 1903. Flora of the southeastern United States. Published by the author. New
York, NY.
1913. Flora of the southeastern Unired Srates. Second edition. Published
by the author. New York, NY. Pp. 122-125.
1 933. Manual of the sotitheastern flora. Univ. of North Carolina Press, Chapel
Hill. Pp. 100-103.
Stape, O. 1 898. Gramineae. In: WT. Thiselton-Dyer. Flora Capensis 7:3 15, 578-589- London.
SwALLEN, J.R. 1941. New United States grasses. J. Wash. Acad. Sci. 31: 348-355.
Taylor, R.J., and C.E.S. Taylor. 1994. An annotated list of the ferns, fern allies, gymno-
sperms and flowering plants of Oklahoma. Third edition. Published by the authors, Durant,
270 Sum 18(1)
Oklahoma. P. 26,
Thomas, R.D., and CM. Allen. 1993. Adas of the vascular llora of Louisiana. Volume I :
Ferns & k-rn alHes, conifers, and monocoryledons. Louisiana Department of Wildlife and
Fisheries, Natural Heritage Program, Baton Rouge, Louisiana. Pp. 163-166.
Wlaklly, A.S. 1998. Flora of the Carolinas and Virginia. Working draft of May 1 998. The
Nature Conservancy, Southeast Regional Office, (Chapel Hill, NC. Pp. 20.61-20.63-
, K.D. Pattlr.son, S. Laxdaai,, M. Pynl, and Othlrs (compilers). 1998. In-
ternational classification of ecological communities: terrestrial vegetation of the south-
eastern United States. Working draft of March 1998. The Nature Conservancy, South-
east Regional Office, Chapel Hill, NC.
CARYOPSIS MORPHOLOGY OF LEPTOCHLOA
SENSU LATO (POACEAE, CHLORIDOIDEAE)
NEIL SNOW^
Department oj Biology
Washington University
Campus Box 1 137
St. Lonis. MO 63130. U.S.A.
P.O. Box 299
Missouri Botanical Garden
St. Louis, MO 63166. U.S.A.
ABSTRACT
Caryopsis morphology of the grass genus Leptochloa was studied regarding the necessity
of some authors for segregating Dtplachne. The data do not support the separation of Dipliulme
from Leptochloa based on a putative bimodal distribution of dorsal and lateral cross-sec-
tional compression. The presence or absence of a prominent sulcus and the relative achia-
tion of the pericarp are the only attributes sufficiently distinct to warrant use as phyloge-
netic markers. However, variations in surface texture and color can be useful regionally as
diagnostic characters at the species level.
Key worixs: caryopsis, Ltptochloa, Diplachne, morphology, systematics, Poaceae, Chloridoideae.
RF.SUMEN
Se estudio la morfologia del cariopside del genero Leptocloloa dada la supuesta necesidad
de algunos autores de segregar Diplachne. Los datos no apoyan la separacion de Diplachne y
Leptochloa basada en una supuesta distribucion bimodal de la compresion dorsal y en seccion
transversal lateral. La jiresencia o ausencia de un surco prominente y la adnacicMi relativa
del j->ericarpo son los unicos atributos suficientemente diferentes para justificar sti uso como
marcadores filogeneticos. Sin embargo, las variaciones en textura y color de la superticie
pLieden ser utiles regionalmente como caracteres cliagnosticos a nivel especitico.
rNTRODi:c;TION
The genus Leptochloa P. Beauv. s.l. (including Diplachne P. Beauv.) has been
the subject of numerous regional systematic studies due to its wide geo-
graphic distribution and the relative abundance of herbarium specimens
(Hitchcock 1903; Parodi 1927; McNeill 1979; Lazarides 1980; Phillips 1982;
Nowack 1994; Nicora 1995). These authors (and others) have disagreed as to
whether Diplachne should be segregated from Leptochloa, thereby mirroring
'Ctirrent Address: Department of Biological Sciences, University of Northern Colorado,
Greeley, CO 80639, U.S.A. Email; nsnowCf; bentley.unco.edu
SiDA 18(1): 271-282. 1998
272 Sum 18(1)
the ditterini,^ opinions of two worldwide generic summaries of grasses (Clayton
& Renvoize 1986; Watson & Dallwitz 1992). A frequently cited source of
evidence to support the segregation of D'tplachm has been differences in caryopsis
features.
Parodi (1 927) apparently was the first to examine cross-sectional shapes
critically. He partitioned four neotropical species into Diplachne or Leptochloa
based on relative compression (dorsal or lateral) of the caryopsis, and the
presence or absence of a hilar groove. He suggested that Leptochloa chloridiformh
was aberrant in Leptochloa because of its lack of a hilar groove (Parodi, I.e.).
Vails (1978) studied the systematic affinities oi Leptochloa dubia in rela-
tion to the generic boundaries of Leptochloa. He illustrated cross-sectional
profiles and profiles from the embryonic and hilar sides for seven species.
The figures revealed a gradation in cross-sectional profile from nearly round
in Diplachne caitclata to somewhat triangular in Leptochloa virgata and L. scabra,
to relatively Battened in L. fascicz/laris (Vails, I.e.: 103). He also concluded
that a hilar depression was a tenuous systematic feature. Despite somewhat
limited sampling, his results suggested strongly that caryopsis features intergrade
too thoroughly to split Leptochloa unambiguously into two genera, although
he acknowledged that "some grouping of species can be achieved on the
basis of caryopsis type" (Vails, I.e.: 105). Unfortunately, his results were
never formally published and have not been cited by subsequent authors.
McNeill (1979: 401) and Nicora (1995: 233) repeated almost verbatim
the observations of Parodi (1927) without adducing additional data or cit-
ing the work of Vails ( 1 978). Lazarides ( 1 980) observed that Australian species
generally could be segregated into Leptochloa or Diplachne on the basis of
caryopsis siiape, with the exception oi^ Leptochloa digitata, with its flattened
shape. Phillips (1982: 144) agreed with Parodi (1927) regarding the util-
ity of caryopsis shape for splitting the genera, but noted exceptions in Diplachne
caudata, Leptochloa obtusiflora, and L. longa. Nowack (1994) provided a cur-
sory review of caryopsis shapes for IMalesian taxa and concluded that the
differences set forth by Parodi (1927) were insufficient to permit recogni-
tion of segregate genera.
Based on a recent monographic treatment (Snow 1997), Leptochloa (in-
cluding Diplachne) represents forty taxa, with one species, L. monticola Chase,
being of dubious inclusion (Vails 1978; Snow 1996). Prior to this study,
relatively few taxa had been examined critically for variation in features of
the caryopsis, and much systematic weight had been placed on the meagre
observations that existed. In light of lingering debates about generic boundaries
between Leptochloa and Diplachne (Jacobs 1987) and the emphasis previous
authors placed on the utility of the caryopsis to segregate these genera, a
survey of all currently recognized taxa of Leptochloa was undertaken to evaluate
whether features of the caryopsis could be useful as phylogenetic markers.
Snow, Caryopsis morphology of Leptochloa 273
MATERIALS AND METHODS
Caryopses of all currently recognized taxa in Leptochloa (Snow 1997) were
removed directly from herbarium specimens (Appendix 1), placed under a
Nikon SJMZ-U dissecting microscope with camera lucida attachment, and
the profiles were traced by hand. Cypholepis yemenicus was included because
it resembles L. eleusine and L. obtusiflora in several respects (Snow 1996),
and was used as an outgroup in preliminary cladistic studies o^ Leptochloa
(Snow 1997; see also van den Borre & Watson 1997). In most cases a mini-
mum of three specimens were examined for variation (Appendix 1). The
following features were observed: 1) caryopsis shape when viewed from the
hilar side ("hilar profile"); 2) caryopsis shape when viewed from a cross-sec-
tion taken at midpoint with the hilar side oriented above ("cross sectional
profile"); 3) the presence or absence of a sulcus or other depression on the
hilar side when viewed in cross-section; 4) ornamentation on the outer coat
(perisperm); 5) relative adnation of the perisperm to the endosperm, and 6)
color of the grain. To standardize the sampled developmental stage, cary-
opses were selected from spikelets in which florets were beginning to disar-
ticulate, a condition that assures their maturity. In virtually all cases the
caryopses from the lowermost floret in the spikelets were selected.
For the sake of precision, descriptive terminology of shapes follows that
of the Systematics Association (1962), whereas that of surface ornamenta-
tion follows Murley (195 1 ). Given that shape is a continuously varying character,
these typologies might not account for the observed and often subtle varia-
tions in shape. For example, a caryopsis might have an intermediate ellip-
tic shape of 2.5:1, which is absent from the diagram. Nonetheless, after
initial analyses, the diagram shapes appeared adequate to standardize and
summarize the majority of both hilar profile and cross-sectional shapes. With
respect to cross-sectional shapes, I accounted for the absence of sharp edges
by prefixing the terms "obtriangular" (3:2), "shallowly obtriangular" (3:2),
and "shallowly obdeltate" with the word "rounded," which more accurately
depicts their shapes. To account for concave inflections of the hilar surface
(always oriented above, Fig. 2) I used the terms "sulcus" and "depression,"
depending on the degree of concavity. As used here, a sulcus is a vertical or
nearly vertically walled groove; a depression refers to any gradual concav-
ity, and will be further modified by the terms shallow, moderate, deep, narrow,
and broad. These subjective terms were deemed necessary to describe the
observed variation.
RESULTS
The hilar profile for most species was some variation ot obovate or ellip-
tic (Fig. 1; Table 1). Only four taxa had ovate hilar profiles {L.fusca subsp.
274
SlDA 18(1)
Narrowly elliptic
A
2
6:1
3:1
2:1
Obovate
2:1
3:2
Elliptic
Ovate
2:1 3:2
Widely elliptic
6:5
3:2
Widely obovate Very widely obovate
6:5
1:1
Fig. 1. Hilar profiles of caryopses observed tor LepUxhioa. Tlie apex is oriented above.
Snow, Caryopsis morphology of Leptochloa
275
Table 1. Variation in hilar profiles and cross-sectional shapes of the caryopsis in l^eptochloa s. 1. The
numbers following each taxon reflect observed variation: numbers to the left of the double bar (|j)
indicate hilar profile shapes (see Fig. I); those to the right of the double bar reflect cross-sectional
shapes (Fig. 2).. Hence, 2,3,8 || 2,9 would indicate a taxon having narrowly elliptic (.3T), elliptic
(2:1), and obovate (2:1) hilar profiles, and oblate and depressed obovate (2:.3) cross-sectional shapes.
Taxa with an asterisk (*) indicate those previoLisly placed in Dtplachm by some authors.
Leptochloa at]uatica .3,9,10 || 8,9,12
L. caudata*l,2,8 || 2
L. chinensis 8,9 || 2,9
L. chloridiformis 2 || 7
L. coerulescens 8,9 || 2,8
L. decipiens subsp. asthenes 3,8 [| 2,8,9
L. decipiens subsp. decipiens 3 |[ 9
L. decipiens subsp. peacockii 3,8 || 3,9
L. digitara 2,3,8 1| 8,12
L. divaricatissima 3 j| 2,9
L, dubia* 2,3,11 || 8
L. eleusine* 3,8,9 || 8,12
L. fusca subsp. fascicularis* 3,8 j] 4
L. fusca subsp. fusca* 9 j| 4,8
L. fusca susbsp. muelleri* 3,9 || 4,8
L. tusca subsj-). uninervia* 3,7,8 jj 4
L. gigantea* 3,6,8 || 8
L. longa 3,8 || 9
L. ligulata 3,4 jj 2,7
L. malayana* 3 || 9,11
L. marquisensis 2,3,6 jj 9,10
L. monticola* 3,8 || 4,8
L. nealleyi 4,9 || 2,9
L. neesii 5,10 j| 1
L. obtusiflora 3 j| 8
L. panicea subsp. brachiata 3,4 || 10,13
L. panicea subps. mucronata 3 |j 1,2,8
L. panicea subsp. panicea 7 [| 1,2
L. |ianicoides* 3,4 j| 8
L. rupestris 3 || 10
L. scabra* 2,3 jj 9
L. southwoodii 3,4,8 jj 1,2
L. squarrosa 2 j| 5,11
I., srilankensis 3 jj 9,10,11
L. tectoneticola* 2 j| 8
L. uniflora 2,6 jj 6,7
L. virgata 2,3,6 j| 6,7,1 1,12,13
L. viscida* 3,8 II 4
L. xerophila 3 jj 2
Cypholepis yemenicus 4 j| 8,9
unimrvia^ L. gigantea, L. uniflora, L. virgata). The widest was the very widely
elliptic shape of a few specimens of L. dubia. The thinnest was the narrowly
elliptic (3T) shape expressed by some specimens ofL. caudata, L. chloridi-
formis, L. dubia, L. digitata, L. scabra, L. squarrosa, L. uniflora, and L. virgata
(the lattermost sensu lato, including L. barbata and L. procera sensu Nicora
1995). Many species were variable, for example having both elliptic (2:1)
and obovate (2:1) shapes. Not surprisingly, the greatest variation in hilar
profile shape occurred in widespread species such as L. dubia and L. virgata.
The cross-sectional shape was considerably more variable than hilar profile
shape (Fig. 2). Overall, the observed variation ranged from dorsally com-
pressed through circular (no compression) to laterally compressed (Fig. 2).
Most taxa had only slight to moderate degrees of lateral or dorsal compres-
sion. As with hilar profiles, many taxa showed infraspecific variation in cross-
sectional shapes (Table 1). Some specimens of L. neesii appeared circular,
whereas others were oblate (Table 1). The greatest degree of dorsal com-
pression was expressed by the depressed obovate (1:2) and transversely el-
liptic (1:2) shapes. With some modifications, these shapes accounted for
some or all of the variation of many species (Table 1). The highest degrees
of lateral compression were the obovate (3:2), rounded shallowly obdeltate
276 SiDA 18(1)
(5:6), and rounded shallowly obtriangular shapes. Only L. squarrosa was obovate
(with a moderate hilar depression), whereas some representatives of L. virgata
were both rotinded shahowly obdeltate (5:6) or rounded shallowly obtriangular
(2:3).
A distinct sulctis was present only for L. n/pestris and L. uniflora, although
a number of taxa had depressions of varying extent on the hilar surface (see
Disctission).
The surface of the pericarp varied from smooth to variously rugose. The
following were at least occasionally somewhat rugose: L. chloridifornt'n, L.
ilecipieiis subsp. decipiens, L. divciriccitissiDUi, L. g/gaiilea, L. hmga, L. //udayai/a,
L. riioiitu'olci, L. riealleyi, L. neesii, L. obtus/flora, L. saibra, L. soi/thwoodii, L.
uniflora. Whereas a smooth pericarp was consistent for many taxa, those that
expressed the rugose condition did so irregularly.
Species with a weakly adnate pericarp (detaching soon after placement
in water at room temperature) included L. chloridiforniis, L. duhia, L. e/e//sh?e,
L.fusai subspecies ///icv/, fascicnhiris, un!)iervia, and L. obtusiflora.
The color of the caryopsis varied from very light brown to dark reddish
or very dark brown, but most were an intermediate shade. Leptochloa longa,
L. obt//sJflora, and L. sqiiarrosa were tisually dark brown. Leptochloa monticola,
a species of dubiotis inclusion in the genus (Clayton & Renvoize 1 986; Vails
1978; Snow 1996, 1997), was usually a dark reddish brown.
DISCUSSION
This simple study of caryopsis morphology has revealed more variation
within and between taxa o( Leptochloa than previously recognized (Parodi
1927; McNeill 1979; McVaugh 1983; Nowack 1994; Nicora 1995). With
two exceptions, features of the caryopsis appear to be of little value in Leptochloa
as phylogenetic niarkers, although some are of diagnostic value in keys. These
results contrast with those of a recent study in Triticeae, which suggested
caryopsis morphology was of systematic value at the tribal level (Terrell &
Peterson 1993).
Parodi's ( 1927) study was limited to four species in Leptochloa and one in
Goutnia Fourn., which represents only ten percent of Leptochloa as currently
circumscribed (Snow 1997). Another disconcerting aspect was his lack of
voucher specimens and tmcertain depth of sampling within taxa, although
this study does not contradict the profiles of the species he illustrated. Overall,
Parodi's sampling underestimated considerably the variation in cross-sec-
tional shape in Leptochloa. This study has revealed nearly continuous varia-
tion in cross-sectional shape, from dorsally compressed and non-compressed
(circular or oblate) to various degrees of lateral compression. The bimodal
compression (lateral or dorsal) ofcaryopses in Leptochloa recognized loy Parodi
Snow, Caryopsis morphology of Leptochloa
277
Circular
Oblate
5:6
Transversely elliptic
3
2:3
4
1:2
Obovate Widely obovate Very Widely obovate
5
3:2
Depressed obovate
1:1
Widely depressed obovate
8
1:2
<^
2:3
"Rounded" "Rounded" shallowly "Rounded" shallowly
obtriangular obtriangular obdeltate
13
3:2
2:3
5:6
Fig. 2. Cross sectional sliapes of caryojises observed tor Leptochloa. The hilar side is oriented
above.
278 SiDA 18(1)
(1927) simply does not exist for the genus as a whole. As such, variation in
caryopsis cross-sectional morphology cannot by itself be invoked as a ge-
neric-level character to segregate Diplachne.
The caryopsis profile as viewed from the hilar side ranges continuously
from narrowly elliptic through ovate to obovate and very widely obovate
(Fig. 1 ). Whereas the extremes ot variation can be useful as diagnostic fea-
tures between some taxa, the continuous variation makes the hilar profile
useless as a phylogenetic marker (Stevens 1991 )•
Various degrees of concavity occur on the hilar side. Broad, shallow de-
pressions (not illustrated) characterize certain taxa fairly well, such as Leptochloa
eleusine, L. ionga, and L. obtustflora. Taxa showing this feature irregularly were
Leptochloa chinensis, L. di/bia, and the related Cyphokphis yernenicns . Shallow,
relatively narrow depressions (not illustrated) occur in other taxa, althotigh
less consistently; these included L. chloridijormis, L.decipiens subsp. decipiem^
L. dtgitata, L. sqiMrrosa, and L, virgcita. At best, the degree of concavity is
useful only as a diagnostic character in regional keys. However, a promi-
nent sulcus, characterized by its vertical or nearly vertical walls, was a con-
sistent character for L. rnpestrh and L. uniflora, and is one of only two char-
acters I consider sufficiently distinct and consistent to be phylogenetically
useful.
The relative adnation of the pericarp is the second character of the cary-
opsis useful for phylogenetic inference. It is well known that the pericarp is
only weakly adnata to the endosperm in some species of Leptochloa (Izaguirre
& Laguardia 1987; Watson & Dallwitz 1992) and some related genera, such
as Eragrostis Wolf (Lazar ides 1997). In such taxa the pericarp will dissociate
from the endosperm quickly when placed in water at room temperature.
iVIost species in Leptochloa have a smooth outer texture. A few can be rug-
ose, althotigh this feature was unreliable within taxa. For example, a coarse
but sparsely rugose surface generally, but not always, characterizes Leptochloa
pantcea stibspecies panicea and mucronata (sensu Snow 1 99Ba, but not sensu
Nowack 1994), which helps to distinguish these from the widespread L.
panicea subsp. brachiata (Snow 199Ha; formerly known as L.jilifornih or L.
mucronata [Snow & Davidse 1993}).
As the color of the caryopsis often varies with the degree of maturity,
only mature specimens should be evaluated for this attribute. Except as a
diagnostic feature in keys, in which a few species are dark brown, color is of
minimal systematic value in Leptochloa.
I return now to cross sectional shape, which has been discussed exten-
sively regarding the separation o'i Diplachne from Leptochloa (Parodi 1927;
McNeill 1979; Phillips 1982; Nicora 1995). As mentioned above, the per-
ceived bimodality of lateral and dorsal compression disctissed by Parodi (1927)
Snow, Caryopsis morphology of Leptochloa 279
has been invoked to segregate Diplachne from Leptochloa. The results of this
study firmly reject such a notion, given the nearly continuous variation of
cross-sectional hilar profiles (Fig. 1).
It also has been suggested that the presence or absence of a distinct keel
on the lemma is positively correlated with cross-sectional shape, and is a
means by which the genera can be separated (Parodi 1927; McNeill 1979;
Nicora 1995). It is true that some taxa with a dorsally compressed cary-
opsis have flat lemmas at maturity, as for example L. fusca subsp. muelleri.
However, others have little or no such positive correlation. Mature fruits of
L. neesii can be round or nearly so in cross section, yet still be borne within
a keeled lemma (e.g. , Lan(^ld283 , CANB). This is also true for some specimens
of L. ligulata, L. nealleyi, L. pankea subsp. panicea, and L. southivoodii. Pre-
liminary cladistic studies have failed to consistently group together taxa
having dorsally flattened caryopses (Snow 1997).
This study upholds and strengthens the unpublished work of Vails (1978),
who concluded that variation in caryopsis shape was too great to support
the recognition of Diplachne. In particular, the data herein have revealed
nearly continuous variation in cross-sectional shape, ranging from dorsal
compression through circular to various degrees of lateral compression (Table
1; Fig. 2). The lateral/dorsal compression dichotomy of Parodi (1927) sim-
ply does not exist for Leptochloa. Moreover, a nearly identical range of cross-
sectional shapes can exist in closely related genera, as illustrated in a recent
revision of Australian £r^^^roj//j- (Lazarides 1997: 176).
The most general observation to emerge from this and other detailed anatomical
and morphological studies of grasses (Davila & Clark 1990; Ellis & Linder
1992; Snow 1996; Lazarides 1997) is that broader sampling regimes gen-
erally reveal additional variation not encountered in narrower surveys. Such
variation cannot be ignored. Future studies therefore should seek both breadth
and depth of sampling in order to minimize the chances of incompletely
characterizing variation. Underestimates of variation leads to errors in the
diagnosis of taxon boundaries and in the accuracy of inferring phylogenetic
relationships, the latter being the very basis by which we make and sup-
port our classifications.
ACKNOWLEDGMENTS
Generous funding was provided by the National Geographic Society (NGS
5594_95) and the Missouri Botanical Garden (Andrew W. Mellon Founda-
tion). A grant from the National Science Foundation to the Population and
Evolutionary Biology Program at Washington University in support of graduate
training is also gratefully acknowledged. My thanks to Dr. John McNeill
and an anonymous reviewer for helping to clarify the manuscript.
280 Sida18(1)
APPENDIX 1
VoucluT specimens aiul lierbariiim of\)n^nn (ueronyms Follow Holmi;ren et til. 1990); chose lack-
ing herbarium designation are housed at MO. For new combinations and new species in Leptucbluct see
Snow 199Ha, b and Snow and Simon 1997.
LeptoMoci .iqihUiai Scribn. & Merr.: Hitchmk 7()<)4 (US); Pr/rii^/e 6664 (US); .Vw/cnYrow 650 (US); MiVa//}^/}
19124 a^S):Siioir662_'i.
L. caiuiata (K. Schum.) N. Snow: Van Somereri AH9575 (US); Smm-den 1429 (US); Bogclciii I jO (UC).
L. chinensis (L.) Nees: Snow et al. 698()\ Kanm 254 (K); Poore 440 (K); Dariclse 7471 (K,MO); Chiytun
5644 (K).
L. chlondifonius (Hack, ex Stuck.) Parodi: Biniheu HO (LP); S//re//s 622 (MICH); Peckrwi 3471 (US);
PeJersefi 2662 (US).
L. coerideueus Sreud.: Adiiw ni75\ At/am 14(>3(>:. Ad/w 5694.
L. decipiens (R. Br.) Stapf ex Maiden subsp. cnthetms (Roem. & Scluilt.) N. Snow: Swiw & Simon 7272;
Snow & Simon 732^; Snow & Simon 7335\ Cnsp et al. 2710 (MEL); 'rimw/Kuni & Slh/r/iv HUGin
(BRI); Hiddhifd 509S(K).
L. decipieus subsp. decipiens: Snow et ul. 724'^; Snow & Simon 732bl; Snow & Snnon 7334; B/ake 22548
(CANB); Roes.n. (MEL, accession 234696); Ljz.nides 5634 (US); Rei^an .un. (CANB).
/.. deap/en.\ subsp. /'tv/c«i// (Maiden & Betclie) N, Snow: S)iow & Simon ^ 323\Siiow & Simon 7329; Snow
& Simon 7330; Snow & Simon '^336; Pi/rdie 3 1 5D (liR\); Boonnan .i.n. (G, accession 8227-(S6); /w/jw.uw
713 (CANB).
/,. digitataiR. Br.) Domur. Snou et ,d. 7224;Snowef al. 7 235;Siiow et al. ^246; B//rhidge 5326 {CAbiW);
Blake 1 1 506 (CANB); Blake 6320 (CANB); Walter & Walter 2590 (B).
L. divaricatisiima S. T. Blake: Snow et al. 7228; Snow et al. ^2)' i; Snow et al. 7236; Snow et al. 7241;
Lloyd y^9 (CANB); Blake 77^7 (BRI).
L. dnbia (Kuni\\)neiis: Snow 5865; Warnock46783 {NCU); Krai 5 1 801; Mearm 1213 (US); Hernandez
& Mathns N-2066 (GH); Gould 12183 (K); Castillon 4356M (GH).
L, eleiisme (Nees) T. A. Cope & N. Snow: Snow et al. 6941; Snow & Burgoyi/e 6)954; Snow & Biirgoyne
6963; Snow et al. 6982; Schweiekerdt 1896 (PRE); Gi/y & Ward ^ (PRE); Drlge s.n. (S, accession 93/
1 94); E.xtenuon 0/fwer 164 1 9 (PRE).
L. /n.\-ea (L.) Kiinth subsp. ///.kv/: Snow et al. 7215; S)iow et al. 7216; Snow et al. 7 222; Snow et al. 7232;
Snow et al. 7234; Snow et al. 723^; Tracy 929? (GH); Pry s.n. (GH).
L. fnsca s.u6-s\\. fascicidaris (Lam.) N. Snow: Snow 5786A; Snow 5800; Snow 5804; Snow 5809H; Snow
581 ! A; Snow & Koster 5824; Snow & Koster 5840; Snow 584 1 ; Snow 5842; Snow 5896; Snou 5900;
Snow 5901 -B; Snow 590 1 -H; Snow 5903; Praser 63 I (MICH); Pringle 9595 (MEXU); Swallen 4265
(BAA); Wooton s.n. (BAA).
/,. Ji/sca subsp. mnelleri (Bcnth.) N. Snow: Patz 1 1 (BRI); Maconochie 13433 (BRI); Ab/st 48^ (CANB).
L. f/isca subsp. unmervia (J. Presl) N. Snow: Snow & Prinzie 648-i;Snou- & Priiizie 6567; Snow & Prinzie
6568; Snow 6598.
L. gigantea (Eaunert) T. A. Cope & N. Snow: Smi/li 4 1 26 (PRE); Smith 13H7 (BRI).
L. ligulata Lazarides: Snow & Simon 7324; Snow 7402; Story & Yapp 25 (CANB).
L. longa Griseb.: Davidse 2612; Soderstrom 10^3 (US); Hitchcock 10377 (US).
L. malayana (C. E. Hubb.)Jansen ex Veldkamp: Yacoh 25883 (P).
L. marqtiisensis (R Br.) P. M. Peterson & Judw.: Perlman 14919; Perlman 15064.
L. monticola C.lvAst: Holdridge 1955 (US); Pkman HI 1874 (US).
L. nealleyi Vasey: Snoir 5793B; Snow 5H05A; Snow 5920; Cory 20298 (GH); Waller & Banml 3096
(GH);Johnston 4872 (MK:H).
L. neesii (Thwaires) Bench.: Snow & Simon ^378; Snotc & Simon 7 384; Davidse & Siimithraarachcht 9180
(MO.US); White 8894 (US).
L. obtiisiftora Hochsc: Milne-lledhead & Taylor 7297 (B); Greenway 10666
L. panicea (Recz.) Ohwi subsp. panicea: Reekmans 4982 (B); Ahmad s.n. (B, accesssion 173193-120);
Hitchcock 194^4 (US); I'leld & Loew 6B (US); Backer 35094 (US).
L. panicea subsp. brachiata (Sceud.) N. Snow: Snow 5905;Snow 5910A;Snow & Prinzie 6529C; Snow &
Prinzie 6558; Snoir 6654; Snow 667 2-A .
L. panicea subsp. nuicronata (Michx.) Nowack: Siiow 5847 A
Snow, Caryopsis morphology of Leptochloa 281
L paniandei (J. Presl) Hicchc: Snmv ':,192A\Sn„w 5SI()A; Snow 6622\Jacob 511 (NCU); Fnimoer &
Williams 41 (ENCB); McDcimd & Rimaihi 18020.
L. rupestris C. E. Hubb.: Wood 2848 (BM); Wood s.n. (K); Gilbert & Phillips 8874 (K).
L. scabni Nets: Snow 5788; Snow 5791A; St/oir 5S/(M; McKenzie & Urhatsch 160 (NY); Nee 31921
(NY); Burger & Ltesner 6958 (NY).
L. southwoodii N. Snow & B. K. Simon: Snow & Stiiuni 7350; Snow & Simon 7362.
L. squarrosa Pilg.: Greenway 2764 (P); Schliebtn 6>940 (P); Milne-Redhead & Taylor 7301 (B).
L. srilankensis N. Snow: Davidse & Siniiithraaracbihi 9066 (K); fnsber^ et al. 50835 (CANB); Clayton
5591 (TABS).
L. tectoneticola {E&cker) ]dnsen ex Veldkamp: Poilam 15394 (P); Smitinand 3418 (CANB); Kerr 20633 (K).
L. uniflora Hochst. in A. Rich.: Trimens 28 (US); Rains 67 (US); Thollon 4093 (US).
L. virgata (L.) P Beauv.: Pohl & Davidse 10603 (UC); Burkart 22139 (SI); Meza 12; Swallen 3109 (US);
Brockmann 18660 (BAA).
L. viscida (Scribn.) Beal: Snow 6597; Riii'z s.n. (PNCB); Palmer 1 789 (CJH); Goodding & Lusher 139-45
(NY).
L. xerophila P. M. Peterson & Judw.: Halle 2098 (P).
Cypholepis yemenciais (SchweinF.) Chiov.: Bally & Melville 1 5783; Verdcourt 3275.
RF.IT.RENCES
Clayton, W.D. and S.A. Ri;NVOiZi:. 19B6. Genera graminum: Grasses of the world. Kew
Bull. Addic. Ser. 1.3.
DAviLA, p. and L.G. Clark. 1 990. Scanning electron microscopy survey of leaf epidermis of
Sorghastruin (Poaccae: Andropogoneae). Amer. J. Bot. 80:1444—1454.
Ellis, R.P. and H.P. Lindlr. 1992. Atlas of the leaf anatomy in Pentaschistis. Mem. Bot.
Surv. S. Africa 60.
Hitchcock, A.S. 1903. North American species oi Leptochloa. U.S.D.A Bull. PI. Industr.
Publ. 13:1-21.
HoLMGRLN, P.K., N.H. HoLMGRLN and L.C. Barxett. 1990. Index herbariorum. Part I:
The herbaria of the world. Eighth Edition. Rcgnum Veg. 120.
IzAGUiRRE, P. and A. Laguardia. 1 987. Un nuevo enfoque hacia la definicion del fruto de
las gramineas. Uruguay Fac. Agron. Bol. Invest. 3:1-15.
Jacobs, S.W.L. 1987. Systematics of the chloridoid grasses. In: T.R. Soderstrom, K.W. Hilu,
C.S. Campbell, and M.E. Barkworth, eels. Grass systematics and evolution. Smithsonian
Institution Press, Washington, D.C. Pp. 277-286.
Lazarides, M. 1980. The genus Leptochloa Beauv. (Poaceae, Eragrostideae) in Australia and
Papua New Guinea. Brunonia 3:247—269-
1997 A revision of Eragrostis (Eragrostideae, Elcusininae, Poaceae) in Aus-
tralia. Aust. Syst. Bot. 10:77-187.
McNeill, G. 1979. Diplachne and Leptochloa (Poaceae) in North America. Brittonia 3 1 :399-
404.
McVAtJGH, R. 1983. Gramineae. Vol. 14. Flora Novo-Cjalaciana: A descriptive accotint of
the vascular plants of western Mexico, W. R. Anderson, editor. The University of Michigan
Press, Ann Arbor.
MuRi.F.Y, M.R. 1951. Seeds of the Cruciferae of northeastern America. Amer. Midi. Natu-
ralist 46:1-81 .
NicoKA, E.G. 1995. Los generos Diplachne y Leptochloa (Gramineae, Eragrosteae) de la Ar-
gentina y pafses limftrofes. Darwiniana 33:233—256.
NowACK, R. 1 994. Revision oi Leptochloa Beauv. (inch Diplachne Beauv.) (Poaceae) in Malesia.
Rheedea 4:79-92.
Parodi, L.R, 1927. Revision de las gramineas argentinas del genero Diplachne. Revista Fac.
Agron. Veterin. 6: 21-43.
282 Sum 18(1)
Piiii.iJFS, S.M. 1982. A numerical analysis ot Eragrosticleac (Gramincae). Kew Bull.
37:133-168.
Snow, N. 1996. The phylogenetic utility of lemmatal micromorphology in Leptochloa and
related genera in subtribe Eleusininae (Poaceae, Chloridoideae, Eragrostideae). Ann. Missouri
Bor. Garden 83:504-529.
I 997. Phylogeny and Systemarics oiLel7tochloa P. Beauv. scnsu lato (Poaceae,
Ghloridoideae). Ph.D. dissertation, Washington University, St. Louis, Missouri.
I 998a. Nomenclatural changes in Leptochloa P. Beauvois sensu lato (Poaceae,
(Chloridoideae). Novon 8:77—80.
I 998b. A new species onAJHochlon (Poaceae, Chloridoideae) from Sri Lanka.
Novon 8:183-186.
Snow, N. and CL Davidsi,. 1993. l.tptucbliki iiiiui-oihilii (Michx.) Kunth is the correct name
for Lt'ptochliki jililoriiiis (Poaceae). Taxon 12:41 3-4 I 7.
, B.K. Simon. 1997. Leptochloa southwoodii (Poaceae, Chloridoideae), a new
species from south-east Queensland. Austrobaileya 5:132-143-
Stfvf.ns, P.P. 1991. Character states, continuous variation, and phylogenetic analysis: A
review. Syst. Bot. 16:553—583.
Systematics Association CoMMin i;i; ior Di:.s(;riptivi; Bioloc;ical Terminoi.oc",y. 1962. 1.
Terminology ot simple symmetrical plane shapes (chart 1). Taxon 11:145—156.
Terrell, E.E. and P.M. Peterson. 1993- (Caryopsis morphology and classification in the
Triticeae (Pooideae: Poaceae). Smithsonian Contr. Bot. 83-
Vai.i.s, J.F.M. 1 97S. A biosystematic study ol Leptochloa with special emphasis on Leptochloa
diibia (Gramineae: Chloridoideae). Ph.D. dissertation, Texas A & M LJniversity, College
Station.
Van 1)i:n Borri:, A. and L. Watson. 1 997. On the classification ol the Chloridoideae (Poaceae).
Australian Syst. Bot. 10:491-53 I .
Veldkamp, J.F. I 97 1 , Notes on Malesian grasses V. New s|iecies and combinations in Phenlochloa,
Hypan'he>/ia dnd Leptochloa. Blumea 19:64.
Watson, L. and M.l. Dallwitz. I 992. The grass genera ol the world. Ci.A.B. International.
XANTHOSTEMON FRUTICOSUS (MYRTACEAE), A
NEW SPECIES FROM THE PHILIPPINES
PETER G. WILSON
National Herbarium of New South Wales
Royal Botanic Gardens
Sydney, N.S.W. 2000, AUSTRALIA
LEONARDO L. CO
Herbarium, Institute of Biology
University of the Philippines
Dilvnan 1101. Quezon City. PHILIPPINES
ABSTRACT
A new species oi'Xdnthostemofi. X.fri/ticosus. presently known only from the Palanan Wilderness
in Isabela Province, Luzon, is described and its relationships discussed.
RESUMEN
Una nueva especie de Xarithostemon, X. fruticosus, en la actualidad solamente conocida de
la Palanan Wilderness en la provincia de Isabela es descrita y sus relaciones con otras especies
discLitidas.
Xanthostemon is a genus of around 45 species that occurs in Australia, New
Caledonia, the Solomon Islands, Papua New Guinea, Indonesia (Irian Jaya,
the Moluccas and Sulawesi) and the Philippines. In the Philippines, only
four species have been named and there has been little published on the
taxonomy of the genus since Merrill (1952) clarified the application of the
names X, speciosus and X. verdugonianus.
In 1991, a preliminary inventory of the Palanan Wilderness was under-
taken as a joint project of Conservation International, Leiden University
and the Isabela State University. A report of this inventory is given by Co
and Tan (1992). One of the vegetation types encountered was a low scrub
that occurred on ultrabasic substrate and was dominated by a shrubby
Xanthostemon species with large, right red flowers. This has proven to be an
undescribed species, most closely related to X. verdugonianus and is here described
as X. fruticosus.
Xanthostemon fruticosus Peter G. Wilson & L. Co, sp. nov. (Fig. 1). Type:
PHILIPPINES. Luzon Island. Isabela Prov.: Aubarede Peninsula, Lanay Spring, ca.
17 km NNW of Palanan Point, 25 May \39\,Uo)utrdoCo '^5S5 (holotype: A; isotypes:
PNH, PUH, CAHUP, ISB, L, KEP, US, K, CANB).
SiDA 18(1): 283-286. 1998
284
SiDA 18(1)
Fk;. 1 . XiinthdStciiiiJii jr/niius/is Peter Ci. Wilson & L. Co. A. Habir B. Immature fruit show-
ing neatly circumscissile hypanthium. (;. Open, mature truit. D. Detail of leat showing
venation. All from holotype. Scale bar: A, D=.7 cm; B, C=l cm.
Wilson and Co, A new Xanthostemon species 285
Xanthostemon verdugiano hypanthio circumscisso valde affinis sed habitu friiricoso foliis
floribusque maioribus pedicellis pedunculisque longioribus et petalis roseis diftert.
Shrub ca. 1 m high; young shoots sparsely appressed-hairy, glabrescent.
Leaves spirally arranged; petiole 7-10 mm long, 2.5-3 mm wide; lamina
obovate, 6-8 cm long, 3.5-4 cm wide, length:breadth ratio 1.6-2.1:1, co-
riaceous, apex rounded and emarginate; oil glands numerous. Inflorescences
crowded toward the apex of seasonal growth unit, subtended by slightly
reduced foliage leaves, axillary, the apical bud apparently always growing
on (i.e. shoot auxotelic). Unit inflorescences triads; peduncles 5-9 mm long,
the upper ones shorter giving a corymb-like appearance to the flowering
shoot; pedicels 6.5-10 mm long. Hypanthium cup-shaped, 5-6 mm in diameter,
4—5 mm deep, exceeding the summit of the ovary, ±glabrous. Petals 5, pink,
oblong to obovate, 5.5-8 mm long, 3.5-6.8 mm wide. Sepals 5, +equal, 2-2.7
mm long, 2.5-3.8 mm wide, ±triangular, apex obtuse to rounded. Stamens
red ±equally spaced, ca. 22 in a single series around the rim of the hypanthium;
filaments 22-25 mm long; anthers 1.2-1.5 mm long, connective broad with
one large gland at the apex and 2-5 other, smaller glands. Ovary partly in-
ferior, (2-)3-locular, glabrous. Style 30-35 mm long, extending 3-5 mm
beyond the stamens; stigma as wide as or slightly narrower than the style,
flat. Fn,?i/ depressed-globular, ca. 8 mm diameter, ca. 6.5 mm high, broadly
lobed at each loculus; hypanthium neatly circumscissile 1.5—2 mm above
the base, the remnant ±flattened under the capsule. Placentas rod-like, horizontal.
Seeds not seen.
Distribution. — Known only from the type locality.
Ecology. — Xanthostemon friittcosNS has been found near the coast in a low
scrub community occuring on exposed sites on soils derived from an ultra-
basic substrate. It is the dominant component of this community type at
Lanay which also includes a number of other genera of spreadiing or erect
shrubs (Co & Tan 1992).
Conservation status. — ^Apparently rare. Only recorded from one of four ultrabasic
sites visited in this part of north-eastern Luzon. Ultrabasic areas are very
marginal for agriculture but are potentially subject to applications for nickel
mining. As a wilderness, the region does have legal status as a 'Protected
Area' but this may not be an impediment to mining.
Etymology. — The epithet is derived from the 'L2itm,fruticosus, shrubby, referring
to the distinctive habit, a characteristic that distinguishes it from the re-
lated X. verdugonianus .
The red-flowered X. speciosus Merr. From Palawan and nearby islands has
a broader, dish-shaped hypanthium with distinct vesicles and is thus not
closely related to the new species but belongs with X. confertiflorus (Sulawesi)
and X. youngii (north-eastern Australia) in sect. Vesicaria of Gugerli (1940).
286 SiDA 18(1)
Xanthoste?nun speciosus is usually a tree of 5 meters or more but small trees or
shrubs of only 1 meter or so in height have been recorded (e.g. Co 3032, A,
PUH, KEP).
Xanthostemon fniticosus is closely related to X, verdugonianus which it re-
sembles in having red flowers with a circumscissile hypanthium. Xanthostemon
verdugonianus is a tree up to 26 m high that ranges from north-eastern iVIindanao
to Sibuyan Island, the latter being over 500 km south of the Palanan area.
Despite its range, X. verdugonianus has a very limited habitat and is much
sought-after for its durable timber, making it a threatened species (Yao &
Ulep 1981, 1983).
The neatly circumscissile hypanthium appears to be unique to this pair
of species. Two species in New Caledonia have hypanthia that split and are
irregularly circumscissile but these are members of an endemic group of
taxa with buUate leaves (sect. Bullata of Gugerli 1940) and do not appear
to be closely related to the Philippine species.
The species may be distinguished as follows:
1. Shrub ca. 1 m tall; lamina 6-8 cm long, 3.5-4 cm wide; petiole 7-10 mm
long; petals pink X. fruticosus
1. Tree to 26 m tali; lamma 3-7 cm long, 1-3.5 mm; petiole 4-7 mm long;
petals red X. verdugonianus
ACKNOWLEDt^MENTS
The first author is grateful to the directors of A and CANB for loans of, or
access to, herbarium specimens examined for this paper. Thanks, also, to
Lesley Elkan for the illustration and Jaime Plaza for assistance with the Spanish
abstract. The second author thanks Conservation International for funding
his field work in the Palanan Wilderness in 1991-
REFERENCES
Co, L.L. and B.C. Tan. 1992. Botanical exploration in Palanan Wilderness, Isabela Prov-
ince, the Philippines: first report. Fl. Males. Bull. 1 1 :49-53.
Gugerli, K. 1940. Monographic der Myrtaceengattung Xanthostemon. Reperc. Spec. Nov.
Regni Veg. Beih. 120:1-149.
Merrfu, E.D. 1952. Notes on Xanthostemon F. Mueller and Kjellbergiodmdnm Burret. J. Arnold
Arbor. 33:150-165.
Yao, C.E. and E.V. Ulep. 1981, Mangkono in Babatngon. Canopy 7:5.
Yao, C.E. and E.V. Ulep. 1983. More on Mangkono. Canopy 9:6-7.
SEASONAL CHANGES IN CONCENTRATION
AND DISTRIBUTION OF HEAVY METALS
IN CREOSOTEBUSH, LARREA TRIDENTATA
(ZYGOPHYLLACEAE), TISSUES IN THE EL PASO,
TX/CIUDAD JUAREZ, MEXICO AREA
WILLIAM P. MACKAY, RICHARD MENA, NICHOLAS E.
PINGITORE JR., KEITH REDETZKE, C. EDWARD FREEMAN,
HAROLD NEWMAN, JOHN GARDEA, and HECTOR NAVARRO
Laboratory for Environmental Biology and
Department of Biological Sciences (WPM RM, CEF, HN.JG, HN, KR)
Department of Geological Sciences (NEP)
University of Texas, El Paso, TX 79968, U.S.A.
ABSTRACT
We compared seasonal changes in concentrations of four elements, arsenic, cadmium,
copper and lead, in samples of various tissues of creosotebush collected in the El Paso, USA/
Ciudadjuarez, Mexico region during 1980-81 and 1994-95. Levels in leaf tissue changed
seasonally, with concentrations dropping in the spring and late fall, corresponding to the
time of leaf drop in the plant. This suggests that most of the heavy metals were simply
deposited on the surface of leaves, although data are presented which indicate that internal
tissues also have significant levels of heavy metals. These seasonal cycles were less pronounced
in the most recent samples. Levels of all four elements in cresotebush were significantly
high in the region, as compared to a distant control area. Concentrations of most elements
dropped below detection limits within 12 to 30 km from areas of highest concentration.
Densities of native Chihuahuan Desert flora and lichens are low in the area, suggesting an
apparent negative impact of industrial pollution on the local vegetation. Cadmium and
lead levels in creosotebush tissues have dropped over the past 15 years, suggesting that
enforcement and strengthening of environmental laws has reduced the air pollution levels
in the El Paso area.
RE.SIJMEN
Comparamos los cambios esracionales de las concentraciones de 4 elementos, arsenico,
cadmio, cobre y plomo, en muestras de varios tejidos de gobernadora colectados en el area
deElPasoy Ciudadjuarez, Mexico durante 1980-81 y 1994-95. Se encontro un cambio
estacional de los metales con las concentraciones disminuyendo en la primavera y otono,
que coresponden al momento de la caida de las hojas de la planta. Esto sugiere que hay
deposicion de los elementos en la superficie de las hojas, aunque presentamos datos de que
hay acumulacion de metales en los tejidos interiores. Estos ciclos estacionales estan menos
pronunciados en los ultimos anos. Los niveles de los elementos son altos en la region, pero
estan concentrados en un area entre 12 y 30 kilometros del area con concentraciones mas
altas. Las densidades de la flora y Ifquenes en el area son bajas, suginendo un impacto negativo
de lapolucion industrial en la vegetacion local. Los niveles de cadmio y plomo en los tejidos
SiDA 18(1): 287-296. 1998
-«8 SiDA 18(1)
ha disminuiclo en los ultimos 1 5 anos, lo que siii^iere que la aplicacion cle las leyes medioambientales
ha reclucido los niveles de polucion en el area de El Paso.
The El Paso/Cd. Juarez area is one of the many regions along the United
States/Mexico border which suffers from environmental contamination. The
area has been subjected to heavy industrial activity, including smelting and
hydrocarbon refining, for over 100 years. This has resulted in an accumula-
tion of heavy metals, especially lead, cadmium, copper and zinc in the local
soils (Barnes 1993; Ndame 1993). Arsenic contamination has been previ-
otisly documented to occur in this region of west Texas (Shields 1991).
These elements and their compounds can cause medical problems in humans
and other animals (Elbahri & Benromdane 1991 ; Louekari et al. 1991). Negative
effects of heavy metals on plants have been documented (D'ltri 1982; Fernandes
& Henriques 1991), including the prevention of the uptake of potassium
and phosphorous by roots. Moreover, copper may damage chlorophyll and
increase the potency of fungal diseases (Connell & Miller 1984). These ef-
fects may eliminate some plant species, with concomitant increases in the
abundances of others, thus changing plant community structure. Plants may
tolerate heavy metal contamination (Connell & Miller 1 984) by excluding
metals from sensitive tissues, modifying metabolic pathways to prevent damage
or assembling specific enzymes to detoxify heavy metals. The specific ef-
fects of heavy metals on the flora of the northern Chihuahuan Desert have
not been documented, but Worthington (1989) previously reported a re-
duction in species richness in native plants in this area, presumably the result
of such contamination.
In this investigation, tissues of the dominant Chihuahuan Desert shrub,
Larrea tridentata were analyzed, to document continued heavy metal con-
tamination in western Texas. This work is part of a larger study on the ef-
fects of heavy metal contamination on the flora and fauna of the northern
Chihuahuan Desert and the investigation of bioaccumulation into higher
trophic levels.
MATERIALS AND METHODS
We selected a site on the campus of the University of Texas (UTEP) in
western El Paso to follow seasonal change in heavy metal concentrations m
creosotebush. The site is located within 2 km of a local smelter, which ap-
pears to be a point source for the high concentrations of heavy metals in the
area. Four elements were included in the analysis: Arsenic, Cadmium, Copper
and Lead. Additional sites used to determine the distribution of heavy metals
in creosotebush tissues in the region included 62 sampling areas near the
border in the United States and six in Mexico within 10 km of the border
and ot the smelter, of which 52 yielded creosotebush specimens (five repli-
Mackay, et al., Heavy metals in Larrea cridencaca 289
cate bushes at each site). Sites were selected which had native Chihuahuan
Desert vegetation located at least 50 meters away from any road. Prelimi-
nary analyses showed higher levels ot lead next to roadways, which decreased
to background levels within a few meters, an effect also noted by Motto et
al. (1970), Gratani et al. (1992) and Lebreton and Thevenot (1992).
Leaves were stripped, and twigs were cut from the plants. Roots and trunk
tissues were also harvested from plants. Tissues were placed in numbered
paper bags and returned to the laboratory. The samples were further cleaned,
removing all foreign matter, but were not washed. Bark and exterior tis-
sues were carefully removed from pieces of trunks, to avoid contamination
of internal tissues. Tissues from 1980—1981 were collected North of the
Education Building on the UTEP campus, as part of another study (Free-
man 1982). The tissues were ground and stored dry in glass vials until they
were analyzed in 1994. Tissues from 1994—1995 were collected through-
out the season from five specific bushes, located in Charlie Davis Park, on
the campus of UTEP. This allowed a comparison of the percentages of heavy
metals in the tissues of each of the five creosote bushes.
Approximately 20 grams of tissue were placed in a crucible and muffled
for three days at 425°C. This relatively low temperature was selected to
avoid vaporizing the four elements or their salts (based on the recommen-
dations of Dr. Jim Rayon and Dr. Ken Dodson of the Environmental Pro-
tection Agency). After ashing, 100 mg of ash were dissolved in 20 ml of
14% nitric acid (V:V), without filtering. Reagent grade acid and double
glass distilled water were used for all solutions. Samples were prepared and
stored in glass scintillation vials with polypropylene-lined caps.
Samples were analyzed on a Beckman SpectraSpan 6 direct current plasma
atomic emission spectrograph (DCP-AES). We followed the protocol of EPA
method 6010 for the inductively coupled plasma emission spectrometer (US
EPA 1986). Three of the wavelengths used are those recommended by EPA:
193-696 nm for arsenic, 213.598 nm for copper, and 220.353 nm for lead.
The fourth wavelength, 228.802 nm, was substituted for the cadmium analysis
to achieve the lowest detection limit. The DCP was calibrated with pre-
pared standard solutions such that the linear calibration curve had an R- of
at least 0.995. The samples were analyzed by DCP three times and reported
the mean and standard deviation of these readings. If the relative standard
deviation was greater than 3%, the sample was reanalyzed. Blanks, dupli-
cate samples and prepared standards (spiked samples) were analyzed at least
once every ten samples for quality assurance/quality control. Cadmium
concentrations in the samples collected in 1994—1995 were close to or be-
low the limits of detection and therefore we have not presented them. Af-
ter analysis, the DCP solution values were converted to ppm in the original
290 SiDA 18(1)
ash. The detection Hmits in the ash for this study were 200 parts per mil-
Hon (ppm) for arsenic, 11 ppm for cadmium, 85 ppm for copper and 130
ppm for lead.
Data were j^lotted with SURFER (Golden Software Inc., Golden Colo-
rado), using the grid method with inverse squared distances and the sur-
fdce module (Keckler 1995).
Rr.SllLIS AND DISCUSSION
Seasonal changi^s. — Levels of all four elements varied seasonally during both
time periods (Fig. 1). Levels were high in the winter and showed a small
drop in the sj^ring when there is a combination of minor leaf drop and strong
spring winds. The levels increased during the summer, especially during
the early sampling period (Figs. 1 a, b, c), but later showed a large drop during
the fall when creosote lose a large proportion of their leaves (Mackay et al.
I 987). (Concentrations increased again during the winter months, especially
during 1980-1981 . Levels of heavy metals were lower in the 1994-1995
samples (Fig. Id & f). Cadmium and lead were both approximately 4 times
higher in 1980-1981, suggesting that enforcement and strengthening of
environmental laws has reduced the air pollution levels in the El Paso area.
Seasonal changes in concentrations of all metals during both sampling
periods were statistically significant (Table 1). The first harmonics from Fourier
analysis of the means (Little & Hills 1972), corresponding to the seasonal
effect, were significant for all years, although the patterns were more pro-
nounced during the first sampling period (Fig. 1) and had correspondingly
higher F values (Table 1). Other harmonics were statistically significant in
1980-198] , showing the importance of leaf drop in lowering heavy metal
concentrations during these years, although there was no pattern in which
harmonic was significant after the first. The lack of significance of higher
harmonics during the second sampling period suggests that leaf drop was
not significant in reducing heavy metal content during those years.
Contani'iucttion of tissues. — It appears that large amounts of the heavy metals
arsenic, copper and lead are deposited on the leaves, but heavy metals are
also incorporated into the tissues of this plant (Fig. 2). Large concentra-
tions of all three elements were found in the roots. The trunks also had high
concentrations of the three elements, which were present in the internal
tissues as well. The branches and the leaves also had high concentrations of
the three elements. These data suggest that heavy metals may be deposited
on the outside of the plant, but is also taken up by the roots and incorpo-
rated into the woody tissue.
Heavy metal geographical distribution: Levels of the four elements (ar-
senic, cadmium, copper and lead) in creosotebush ash were elevated adja-
cent to the LJnited States/Mexico border on the west side of El Paso (Fig.
Ma
;kay, I
500
400
300
200
100
<
180
160
140
120
100
^H
80
Plh
60
40
20
6000
5000
4000
3000
2000
1000
T Al,., Heavy metals in Larrea tndentata
291
1980-1981
400 -
J
1994-1995
300 -
\
1
-p T
200 -
\
I
.<^i
100 -
\i
-i^
V
'^ J_ I?
-
As
1 1 1 I
^ d)
1 1
1980-1981
uuwu
T 1994 - 199f
"
5000 ^
1
,
4000 -
I 1
3000 -
*N
k T <
1 1
Ai
\}
^^^
2000 -
1000 -
1
Cu
1
•— ir*
r~
r ^1
1
e)
1 1
Pb c)
n \ 1 1 1 1 r
FAJAODFA
1994-1995
f)
A
O D
"T"
F
MONTHS
Fir,. I: Seasonal changes in heavy metal concentrations in the leaves of the creosotebiish,
Larrea tridentata dnrmg 1980-1981 and 1994-1995. The gap in 1994-1995 is due to
lost samples. Error bars represent standard error of the mean, and are not included in a, b,
and c as many ol the data points are based on a single sample. The months are abbreviated
by a single letter on the x axis.
3). Levels of cadmium were as high as 190 ppm in ash, chose of copper reached
5200 ppm and lead levels were as high as 1200 ppm (Fig. 3, note that the
surfaces in the figures are close to average values and do not extend to these
extreme values). The highest levels of the three elements were recorded on
the east side of Mount Cristo Rey {Peak 1 in Fig. 3a), on a small mesa west
of McNutt Road (Peak 2 in Fig. 3a), and on the UTEP campus (Peak 3 in
Fig. 3a). Levels of all three elements were lower on the southwest side of
Mount Cristo Rey, perhaps due to a wind-shadow effect. Levels of all three
elements rapidly decreased to the east and fell below detection limits at
distances of between 12 and 30 km (Fig. 3).
292
SlDA 18(1)
Tabu; 1 . Resulrs oFFourier Analysis (Little & Hills 1972) of'tht' periodic tunctions in Figure 1 . Meuns
were used m the analysis.
Source
df Mean Squares
Arsenic, ] 980-198 I
F Values
First harmonic
error
Second harmonic
error
Third harmonic
error
Fourth harmonic
error
Fifth harmonic
error
2
12
2
10
2
8
2
6
2
4
108121.6
4880.5
9086.0
40.19.1
10400.7
2449.1
5426.9
1456.5
1.361.7
1503.8
22,2***
2.2 ns
4.2*
3.7 ns
0.9 ns
Cadmium, 1980-1981
First harmonic
error
Second harmonic
error
Third harmonic
error
Fourth harmonic
error
Filth harmonic
error
I'lrst harmonic
error
Second harmonic
error
Third harmonic
error
Foiu'th harmonic
error
Filth harmonic
2
8798.9
12
940.7
2
2910.9
10
546.7
2
28.6
8
676.3
2
1051.9
6
551.1
2
1627.3
4
1.1.0
Lead, 1980-1981
2
12113720.8
12
608560.6
2
180191 1.0
10
369890.5
2
292706.7
8
389186.5
2
423497.9
6
377749.3
2
732260.3
4
200493.9
9.4**
5.3*
ns
1.9 ns
[25.5***
IC) C)***
4.9*
0.8 ns
1.1 ns
3.7 ns
Arsenic, 1994-1995
First harmonic
error
Second harmonic
error
Third harmonic
error
Fourth harmonic
error
Fifth harmonic
error
2
12
10
2
8
2
6
9
19826.3
i362.4
1 159.2
3.991.7
5697.3
3309.7
1 123.4
4733.9
61 1 1.7
1978.4
5.9*
0.3 ns
1.7 ns
0.2 ns
3. 1 ns
Mackay, 1-t al., Heavy metals in Larrea tridencata 293
Tabu: I . continued.
Source df Mean Squares F Values
Arsenic, 19.S0-I981
Copper, 199i
-1995
I'irst liarmonic
2
1 168398.7
5.0*
error
12
23296.3.0
Second harmonic
2
352562.1
1.8 ns
error
10
198791.8
Third harmonic
2
231261.4
1.2 ns
error
8
185804.0
Fourth liarmonic
2
1 18186.9
0.5 ns
error
6
230882.1
Fiftii harmonic
2
267145.6
1.7 ns
error
4
Lead, 1994-
158355.2
■1995
First harmonic
2
68884.3
7.1*
error
12
9648.9
Second harmonic
2
8773.5
0.9 ns
error
10
9899.0
Tiiird liarmonic
2
8648.6
0.8 ns
error
8
10399.2
I'ourth harmonic
2
2035.2
0.1 ns
error
6
15975.2
Fiftli harmonic
2
15531.3
0.9 ns
error
4
16863.0
* = F value of harmonic
sit,'nificant ;
ut 0.05 level, **
= F significant at 0.01 level
, *** = Fsignifi-
cant at ().()() 1 level, ns -
F" vaJLie not
significant.
The areas with the highest values of As, Cu, Cd, and Pb in creosotebush
(peaks 1-3, Fig. 3) coincide with the sites of the highest concentrations of
heavy metals in the soils (Barnes 1 993; Ndame 1 993) and in fluff grass (MacKay
et al. 1998). In this locale, Pb in the soils exceeds the EPA TCLP (Toxicity
Characteristics Leaching Procedures) regulatory limit at a number of sites,
and Cd is reported quite close to the limit. The spatial correspondence between
elevated metal levels in the soil and the flora is not unexpected. The mechanism
of uptake of heavy metals by creosotebush remains to be elucidated.
Densities of native Chihuahuan Desert flora and lichens are low in the
area (Worthington 1989, MacKay et al. in prep.), suggesting an apparent
negative impact of industrial pollution on the local vegetation. The spe-
cific effects of heavy metals on the flora of the northern Chihuahuan Desert
are currently being documented. The demonstrated reduction in species richness
in native plants in this area is presumably the result of heavy metal con-
tamination from the smelter. These effects may eliminate some species and
increase the abundance of others.
294
SiOA 18(1)
I I Roots
y//////A Trunk
^^ Branches
Nxxxxxx?! Leaves
I I Bare Trunk
Fic,. 2: The- tlisrnbutions of lead, arsenic and copper in specific tissues of the creosotebush
from Ciiarlie Davis park on the University of Texas, El Paso campus. El Paso, TX.
ac:knowlf.dgments
We would like to thank the United States Department of Agriculture,
especially J. A. Vigil and J. Underwood, for permission to import plant tis-
sues from Mexico. The Texas Department of Parks and Wildlife, especially
David Riskind, allowed tis to collect plant tissues in the Franklin Moun-
tains State Park. Ken Dodson and Jim Rayon offered suggestions regard-
ing sample preparation. Our research was supported by the Center for
InterAmerican and Border Studies of the University of Texas and by the
National Science Foundation (IIRD 9253021).
REFERENCES
Barnes, B. 199.t. An evaluation of metals concentrations in siirficial soils. El PasoCotmty,
Texas. Unpublished Master's Thesis, University of Texas at El Paso.
Mackay, et Ai., Heavy metals in Larrea triclentata
295
II -i DISTRIBUTION OF ARSENIC
DISTRIBUTION OF CADMIUM
y. ^-C***
DISTRIBUTION OF COPPER
^--'■*.^
'"^l
■^g^
DISTRIBUTION OF LEAD
Fig. 3: The distribution of arsenic, cadmium, c()|")per and lead in the Ei Paso/Ciudad Juarez
area. Charley Davis Park on the UTEP campus is located at the origin in the x-y coordi-
nate system (0,0). Peak one corresponds to tiie east side of Mount Cristo Rey, Peak 2 to a
mesa above (west of) McNutt Road and Peak 3 is on the UTEP campLis. Detection limits
were 200 ppm for arsenic, 1 1 .2 ppm for cadmiimi, 84 ppm for copper and 132 ppm for
lead.
CoNNF.i.i., D.W. and G.J. Miller. 1984. Cihemistry and ecotoxicology of polhition. Jolm
Wiley and Sons, New York.
D'llr], F.M. 1982. Acid precipitation: effects on ecological systems. Ann Arbor Science,
Ann Arbor, MI.
Ei.BAHKi, L. and S. Benromdane. 1991. Arsenic poisoning in livestock. Veterinary Human
Toxicol. 33:259-264.
Fernandes, J.C. and F.S. Henriques. 1991. Biochemical, physiological and structural ef-
fects of excess copper in plants. Boc. Rev. 57:246—262.
Freeman, C.E. 1982. Seasonal variation in leal nitrogen in creosotebLish {Larrea tridentata
{DC.} Gov.: Zygophyllaceae). SouthW. JMatutalist 27:354-356.
Gratani, L., S. Ta(;li()ni, and M.F. Crescen'it,. 1992. The accumulation of lead in agricul-
tural soil and vegetation along a highway. Chemosphere 24:941—949.
Keckler, D. 1995. Surfer for Windows, Ver 6.01, Golden Software, Inc., Golden Colo-
rado.
Lebreton, L. and D.R. Tiievenot. 1992 Polkition metallique relargable par les aerosols
d'origine autoroutiere. Environ. Technol. 13:35-44.
LrnxE, T.M. and F. Hills. 1972. Statistical methods in agriculrural research. Agric. Ext.
Service, Univ. California.
296 SiDA 18(1)
LouHKAR), K., S. Valkonhn, S. Pcmsi, and L. Virtani;n. 1991 . Estimated dietary intake of
lead and cadmium and tlieir concentration in blood. Sci. Total Envnx)n. 105:87—99.
iViACKAY, W.F'., S. Silva, S. Lorinc, and W. Whitford. 1987. The role of subterranean ter-
mites in the decomposition of above ground creosotebtish litter. Sociobiohjgy 1 3:235—
239.
JVIackay, W.P., L. l^E La Rosa, M. Zavala, P. Suhton, J. Gari)[;a, R. Mi;na, N. Pinc;ii()ri;,
and G. C^RAWfORD. 1998. The distribution ot heavy metal contamination m the El Paso,
TX/Ciudad Juarez, Mexico area, usmg fluff grass, Er/one/in))i juilchelli/rti, as an indicator
organism. Proc. Chih. Desert Res. Inst. (In Press).
Morw) I I.E., R.H. Dainhs, D.M. Ciiii.ko, and C.K. Mcrrro. 1970. bead in soils and plants:
its relationship to tralfic volume and proximity to highways. Environ. Sci. Technol. 4:231 —
237.
Ndamb, E.G. 1993. Heavy metals in soils in the vicinity of the University ol Texas at El
Paso campus (El Paso, Texas). LJnpubl. Master's Thesis, LJniversity of Texas, El Paso.
Shields, J. 199 1 . Ambient air arsenic levels along the Texas - Mexico border. J. Air Waste
Manag. Assoc. 41:827-831.
LIS Environmental Proiec-iion Ac;ency. 1986. Test methods for evaluating solid waste
(third edition). Washington, D.C.
WoKiiiiNCiTON, R.D. 1989. Effects of El Paso pollutants on the lichen, moss, and winter
annual flora on andesite rock formations. In: P. Ganster and H. Walter, eds. Environ-
mental hazards and bioresource management in the United States/Mexico borderlands,
UCLA Latin American Center Publications. Los Angeles. Pp. 105 — 115.
NEW PLANT RECORDS FOR DOMINICA,
LESSER ANTILLES
STEVEN R. HILL
Center jor B wdwer.uty
Illinois Natural History Survey
607 E. Peabody Drive
Champaign. IL 61820. U.S.A.
ARLINGTON JAMES
Division of Forestry and Wildlife
Botanic Gardens
Roseau. COMMONWEALTH OF DOMINICA
ABSTRACT
Twenty-six vascular plant species are reported here for the flora of Dominica, in the Lesser
Antilles. These include two pteridophytes, 13 monocotyledons, and 11 dicotyledons. Four
of these species arc reported as new to the Lesser Antilles: Mar.ulea polycarjm Hook. & Grev.,
Dipldcriim Ccipttciliini (WiUd.) Boeck., Rhy)ichoipi)rci niceiiiosn Wright ex Sauvalle, and Phariis
lLlppnlclCt!lS Aubl.
RF.SUMEN
Se citan 26 plantas vasculares adicionales para la flora de Dominica en las Antillas JMenores.
Estas especies inckiyen dos pteridolitos, 1 3 monocotiledoncas y 1 1 dicotiledoneas. Cuatro
de estas especies se presentan como nncvas para la flora de las Antillas iVIenorcs: iWarsilea
polycarpa Hook. & Grev., Diplacr//iii aipiu/i/nii (Willd.) Boeck., Rhyrichospora racemout Wright
ex Sauvalle, y Vloarin lappulaceus Aubl.
INTROni'CTION
Dominica is a volcanic island located between the French islands of Guadeloupe
and Martinique in the Lesser Antilles. It is about 45 km long and 24 km
wide. While it is a small island (1088 sq. km), it presents a diversity of
habitats along altitudinal and moisture gradients because of its rugged
mountainous topography. Several of the mountain peaks exceed 1200 m
(MorneDiablotins 1433 m, Morne Trois Pitons 1402 m, Morne Watt 1242
m, Morne Anglais 1223 m). Water is abundant on the upper slopes of the
mountains (e.g. at Freshwater Lake, 8459 mm of rain/yr. has been recorded)
but rainfall on the western (Caribbean) coast is severely restricted by a rain
shadow (< 2000 mm/yr.) and a dry scrub forest prevails. Severe damage caused
by hurricanes and agriculture has also provided opportunities for opportu-
nistic species, increasing the diversity. The coastline is very steep, and the
SiDA 18(1): 297-305. 1998
298 Sida18(I)
cliffs generally continue precipitously into the ocean. Level land is prima-
rily restricted to river deltas in a few narrow bands near the coast, where
most citizens live.
Botanically, Dominica is rather well-known. The dicotyledonous flora
was treated by Nicolson ( 1 99 1 ) who estimated a vascular flora of 1226 spe-
cies. The most recent treatment of its pteridophytes and monocots Wiis Howard's
?,\x\o[ivm(^ Flora of the Lesser Anttlles {\91 A, 1977, 1979, 1988, 1989a, 1989b)
which included detailed distribution notes within that region, and treated
the dicots as well. Whitefoord (1989) added 40 phanerogams and six ferns
to the flora. This paper reports twenty-six vascular plant species newly dis-
covered and vouchered for Dominica.
METHODS
The plants reported here as new to Dominica were discovered during the
period 1990-1997. Approximately 2()()() collections were made by the first
author lor the purpose of assembling a reference herbarium at the Spring-
field Estate, formerly called the Archbold Tropical Research Center after
the late John Archbold who donated the property to Clemson University
in 1989. The Springfield Estate is currently leased from Clemson Univer-
sity by a Dominican non-government organization, the Springfield Centre
for Environmental Protection, Research, and Education (SCEPTRE). The
herbarium there has been informally designated with the acronym "atrc."
Specimens donated by several other collectors have also been incorporated
into the herbarium.
Collections (2005 numbers) have been made on Dominica during eight
visits to the island by Hill as follows: 20 March-26 March 1990 (collec-
tion numbers 27230-27.379), 4 March-23 March 1991 i22()()9-22]79),
26july-5 August 1992 (23924-24171), 5 March-16 March 1993 (24612-
24847, w/J. Gable & B. Dorsey), 7 December-1 9 December 1 993 (25309-
23363, w/l.Renne & D.Bradshaw), 28 May-6 June 1994 (23610-23872),
24 May-9 Jtme 1996 (27861-28262), and 23 Eebruary 1997-14 March
1997 (28868-29109, w/L.R. Phillippe). Sets of specimens have been de-
posited primarily in the Dominican herbarium (atrc), Clemson University
(CLEMS), the Illinois Natural History Survey (ILLS), and the Smithsonian
Institution (US). Additional duplicates have been distributed to herbaria
cited after the individual specimens.
New records of species generally considered to be indigenous to the Lesser
Antilles (according to Howard 1974, 1977, 1979, 1988, 1989a, 1989b)
are reported here. Some may have actually been introduced to Dominica by
human activities, but direct evidence is lacking.
Hill and James, Plant records for Dominica 299
ANNOTATED PLANT LIST
PTERIDOPHYTES
HYMENOPHYLLACEAE
Trichomanei crinitum Sw. An epiphyte similar to Trkhomafies crispum L. but
differing in its lobed or pinnatifid, not entire, pinnae. Its general distribu-
tion is northern South America to Jamaica, and it has been reported from
Guadeloupe, St. Vincent, and Grenada in the Lesser Antilles. We thank A.R.
Smith for the identification.
St. George Parish: elfin forest on windswept ridge between Breakfast River and Valley
of Desolation, 6 Mar 1997, Hill & Phillippe 29070, (atrc, ILLS, UC).
MARSILEACEAE
Marsilea polycarpa Hook. & Grev. A trailing perennial of temporary ponds,
this species of northern South America has not been previously reported in
the Lesser Antilles. It has several globose sporocarps arranged and attached
in a linear fashion on the frond stipes rather than the single basal sporocarp
seen in most other species of the genus. Neither the genus nor family have
been previously reported from Dominica.
St. Andrew Parish: locally abundant in and around temporary ponds at E end of Melville
Hall Airport, Londonderry Bay, 23 Feb 1997, Hill & Phillippe 28868 (atrc, BRIT, ILLS,
MO, NY, UC, VT).
FLOWERING PLANTS
LILIOPSIDA
AGAVACEAE
Agave dmstana Trel. The century plants are seldom collected, and none were
reported from Dominica by Howard (1979)- This species, which is endemic
to the Lesser Antilles, is very conspicuous in the dry scrub on the dry rocky
hills of the NW coast of Dominica in areas of very low rainfall, growing
mostly with cacti. Previous collections are known from St. Barts, Antigua,
Montserrat, Guadeloupe, and Martinique.
St. Joseph Parish: Morne Raquette (Rachette), Au Piton Road, 18 Dec 1993, H/7/ 2532-^
(atrc, NY). St. Peter Parish: Clement James' property, Anse a Liane trail to coast, 0.9 km
N of Colihaut, 2 Mar 1997, Hill & Phillippe 28988 (atrc, BRIT, ILLS, MO, NY, TAES,
USF, VT).
CYPERACEAE
Diplacr/iiii cctptlalnm (Willd.) Boeck. This species is rather common in Ven-
ezuela and ranges from Panama to Bolivia as well. This appears to be the
first record from the Lesser Antilles. Howard (1979) listed only Diplacrum
longifoliiim (Griseb.) C.B. Clarke ex Dur. & Schinz. for the Lesser Antilles,
from St. Paul Parish, Dominica. We thank JVL Strong for the identification.
300 SlDA IS(l)
St. Joseph Parish: Imperial Higiiway, Central I'Oresi Reserve, Dleu Gommier area, 6
Jun 199 1, ///// JXSY}^ (acre, NY, US, USF).
i'liNhristylis cymosci R. Br. subsp. spcithitceci (Roch) T. Koyama. This is a char-
acteristic, wiry-leaved and toti^di-rooted, sedge of the windswept flats of
the E and NE coasts of Dominica. Pantropical in cUstribution, it has been
collected previously on most of the Lesser Antilles, including Gtiadelc:>upe
and Martinitjue.
St. Andrew Parish: roaclsiele 0.5 mi N ol Melville Hall Airport, Londonderry Estate,
16 Mar 1993, ///// 24^42 (acre, CLEMS, MO, NY, TAES, USF).
Rhyiich(is[>i>rc/ nwei/iosa Wright ex Sauvalle. This is the first report of this species
for the Lesser Antilles. It is relatively common in the Greater Antilles. We
thank M. Strong for the identificatit)n.
St. Peter Parish: Morne Espagnol, Irom (xiasral Hii^iiway to stimmit, 4 Aiii,^ 1 992, /////
24/49 (CLEMS, NY).
ORCHIDACEAE
Brachiomdinvi parvnm Cogn. This tiny epiphyte of the upper montane rainforests
was previously known from Venezuela and from Guadeloupe, Martinique,
and Grenada in the Lesser Antilles. It was to be expected in Dominica but
probably has been overlooked because of its small size (< 5 cm tall). We
thank D. Nicolson for the identification.
St. Peter Parish: NW side of Morne Oiahlotins peak, elfin forest, 6 Mar 1993, /////
24(^47 (ariT, CLEMS). St. George Parish: Lacidat, ea. I km W of Titou Gorge, 4 Jun I 996,
///// 2HI77 (ati-e, ILLS, USE).
POACEAE
khiianthns mnioroms (Sw.) Doell. van nvartzii K.E.Rogers. Generally distributed
in the Antilles and Trinidad, this trailing rainforest species has been col-
lected previously on St. Kitts, Nevis, Montserrat, iVlartinique, St. Vincent,
and Grenada in the Lesser Antilles.
St. George Parish: Morne Anglais trail to snmmir, S of Giracidel, 1 Jun 1996, Hill
28124 (acre, ILLS).
l.\achiiccii/i:^//.stif(jlui Nash. This tiiicket-forming grass which often climbs over
other plants has been known previously from Puerto Rico, and from Guadeloupe,
Martinique and St. Kitts in the Lesser Antilles. The new record is an ex-
pected range extension. We thank G. Davidse for the identification.
St. George Parish: Morne Anglais crail at siimmic, S ol Giraudel, 2 Jun 1996, /////
2.S'077 (acre, ILLS, TAES).
Iscichne ctnnidnuiceci (Sw.) Griseb. This is another clambering grass of exposed
wet mountain slopes, and its general range is Jamaica to Trinidad S to Bo-
livia and N to Mexico. In the Lesser Antilles it has been previously col-
lected on St. Kitts, Guadeloupe, and (irenada. We thank G. Davidse for
the identification.
Hill and James, Plant records for Dominica 301
St. Paul Parish: trail to summit of Mornc Trois Pitons, N side, just NE of Pont Casse,
9 Dec 1993, HjH 23340 (atrc, iVIO, VT).
Leptochloa virgata (L.) Beauv. A very widely distributed grass which ranges
from the southern United States through the Caribbean and Central America
to South America. It has been collected on many of the Lesser Antilles, m-
cluding nearby Marie Galante and Martinique, and was to be expected on
Dominica.
St. Peter Parish: Clement James' property, Anse a Liane trail to coast, 0.9 km N of
Colihaut, 26 Feb 1997, Htll & Phillippe 28962 (atrc, ILLS, MO, TAES).
Pharus lappulaceus Aubl. This wide-ranging, wide-leaved grass is found in
forests from the Caribbean and Central America to southern South America.
Howard (1979) noted the similar?. glaberKvmth from Dominica. We thank
T.S. Filgueiras for the identification.
St. Mark Parish: Petit Coulibri,Morne Pons, 15 Mar 1993, H/7/ 24796 (atrc, CLEMS).
Phragmites amtralh (Cav.) Trin. ex Steud. Probably due to the general lack
of suitable habitat, the common reed has not previously been collected on
Dominica. Otherwise, it is widespread in both tropical and temperate por-
tions of the world, and has been collected previously in the Lesser Antilles
on Guadeloupe, Martinique, and St. Lucia.
St. John Parish: marsli at N side of Coconut Beach Hotel, Prince Rupert Bay, Ports-
mouth, 29 May 1996, Hill 27993 (atrc, ILLS).
Spartma patem (Ait.) Muhl. This grass, usually found in salt marshes and
flats along the coast, was a surprising find in Dominica where these habi-
tats are essentially lacking. The species generally ranges from the E coast of
North America from Quebec to Mexico, Central America, and the Carib-
bean. In the Lesser Antilles it has been collected previously on St. Martin,
Antigua, and Guadeloupe. We thank D. Nicolson for the identification.
St. Andrew Parish: just N of the mouth of the Londonderry (Tweed) River, Londonderry
Estate, coastal rock ledges, 14 Dec 1993, Hill & James 2547 3 A (atrc, VT).
Urochloaplantaginea (Link) Webster. Often reported as Brachtaria plantaginea
(Link) Hitchc, this grass of unstable areas ranges widely from Mexico and
the Caribbean to Brazil and Bolivia. As with Phragmites, it may be a recent
introduction, though Dominica falls within its general range. It has been
previously collected in the Lesser Antilles only on St. Lucia. We thank T.S.
Filgueiras for the identification.
St. Paul Parish: Morne Daniel Road, Canefield, 10 Mar 1993, 24701 (atrc, CLEMS).
MAGNOLIOPSIDA
COMBRETACEAE
Conocarpus erecta L. The button mangrove (buttonwood) is widely distrib-
uted in the American tropics along the coasts of Florida, USA, S through
302 SiDA 18(1)
Mexico and Central America ro the Caribbean and tropical South America
and it has also been collected in West Africa. Buttonwood has been found
on nearly all of the other Lesser Antilles. Hodge (1964) noted: "the com-
mon genera of mangrove {Rh'izophora, Aviceimia, Lagunaihma, and Conocarpus)
are absent from Dominica, a fact easily accounted for by the lack of suffi-
cient lowland sites on an island where the coastline is very precipitous."
Diligent searching by Arlington James and other members of the Division
of Forestry and Wildlife have resulted in the verification of the occurrence
on Dominica of three of these genera, two of which {Aviceuuia and Conocarptti)
are reported here as new. Of the four genera listed by Hodge, only Rhiznphora
has not yet been found on Dommica. Lagunculana was reported by Nicolson
(1991). Only one individual o'^ Conocarpus has been found.
St. Andrew Parish: Ballyhoo Ik-acii, Baribou Bay, Hampstt-ad, I I Dec 1993, Will &
Jcimes 254U. (acre, CLEMS).
CONVOLVULACEAE
Convolvnlm nodiflorus Desr. This species is widespread in dry scrub in the
West Indies, Central and South America. It has previously been collected
on the majority of the Lesser Antilles, including nearby Guadeloupe and
Martiniqtie, and was to be expected on Dominica. We thank D. Nicolson
for the identification.
St. Joseph Parish: iMorne Raquern- (Rachctte), Au Picon Road, 1 S Dec 1 993, H/7/ 25^26
(atrc, GH, MO, NY, US, USF). St. Peter Parish: S of Morne Espagnol between Bioche
and Colihaut, coastal highway, 7 Jun 1996, Hill 28248 (atrc, MO, US).
EUPHORBIACEAE
Uiiphorbui balhisii- Boiss. A species endemic to the Lesser Antilles, also known
as Chamaesyce halbisii (Boiss.) Millsp., E. balbim was previously known only
from Barbuda, Antigua, Guadeloupe, and La Desirade. It is a prostrate spe-
cies of wind-swept salt-sprayed coastal cliffs in Dominica.
St. Andrew Parish: Woodford Hill Estate, jtist E of Pte. La Sole E of an old quarry, 14
Dec 1993, ///// &Ju»m 25464 (atrc, NY).
FABACEAE
Dalbergia monetarhi L.f Generally distributed from Jamaica to trojMcal South
America, Howard (1988) reported that the species previously had been collected
in Guadeloupe, Martinique, and St. Vincent. Velez (1 957) reported this species
as present on Dominica, but Nicolson ( 1 99 1 ) dismissed this iis a misidentification.
Differing from D. ecaslaphylh/n/ (L.) Taubert (which is common on Dominica
and has unifoliolate leaves) by its pinnately compound leaves (3-5 leaflets),
it is rather common locally along the banks of the Indian river. The identi-
fication was confirmed by V. Rudd (SFV).
St. John Parish: hidian River banks, Portsmouth, 5 Mar 1 99^, ///// 6 Phillippv 29056
(atrc, BRIT, ILLS, MO, NY, US, USE, VT).
Him. and Ja.\[F,s, Plant records for Dominica 303
MALVACEAE
Si-da jamaicensis L. This wide-ranging species is known from Mexico, the
Caribbean, Central America, and northern Soutli America. In the Lesser Antilles
it has previously been collected on St. Barts, Antigua, Montserrat, Guadeloupe,
and Les Saintes in the dry scrub forests. We thank Paul Fryxell for the iden-
tification.
St. Joseph Parish: Morne Raquetre (Rachette), Au Piron Road, 4 Jun 1994, Hill 2580H
(atrc, NY, TEX, USF).
NYMPHAEACEAE
Nymphaea amazonum C.Mart. &c Zucc. This night-flowering water lily ranges
throughout the Caribbean and from Mexico to southern Brazil. It previ-
ously has been collected in Guadeloupe, Marie Galante, Martinique and St.
Lucia in the Lesser Antilles. Suitable habitat for the species is very scarce
on the island. We thank J. Wiersema for the identification.
St. Andrew Parish: roadside 0.5 mi N ol Melville Hall Airport, Londonderry Estate,
16 Mar 1993, Hill 24844 (atrc, CLEMS).
OLEACEAE
Forestiera segregata (Jacq.) Krug & Urb. Another new discovery in the dry
scrub forests of western Dominica, this species ranges from the coastal SE
USA, S to the Lesser Antilles. It has been collected previously on Anguilla,
Barbuda, Antigua, La Desiracle, and Marie Galante in the Lesser Antilles.
We thank A. Goldberg and D. Nicolson (US) for the determination.
St. Joseph Parish: Morne Raquette (Rachette), Au Piton Road, 18 Dec 1993, Hill 25530
(atrc, GH, NY, US).
RUBIACEAE
Randia nitida (H.B.K) DC. This spiny coastal shrub is distributed from Mexico
to South America and the Caribbean. In the Lesser Antilles it has been re-
ported from Martinique, St. Lucia, St. Vincent, and Grenada. It is distin-
guished from the more frequent /?. aculeata L. by its flower size (3-.3.8 cm
vs. 2 cm long in R. aculeata) and fruit size (3.2-5 cm vs. 1-1.2 cm long in
R. aculeata). We thank D. Nicolson (US) for the identification.
St. Paul Parish: Rodney's Rock on the coast, Hertford Estate, 1 3 Dec 1 993, Hill 23423
(atrc).
RUTACEAE
Pilocarpus race?nosus Vahl. This Caribbean endemic ranges from Cuba to the
Lesser Antilles. It has been collected previously on Montserrat, Guadeloupe,
and Martinique in the Lesser Antilles and was to be expected on Dominica.
It is restricted to windswept coastal forests and is associated with the palm
Rhyticocos on the N coast of Dominica. We thank D. Nicolson for the iden-
tification.
304 SiDA lcS(l)
St. Andrew Parish: clitLs at W side of Bailyboo Ik-acli, BatiboLi Bay, 1 lampsreatl, 16
Mar 1993, Hi// 24823 (atrc, CLEMS, NY, TAHS, US, LJSF); same locality, M Dec 1993,
///// & James 25433 (CLRMS).
VERBENACEAE
Avicennia germinans (L.) L. The black mangrove was a particularly interest-
ing find on Dominica. While it was thought to be absent by previous au-
thors (see comments under Cimocarpiis, above) it was located by the second
author in recent years at several places, the most unusual one in elevated
cliffside sedge-dominated seeps on the NE coast of the island. Other popu-
lations are restricted to a few square meters of level land at the mouth of
small rivers. Like Couocarpi/s, Ai'icennia is widely distributed along coasts
from Florida and Texas, USA, to northern South America. It has been found
on nearly all of the other Lesser Antilles.
St. David Parish: mouth of the Ouayaperi River, Corossol Beach, 0.6 km N of La Plaine,
12 Dec 1993, Hi// 23390 (atrc, CLEMS, BRIT, ILLS, MO, NY, US, USE, VT). St. An-
drew Parish: Wooclhird Hill Estate, just E of Pte. La Soie E of an old quarry in sedge
seeps, M Dec 1993, ///// OJuwes 254C^9 (atrc); motith of Eden River, Eden Estate, S of
Crompton Point, 11 Dec 1993, Hi// & James 23473 (atrc, GH, NY, US, LJSF).
VITACEAE
Ctssus obovata M. Vahl. A vine endemic to the Caribbean, C. obovata is eas-
ily distinguished from the more common C. vertiallata (L.) Nicolson & Jarvis
by its trifoliolate rather than simple leaves. It ranges from Cuba, S to the
northern Lesser Antilles. Collections previously have been made in the Lesser
Antilles on St. Martin, St. Barts, and Barbuda.
St. George Parish: head ol trail to Trafalgar Falls, Trafalgar E of hydroelectric facility,
25 M-ay 1996, f//// 27<S7 / (atrc).
ACKNOWLF.DCMIiNTS
We would like to acknowledge the continuing kind cooperation and support
of the Ministry of Agriculture of the Commonwealth of Dominica, and es-
pecially to the helpful staff of its Division of Forestry and Wildlife. Finan-
cial support ior this research has been obtained primarily through various
units of Clemson University (particularly the Archbold Tropical Research
Center), Texas A&M University, and The LJniversity of Maine at Orono.
The senior author was also supported by grants from the Illinois Depart-
ment of Transportation (IDOT) and the Critical Trends Assessment Pro-
gram (CTAP). Special thanks are extended to the late John Archbold, whose
generosity made the research possible, to Dan Nicolson and his colleagues
at the Smithsonian Institution lor their generous expenditure of time in
working with several of the identifications, to our other colleagues (cited
in the text) who assisted with identifications and collecting, and to Dr. Thomas
Lacher, former Director of the Archbold Tropical Research Center, for his
assistance and commitment throughout this study.
Hill and James, Plant records for Dommica 305
REFERENCES
HODCE, W.H. 1954. Flora of Dominica, B.W.I. Parr 1 , Lloydia, 17:1-238.
Howard, R.A.I 974. Flora of the Lesser Antilles, Leeward and Windward Lslands: Orchidaceae.
Arnold Arboretum, Harvard University, Cambridge, MA.
Howard, R.A. 1977. Floraof the Lesser Antilles, Leeward and Windward Islands: Volume
2 Pteridophyta. Arnold Arboretum, Harvard University, Cambridge, MA.
Howard, R.A. 1979. Floraof the Lesser Antilles, Leeward and Windward Islands: Volume
3 Monocotyledoneae. Arnold Arboretum, Harvard University, Cambridge, MA.
Howard, R.A. 1988. Floraof the Lesser Antilles, Leeward and Windward Islands: Volume
4 Dicotyledoneae-Part 1. Arnold Arboretum, Harvard University, Cambridge, MA.
Howard, R.A. 1989a. Flora of the Lesser Antilles, Leeward and Windward Islands: Vol-
ume 5 Dicotyledoneae-Part 2. Arnold Arboretum, Harvard University, Cambridge, MA.
Howard, R.A. 1989b. Flora of the Lesser Antilles, Leeward and Windward Islands: Vol-
ume 6 Dicotyledoneae-Part 3. Arnold Arboretum, Harvard University, Cambridge, MA.
NicoLSON, D.H. 1991. Flora of Dominica, Part 2: Dicotyledoneae. Smiths. Contr. Bot. 77:1-
274.
Velez, I. 1957. Herbaceous angiosperms of the Lesser Antilles. San German, Puerto Rico.
Whitefoord, C. 1989. Recent plant collections from Dominica. J. Arnold Arbor. 70:l43-
152.
A NEW ADDER'S-TONGUE (OPHIOGLOSSUM:
OPHIOGLOSSACEAE) FOR NORTH AMERICA
JAMES C. ZECH and PATRICIA R. MANNING
Department of Biology
Sul Ross State University
Alpine, TX 79832. U.S.A.
WARREN HERB WAGNER JR.
Department of Biology
University of Michigan
Ann Arbor. All 48109. U.S.A.
ABSTRACT
The widespread Old World adder's-tongue, 0/)h/oi^/oss//w poiyphyttum A. Br. is reported
for the first time in Nortli America. Formerly confused with 0, engelmannii Prantl, we now
recognize as 0. piilyphyll/nn collections from U.S.A. (T-xas and Arizona) and Mexico (Hidalgo,
Oaxaca, San Luis Porosf, and Zacatecas). The earliest confirmed collection was in 1<S74.
The two species are keyed out and illustrated, and specimens of 0. polyployllum are listetl.
The known county distribution of the species in Texas is mapped, and the habitat and phenology
are briefly described.
RESUMnN
La lengua de serpiente del Viejo Mundo, muy frecuente, Ophioglossiim polyphylli/Di A. Br.
se cita pot primera vez en Norte America. Previamente confundida con 0. engelviannii Prantl,
reconocemos ahora como 0. pulyphylliiin colectas de E.E.U.U. (Texas y Arizona) y Mexico
(Hidalgo, Oaxaca, San Luis Potosi, y Zacatecas). La primera colecta confirmada fue en 1874.
Se ofrece tma clave de identificacion y se ilustran las dos especies, y se listan los especimenes
de 0. polyphylli/m. Se cartografia la distribucion conocida per condados en Texas, y se describen
brevcmente el habitat y la fenologfa.
The adder's-congues, Ophioglossum (Ophioglossaceae) are a small group
of striking and unusual species, with the highest known chromosome numbers
in the plant kingdom. The gametophytes are subterranean, the roots lack
root hairs, and the leaf is made up of two parts: the sterile blade portion
(trophophore) and the spore-bearing portion (sporophore). The trophophore
has complex reticulate venation and is believed to be phyllodial in origin
(Wagner 1979). The sporangia are deeply sunken in the apical portion of
the sporophore. There are probably as many as 35 species worldwide but
two areas, India and Africa, seem to be regions of maximum diversity. Many
of the species occur in open arid sites and remain underground until sea-
sonal or sporadic rains stimulate the leaves to push up through the soil and
SiDA 18(1): 307-31.^. 1998
308 SiDA ISd)
grow CO full size before discharging the spores. In America norrh of Mexico
seven terrestrial species are traditionally recognized: Ophioglossinii piisiUnni
Raf., 0. vNli^cttuDi L., 0. ccilijoruiciini Prantl, 0. uitdicaiik L.h, 0, pdioUitiim
Hook., 0. crotalophoroules Walter, and 0. engelvunDi'ii Prantl. However, we
have recently discovered another species heretofore unknown from the New
World. We originally interpreted the plants in question as a narrow form
of 0. engelmannii (Zech & Manning 1996), but subsequent study reveals a
number of differences from that species, and we now identify it confidently
as 0. polyphylliim A. Br. The earliest collection was in 1874. Clausen's monograph
(1938) does not even recognize this species anywhere in the world; he merely
synonymized it with 0. vulgatum. Nevertheless, we have come a long way
taxonomically from the first half of this centtiry, not only in regard to
Ophioglossaceae, but Lycopodiaceae, Isoetaceae, Thelypteridaceae, and many
other pteridophyte groups. Lellinger's (1985) manual of North America
pteridophytes does not list 0, polyphyllum , nor does Flora North America
north of Mexico (Wagner & Wagner 1993).
This species has an enormous range: Africa, Asia (e.g., S. China, India),
Polynesia (e.g. Hawaii), and now North America. It is probably made up of
a number of subspecies, but these are held together by a set of distinctive
characters, described below. There is no question that it is most closely re-
lated to 0, eugelnunin'ii. Its main point of resemblance involves rhe venation
pattern in which the major areoles include minor areoles, a condition com-
monly referred to as bireticulate. In most species the major areoles of adder's-
tongues possess only free included veinlets that themselves do not inter-
connect. Large specimens of 0. uudicaule (the form known as 0. dlipt'icmn
Hook. & Grev.) usually possess bireticulate veins (Wagner et al. 1984), as
does rhe Old World 0. costcitmu A. Br. (Burrows 1996). Both Oph/ogioss//w
viilgcitu))t and 0. ydiciilatiini show a tendency toward bireticulation, but the
relatively few areoles are coarser and more open. The very fine bireticulation
in 0, oigelnuinu'ii and 0. polyphyllum is cjuite distinctive.
Ophiogloss/nii eiigeln/ciiinii is apparently a strictly New World taxon, and
limited to North America, including Mexico, and Central America. It is
commonly referred to as "Limestone adders-tongue" for its tendency to grow
on soil layers above limestone. In the United States it tisually appears dur-
ing and after rains, but it may reappear in the same places during later high
precipitation periods. Dean P. Whittier of Vandcrbilt LIniversity has actu-
ally forced a new "brood" of leaves to form in late summer by watering a
site where the plants normally appear only in the spring (pers. comm.).
A key to the two species is presented below. To see the venation of a dried
specimen more clearly, a drop of 95% alcohol placed on the laminar surface
will, after soaking in, clear the leaf temporarily and will not damage the
ZncH, ET AL., A new Ophioglossum for North America 309
Specimen. Collectors should be warned not to break off or remove the char-
acteristic elongate brown leaf sheaths that surround the green leaf bases:
collecting can be accomplished best by carefully digging up the specimens.
This has the added advantage of showing the roots, and their numbers and
relative sizes.
Roots 25(15-30), thinner, mostly 0.6(0.3-0.9) mm thick, yellow to dark brown,
commonly somewhat crooked (dried); persistent old leaf bases numerous, 5
(2—9); trophophore narrowly ovate, attenuate proximally and distally, length/
width 4, 4.8(3.2-7.7) x 1.2(0.7-1.9) cm; sporangial clusters 1.8(0.6-2.6)
X 0.25(0.2 -0.3) cm; apicula 1.0(0.3-1 .<S) mm O. polyphyllum
Roots 15(6-24), thicker, 0.9-1.1(0.3-2.0) mm, often darker or blackish, usu-
ally straight (dried); persistent old leaf bases few 2(1—4); trophophore
broadly ovate to oblong, length/width = 3,6.0(4.5-8.5) x 2.0(1 .5-2.5); sporan-
gial clusters 2.5(2.0-4.0) x 0.25(0.2-0.4) cm; apicula 0.6(0.0-1.3) mm.
O. engelmannii (Fig. 1)
The county distribution oii OphiogUnsmn polyphyllum in Texas is shown in
Figure 2 (triangles) in comparison with 0. engelmanni {dots). The map shows
a dramatic difference in range between the two species. Most of the collec-
tions of 0. polyphyllum were made along U.S. Highway 90 or on roads nearby:
in Brewster County near Alpine and Marathon; in Jeff Davis County near
Valentine; and in Presidio County near Marfa. On the basis of our experi-
ence, we believe that an effort ought to be made to explore much further
afield in other Texas (as well as New Mexico and Arizona) counties, e.g.,
Hudspeth, Culberson, Reeves, Ward, Pecos, and Terrell in Texas, and also
areas to the south in Mexico (e.g., Coahuila, Chihuahua, and Sonora states
and southwardly adjacent states).
There are many reasons why 0. polyphyllum was not heretofore collected,
among the most important being its occurrence among grasses, the shape
of the leaf resembling some monocots, and especially the very short period
of appearance above ground, which after strong rains probably rarely ex-
ceeds two weeks, the latter week presenting only dying straw colored and
withered blades.
Plants are found in shallow ditches and troughs along the roadsides (Fig.
3). The adjacent landscape is open rangeland. While no plant species occur
consistently with 0. polyphyllum, the two most prevalent are the grasses Buchlbe
dactyloides and Bouteloua curtipendula. Other, less consistent associates are Bouteloua
gracilis, Chloris verticillata, Hilaria mutica, Panicum obtusatum, and Bothriochloa
sp. Among the forbs are Asclepias oenotheroides, Berlandiera lyrata, Cirsium
texanum, Cooperia drummondii, CrotonVott'3Jn,Engelmanniapinnatifida, Ratibida
columnaris, and Thymophylla sp. The vegetation is typical Chihuahuan Desert
grassland, and includes such cactus genera as Coryphantha, Echinocereus , and
Opuntia. The substrate involves various clays that are usually deep and allu-
^^ 1
SiDA 18(1)
F1C7. 1. A semi-cliagranimatic comixinson ot (). jyil)ph)H/iiii (It-Fr) and 0, c^/^t^'t/wi/ww// (right).
Scale bar = 2 cm.
vial. Plants within ackler's-tongue colonies vary from several to hundreds.
Vegetative reproduction is accomplished by horizontal roots, the prolifera-
tions producing plantlets as far as ten centimeters from the genet.
Our records outside of Texas are very few, and all were previously identi-
fied as 0. engelmannii. In Arizona it was found in "mesas around the Mus-
tang Mts." The habitats in Mexico (what little is recorded on the labels)
appear to be more or less similar to those m Texas. The Rollins and Tryon
collection listed below was found in "clay soil over limestone, in shade of
small arborescent Opiiiitut in Larrea and Prosopis desert," and the Reeder et
Zech, et al., a new Ophioglossum for North America
311
Fic. 2. Presently confirmed cotinty records in Texas for 0. po!yphyll//ii/ (:riant,des) and O.
vugelmannii (dots).
al. collection in "thorn savannah in the open among E/ahloe dacty bides and
Bouteloua stolonifera." The Pringle and Purpus Mexican collections lack habitat
data.
Collection citations: U.S.A. ARIZONA: MustanL; Mountams, 13 Sep, C. G. Pringle in
1884, (GRAY). TEXAS. (Collections made along Hwy 90 and deposited in SRSC, tmless
otherwise indicated. Brewster Co.: 5.5 mi E ofMarathon N of Hwy 90, 7 Jun 1995, Manning
896; 5.9 mi W ofMarathon S of Hwy 90, 20 Jun 1995, Manning 897; 6.5 mi E of Alpine
Sof Hwy 90, H Jul 1993. Manning 914; 12 mi W of Marathon S of Hwy 90, 5 Jul 1995,
Mann/ng s.n.; 6.5 mi W ofMarathon N of Hwy 90, 24 Jun 1996, Mann/ng 959; 5.7 mi E
ofMarathon, 1 1 Jun 1995. A.M. Powell & S.A. Powell 6085; 6.0 mi E ofMarathon N of
Hwy 90, 1.^ Jun I 997, Manning 1013; 17 Jun 1997, Manning & Zech 1017 A (xVIICH); 24
Jun 1997, Manning & Zech 10I7B (MICH); 6.2 mi E ofMarathon S of Hwy 90, 14 Jul
1997, Hansen et al. 40; 6.3 mi S of Brewscer/Pecos Co. line, 30 Jun 1997, Manning 1034.
Jeff Davis Co.: 7.8 mi W of Valenrine E of Hwy 90, 1 I Jul 1995, Manning 915; 25 Jun
1997, Mann/ng & Zech 7 020 (MICH); 0.2 mi SE of 505/166 jct. on loop 166, 14 Aug 1971,
Ke/l 1699 (GRAY); 25 Jun 1997, Manning & Zech 1031 (MICH). Presidio Co.: 0.1 mi E
of Marfa, 26 Jul 1995, Manning 925.
^12
SiiiA 18(1)
, ■-4.."BT-1|i*;'^^,
jsft
.0. ■'^Rk"*'"^^
1^
*•••
Fici. 3. Habitat of 0. polylihylli/in in sliallow roadside ditch. TX: )eff Davis C^o., alon^ loop
166.
MEXICO. Hidalgo: Hill, near El Ciallo, 7{)0() ft, 17 Aug 1 902 C.G. Pniii^li(GU). Oaxaca:
Los Maranjos, vicinity of'San Lins Ttikiclanapa, Ptiebla, near Oaxaca, May 1908, C.A. Purpm
3152 (GH). San Luis Potosi: 15 mi NE of San Luis Potosf, 6500 ft, 4 Sep 1961, _/./<?.
Reeckr. C.G. ReeJer. & T.R. Sodentrom 3.^00 (GH). Zacatecas: 32 mi N of Fresnillo, 25 Nov
1958, 7000 ft, R.C. RoUim ,niJ R.M. Tiyw (GH).
There is no easily projecred certainty about when the plants will appear.
Continuous clrout^ht will prevent the leaves from expandin^^ and extending
above the soil. However, the best months in general arejune and July, whenever
several rains occur. Apparently, chance rainy periods at any other time, however,
will bring forth an "out-of-season" growth, and we now have records from
May, August, September, October, November, and December. Our impression
is that 0. polyphyllnni. "lies in wait" for rain, emerges through tightly com-
pacted soil that has been softened by the water, and then has a fairly short
time span to reproduce and disperse spores. However, based upon green-
house observations, even when water availability is not a factor, trophophores
will tend to die back following spore production and dispersal.
It should be noted that 0, polyphyllimi is ecologically a "loner," in the
sense that it does not occur microsympatrically along with other species of
Ophioglos.u/iv (or Botrychtum). Other Ophioglossaceae are noted for forming
"genus communities," members of the same or closely related genera that
ZncH, ET AL., A new Ophioglossum for North America 313
co-occur syntopically (Wagner & Wagner 1 983). To illustrate the usual situation,
in the United States in the region from the Carolinas to Arkansas and southward,
we find 0, nudkaule. 0. crotalophoroides , and 0. petiolatum (plus the ophioglossaceous
Botrychium bitematum and B. lunanoides) together in the same habitat, sometimes
side by side and intermixed with each other. But this is evidently not the
case with 0. poly phy Hum according to Burrows (1996) who writes that in
Africa O.polyphyllum is "The one species oi Ophioglossum that is rarely found
as a component of genus communities."
ACKNOWLEDGMENTS
This study could not have been completed without the help of several
herbaria, including Gray Herbarium, Harvard (GH), Lundell Herbarium
(LL), University of Michigan (MICH), University of California Berkeley (UC),
New Mexico State (NMS), Sul Ross State (SRSC), Southwest Texas State
(SWT), Texas Tech (TTC), University of Texas at El Paso (UTEP), Botani-
cal Research Institute of Texas (BRIT), Texas A&M University (TAMU),
and the help of Christopher Ruggia who kindly provided the distribution
map. Alan R. Smith and R.D. Thomas made useful suggestions for the
manuscript.
REFERENCES
Burrows, J. E. 1996. The genus Ophioglossinii L. in southern central Africa. In: J.M. Camus,
M. GiBBY, and R.J.Johns, eds. Pteridological Perspectives. Royal Bot Gard. Kew. Pp..329-336.
Clausen, R.T. 1938. A monograph of the Ophioglossaceae. Mem. Torrey Bot. Club. 19: 1-177.
Clute, W. 1901 . Our ferns m their haunts. F. Stokes Co. NY.
Lellinger, D.B. 1985. A field manual of the ferns & fern allies of the United States & Canada.
Smithsonian Institution Press, Washington, DC.
Wagner, W.H. 1979- Reticulate veins in the systematics of modern ferns. Taxon 28:87-95.
Wagner, W.H., CM. Allen, and G.P. Landry. 1 984. Ophiogloisnni ellipticum Hook. & Grev.
in Louisiana and the taxonomy of 0. nudicauk L. f. Castanea 49:99—1 10.
Wagner, W.H. and F.S. Wagner. 1983. Genus communities as a systematic tool in the
study ot New World Botrychium (Opiuoglossaceae). Taxon 32:5 1—63.
Wac;ner, W.H. and F.S. Wac;ner. 1993- Ophioglossaceae C. Agardh. Adder's-tongue Family.
In: Flora of North America Editorial C^ommittee, eds. 1993 <■. Flora of North America
north of Mexico, 3+ vols. New York anti Oxtord. Vol. 2, jip. 85—106.
Zech, J.C. and PR. Manning. 1996. Opbioglossi/m engelmannii in Trans-Pecos, Texas. Amer.
Fern J. 86:102.
NOTES ON THE FLORA OF TEXAS WITH ADDI-
TIONS AND OTHER SIGNIFICANT RECORDS
LARRY E. BROWN
Houston Comvuinity College
1300 Holman. Houston. TX 77007, U.S.A.
and
Spring Branch Science Center Herbarium
8856 Westvieiv Drive
Houston, TX 77033. U.S.A. plantman@flash.net
STUART J. MARCUS
Trinity River National Wildlife Refuge
R 0. Box 10013
Liberty. TX 77373. U.S.A. r2rw_tr@fws.gov
ABSTRACT
The following 15 taxaaredocLimentctI new to Texas: Alterriantlxra ses.ulis, Cyperiis ercigroUis,
Urochloci mmosa, Lindernia criistaceci, Phyllciiith/is Irdttriu/s, Stellnria parva, Alysiairp/is vcigimdis,
Cardamim dehilis, Rhynchospora ckhU'n, Lilaeopsis chinensis, Rumex paraguayensis, Saaharum
coarctatum, Utriadaria foltosa, Rolygoniini mehnenaninn, and Cuiciita polygononm. Significant
new collections records are provided for five: Sarp/is c/iheiue. Polygonum cespitos/ini, Ployllanthi/s
iiiyiiri, Ludwtgia m'uTuairpci, ixnA Aruionnhi lacintris. Data are presented to c]i.iestion the presence
oi Eleodoaris compressa in Texas. Taxa are listed alphabetically by himily, genus, and species
with annotations and citation of voucher specimens.
RESliiMF.N
Los sigLiientes 1 5 taxa se citan como nnevos para Texas: Alternaiitlxra sessilis, Cyperus eragrostis,
LJrocloloci ramosa, Lindernia irusUicen, Pl)yllcintl}//sjraternus, Stdlarici parva., Alysicarpus vciginalh,
Cardamine de!)i!is, Rl?ynchospora dehilis, Lilaeopsis chiriensis, Rumex paraguayensis, Saaharum
coarctat//iii, Lltric/daria foliosa, Polygoni/m meisiierniuum, y Cuscuta polygonorum. Se ofrecen nuevas
citas importantes para cinco: Scirpus cuk'use, Polygonunj cespitiisum, Phyllanthus niruri, Ludivigia
microcarpa, y Armoracia lacustns. Se presentan datos |-)ara ctiestionar la presencia de Eleocharis
compressa en Texas. Los taxa estan listados altabeticamente por familia, genero, y especie
con anotaciones y citas de los pliegos tesrigo.
INTRODUCTION
Floristic work in the Houston area has resulted m a mimeographed plant
checklist for: Brazos Bend State Park, Fort Bend County by Larry E. Brown,
Frank Gregg, and Kay Lewis; Davis Hill State Natural Area, Liberty County
by Larry E. Brown, Charles D. Peterson, and Joe Liggio; and the entire Houston
area (Harris and adjacent counties) by Larry E. Brown. In addition, the present
authors have just completed a plant checklist for the Trinity River National
SiDA 18(1): 315-324. 1998
316 SiDA 18(1)
Wildlife Refuge, Liberty County. This work coupled with plants sent to
the SBSC^ herbarium for identification has revealed a number of plants new
to Texas and significant new records for others. Some of these records were
provided to Stanley D. Jones prior to the publication of his book, Vascular
Plants oj Texas (Jones et al. 1997). Here we give data to support inclusion of
these plants in this new publication.
Unless otherwise indicated, all collections cited in this paper are in the
Spring Branch Science Center Herbarium (SBSC), Houston, Texas. Dupli-
cates of some collections are at TEX or TAES. Herbarium acronyms follow
Holmgren etal.( 1990).
AMARANTHACEAE
Alternanthera sessilis (L.) DC. — A few plants of this pantropical and federal
noxious weed were found growing in the bottomland of the Trinity River
hi the Davis Hill State Natural Area; 4 Jul 199-1, Brown 18040. Clewell
(1985) reported it for Escambia County in the Florida Panhandle and Tho-
mas and Allen (1996) mapped it for ten, mostly central, Louisiana parishes.
Nelson et al.(1997) reported it new for South Carolina and cited published
reports for its occurrence in Alabama and Mississippi. The key in Mears
(1977) can distinguish the Texas species oi Alternanthera, including A. sessilis.
APIACEAE
Lilaeopsis chtmnsis (L.) Kuntze — Affolter (1 985) mapped this species along
the Atlantic and Gulf Coast from Nova Scotia south to Florida and west-
ward along the Gulf Coast to Louisiana (one collection). Thomas and Allen
(1996) mapped it for eight coastal parishes west to Vermillion Parish. The
only Texas collection is from Galveston Bay below the Abshire House in
the Abshire Wildlife Management Area at Smith Point, Chambers County,
7 May 1993, Brown 16926. These plants were submerged at high tide and
exposed during low tide.
BRASSICACEAE
Arn/oracia lacustris (A. Gray) Al-Shehbaz & Bates (lake cress) — Correll and
Johnston ( 1 970) did not report this white flowered aquatic crucifer for Texas.
The first record appears to be in Al-Shehbaz and Bates (1987) where it is
mapped in Tyler County. We found it to be infrequent in standing water in
the Davis Hill State Natural Area, Liberty County (3 1 May 1996, Brown &
Liggio 19172) and in the bottomland of the Trinity River National Wild-
life Refuge, Liberty County (26 Apr 1 997, Brown & Marcus 20262). These
plants produced numerous flowers but had poor fruit development. Mo-
lecular studies based upon DNA (Les 1 994) indicate that lake cress should
be placed into a monotypic genus Neoheckia as N. aqnatica (Eaton) Greene
and that the cause for a poor seed set is suggested by the discovery of a trip-
loid chromosome number of 2)i = 2H (Les et al. 1995).
Brown and Marcus, Notes on the flora of Texas 317
Cardamine deUlis D. Don — Plants tentatively identified as this non-native
species have been collected in and around nurseries, greenhouses, and yards in
the Houston area. These are similar to the native C. pennsylvanka Willd. and
another introduced species mostly present m the norteast U.S. and Canada, C.
flexuosa Withering. Rollins (1993) separated C. debilis from the other two species
by its fibrillose roots and fruits <1 mm wide. Botanists may wish to reex-
amine their collections, especially those identified as C. pennsylvanka, to see if
they may fit the description of C debilis. It also would be helpful to search for
additional characters to separate these three similar species. Lipscomb (1978) cited
the Shinners collection as a voucher for the presence of C pemtsylvanica in Texas.
Voucher specimens: Dallas Co.: weed under lath, mostly along walks in clay, Northaven
Gardens, north Dallas, 6 Nov 1 960, Sh'niiwn 291 54 (TEX). Dewitt Co.: in Sandies Creek,
2 mi E of Westhoff, 5 Mar 1995, D. Musthaiek j.«. (SBSC). Galveston Co.: weed in nurs-
ery in Kemah, 1 Apr 1975, Waller & Baiiiiil 3460 (TEX). Harris Co.: weed around the
greenhouse in the Mercer Arboretum and Botanical Gardens, off of Aldine Westheid Rd,
I Nov 1997, Brou'/i 21713 (SBSC); growing in pots at the Edith Moore Sanctuary off of
Memorial Avenue in Houston, 27 Mar 1992, Bri)U7/ I y86H (SBSC); weed at edge of San
Jacinto Bldg at HCCS Central College, 1300 Holman Street in Houston, 25 Nov 1997,
Brown 2 1742, (SBSC).
CARYOPHYLLACEAE
Stellaria parva Pedersen — Landry et ai. ( 1 988) reported this chickweed new
to North America from Acadia and Jeff Davis Parishes in Louisiana. Landry
et al.(I989) provided a more complete discussion of its presence in Louisi-
ana and cited an additional collection from St. Landry Parish. Thomas and
Allen (1996) added Lafayette Parish to the Louisiana distribution. We dis-
covered this species on a few disturbed sandy sites in the Trinity River National
Wildlife Refuge, Liberty County, 1 1 Mar 1997, Brown & Marcus 20104; 6
Apr 1997, Brown & Marcus 20216. Since the original description is from
Argentina in I96I and the first Louisiana collection is in 1966, Landry et
al. (1989) suggest it may have arrived in Louisiana and Argentina from an
unknown native source. However, they failed to note that Pedersen (1961)
in the protologue reported collections from Argentina and Paraguay in 1869
and 1872 and up to 1957.
CUSCUTACEAE (CONVOLVULACEAE)
Cuscuta polygonorum Engelmann — Correll and Johnston (1970), Hatch et
al.d 990), Johnston (1990), and Jones et al. (1997) all include this dodder
for Texas, apparently from Yuncker (in Lundell 1 943), who included it fide
Small. Even though Small (1933) gave its range from Maryland to Texas,
the first Louisiana collection of it was in 1979 with a second collection in
1 98 1 (Gandhi & Thomas 1983; Gandhi et al. 1987). The first Texas collec-
tion appears to be from Brazos County where it was growing on smartweed
in the first bottomland along Highway 30, east of the Highway 6 bypass of
Bryan/College Station, Sep 1996, Cheathaiu. Marshall & Jones s.n.
31S SiDA 18(1)
CYPERACEAE
Cyperus eraf^rost/s Lam. — Tucker ( 19H7, 1994) reported char this species has
become naturahzed in southeastern Texas but he cited no collections nor
furnished details about its status in Texas. Tucker ( 1 9(S7) indicated it is native
to the Pacific (x)ast ol the U.S., a wail in South Carolina, and naturalized in
southern Europe. It is similar to C. ochvaceiis Vahl and may be under this
name in otlier Texas herbaria. The key in Denton ( 1 978) can separate these
two taxa and other similar species. The larger number of collections from
LaPorte near the Barbours Cut Terminal of the Port of Houston suggest it
may have arrived in the Houston area by ship.
Voucher specimens: Galveston Co.: ca. 100 m Irom GuifOf Mexico on Iiast Beach in
Cialvesron, 1 1 Jul 1 981 , A't'j'.i7t'/--^6J'/ (TAES). Harris Co.: in roadside diich olBarrleground
Rd, ca, 0.5 mi S ol Hwy 225,18 Jiin 198i, llruini 1201 (SBSC); in the Marshall Tract of"
the Ciompaq Computer Corp at the intersection of I Iwy 249 and Louetta Rd, S of Tomball,
23 May 1992, Brown 16104 (SBSC); weedy plants in the Park Harbor development along
Barker-Cypress Rd N ot Addicks Reservoir, W oi Hotiston, May 1 99 1, collector unknown
(SBSC); a single plant in a prairie at end of Ramsey Rd, S of Keith Rd, E of Beltway <S in
Pasadena, 1 Jun 199-1, Brotn/ /79^2 (SBSC); small population on verge of Barbours Cait
Rd between hwy li6 and Barbours Cut Terminal on Galveston Bay in LaPorte, 16 ]im
199-1, Broun ISOJJ (SBSC); frequent in ditch along Hwy 1 16 on W side across from en-
trance to Barbours C^it Rd in LaPorte, 27 Jtm I 994, Brintii IS023 (SBSC); weedy plant at
the Texas Parks and Wildlile Department office, near business Hwy 146 in LaPorte, 24
Jan 1 995, Brown 1 1^403 (SBSC]); in low waste areas along Southern Pacific Railroad Tracts
just L of Llwy 146 in LaPorte, S of the Baytown Tunnel and Bridge complex, 19 |uly 1995,
Brotn/ /<S'6,S'9/;(SBSC).
Eleochans a))uj>}xssci Sullivan — Correll and Johnston (1970) separate E. conipressa
and E. aciitiiquamata Buckley, by the wider compressed culms in E. compressa
and the more narrow non-compressed culms m E. aciitnquamatci . Further, they
reported /:. conipressa only for San Augustine County in East Texas and re-
stricted E. cicntisquamata to the Edwards Plateau, the north cental, and southeast
(Refugio County) portions of the state. In addition, tliey indicated (page 275)
that they are probably conspecific whicli is suggested by the twerlapping culms
width measurements used to separate them in their key. We were unable to
borrow Texas and non-Texas material ol both entities from TEX for they are
out on loan to S. Cialen Smith for the Idora of North America project. How-
ever, in a survey ol collections at SBSC and ASTC, we found only specimens
referable to E. aaitisquaviata Buckley in East Texas (San Augustine and Sabine
counties). From the specimens available to us, both entities can be readily
separated by the nearly round culms (7-9 angled) of Z:. aciitisqiianutta that are
up to 25 cm tall and 0.5 mm wide in contrast to the flat culms of £. compressa
that are up to 42 cm tall and 2 mm wide.
Rhynchospoya dehilis Gale — Gale (19-11) described this taxon as a new spe-
cies and noted its similarity to R . fascicularis (Michx.) Vahl. Both of these
Brown and Marcus, Notes on the flora of Texas 3 1 9
Species were growing together at the edge of Galveston Bay on a sandy bench
below the bluff next to the Abshire House in the Abshire Wildlife Man-
agement Area at Smith Point, Chambers County, 7 May 1993, Brown 16932;
27 May 1996, Brown & Liggto 19138. Here they may be distinguished by
the taller (> 35 cm), thicker culms (> 1 .5 mm wide at culm base), and flat
leaves (> 1.0 mm wide) oHl. fasciadciris in contrast to the shorter (ca. 15
cm) filiform culms (< 1 mm wide at culm base) and filiform leaves (< 0.5
wide) of/?, ckbilis.
Sarpus cubemis Poeppig & Kunth \Oxycarynm a/Z'em^ (Poeppig & Kunth) Lye} —
The first Texas collection of this sedge was in 1958 from Eagle Nest Lake
m Brazoria County, Hotchktss 7633 (TEX). The second Texas collection is
from Fort Bend County where is was somewhat frequent in a floating mat
of vegetation in Elm Lake at Brazos Bend State Park, 7 Nov 1997, Brown
21737 . Thomas and Allen (1993) mapped it for eight central and eastern
Louisiana parishes, none near the Texas border.
EUPHORBIACEAE
Phyllanthiisfraternm G.L. Webster — The first Texas collection is from downtown
Houston growing on bare ground among planted shrubs; 17 Sep \99'b. Brown
1 87 33. Webster (1970) indicated it is native to Pakistan and India and spo-
radically introduced into Africa and America. He cited only three U.S. col-
lections, one from an abandoned field in Seminole County, Florida, and the
other two from New Orleans, Louisiana (including the first U.S. collection).
It is now mapped for nine Louisiana parishes mostly in the New Orleans area
(Thomas & Allen 1996). It may also expand its range in the Houston area.
Phylianthus nin/ri L. subsp. lathyroides (Kunth) G.L. Webster — Correll and
Johnston (1970) indicated this taxon may be extinct in Texas (the only U.S.
records for this widespread tropical American species are from Texas) for
they reported it only from the Ottine Swamp in Gonzales County, prob-
ably based upon the 1935 Tharp collection at TEX. The only collections
since 1935 are the following.
Dewitt Co.: a group of plants ca. 2 ft tall, on sand under hridt^e on Thomaston Rd, W
of Hwy 87 along the Guadalupe River, ca. 2 mi W of Thomaston Community, 11 Oct
1994, Miischalek j-.k.(SBSC). Fayette Co.: S bank of the Colorado River at foot of bluff, N
of Monument Hill State Historic Sue, 1 1 Oct 1987, Can 7867 (THX).
The only known U.S. collections are two in the nineteenth century by
Wright, one of which is from the Colorado River bottomland, the one in
1935 by Tharp, and the two recent ones reported here.
FAB ACE AE
AlysicarpiLS vaginalis (L.) DC. — Isely (1990) mentions this native from the
Old World tropics as occurring in Texas. He has examined Fabaceae collec-
320 SiDA 18(1)
tions at TEX and probably discovered this Hardin County specimen, 4 mi
5 of Kountze along roadside, 22 Sep 1960, McLeoci s.n. We have collected
plants ol this legume along a pipeline right-of-way in the Trinity River
National Wildlife Refuge, Liberty County, 13 Oct 1996, Broivn & Marcus
19731.
LENTIBULARIACEAE
Utr/c7//ar/a/o//osa L. — This large flowered, robust, floating bladderwort was
not reported for Texas by Correll and Johnston (1970), possibly because Small
(1933) indicated that it ranged from Florida to Louisiana. The first report
seems to be in Taylor (1989), where it is listed for North Carolina, Georgia,
Florida, Mississippi, Louisiana, and Texas. However, Taylor does not indi-
cate its distribution in Texas nor provide specimen citations. The majority
of the TEX and LL specimens were identified as the robust U. vulgaris L.
Taylor (1 9cS9) considers the North American representatives of U . vulgaris
a distinct species, U . iiiacrorhiza Le (^onte, and indicates its absence from
eastern Texas and most of the southeastern LJ.S. Coastal Plain.
Voucher specimens: Brazoria Co.: iloaring aquatic in water in the Brazoria National
Wildlife Refuge S of Anglcton, 26 Apr 1992, Brotrn 16020 (SBSC); Muldoon Club be-
tween Austin Bayou and Danbury, I 9 May 1 97 1 , Fleetwood 10029 (TEX). Chambers Co.:
present in Anahuac National Wildlife, Jul 1990, Near///e s.//.{SV,SC). Fort Bend Co.: Brazos
Bend State Park in 40 Acre Lake beside dike between 40 Acre Lake and Pilant Lake, May
1994, Gnxii s.n.(SBSC); 29 May 1993, Greiif^ s.n. (SBSC). Hardin Co.: 28 Jun 1952, Tharp
6 Tyson j.^/.(TLX); Ariola Rd, S of Kountze, home of Jack Lloyd, 9 May 1957, Whitehouse
52^HI (TEX). Llenderson Co.: Code Lake, 27 May I940, Sie^ler 2000 (TEX). Jefferson
Co.: tloatmg in canal along rd,4 mi N of Port Authur, 19 Jul 1946, Correll 13357 (LL);
J.D. Murphree Wildlife Mangement Area, 1-10 mi SW of Port Arthur, Big Hill Bayou,
submerged, 20 Oct I 966, We.s/ 127 (TEX). Liberty Co.: floating on water in bayou about
7 mi E of Cleveland, Rte. 321, 12 Sep 196S, Correll 36458 (LL).
ONAGRACEAE
Ludungia /uicrocarpa Michx. — Bridges and Orzell (1989) reported this spe-
cies new to Texas from two roadside collections in Hardin County. They
state these collections are within former extensive wetland pine savannahs
on the Montgomery and Beaumont Formations of the Pleistocene Epoch.
A new county collection, perhaps from a younger formation, is from Chambers
County where it was frequent along the sandy roadside of Highway 562 ca.
2.8 mi east of the road to Robbins Point Park, 27 May 1996, Broivn & Liggio
191.]
1
The h)llowing collections at SBSC complete the documentation of this
species in Texas. These roadside collections suggest it may be spreading westward
via highways.
Voucher specimens: Hardin Co.: frecjuent in a dry roadside ditch along hwy 69/287 ca.
1 mi N of Kountze, associated with Rbyinhnspora dheri^eiis and Viiirena hreviseta, 2 Aug 1986,
Broun I05HI\ treqLienr in a roadside ditch at etige of a sandy acid area along Hwy 770, ca.
Brown and Marcus, Notes on the flora of Texas 321
0.4 mi W of intersection with Hwy 1003 between Saratoga and Kountze, 16 Aug 1986,
Brown 1()64J', on acid sandy soil with many bog plants along Hwy 421 ca. 3 mi E of inter-
section with Hwy 326 N of Sour Lake, 9 Jun 1996, Brown 19202.
POACEAE
Saccharum coarctatum (Fern.) R. Webster — Webster and Shaw (1995) did not
consider the presence {Erianthus) or absence {Saccharum) of an upper lemma
awn sufficient to separate these two genera. They, tlierefore, merged Erianthus
under Saccharum and made the appropriate combinations under Saccharum.
They cHd not cite any Texas collections oi S. coarctatum., possibly because
they did not examine sufficient material from Texas herbaria (the only Texas
herbarium cited is BRIT). The following collections at SBSC document its
presence in the state. The senior author first iclentifed these plants as E.
brevibarbis Michx. and then annotated them to S. hrevibarbis (Michx.) Pers.
var. brevibarbis. Joseph K. Wipff, then at TAES, identified them to S. coarctatum.
Voucher specimens: Liberty Co.: tall plants in moist ditch along Hwy 90, 5.4 mi E of
Hwy 61 in Devers, 25 Sep 1993, Brown 17493', tall plants with brownish inflorescence
along Hwy 90, 4.9 mi E of Hwy 61 in Devers, 7 Nov 1994, Brown 17618. Tyler Co.: large
colony on low moist acid soil within Kirby State Forest off of Hwy 69, S of Warren, 8 Oct
19H5, Brown 6674.
Uroch/oa ramosa (L.) Webster {Panicum ramostim L.; Bachiaria ramosa (L.) Stapf} —
This native of tropical Asia has been planted in Texas and other states for
erosion control and wildlife food but has not been included in any Texas
floristic publications. It is reported for 31 Louisiana Parishes (Allen 1992).
An adventive Texas collection is from Chambers County where it was growing
at the edge of the Highway 1663 bridge over Whites Bayou northeast of
Hankamer, 25 Sep 1993, Brown 17488.
POLYGONACEAE
Polygonum cespitosum Blume — Carr and Hernandez (1993) reported this Asian
species new to Texas (first collection from Jasper County in 1992) using
the spelling caespitosum. We are following the spelling in Kartesz (1994).
We discovered this species on moist to dry disturbed sites in the Trinity
River National Wildlife Refuge. Here the reddish flowers stand out in contrast
to the whitish flowers of P. hydropiperoides Michx. and other similar species.
Voucher specimens: Liberty Co.: 15 Sep 1996, Brown 19701; 13 Oct 1996, Broirn &
Marcin 19732; 9 Nov 1996, Broivn 19787; 21 May 1997, Brown 20445. The following
two collections, previously identified as P.perskarza L., are at TAES: Galveston Co.: Galveston
Island State Park, 15 May 1997, Rosen 0316. Liberty Co.: ca. 4.5 mi S of Dayton along
Hwy 146, 21 Oct 1987, N/xon 16050.
Additional state collections may be masquerading under this name in
other Texas herbaria.
Polygonum meisnerianum Cham. & Schlecht. var. beyrichianmn (Cham. & Schlecht.)
Meisn. — Correll and Johnston (1970) indicated this species has been re-
322 Sida18(1)
ported for Texas but saw no specimens. Park (19fiS) also saw no Texas col-
lections but reported U.S. collections from Florida and South Carolina and
from Plaquemines and St. Tammany Parishes in Louisiana. The only Texas
collection known to us is from Orange County where it was growing in a
slight swale under a canopy of Chinese tallow trees adjacent to a fallow rice
field near Nederland, Knowles sji. The senior author originally identified
this specimen as P. siigittatuvi L. and additional collections may be filed under
this name in Texas herbaria.
Rumex paraguayensn Parodi — This native of southern South America (natu-
ralized in Florida and Louisiana) is now in Texas (Chambers County, present
in the East Unit of the Anahtiac National Wildlife Refuge off of Hwy 19H5,
SE of Anahuac, Spring 1 99 1 , Nei'ille 274). The key in Godfrey and Wooten
(19<^ 1 ) can separate this species from R. obovatus Danser, a similar natural-
ized South American species in northern Florida and Louisiana. Thieret (1969)
reported both species new to North America and speculated that R. obovcit//s
almost certainly occurs in Texas because one collection of it was made within
150 feet of the Texas border. However, based on Ciodfrey and Wooten's key,
our specimen is R. paragiuiyensis.
SCROPHULARIACEAE
Lifickrnui Crustacea (L.) F. Muell. — This Indo-Malaya native ranges in the
U.S. from South Carolina, south to Florida, and along the Gulf Coastal Plain
to Louisiana (Godfrey & Wooten 1981). Vincent (1982) mapped it for 1 5,
mostly south central and eastern, Louisiana Parishes. He indicated it is spreading
westward and that it may eventually appear in East Texas. We found this
short blue flowered species on two disturbed sites in the Trinity River National
Wildlife Refuge, Librty County, 29 Sep 1996, Brown & Marcus 19703; 13
July 1997, Broin/ 20581.
ACKN()Wl.F,DC;iMi;N'tS
We wish to thank the following; Tom Wendt (TEX), James VanKley (ASTC)
and Stephan Hatch (TAES) for permitting access to their collections, for
the loan of herbarium material, and for sending copies of label data from
others; Guy Nesom and an anonymous reviewer for their careful reading
and improvement of the paper; all of the Houston area and state-wide ama-
teur botanists who have added collections to our local herbarium (SBSC)
and for their help in numerotis other ways; and the Texas Parks and Wild-
life Department and local National Wildlife Refuges for permitting col-
lections of plants from their respective lands. Work on tlie Trinity River
National Wildlife Refuge was sup[iorted by Challenge C^ost-Share Agree-
ment with the refuue.
Brown and Marcus, Notes on the flora of Texas 323
REFERENCES
Affoltf.r, J.M. 1985. A monograph of the genus Lilaeopsis (Umbelliferae). Syst. Bot. Monogr.
6:1-140.
Allhn, C.A. 1 992. Grasses of Louisiana. Second cd. Cajun Prairie Habitat Preservation Society.
Eunice, LA.
Al-Shhiiba/, la. and V. Ba'ii'.s. 19H7. Arnioraaa lacustris (Brassicaceae), the correct name
for the North American lake cress. J. Arnold Arbor. 68:357-359-
Bridcf.s, E.L. and S.L. Orzell. 1989. Additions and noteworthy vascular plant
collections from Texas and Louisiana, with historical, ecological and geographical notes.
Phytologia 66:12-69.
Carr, B. and D. Hernandez. 1 993. Polygonum caes[vtos/tm var. longisetmn (Polygonaceae) new
to Texas. Sida 15:656.
Cleweel, A.R 1985. Guide to the vascular plants of the Florida Panhandle. Univ. Presses
of Florida, Tallahassee.
Correll, D.S. and M.C. Johnston. 1970. Manual of the vascular plants of Texas. Texas
Research Foundation, Rennet, TX.
Denton, M.F. 1978. A taxonomic cteatment of the Luzulae group oiCypems. Contr. Univ.
Michigan Herb. 11:197-271.
Gale, S. 1944. Rhynchospora , section Eurhynchospora , in Canada, the United States and the
West Indies. Rhodora 46:88-134, 159-197, 207-249, 255-278.
Gandhi, K.N. and R.D. Thomas. 1983. Cnscuta polgonorum Engelm. new to Louisiana and
comments about its antestaminai scales. Phytologia 53:181—186.
Gandhi, K.N., R.D. Thc:»mas, and S.L. Hatch. 1987. Cuscutaceaeol Louisiana. Sida 12:361-
379.
Godfrey, R.K. and J.W. Wooten. 1981. Aquatic and wetland plants of southeastern United
States, dicotyledons. Univ. of Georgia Press, Atlncns.
Hatch, S.L., K.N. Gandhi, and L.E. Brown. 1 990. Checklist of the vascular plants of Texas.
Agric. Exp. Sta. MP-1655. College Station, TX.
Holmoren,P.K., N.H. Hc:)EMc;rfn, and L.C:. BARNirrT. 1990. Index Herbariorum. Eight
ed. New York Botanical Gatden, Btonx.
Isely, D. 1990. Vascular flora of the southeastern United States, Vol. 3(2), Leguminosae
(Fabaceae). Univ. of North Carolina Press, Chapel Hill.
Johnston, M.C. 1990. The vascular plants of Texas, a list up-dating the manual of the
vascular plants of Texas. Second etl. Ptivately published by the author. Austin, TX.
Jones, S.D., J.K. Wipef, and P.M. Montoomkry. 1997. Vascular plants of Texas. A com-
prehensive checklist including synonymy, bibliography, and index. University of Texas
Press, Austin.
Kar-im-sz, J.T 1994. A synonymized checklist of the vascular flora of the United States,
Canada, and Greenland. 2nd ed. vol.1, (Checklist. Timber Press, Portland, OR.
Landry, G.P., W.D. Ri;ese, and CM. Allen. 1988. Stdlaria parva Pederson new to North
America. Phytologia 64:497.
Landry, G.R, W.D. Reese, and K. Vincip.nt. 1 989. Stellana parva Pederson (Caryophyllaceae)
in North America. Sida 13:373-375.
Les, D.H. 1994. Molecular systematics and taxonomy of lake cress (Neoheck/aaq/iatica, Brassicaceae),
an imperiled aquatic mustard. Aquatic Bot. 49:149—165.
, G.J. Anderson, and M.A. Cleland. 1995. Sterility in the North American lake
cress Neoheckia aquatica (Brassicaceae): inferences from chromosome number. Rhodora
97:185-200.
324 SiDA 18(1)
LiPSc;()MB, l^L. 1978. Additions to rhe Texas llora. Sida 7:392-394.
LUNDCI.L, C.L. 1943. Flora of Texas, vol. 3, part 11. Sourhern Methodist Univ. Press. Dallas,
TX.
Mears, J. A. 1 977. The nomcncJattire and type collections oi the widespread taxa oiAltenianthera
(Amaranrhaceae). Proc. Acad. Nat. Sci. Philadelphia 129:1-21.
Nhi^on, J.B., K.B. Kelly, Jr., and A.C;, iVIoom;. 1 997. Noteworthy collections, South Carolina.
Castanea 62:283-288.
Park, Chonc-Wook. 1988. Taxonomy ot Polygouiuu section Echmocciulini (Polygonaceae).
Mem. New York Bot. Gard. 47:1-81.
Pedersi^n, T.M. i 9<3l . New species ai Hyclrocleis, Scirpi/s, and Stellciria. Bot. Tidsskr. 57:38-
48.
Rollins, R.C. 1993. The Crucilcrac oi continental North America. Stanford Univ. Press,
Stanford, C^A.
Small, J. K. 1 933. Manual of the sontheastern flora. Privately published by the atithor. New
York, NY.
Taylor, P. 1989- The ^t^enus UtriaiLir'hi-:\ taxonomic monotjraph. Kew Bull. 14:1-724.
Thieret, J.W. 1969. Riimex obovcitiis and Riinmx pdycigiuiyoisis (Poly^onaceae) in Lotiisiana:
new to North America. Sida 3:445-446.
Thomas, R.D. and CM. Allen. 1993. Atlas of the vascular flora of Louisiana. Vol. 1: ferns
and fern allies, conifers, and monocotyledons. Lotusiana Department of Wildlife & Fisheries,
Baton Rouge.
TiiOMA.s, R.D. and CM. Allen. 1996. Atlas of the vascular flora of Louisiana. Vol. II: di-
cotyledons, Acanthaceae-Euphorbiaceae. Louisiana Department of Wildlife <Sc Fisher-
ies, Baton Rouge.
Tucker, Ci.C7 1987. The genera of (^yperaceae in the southeastern United States. J. Arnold
Arbor. 68:361-445.
Tucker, Cj.C 1994. Revision of the Mexican species of Cyperin (Cyperaceae). Syst. Bot.
Monogr. 43:1-213.
Wlb.stlr, G.L. 1970. A revision of Pb)Uct>ttl)iis (Ftiphorbiaceae) in the continental United
States. Brittonia 22:44-76.
Web.ster, R.D. and R.B. Shaw. 1995. Taxonomy of the native North American species of
5^/(r/wr//w (Poaceae: Andropogoneae). Sida 16:55 1—580
Vinc;ln'1, K.A. 1982. Scrophulariaceae of Lotiisiana, M.S. Thesis. The Lhiiversity of South-
western Lotiisiana, Lafayette.
CYPERUS FUSCUS (CYPERACEAE), NEW TO MIS-
SOURI AND NEVADA, WITH COMMENTS ON ITS
OCCURRENCE IN NORTH AMERICA
PAUL M. MCKENZIE BRAD JACOBS
U.S. Fish and Wildlife Service Missouri Department of Conservation
608 E. Cherry St. Room 200 P.O. Box 180
Columbia, MO 63201. U.S.A. Jeffrson City. MO 63102-0180. U.S.A.
CHARLES T. BRYSON
USD A. ARS. Southern Weed Science Research Unit
P.O. Box 530
Stoneville. MS 58116-0530. U.S.A.
GORDON C. TUCKER RICHARD CARTER
Stover-Ehinger Herbarium. Herbarium, Department of Biology
Botany Department, Eastern Illinois University Valdosta State University
Charleston. IL 61920-5099. U.S.A. Valdosta. GA 51698. U.S.A.
ABSTRACT
Field and herbarium studies have documented Cyperus f//scus as new for Missouri and
Nevada. Localities, habitat data, lists of associated species, description, illustrations, photo
oF habit, and discussion of weedy potential are |iresented.
RESUMEN
Los estudios de campo y de herbario han documentado Cyperus fusa/s como nuevo para
Missouri y Nevada. Se aportan localidades, datos del habitat, listas de especies asociadas,
descripcion, ikistraciones, foto del habito, y discusion de su potencial como mala hierba.
IKTRODUCTION
Cyperus fuscus L. [section Fusci (Kunth) Clarke} is native to Eurasia and
the Mediterranean Region of northern Africa, extending from Greenland
and Iceland to China south to Spain, Iran, Egypt, Algeria, and northern
India (Kiikenthal 1936; Tutin et al. 1980). The common name for this spe-
cies has been listed as "galingale," "brown galingale," or "black galingale"
(Weedon & Stephens 1969)- It apparently was first discovered in North America
based on specimens collected by Herbert A. Young along Revere Beach in
Essex County, Massachusetts in 1877 (Knowlton et al. 1911; McGivney
1938). Cyperus fuscus was subsequently documented in California (McGivney
SiDA 18(1); 325-333. 1998
326 SioA 18(1)
1938; Tucker 1993), Connecticut (McGivney 1938;Dowhan 1979; Tucker
1 995), Maryland, (Kiikenthal 1 936), Nebraska (Weedon & Stephens 1969;
Kolstad 1986;Rolfsmeier 1995), New Jersey (Britton 1886; McGivney 1938),
Pennsylvania {Rhoads & Klein 1993), South Dakota (Weedon & Stephens
1969), and Virt^inia (Hitchcock & Standley 1919; Kolstad 1986; Tucker
1987). Althougli the species has also been recorded for New York (e.g., Weedon
& Stephens 1969; Kolstad 1986), no specimens have been located and the
only ones so identified were actually C. diandrns Torr. (Mitchell & Tucker
1997). Although C. J//SC//S has been present on the North American Conti-
nent since at least 1877, its spread apparently has been slow, and outside of
brief accounts related to its increase in distribution, there has been little
attention <dven to it in the New World literature.
Given the great attention to new records of Cyperus in North America by
Charles Bryson, Richard Carter, Stanley Jones, Barney Lipscomb, Gordon
Tucker, and others in the last 20 years (e.g., Lipscomb 1978, 1980; Tyndall
1983; Carter et al. 1987; Carr 1 988; Carter 1988; Sundell & Thomas 1988;
Carter & Bryson 1991; Webb et al. 1991; Bryson & Carter 1992; Jones et
al. 1 993; Bryson & Carter 1994; Tucker 1 994; Bryson et al. 1 996; Carter et
al. 1996), it is interesting and somewhat amazing that the species has not
been discovered at more localities throughout the United States, especially
as Weedon and Stephens (1969) noted that it was a weed of rice fields in
the Old World. Outside the United States, the species has been discovered
in Canada (GiUett 1970), but it has not been found in Mexico (Tucker 1994)
or Central America (Tucker 1983).
DESCRIFllON
The following is a description of C fiiscin based on Kiikenthal (1936),
McGivney (1938), Fernald (1950), Kolstad (1986), and our observations.
Cyperiii jiisciii (Figs. 1, 2): a caesj-jitose annual with few to numerous culms
and reddish-brown fibrous roots; culms upright, inclined, or decumbent,
2—30 (rarely 50) cm long, 1.3—3.0 mm thick, smooth, flaccid, and triangu-
lar in cross-section; 2—4 basal leaves per culm, 1—25 cm long, 1—4 mm wide,
flat, flaccid, and minutely scabrous toward the acute apex; 2 or 3 leaf-like
bracts subtend each inflorescence, varying in length, 2-25 cm long, l-3(—
5) mm wide, spreading, flat, flaccid, and minutely scabrous toward the acute
apex; 6— 24(— 80) densely or loosely congested spikelets per spike; spikelets
8—1 8(— 40) flowered, 3— 8(— 12) mm long, 1 — 1 .5 mm wide, linear, and com-
pressed; rachillas 0.3 mm wide, 0.2 mm thick, dark brown, straight or slightly
curved, and wingless; scales (sometimes called "glumes") subtending the
flowers O.H— 1.2 mm long, 0.8—1 .5 mm wide, broader than long, 3-nerved,
tan or light brown at the center with narrowly hyaline margins, and the
McKenzic, et al., Cyperus fuscus in North America
327
Fic;. 1. Cyperus fuscus. A. Habit (top and side views of clump and side view of two erect
plants). B. Inflorescence. C. Spikelet. D. Scale. E. Achene (including cross-sectional view
and side view of 3-cleft style). (Based on McKenzie 1802 with Jacobs; McKenzie 1804; McKenzie
1807 with Jacobs and joh>ison; illustrated by Charles T. Bryson).
328 SiDAlSd)
surfaces primarily dark reddish-brown to purple; the tips minutely apicu-
late; achenes 0.9-1 -1 mm long, 0.4-0.5 mm wide, gray to almost white,
trigonous, and narrowly ovoid; styles 0.4—0.6 mm long, 3-cleh, and not
persistent; stamens 2 per flower, 0.7-0.8 mm long; anthers 0.4-0.5 mm
long, tan or yellowisli, and linear oblong.
BIOLOCY, F.COLOCY, AND DISTRIBl :TIC)N
On 9 September 1997, while conducting a search for state-listed species
ofCyper//s, Schoenoplectus, and Lipocarpha along mudflats of the Missouri River
in Cooper Coimty, McKenzie and Jacobs discovered a population of Cyperus
sp. unfamiliar to them. This Cyperits was semiprostrate with the culms ra-
diating like spokes in a wheel and leaning mostly horizontal to the ground
{Fig. 2). The most diagnostic features of the sedge were its dark purple to
reddish-brown scales, its strongly triangular stems, its bright rusty red roots,
its small spikelets, and its pale, trigonous achenes. Specimens were subse-
quently determined as Cyperm fuscin. McKenzie and Jacobs returned to the
site on 28 September 1997 and counted 1 10 plants scattered along the silty
mudflats and shoreline of the Missouri River, associated with the following
species: A))i))kniui coccinea Rottb., Cyper//s diandrm Torrey, C. e)jthrorhizos Muhl.,
C. odovcttus L., C sqiicirrosiis L. (C, anstatus Rottb.; C nifhxiis Muhlenb.),
Eclipta prostrate! (L.) L., Eragrost/s hypiioides (Lam.) Brirton, Sterns, & Pogg.,
Vi))d)rist)lis vahlit (Lam.) Link, Le/zcospora niultifida (Michx.) Nutt., Lipocarpha
Diicrantha (Vahl) G. Tucker, VolygoniiDi spp. and Rcnuniciilin schratiis L. Al-
though the majority of plants were in full sunlight, some extended into the
partial shade of the developing seedlings o'l Salix spp. and Popiilus ddto'uks
Bartram ex Marshall.
Subsequent to this discovery, McKenzie and Jacobs were asked by Rhett
Johnson of the Missouri Department of Conservation to identify an unknown
specimen of Cyperin that had been collected from another site along the Missouri
River in adjacent Howard County, approximately 32 km WNW of the original
discovery site. McKenzie and Jacobs identified the specimen as C. fiisciis
and visited the site with Johnson on 1 1 October 1 997, where they counted
approximately 70 plants scattered along moist, sandy and silty flats adja-
cent to a chute that been formed during the 1 995 flood of the Missouri River.
Because the discovery of C, jusais in two adjacent counties along the Missotiri
River suggested that the species could be widely distributed along the river,
Jacobs searched locations downstream of the original discover)' site and discovered
it in Boone, C^allaway, and Cole counties. A lack of time due to the season's
first killing frosts prevented additional searches in other counties border-
ing the Missouri River. With the exception of Fiiuhristylis vahlii, associated
plants at the second, third, fourth, and fifth sites were mostly identical to
those previously mentioned for the original discovery site.
McKenzie, et al., Cyperus fuscus in North America
329
Fic; 2. Photograph of Cyperus J //sens taken from T.isbon Bottoms, Big Muddy National Fish
and WildHfe Refuge, Howard Co., Missouri, 1 1 Oct 1 997 (photographed by Paul M. McKenzie).
Although the initial North American discovery of C. fuscus In Massa-
chusetts was apparently along a sandy beach, many subsequent records along
the Atlantic seaboard were associated with "ballast sand" (Britton 1886),
or "waste ground, ballast, and wharf areas" (Rhoads & Klein 1993). The
Connecticut record comes from a "sandy river shore" (Tucker 1995) and habitat
for the species in California has been reported its "damp, disturbed soil, receding
shorelines, and puddles" (Tucker 1993). In the Midwest, the initial discov-
eries of this sedge were made along "wet open sandy flats" of the South Platte
River in Lincoln County, Nebraska and "wet sandy clay soil" along the banks
of the Keya Paha River in Tripp County, South Dakota (Weedon & Stephens
1969). The accounts by Britton (1886) and Rhoads and Klem (1993) sug-
gest that C. fuscus was initially introduced to the New England coast acci-
dentally via achenes that were in ship ballast water that originated in Eu-
rope. Introductions into other areas of the country, however, are more difficult
to assess. It is possible that waterfowl were responsible for the species' in-
troduction into Nebraska, South Dakota, and elsewhere. Dunn and Knauer
(1975) postulated that waterfowl were responsible for the introductions of
Cyperus flavkomus Mich. [C. albomarginatus (C. Martius & Schrader ex Nees)
Steudel], Fivibristylis miliacea (L.) Vahl, and Aeschynomone indica L. into the
Mingo National Wildlife Refuge in southeastern Missouri.
330 SiDA 18(1)
While the source of introduction of C. fuscus into Missouri is unknown,
the recent and apparent widespread distribution of this sedge along the Missouri
River suggests that the species may have become established soon after the
1993 or 1995 floods. At the original discovery site in Cooper County, plants
were located at the spot where a large levee break had occurred in 1995.
Rolfsmeier (1995) reported that the two Nebraska records oiC. fusais were
from separate locations along the Platte River. Because the Platte River empties
into the Missouri River, and the Keya Paha eventually does the same, it is
feasible that Nebraska or South Dakota may have been the seed source for
the Missouri populations that became established along the Missouri River
following the 1993 and/or 1995 floods. This introduction may be due to
soil movement and sedimentation within the Missouri River floodplain.
Rice growers in the United States should be alerted to the potential of
this sedge becoming a troublesome weed. Holm et al. (1979) listed C. fuscus
as a principal weed in Portugal and as occurring in Afghanistan and Israel.
Based on the rapid and aggressive spread of its relative Cyperus diffornm L.
in North American rice fields (Lipscomb 1980; Tyndall 1983; Bryson et al.
1996), and because C. fuscus is a nee weed in the Old World (Weedon &
Stephens 1969; Mingyuan & Dehu 1970; Holm et al. 1979), this species
should be looked for in rice producing areas of Arkansas, California, Loui-
siana, Mississippi, Missouri, Tennessee, and Texas.
Due to the combination of its semiprostrate habit with the culms radi-
ating like spokes in a wheel (Fig. 2), its dark purple to reddish-brown scales,
its strongly triangular stems, its bright rusty red roots, its small spikelets,
and its pale (almost white), trigonous achenes, this Cyperus is unlikely to be
confused with any other North American member of the genus. The con-
spicuous rusty red roots and small trigonous achenes are similar to C erythrorhizos ,
but the dark purplish scales, strongly triangular stems, apiculate achenes,
and stamen number (2 vs. 3 in C. erythrorhizos) easily distinguish it from
that species. The purplish-tinged scales of C. fuscus are somewhat reminis-
cent of the scales of C, diandrus, C. bipartttus Torrey, or C. flavicovius. It can
easily be separated from C. diandrus and C bipartitus by its trigonous vs.
lenticular achenes, and by its smaller (ca. 1.0 mm vs. 1.8 mm) scales. It
differs from C flavicomus by its flattened vs. erect habit, by the lack of white
margins on the scales, and by its trigonous vs. lenticular achenes.
The authors noted that bruised fresh and herbarium material of C, fuscus
possesses a fragrance similar to, but not as strong as, C. scpiarrosus or C. setigerus
Torr. & Hook. Steyermark (1963) described this odor as that of slippery
elm {Ulm//s rubra Mtihl.). Baihl (1 995) stated that "(i)n a few genera of Cyperaceae,
a fenugreek ifCrigonella fomuju-grciecum) odour is readily detectable in fresh
and (especially) herbarium material."
There is apparently some disagreement in the literature regarding the
McKenzie, et al., Cyperus fuscus in North America 331
number of stamens of each floret of C. jusciis. In the Vlora of the Great Flams
(Kolstad 1986), two stamens are given in the key but three in the species
account. One (McGivney 1938), rarely two (Kiikenthal 1936), or two (Tutin
et al. 1980) stamens are also cited elsewhere in the literature. George Yatskievych
(JVIissouri Botanical Garden, pers. comm.) examined all voucher specimens
from Missouri and noted that florets had only two stamens.
Given the apparent rapid spread of this species along the Missouri River in
Missouri and the weedy nature of many Cyperus in the Old World, it is likely
that this species will be discovered with sufficient effort at other Midwest loca-
tions and possibly elsewhere in North America. While studying Cyperus speci-
mens at the Missouri Botanical Garden in 1 988, Carter discovered a misidentified
specimen of C. fuscus from western Nevada that originally was determined
as C. acuminatus Torr. & Hook. The "exposed mudflat" habitat at the Ne-
vada site is apparently similar to that in the San Joaquin Valley of adjacent
California described by Tucker (1993). Cyperus fuscus wd& not listed byCronquist
et al. (1977) in the Intermountain Flora, nor in Kartesz's (1987) dissertation
on the Flora of Nevada. As with California, Missouri, Nebraska, and South
Dakota, waterfowl may have been responsible for the introduction of achenes
of C. Juscus into Nevada. The following data provide documentation for C.
fuscus in Missouri and Nevada with herbarium abbreviations following Holmgren
et al. (1990), except ctb (pers. herb, of Charles T. Bryson).
Vouciicr specimens: U.S.A. Missouri. Cooper Co.: Big Miickly National Fish and WildHfe
Refuge-Overton Bottoms, ca. 1.9 mi NW of Overton, T49NR1 5WSect.3 lSESENWl/4,
Rocheport 7.5' Quad., 9 Sep 1991 , McKenzte 1802 whh Bracijacohs (ctb, EIU, MO, SWSL,VSC);
Jacobs 97-67 with Pa/fi AkKenz/e (MO); 28 Sep 1997, McKenzw 1804 icith Brad Jacobs (ctb,
MICH, SWSL, UMO); Howard Co.: Big Muddy National Fish and Wildlife Refuge-Lisbon
Bottoms, ca. 1.5 mi NW of Lisbon, T50NR18WSect. 18, Arrow Rock 7.5' Quad., 11 Oct
1997, McKenzie 1807 ivlth Brad Jacobs and Rhett Johnson (ctb, MICH, MO, SWSL, UMN,
UMO, VDB, NSC); Jacobs 97-40 with Paul McKenzie and Rhett Johnson (ctb, MO, MICH,
SWSL, UMN, UMO). Boone Co.: McBaine, W of Columbia waste water treatment la-
goon # 3, T47NR13WSect. 7SWSESW1/4, Hunstdale 7.5' Quad., 13 Oct 1991, Jacobs
97-57 with Tim Smith (MO, UMN, UMO). Callaway Co.: Notth Jefferson City, along N
side of the Missouri River, approximately 400—500 m downstream from the Jefferson City
bridge over the Missouri River, T44N Rl IWSect. 15SWSW 1/4, Jefferson City 7.5' Quad.,
14 Oct 1991, Jacobs 97-58 (ctb, MO, \]UO), Jacobs 97-44 with Chris Dietrich (MO). Cole
Co.: Jefferson City, along the S edge of the Missouri River, approximately 400-500 m downstream
from the Jefferson City Btidge over the Missouri River, just W of Wears Creek, T44NR1 IWSect.
7NENE 1/4, Jefferson City 7.5' Quad,, 15 Oct \991 , Jacobs 97-59 (MO, NEMO, UMO);
Jacobs 97-46 (MO). Nevada. Lyon Co.: Lahontan Reservoir, mudflats at the SE end of the
reservoir, Tl 7NR26ESect.8, 4150 ft, plants growing on exposed mudflats, 1 Sep 1981,
Tiehni 6769 (MO).
ACKNOWLEDGMENTS
We greatly appreciate the assistance of George Yatskievych, Missouri
Department of Conservation, Flora of Missouri Project, Missouri Botanical
332 SiDA 18(1)
Garden, for initially identifying the Missouri specimens and providing additional
information on C.jiiKia. We thank Connie Rutherford, U.S. Fish and Wildlife
Service, Ventura, California, for providing information on the species' oc-
currence in California. Steve Rolfsmeier of Milford, Nebraska, graciously
supplied data on the Nebraska records. The helpful suggestions of George
Yatskievych and Robert Krai improved the quality of the manuscript.
REFERENCES
Biui'iON, N.L. 1886. A preliminary list of N.A. species ol Cyperin, wich descriptions of
new forms. Btill. Torrey Bot. Cllub 13:205-216.
Biu'iii., J.J. 1995. Sedge genera ot the world: relarionshi|is and a new classiticarion of the
Cyperaceae. Austral. Sysr. Bot. 8:125-305.
Bryson, C.T. and R. Cakti-k. I 992. Notes on Cyper/t.s and Kyllingct (Cyi^ieraceae) in Missis-
sippi with records of six species new to the state. Sida 15:1 19—124.
Bkyson, C.T. and R. Carti'.r. 1994. Nt)tes on Carex, Cyperus, and Kyllinga (Cyperaceae) in
Mississippi with records of eight species previously unreported to the state. Sida 16: 17 1 —
182.
Bryson, C.T.,J.R. MacDoxaii), R. (latter, and S.D.Jones. 1 996. Noteworthy Carex, Cypcr/is,
Eleochcirii, Kyllinga, and Oxja^^v/w (Cyperaceae) from Alabama, Arkansas, Georgia, Louisiana,
Mississippi, North Carolina, Tennessee, and Texas. Sida 17:501—518.
(Iarr, B. 1988. Cyperiis ctifloriiiis L. (Cyperaceae) new to Texas. Sida 13:25 5—256.
Car'i'er, R. 1988. Cypiri/s hystncnu/s (Cyperaceae) new to Florida. Sida 13:1 18-119.
Carter, R. and C.T. Bryson. 1991- A report of Cypen/s grayioiJe.s and Cyperus retrofkxus
(Cyperaceae) new to Missouri and notes on other selected Missouri Cyperus. Sida 14:475—
481.
(lARTiiR, R., C.T. Bry.son, antl B.L. Lipscomb. 1987. Cyper/is //>iiflnri/s (Cyperaceae) cast of
the Mississippi River. Sida I 2:250.
Car] i;k, R., R.L. Mi:ars, K.C. Bcrks, and C.T. Bry.son. 1996. A report of four exotic Cyperi/s
((A'peraceac) species new to Lloricla, U.S.A. Sida 17:275—281.
Cronqi LSI, A., A.H. Hoi.MCRi-.N, N.H. HolmciREn, J.L. Rhvbai., and P.K. Holmgren. 1997.
IntermoLintain flora. Vol. 6. Columbia University Press. New York.
Dowiian, J.J. 1979. Preliminary checklist of the vascular flora of Connecticut. Stare geo-
logical and natural history survey of Connecticut. Dei")t. ol Env, Protection— Nat. Re-
sources Center Rept. ol Investigations No. 8.
DiiNN, D.B. and D.F. Knaiii;r. I 975. Plant introductions by waterfowl to Mingo National
Wildlife Refuge Missouri. 'iVans. Missouri Acad, of Sci. 9:27-28.
Fernaei^, M.L. 1950. Gniy's manual of botany, 8th ed. D. Van Nostrand Co., New York.
GiEEEi'r, J.M. 1970. Cyperus jiiscus L.-new to Canada. Canad. Field Nat. 85:190.
L^n(;lK:o(:K, A.S. and P.C. S'EANIM.I'Y. 1919. Flora of the District of Columbia and vicinity.
Contr, U.S. Natl. Herb. 21:1-329, pi. 1-42.
LloEM, L, J.V. Pancik), J.P. Herhercer, and D.L. Plickni ri . I 979. A geographical atlas of
world weeds. John Wiley and Sons, New York.
HoLMCiREN, P.K., N.H. HoEMCREN, and L.C. Barni.ri I, eds. 1990. index Herbariorinn.
Part 1: 'Fhe herbaria ol the world, 8th ed. New York Botanical Garden, Bronx, New
York.
JoNE.s, S.D., A. A. Reznicek, and (i. D.Jones. 1993. C')'/«'r/Av .tt'//(,'£'r//.f (Cy|:)eraceae), new for
New Mexcio. Sida 15:655.
McKENzrE, ET AL., Cyperus fuscus in North America 333
Kartesz, J.T. 1 987. A flora of Nevada. Unpublished Ph.D. dissertation. University of Nevada,
Reno.
Knowlton, C.H., J.A. Cushman, W. Deane, and A.K. Harrison. 191 1. Reports on the
flora of Boston District-VIII. Rhodora 13:27-29.
Kolstad, K.O. 1986. Cyperaceae. In: R.L. McGregor, Coordinator; T.M. Barkley, R.E. Brooks,
and E.K. Schofield, eds. Flora ol the Great Plains. University Press of Kansas, Lawrence.
Pp. 1059-1113.
KuKENTHAl., G. 1936. Cyperaceae-Scirpoideae-Cypereae. In: L. Diels, ed. Pflanzenreich IV.
20(Heft) 101:1-671.
Lipscomb, B.L. 1978. Cypems snrhiamensis (Cyperaceae): new to Arkansas, Kansas, and Oklahoma.
Sida 7:307.
Lipscomb, B.L. 1980. Cyperm difformh L. (Cyperaceae) in North America. Sida 8:320-327.
McGlVNEY, M.V. 1938. A revision of the subgenus Eiicyperia found in the United States.
Cathol.Univ. Amer. Biol. Ser. 26:1-74.
MiNGYUAN, X. and M. Deiii;. 1970. Farmland weeds in China: a collection of coloured
illustrated plates. Agricultural Publishing House. Beijing, China.
Mitchell, R.S. and G.C. Tucker. 1997. Revised checklist of New York State plants. New
York State Museum Bulletin 490.
RnoADS, A.F. and W.M. Klein, Jr. 1993. The vascular flora of Pennsylvanja-annotated
checklist and atlas. American Philosophical Society, Philadelphia.
Rolfsmeier, S.B. 1995. Keys and distributional maps for Nebraska Cyperaceae, Part 1: Bulbostylis,
Cyperus, DiilichiiDii, Elerjcharis, Eriophori/iii, Fimhristylis, Viiire^ia, Lipocarpha, and Scirpi/s.
Trans. Nebraska Acad. Sci. 22:27-42.
SuNOELL, E. and R.D. Thomas. 1988. Four new records oi Cyperus (Cyperaceae) in Arkan-
sas. Sida 13:259-261.
Steyermark, J.A. 1963. Flora of Missouri. Iowa State University Press, Ames, Iowa.
Tucker, G.C. 1983. The taxonomy o^ Cyperus (Cyperaceae) in Costa Rica and Panama. Syst.
Bot. Mongr. 43:1-85.
Tucker, G.C. 1987. The genera of Cyperaceae in the southeastern United States. J. Arnold
Arbor. 68:361-445.
Tucker, G.C. 1993. Cyperus. In: J.C. Hickman, ed. The Jepson manual: Higher plants of
California. University of California Press, Berkeley. Pp. 1138-1141.
Tucker, G.C. 1994. Revision of the Mexican Species of Cyperus (Cyperaceae). Syst. Bot.
Mongr. 43:1-213.
Tuc:ker, G.C. 1995. The vascular flora of southeastern Connecticut. Mem. Connecticut
Bot, Soc. 3:1-205.
TuTiN, T.G., V.H. Heywood, N.A. Burc.es, D.M. Moore, D.H. Valentine, S.M. Walters,
and D.A. Webb, eds. 1980. Flora Europaea-Voliime 5, Alismataceae to Orchidaceae
(Monocotyledons). Cambridge University Press, Cambridge.
Tyndall, R.W. 1983. Distribution of Cyperus difformis L. (Cyperaceae) in the southeastern
United States. Castanea 48:277-280.
Webb, D.H., W.M. Dennis, and TS. Patrick. 1991. Distribution and naturalization of
Cyperus brevifoltoides (Cyperaceae) in eastern United States. Sida 9: 188—190.
Weedon, R.R. and H.A. Steeiii;\s. 1969. Cyperus juscus in Nebraska and South Dakota.
Rhodora 71:433.
THE USE OF ANIMAL-DISPERSED SEEDS AND
FRUITS IN FORENSIC BOTANY
BARNEY L. LIPSCOMB
Botanical Research Institute of Texas
309 Pecan Street
Fort Worth, TX 76102. U.S.A. barney@hrtt.org
GEORGE M. DIGGS, JR.
Department of Biology
Austin College. Sherman. TX 75090, U.S.A.
& Botanical Research Institute ol Texas, gdiggs@austinc.edu
abstrac:t
A specific case of the forensic use of animal-dispersed propagiiles is presented, and it is
suggested that this type of evidence deserves wider utilization by the hvw enforcement community.
Animal dispersed seeds and fruits are ubiquitous, otten cling tenaciously to clothes or other
marerials worn or used by suspects, and are small and frequently go unnoticed. Further-
more, their identification is relatively inexpensive and technically straightforward, and their
presentation as evidence is visually and intuitively obvious, making it ideal for the court-
room. It is also suggested that forensic botany is an excellent topic to use as a case study in
college botany or biology classes because of its inherent interest and integrative nature. In
order to facilitate such usage, a brief review of some aspects of forensic botany is presented
including references to pertinent literature.
RESUMEN
Se presenta un caso especffico de propagulos diseminados por animales en uso forense, y
se sugiere que este tipo de evidencia puede tenet mayor utilizacion en varios aspectos le-
gales. Las semillas y frutos dispersados por animales estan por todas partes, a menudo se
enganchan tenazmente a las ropas u otros materiales llcvados o usados por sospechosos, y
por ser pequefios pasan frecuentemente inadvertidos. ademas, su identificacion es relativamente
barata y tecnicamente senciUa, y su presentacion como prueba es obvia visual e intuitivamente,
convirtiendose en ideal para los juicios. Se sugiere tambien que la botanica forense es un
tema excelente para ser usado como caso pracrico en las clases de biologfa por su interes
inherente y naturaleza integrativa. Para facilitar ese uso se hace una breve revision de algunos
aspectos de la botanica forense incluycndo las referencias bibliograficas pertinentes.
The use of plants in justice and legal systems is thousands of years old,
probably beginning in such ways as trials by ordeal (Talalaj et al. 1991;
Mabberley 1997). In these cases, suspects were forced to eat poisonous plants
and guilt or innocence was determined by survival. Presumably, this was
based on a psychological effect of guilt on the vomiting reflex — suppos-
edly, innocent individuals would expel the poisonous material, while the
SiDA 18(1): 335-346. 1998
i.io
SlDA 1 8( 1 )
guilty would retain the poison and thus die; the efficacy of such a tech-
nique is obviously questionable. The use of plant material as evidence has
also appeared in fictional works, such as the series of books by Ellis Peters
about the medieval Welsh monk/herbalist/sleuth, Brother Cadfael (Whiteman
1 995) and the works by H.C. Bailey about the detective, Reginald Fortune
(Bailey 1936, 1943).
The modern use of plants as evidence in a cotirt case dates to the famous
1930s Lindbergh kidnapping case in which the son of Charles and Anne
Morrow Lmdbergh was kidnapped and murdered. Largely through evidence
provided by botanist Arthur Koehler, Bruno Hatiptmann was convicted of
the crime in 1935. Koehler demonstrated that the ladder used in the kid-
napping was built in part from wood taken from the attic of Hauptmann's
residence (Tippo & Stern 1977; Baden 1 983; Haag 1983; Lane et al. 1990;
Graham 1997). Koehler's evidence included identification of the wood based
on anatomical characteristics, matching of annual growth rings, and unique
markings made on the wood by tools including a lumber yard planer and a
hand plane. Detailed accotints of the botanical evidence including photo-
graphs and graphics can be found in Tippo and Stern (1977), Haag (1983),
and Graham (1997). Graham (1997) gave an extensive list of references.
This was a landmark case, not only because it lead to the formation of fed-
eral kidnapping laws (Bock & Norris 1997), but also because the obvious
value of the evidence provided by Koehler set the stage for future forensic
uses of botanical information.
Since that time, there have been numerous other examples of forensic
botany (and other biological disciplines such as entomology — e.g., Rozen
and Eickwort 1997). The use of plant fragments, pollen grains or fungal
spores, plant trichomes (hairs), anatomical evidence from indigestible cell
wall material from the stomach contents of crime victims, molectilar evi-
dence utilizing DNA, and ecological evidence useful in locating hidden graves,
crime sites, or dating when a crime occurred are just a few examples (Bock
etal. 1988; Lane etal. 1990; IMestel 1993; Yoon 1993;Blaney 1995; Bates
etal. 1997; Bock & Norris 1997; Graham 1997; Lewis 1997;Linden 1997).
The following specific cases show some of the diversity in the field of bo-
tanical forensics. Lane et al. (199(^) discussed a rape case in which leaves
and bark fragments were found in the pants cuffs of a suspect. The material
had gotten into his cuffs while he was climbing a tree to irain access to a
window of the victim's house. His claim, that the victim had let him in
through a door, did not match the botanical evidence. Another example
used by Lane et al. (1990) involved a child abuse case. The parents claimed
that the child had been fed fruit cocktail just prior to dying. However, their
story was contradicted when his stomach contents showed no evidence of
the anatomically characteristic materials expected from such a meal (e.g.,
Lipscomb and DicKis, Forensic botany 337
stone cells from pears or needle-shaped crystals from pineapples). In the
case of a 1989 plane crash near Ruidoso, New Mexico, it was alleged that
an engine design flaw had allowed particulate matter (pollen) to build up
in the engine and cause the crash. However, it was shown that smce the
pollen was in fresh condition and had normal cytoplasm and cell walls as
seen by electron microscopy, it could not have been exposed to the high
temperatures present during engine operation or in the post-crash fire that
distorted even aluminum. Further, the pollen was that of insect-pollinated
plants found near the storage site of the plane wreckage; such pollen grains
are unlikely to be found in any quantity in the atmosphere. Based on the
evidence from forensic palynology, it was concluded that the pollen had gotten
into the wreckage post-crash during several months of storage and there-
fore had nothing to do with the accident (Blaney 1995; Brunk 1 997; Gra-
ham 1997; Lewis 1997). A final example is the use of molecular evidence
linking a murder suspect to a palo verde tree {Parkinsoma aadeata L., Fabaceae)
at an Arizona crime site where he allegedly dumped the body of a victim.
Plant geneticist Tim Helentjaris of the University of Arizona demonstrated
that two seed pods found in the suspect's truck came from a specific palo
verde tree scraped by the suspect's truck at the crime scene. This example is
important because it was the first in which plant DNA was used in a crimi-
nal case (JMestel 1993; Yoon 1993). Overviews of forensic botany were pro-
vided by Lane et al. (1990) in the general science literature and by Bock
and Norris (1997) in the forensic science literature.
Our interest in this topic has developed over the course of a number of
years during which as plant taxonomists we have been called upon numer-
ous times by poison centers, hospitals, and law enforcement agencies to identify
plants or their fragments. We agree with Bock and Norris (1997) that fo-
rensic botany is a resource underutilized by the law enforcement commu-
nity. Further, we believe that forensic botany can be very effectively used in
botany or biology courses to show the importance, applicability, and inte-
grative nature of botany. Because forensic botany cuts across all botanical
disciplines and because a given case may require many research approaches
and techniques, it is a discipline that can stress the integrative nature of
botany and science as a whole. Additionally, it is an excellent topic to use in
emphasizing problem solving and critical thinking skills. The purpose of
the present article is thus two-fold. First, based on our successful use of such
information in college teaching, we want to provide in an easily accessible
botanical journal a brief review of forensic botany and appropriate refer-
ences in order to encourage further such usage. Second, we present a spe-
cific case of the forensic use of animal-dispersed propagules and suggest that
this type of evidence has the potential to be more widely used in criminal
investigations.
3.^,8 SiDAlHd)
On July 12, 1995, a sleeping two year old girl was pulled from the first
floor window of an apartment in Fort Worth, Texas. The child was sexually
molested, but fortunately was found alive about three hours later in a weedy
area several hundred meters from where she was abducted. Assorted evi-
dence was used in the case including fingerprints and DNA from semen.
However, the easily understandable botanical evidence was an important
factor in convincing the jury of the suspect's guilt. Because the Botanical
Research Institute of Texas (BRIT) is a well known source of botanical in-
formation in the local community, we were contacted by the district attorney's
office to identify tiny plant fragments taken from the shoelaces of the sus-
pect. In addition, we were provided with a bag of assorted plant material
that had been collected from the crime scene where the child was left (Fig.
1). Upon opening the evidence envelope, we immediately recognized the
ca. 4 mm long plant fragments as single-seeded mericarps from a member
of the Apiaceae (carrot family). Using a dissecting scope and authenticated
specimens in the Botanical Research Institute of Texas herbarium, these were
identified as mericarps oi'Tor/I/s arvensis (Huds.) Link, an introduced spe-
cies commonly known as hedge parsley. The bag of material from the crime
scene was then examined and a mature, fruit-bearing plant of hedge pars-
ley was found. Under a dissecting scope, the small mericarps of this species
(Fig. 2) are very distinctive. They are densely covered with bristles tipped
with microscopic barbs that enhance their attachment to fur or clothing.
They also have several very characteristic lines of closely appressed hairs between
the bristles. Large photographs of mericarps from both the suspect's shoe-
laces and the crime scene (Fig. 3) were presented in court by one of us (BLL).
Like fingerprints, this was distmctive visual evidence, more easily under-
stood than the scientifically sound but conceptually complex evidence pro-
vided by molecular techniques such as DNA analysis. Because the suspect
could have possibly picked up the mericarps from some other location, the
botanical evidence alone would not have been sufficient for a conviction.
However, it firmly linked the suspect with the crime site and in combina-
tion with other evidence was successfully utilized by prosecutors Sharon
McLauchlin and Larry Thompson and criminal investigator Dennis Timmons.
The suspect, David Noel Saddlemire, was convicted of aggravated kidnap-
ping with the jury taking only 55 minutes to reach their verdict (Fig. 4).
He was subsequently sentenced to 99 years in the Texas state prison system.
Ectozoochory, the transport of a diaspore or propagule on the outside of
an animal, is a common mechanism of dispersal among flowering plants
(van der Pjil 1982). While there are various types of diaspores (e.g., vegeta-
tive bulbils, fragments of the parent plant), the most common types are
seeds, whole fruits, or fruit segments (e.g., mericarps as in the above ex-
ample). Various methods of attachment are known, ranging from viscous
Lipscomb and Diggs, Forensic botany 339
FORT WORTH POLICE DEPAflDlENT
CRIME UBORATORI
RSCIIPT
CASl NO. 141868
TYFS r. i.SE Kidnapping/Sexual Assaulc Chi ld
OFFKNSI NO. 95 364266
VICTIM A. S.
SUSPSCT David Saddlemire
ic.iv.d fromt 'l^(^/->0 ^^ L; p<.£r^^ 1^ ^'^.
Rac
DAT! "^ • ^b TIM i I Z 3^ AM FM ^^
DISCRIPTIONt
Received botanical evidence as listed below:
A tape sealed brown paper bag containing:
1 . A tape sealed bag holding plant material
(II - Invoice #95 C03400) .
2. A tape sealed envelope holding:
A. A tape sealed envelope holding plant
material collected from right shoe
)?22 (Invoice ?/95C03410).
B. A tape sealed envelope holding plant
material collected from left shoe
#23 (Invoice #95C03410) .
C. A tape sealed envelope holding a
subsample of plant material
(28 - Invoice #95C03471).
3. A tape sealed bag holding plant material
(28 - Invoice //95C03471). < „ n
~^^ PRINT GtLjgi^ Uion VQ^^Ctu
ISLEPHONl NO. <?/^ %1^ '6D^V
001-35506-025
8-1-77
Fig. 1. Receipt of botanical evidence from the Fort Worth Police Department Crime
Laboratory.
340
SiDA lcS(l)
■yy?
Fic. 2. Torilis Lini'iisis. A) habit; B) llower; C) iruit showing mericarps with iiiicinace bristles;
D) cross-SL'crion of frLiir (drawn by Linny Heagy).
Lipscomb and Diggs, Forensic botany
341
Left Shoe Right Shoe
<—lcm~- > <— icm— >
Known
< — 1cm — >
Fig. 3. Photographs of mericarps from botli the suspect's shoelaces and the crime scene
(Photos by Larry A. Reynolds, courtesy of Tarrant County Medical Examiner).
or mucilaginous substances that cause the propagule to stick to the dis-
persal agent, to very sharp, barbed or recurved hooks, spines, or avv'ns (van
der Pjil 1982). We believe that many of these examples are potentially valuable
to law enforcement agencies. From many field trips both for research and
with students, it has been our experience that it is rare to rettirn from the
field without attached plant material either on the clothes, in pant cuffs,
on socks, or embedded in shoelaces. Some of these are merely annoying,
while others {e.g., Aristida species — threeawn grasses) are extremely irri-
tating when penetrating socks or pants. Many of these seeds or fruits fre-
quently get transferred to the interior of cars on carpeting or upholstery.
One does not have to go far from the sidewalk to encounter such plant materials.
Because many abundant weedy species are animal-dispersed, yards, aban-
doned lots, and virtually any weedy or disturbed site will have some such
species. Becatise these plant materials are so frequently encountered and because
different plant species are found predominantly in specific habitats and during
particular seasons, they are potentially valuable sources of evidence chat can
link suspects with crime scenes both spatially and temporally. Also, many
(e.g., tiny fruit segments of some Apiaceae or tiny fruits of some grasses)
342
SlDA 18(1)
TIM CURRY
CRIMINAL DISTRICT ATTORNEY
817/884-1400
TARRANT COUNTY
OFFICE OF THE
CRIMINAL DISTRICT ATTORNEY
JUSTICE CENTER
401 W. BELKNAP
FORT WORTH, TS 76196.0201
Apnl21, 1997
Mr. Barney Lipscomb
Botanical Research Institute of Texas
509 Pecan Street
Fon Worth, Texas 76102
Dear iVIr. Lipscomb:
Thank you for your expertise and assistance in the David Saddlemire trial. The case was
a complex one and your testimony was very helpful in explaining to the jury one of the
circumstances surrounding the offense.
The defendant was found guilty and sentenced to 99 years in the penitentiary. Again,
thank you for your assistance. This case was an important one to the State of Texas and this
community.
n
.(j^-^^llciji2a^^d^
Sharon McLauchlin
Assistant Criminal Distnct Attorney
Fig. 4. Letter Irom Tarrant County Olfice of the Criminal Di.strict Attorney confirming
the jury conviction of David Noel Saddlemire.
become deeply imbedded in cloth or carpeting, go virtually unnoticed, and
often remain attached even after repeated washings or other types of clean-
ing. Further, seeds and fruits are easily and inexpensively identified by trained
botanists using nothing more than a lOX hand lens or inexpensive dissect-
ing scope, taxonomic literature, appropriate illustrations, and herbarium
specimens. While molecular forensic techniques can potentially provide
very definitive information, they are much more expensive, recjuire sophis-
ticated laboratory facilities, and are less intuitively obvious for courtroom
presentation.
Lipscomb and Diggs, Forensic botany 343
Sometimes the attachment of ectozoochorous propagules can be quite
tenacious. This means that they could be attached to a perpetrator's cloth-
ing for a very long time, or even enter the flesh. The following are specific
examples from mammals that illustrate the point. Sharp-pointed structures
such as fruits or awns can penetrate the mouth or other tissues of livestock
(and have to be extracted by pliers) or even become subcutaneous and re-
quire surgical removal. Veterinarians (John Brakebill, Larry Edwards, Ken
Lawrence, pers. comm.) indicate that it is not uncommon to find grassburs
{Cenchrus species), awns (e.g., from Hordeum species — foxtail) or the pointed
fruits of needle, spear, or threeawn grasses {Nassella or Aristidci species) in
animals. For example, in North Central Texas, Cenchrus burs are often found
embedded between the toes of dogs and grass fruits are known to penetrate
buccal tissues including the tongue causing serious problems (Ken Lawrence,
pers. comm.). Perhaps more striking is the ability of the sharp fruits of needle
grass (apparently, Nassella leucotrkha (Trin. & Rupr.) Barkworth {formerly
known as Stipa leucotrkha Trin. & Rupr.} — commonly called winter grass,
Texas winter grass, spear grass, or Texas needle grass) to deeply penetrate
flesh. These can become subcutaneous and require surgical removal; for example,
they can enter between the toes of dogs and sometimes migrate long dis-
tances subcutaneously causing draining tracts that will not heal until the
fruit is removed (Larry Edwards, pers. comm.). The most extreme case we
are aware of involved a fruit that penetrated through the skin and then the
chest wall of a dog, eventually becoming embedded in a lung and causing
a fatal case of pneumonia (John Brakebill, pers. comm.). Also tenacious are
the spiny fruits oiTribidus terrestris L., puncture vine, of the Zygophyllaceae.
These are very painful to both animal and human feet, damage even tires,
and are occasionally faral to livestock if eaten (Correll & Johnston 1970); it
would not be surprising to find them attached to various objects and pos-
sibly even the tires of a suspect's vehicle.
Locally in North Central Texas, we believe Soliva pterosperma (Juss.) Less.,
lawn burweed, (Asteraceae) collected from a soccer field near Arlington, Tarrant
Co. (1995), was possibly spread from eastern Texas by athletic shoes; its
fruit is easily, and painfully, attached by its persistent, spine-like style (Diggs
et al., forthcoming). Such propagules could remain attached to a suspect's
clothing or shoes for long periods of time. Other Asteraceae are well-known
as being animal-dispersed with the pappus of many species being modified
into an attachment structure. The retrorsely barbed awns oiBidens species,
beggar's ticks, are strikingly effective. In another composite genus, Xanthium,
cocklebur, the surface of the bur is conspicuously covered with stiff, hooked
prickles ca. 5 mm long and the bur is also terminated by two prominent
spines. The attachment of the hooked prickles to clothing or shoelaces is
344 SiDAlSd)
very effective and they can also easily penetrate human skin. The hooks are
strikingly reminiscent of velcro. According to the VELCRO® Industries
homepage (www.velcro.com), in the early 1940s, a Swiss inventor, George
de Mestral, after a walk noticed "cockleburrs" {presumably X.anthiuvi\ on
his dog and his pants. He examined the hooked prickles under a micro-
scope and derived the idea for the well known two-sided fastener — one side
with stiff, cocklebur-like "hooks" and the other side with soft "loops" like
the cloth of his pants. The word velcro comes from the French words ve-
lours, velvet, and croche, hooked.
While some of the examples above were presented to show the tenacity
with which diaspores attach, commonly the seeds or fruits are small and
merely cling to the fur, feathers, feet, beak, etc. of the dispersal agent with
little or no adverse effect. Because they are often small and inconspicuous,
they may be particularly valuable from the forensic standpoint. A well known
such member of the Fabaceae (legume family) is the genus Desmodium, tickseeds.
The fruits or loments are jointed and break apart into 1 -seeded flat seg-
ments that are the dispersal units; they easily attach to hair or clothing. In
North Central Texas for example, there are 12 members of this genus, a
number of them occupying rather specific habitats (Diggs et al., forthcom-
ing). Other well known examples are rhe numerous Apiaceae that have small
schizocarps (a fruit that splits between carpels into one-seeded portions called
mericarps) whose mericarps are covered with bristles or hairs and become
readily attached to many surfaces. These are particularly well known to owners
of long-haired dogs because large numbers of the mericarps become entangled
in the fur — sometimes the situation is so severe that the only recourse is to
shave the dog. The final example given here is the legume genus Medicago,
commonly known as bur-clovers. There are numerous introduced species
of this genus, many of which have prickly fruits. These fruits can be some-
what larger than those mentioned previously, but are still effective at at-
taching to dispersal agents. Numerous other examples could be given which
have potential use forensically. Fortunately, most are easily identified by
experienced field botanists because such researchers have encountered them
many times on their own clothing or equipment.
Other less obvious diaspores could also be potentially useful. For example,
at the present time, several invasive aquatic species (e.g., Hydrilla verticillata
(L.f.) Royle, in the Hydrocharitaceae) are spreading in North Central Texas,
apparently by power boars transporting vegetative propagules (plant frag-
ments). In areas of the country where there are numerous relatively new
reservoirs and where the distributions of many aquatics, especially intro-
duced species, are spotty, plant material of a given species could easily be
used in linking a suspect with a given body of water.
Lipscomb and Diggs, Forensic botany 345
SUMMARY
Forensic botany is a developing discipline that potentially has broader
applicability than is seen at present. Technically simple, visually obvious,
easily understood, and inexpensive methods such as the use of animal-dis-
persed seeds and fruits are particularly worthy of further consideration. Because
of the inherent interest in criminal cases, the potential for emphasizing problem
solving and critical thinking skills, and the integrative nature of the sub-
ject, forensic botany is a field that can be useful in botany and biology edu-
cation.
ACKNOWLEDGMENTS
We would like to thank our colleagues at various law enforcement agen-
cies, poison centers, and hospitals for their cooperation and hard work. Special
thanks to Dennis Timmons, Criminal Investigator of the Tarrant County
Office of the Criminal District Attorney, and Larry A. Reynolds, Forensic
Photographer, for the photographs, which are used courtesy of the Tarrant
County Medical Examiner. Thanks also to John Brakebill, D.V.M., Larry
Edwards, D.V.M., and Ken Lawrence, D.V.M., veterinarians in Sherman,
Texas, John Lanzalotti, M.D. and Bill Vande Water, BSI, for information on
fictional uses of plants as evidence, and Kenna Pirkle, an Austin College stu-
dent, whose Plant Biology class paper on forensic botany prompted us to
write this manuscript. Linny Heagy provided the illustration oiTorilis cirvtiisn.
REFERENCES
Badi:N, M.M. 1983. Plenary session: The LinclberL;h kidnapping revisited: Forensic sci-
ences then and now. J. Forensic Sci. 8:1035—1037.
Baii.IiY, H.C. 1936. A clue for Mr. Fortune: Reggie at his best. The Sun Dial Press, Garden
City, NY.
1943. The thistle down. Fllery Qi.ieen's Mystery JVlagazine 4:5-1 5.
Batp.s, D.M., G.J. Ander.son, and R.D. Li.i;. 1997. Forensic botany: Tnchome evidence. J.
Forensic Sci. 42:380-386.
Blaney, C. 1995. Who ya gonna call:-' Bioscience 45:746-747.
Bock, J. H., M. Lank, and D, Norris. 1 988. Identifying plant food cells in gastric contents
for use in forensic investigations — a laboratory manual. U.S. Dej:iartment of Justice, National
Institute of Justice, Washington, D.C.
Bock, J. H. and D.O. Norris. 1997. Forensic botany: An under-utilized resource. J. Foren-
sic Sci. 42:364-367.
Brunk, S.K. 1997. The Ruidoso plane crash — the background and the trial verdict. J. Forensic
Sci. 42:378-379.
CoRRi;i.i,, D.S. and M.CJ. JoiixsroN. 1970. Manual of the vasculat plants of Texas. Texas
Research Foundation, Rennet.
Diggs, G.M. Jr., B.L. Lipscomb, and R.J. O'Kennon. Shinners & Mahler's illustrated plants
of North Central Texas. To be published in late 1998 by the Botanical Research Insti-
tute of Texas, Fort Worth.
346 SiDA 18(1)
Graiia.\[, a. 1997. Forensic palynoloi^y and the Ruicloso, New Mexico plane crash — the
pollen evidence II. J. Forensic Science 12:391—393.
Graham, S.A. 1997. Anatomy of the Lindbergh kidnapping. J. Forensic Sci. 42:368-377.
Haa(,, L.C. 1983. The investigation of the Lindbergh kichiapping case. J. Forensic Sci.
«: 1044-1048.
LIoRNAN, J.J. 1983. The investigation ol the Lintlbergh kidnapping case. J. Forensic Sci.
8: 1040-1 043
Lani:, M.A., L.C. Andiirson, 'J'.M. Barkli;v, J.Ll. Bck;k, E.JVl. GitioRn, D.W. F^Al,I., D.O.
NoRRis, TL. Ro.ST, and W.L. Sti:rn. 1990. Forensic botany. Bioscience 40:34-39.
Li'Wis, W. H. 1997. Pollen composition in a crashed plane's engine. J. Forensic Sci. 12:387-
390.
LiNDi-M., CM. 1997. Field sampling and chemical analysis. J. Forensic Sci. 42:398-400.
Mahimri.i:^', D.J. 1987. The plant book, a portable dictionary of the higher plants. Re|M-inted
with corrections in 1989. Cambridge Univ. Press, Cambridge, Unitetl Kingdom.
. 1997. The plant book, a portable dictionary ol the higher plants, 2nd ed.
Cambridge Univ. Press, Cambridge, United Kingck)m.
Me.stql, R. 1993. Murder trial features tree's genetic Fingerprint. New Scientist 138:6.
Ro/KN, J.Ci. Jr. and G.C. Eickwort. 1997. The entomological evidence. J. Forensic Sci.
42:394-397.
TAi.AtAi, S., D. Talalaj, and J. Taiai.ai. 1 991 . The strangest plants in the world. iliU of
Content Publishing Co., Melbourne, Australia.
TiFi'o, C). and 'W.L. Stern. 1977. Fkmianistic botany. 'W.'W. Norton & Co., New York.
VAN Der Pijl, L. 1982. Principles of dispersal in higher plants. Springer- Verlag, Berlin.
WiiiTEMAN, R. 1995. The Cadfael companion: The world of Brother Cadfael. Mysterious
Press Books, New York.
YooN, C^.K. 1993. Botanical witness for the prosecution. Science 260:894-895.
NOTES
GLAUCIUM CORNICU LATUM (PAPAVERACEAE) IN TEXAS
In May 1993, the senior author was informed by her friends, Don and Jo
Robison, about an unusual population of wildflowers growing on their ranch.
The population has been growing in the same field since the early 1960s
when it first appeared with a culrivated batch of cotton seed (D. Robison,
pers. comm.). The cotton was eventually replaced wirh sorghum alum and
other grasses in order to graze cattle and the plant has continued to remain
despite grazing upon by cattle during periods of severe drought.
Thousands of these individuals were observed by the senior author growing
in a sandy field. The majority of the plants, however, were toward the end
of their blooming period and had already developed their fruits (long lin-
ear capsules full of tiny black seeds). The senior author took the plant to be
a poppy, however, one that she was unfamiliar with.
This plant was later identified as Glaiiciiim corniculatum (L.) Rudolph by
the junior author by comparison with herbarium collections at TEX. For-
tunately, there was already a collection (0 'Kennon & Cheatham s. n. ) oiGlaucium
from Texas. This specimen, however, was incorrectly identified (as G. flavum,
a perennial species with yellow petals and glabrous capsules).
In the years since 1993, Texas experienced a drought and few, if any, of
the Glaucium in the Garza population had bloomed, preventing further
observations by the senior author. In April 1997 sufficient rains brought
the Glaucium population to bloom allowing the senior author to both ob-
serve and photograph (Fig. 1) the population. Later, perhaps as a result of
the rains, a third population v,{ Glaucium was brought to the attention of
the junior author (Bill Henderson, pers. comm.). This population was col-
lected at the Balcones Canyonlands National Refuge. An estimated 100 plants
were observed growing in a circular area of about 30 feet in diameter. Two
individuals were blooming and five were in fruit, but the majority of the
plants were in a juvenile phase too young to flower.
The genus Glaucium is native to Southern Europe, the Mediterranean,
Hungary and South Central Russia. It has become adventive in California,
Kansas, Montana, Nevada, New York, Oregon and Pennsylvania (Barkley
1986; Hickman 1993; Kiger 1997). It is immediately distinguished from
other Texas genera in the Papaveraceae by its fruits which are straight, unribbed,
linear, capsules up to 25 cm long. Glaucium corniculatum is a wholly pubes-
cent annual, 30-40 cm tall, caulescent, with clear yellow sap, cauline deeply
pinnate unprickled leaves, inflorescence of a solitary flower that has two
free sepals and four petals. The petals are a vivid scarlet color, two of them
SiDA 18(1): 347. 1998
348 SiDA 18(1)
are wider than the other but all are oi equal length (3—4 cm). At the base of
each petal is a purple pattern that extends toward the tip about 1/3 the
length. The pattern is oval, 10 mm wide and lined with a creamy feather-
like design (Fig. 1).
Although collected in Texas as early as 1986 {Keeney 3[)()6) and recently
reported in Texas (Jones et al. 1997), and North America (Kiger 1997),
Glaucuint cin-iiiciihituDi has yet to be officially documented in the state. Pre-
sented here is an updated key to the genera of Papaveraceae in Texas, a lisr-
ing of known voucher specimens, and a photo of the flower (Fig. 1).
Below is a revised key to the genera of Papaveraceae in the Manual of the
vascular plants of Texas (Correll & Johnston 1970).
1 . Acaulesccnt lu-rbs; petals 8 or more, not cTumpJL'cl in the bud; stii;mas over
the valves ol the ca|rsLile Sanguinaria
1 . Caulescent herbs or (ol siibacaiilescent) conlinecl to the Trans-Pecos; petals
4 to 6, mostly crumpled in the bud; stigmas over the placentae 2
2. Perianth and stamens borne on the rim of the hypanthiumlike expansion
ol the receptacle; sepals united into a calyptra; Iruit conspiCLiou.sly ribbed.
Eschschcjlzia
2. Perianth and stamens strictly hypogynotis; sepals not united into a calyp-
tra; fruit not ribbed 3
3. Herbage |-irickly; llowers on short |x'dicels; capSLile dehiscing from the
apex by i to 6 valves Argemone
3. Herbage not prickly; flowers on long pedicels 4
4. Stigmatic disk present; capsules short and turgid, to ^ cm long,
dehiscing by means of small openings jtist beneath the truncate
summit Papaver
4. Stigmatic disk absent; capsules linear tip to 25 cm long, dehisc-
ing longitudinally from the apex Glaucium
Glauciumcorniculatum(L.) Rudolph, FL Jen., 13. 1781 . CbduloniiiDicitniiaiLninu
L., Sp. PI. 1:56. 1753.
Voucher s|-iecimens. TEXAS. Garza Co.: community of Jiisticeburg, on harm Rd. 35 19
on the way to Lake Alan 1 lenry, Don and jo Robison Ranch, SE corner of Ciarza C^o., ca. 3
mi Irom Kent Co. line, in field on N side ofdirr road, 10 May 1993, Ktrkpatrick. s.ii. (TEX).
Kerr Co.: W of Hunt on TX 39, dry creek bed on N side of road, 2 May 1 993, O'Keiinon &
Checithcim s.n. (TEX). Travis Co.: Balcones Canyoniands National Refuge, 0.2 mi along
dirt road NE from intersection with Cjiw (j'eek road, dirt road is 3.4 mi N along Cow
Creek Road from 1431, just after large yellow house (30° 33' 50" N, 98" 07' 70" W), 3
May 1991, J. K. Willuims et ul. 150 (TEX). San Saba Co.: along the Cherokee Creek on the
William Clark (arm at Rend, on the Q'\^i:! of an over grazed field, I 1 May 1986, Keeney
5906 (BRIT); SE of the William Clark house on the east side of the Cherokee Creek at
Bend (1654 plants iiresent), 9 May 1987, Keony 6642 (BRIT).
The following partial list of species from the Travis county population
SiOA 18(1): 348. 1998
NOTliS
349
Fic",. 1 . Photo o'i Glciucium coyniadatiaii (L.) J.H. RLiclolph. l-'hoto by Z. Kirkpacrick.
emphasizes the weedy nature of the vegetation at this site: Argemone albiflora
Hornem., Buglossoicks arvense (L.) I.M. Johnston*, Callirhoe involucrata (Torr.)
A. Gray, Centaurea melitemis L.*, Cirniim texanum Buckl., Convolvulus equitans
Benth., Ciicurbita foetidmima Kunth, Evax sp., Gauracalckola Raven & Gregory,
Glandularia hipinnattfida (Nutt.) Umber, Lamium amplexicaule'L^', Lepidium
virginicum L., Marruhium vulgare L.*, Medicago lupulina L.*, Hill., Oenothera
trilobata Nutt., Physalis cinarescens (Dunal) A. Hitch., Plantago rhodospermum
Dene., Rapistri/m rugosum (L.) AIL*, Ratihida colunmaris (Sims.) D. Don., Salvia
farinacea Benth., Verbena halei SmaU. * = naturalized species.
— Zoe Kirkpatrick Box 696. Post, TX 79336, U.S.A. andJ.K. Williams, De-
partment of Botany, University of Texas. Austin. TX 78713. U.S.A.
REFFJU-NCES
Barki,ey, T.M., ed. 19S6. Flora of the Great Plains. University Press of Kansas, Lawrence.
CoRKiii.L, D.S. and M.C. Johnston. 1970. Manual of the vascular plants of Texas. Texas
Research Foundation, Renner, Texas,
Hickman, J.C, ed. 1993. The Jepson mantiah Hi^dier plants of C^alifornia. University of
California Press, Berkeley.
Jones, S., J. Wipff, and P.M. Montgomery, 1 997. Vascular plants of Texas: A comprehensive
checklist including synonymy, bibliography, and index. University of Texas Press, Atistin.
KiGER, R.W. 1997. Glciiicium. In: Flora North America Editorial Committee, eds. Flora of
North America: Volume 3. Magnoliojihyta: Magnoliidaeand Hamamelidae. Oxford University
Press, Oxford. Pp. 302-304.
Sum 18(1): 349. 1998
350 SiDA 18(1)
HOVENIA DULCIS (RHAMNACEAE) NATURALIZED
IN CENTRAL TEXAS
In the lace summer of 1997 an unusual tree about 12 m tall was observed at
the base of a high limestone cliff above Town Lake on the Colorado River in
Austin. It had alternate leaves with prominent veins, and appeared to have
strange fruiting structures high on the branches, these structures apparently
deformed with galls. The tree was identified as a member of the Rhamnaceae
based on the characteristic three-carpellate fruit with a remnant disk at the
base, very similar to Coliibnnci. Comparison with the Rhamnaceae collec-
tion at TEX revealed the tree to be Hovenia diilcis Thunberg (Eig. 1), the
Chinese raisin tree, which is known to be cultivated in Texas (Jones et al.
1997). Mabberley (1997) suggested Hovenia to be of close affinity to Colubrina,
while recent work by Richardson (unpublished data) suggest that it is more
closely-allied with Ziziph/is. What appeared to be galls are actually peduncles
that swell naturally as the fruit develops.
This species is native to mesic forests in China and was introduced into
India and Japan for its edible peduncles (Hooker 1 872; Rehder 1 940; Roxburgh
1975; Sargent 1916). In China the swollen peduncles of the cymose inflo-
rescences have long been used medicinally "to offset the effects of over-in-
dulgence in wine" (Sargent 1916). The peduncles are especially palatable
after frost when they redden and the juice sweetens with pear-like flavor
(Mabberly 1990; Reich 1991). The frtut itself is not edible.
This tree is known for its cold-hardiness and is cultivated in USDA cli-
mate zone 5, with minimum temperatures of -20"F (Dirr 1990; Staff of the
L.H. Bailey Hortorium 1976), the equivalent of Milwaukee, WI or Binghamton,
NY. It is therefore surprising to find it naturalized in Austin, TX, an area
that is much warmer, and more importantly, significantly drier than much
of zone 5. The locality where it grows in Austin is a cool, moist microcli-
mate, at the base of a cliff of Edwards limestone and dolomite perhaps 40
m high, facing directly to the northeast. This cliff reaches almost to lake
level, except for a very short but steep talus slope of boulders and loamy
soil. Vegetation growing at the base of this northeast-facing cliff is almost
completely protected from mid-day and afternoon sun, and the presence of
a body of water surely modifies the microclimate further. Eurther explora-
tion via boat several weeks later resulted in the discovery of a larger tree
perhaps 100 m upstream, and a smaller tree perhaps 100 m downstream,
both at the base of the same cliff. Elowering specimens were later collected
in May 1998 from the first tree discovered.
It is possible that this small population off/, dulcis is descended from
cultivated specimens that existed at the University of Texas College of Pharmacy
SiDA 18(1): 350. 1998
Notes
351
Fig. 1. Fruiting branch oi Hovenia diilch Tiiunb., showing capsules and swollen peduncles.
Bar = 1 cm.
Drug Garden, the only other specimens known from the Austin area. Prior
to the mid- 1940s this garden existed about four miles to the north-east of
the collection site, and was subsequently abandoned and never reconstructed
(Beryl Simpson, pers. comm.). Two specimens exist at TEX from this gar-
den, dated 1940 and 1946. It is possible that while H. dulcis was cultivated
in this garden, birds ate the fruit, roosted on the cliffs and trees by the river,
and defecated the seed to the forest soil far below. Until the early 1960's
this portion of the Colorado river was still free-flowing, and large stretches
of loamier soil, which is preferred by H. dukis (Dirr 1990; Staff of the L.H.
Bailey Hortoritim 1976), likely existed. When Town Lake was createql in
the early 1960's, perhaps a much larger population of this species was sub-
merged and destroyed, leaving only a small number of individuals grow-
ing near the top of the appropriate habitat. During further explorations, no
seedlings were observed, and seedling success is probably quite low because
numerous other species compete for space on this very thin ribbon of land.
Furthermore, the majority of fruiting branches overhang the water, so most
Sum 18(1): 351. 1998
352 Si HA 18(1)
fruit falls directly to the bottom of the lake. Similar habitats exist within
the region, so it is possible that H. dulcn occurs unobserved elsewhere.
Voucher specimens: TEXAS. Travis Co.: tree 1() it tall, growini; ar the base of a high,
NE-hicmg limestone cliff along the edge of Town Lake, with Cani/is clr/niiwoiulii. PLitatms
occidentalism and SniiUix Immi-nox, across the lake h-om the boat ramp at the University of
Texas Brackenridge Field Lab, Austin, 30 Sep 1997, D. Goldman with J. Criitchjidd 1105
(BLI, BRIT, TLX); same locality but with flowers, 29 May 1998, D. Goldman with P. Griffith
1 200 (BH, BRIT, TEX).
either specimens examined: TEXAS. Travis Co.: ciiicivated, Austin, 16 Sep I94(), C.C.
Alhers s.n. (TEX); ex cultivation; Drug Ciarden, University ol Texas, 24 May 1946, C.C.
All?en46ph060 (TEX).
Thanks is given to John Crutchfield and Patrick Griffith for their assis-
tance in obtaining specimens of this plant, and Hobbes Goldman, Beryl
Simpson, Billie Turner, Justin Williams, and Tom Wendt for their assis-
tance with the manuscript. — Dougla.sH. Golcliiuiu. Departmmt of Botany. Vmvmity
of Texas. Austin, TX 78713-7640. U.S.A.
Rr.I'HRFNCnS
DiRR, M.A. 1990. Manual of woody landscape plants: Their identilication, ornamental
characteristic s, ctilture, propagation and uses. Fourth edition. Stipes Publishing, Cdiampaign.
HooKiiR, J.l^. 1872. The flora of British India, part I. L. Reeve & CO., London.
JoNHS, S.D., J.K. WiPrf, and P.M. Monk.omp.ry. 1997. Vasctilar |dants of Texas. A com-
prehensive checklist. University ot Texas Press, Austin.
MabbeiU-EY, D.J. 1997. The plant book. Second edition. Cambridge University Press, (Cambridge.
Rehder, A. 1940. Manual of cultivated trees and shrubs. Macmillan Publishing Co., New
York.
Reich, L. 1991. Uncommon fruits worthy ol attention. Addison-Wesley, Reading.
Roxburc^tH, W. 1824. Flora Indica. Reprinted by Oriole Editions, 1975, New York.
Sakcen'I', C]. S. 1916. Plantae Wilsonianae, vol 2. Cambridge University Press, C^ambridge.
Si'Ai'i' Ol- ■nii: L.l L Bailey Horiorium. 1976. tlortus third. Macmillan Publishing Co., New
York.
SiDA 18(1): ^52. 1998
Notes 353
HABRANTHUS TUBISPATHUS (LILIACEAE) NEW
TO THE FLORA OF ALABAMA
The Copper Lily Habranthus tuhhpathm (L'Her.) Traub was first collected in
Alabama in the northeast part of the city of Troy in Pike County on 1 5 July
1988 during field work for a Flora of Pike County. This species was then
collected in the community of China Grove in the northern part of Pike
County on 17 July 1995 and in rural western Bullock County near the area
referred to as High Ridge on 12 July 1996. A fourth site southwest of the
city of Troy in Pike County was discovered in August of 1997 (Fig, 1). These
are the first records of this species from Alabama, and the second record of
this species in North America outside of Texas and Louisiana according to
Holmes and Wells (1980) and Burkhalter (1984),
Habranthus tuhispathus (L'Her.) Traub (Liliaceae), also known as Zephyrantlm
texana Herbert or Habranthus texanus (Herbert) Herbert ex Steud,, is a sea-
pose perennial from a small bulb. Each 10-20 cm scape bears a single yel-
lowish-orange flower produced during the summer months, usually following
a rain. The leaves are linear, 3-5 mm wide and up to 25 cm in length, and
appear in autumn and wither by early spring. The fruit is ovoid and three
lobed containing numerous flat, black papery seed at maturity. The genus
Habranthus is separated from the genus Zephyranthes by Sealy (19.37) based
upon the filaments oi Habranthus being of four lengths and the anthers af-
fixed at the middle, while in Zephyranthes the filaments are of alternate lengths
and the anthers affixed below the middle.
In the city of Troy, Copper Lily is found growing in lawns, along side
walks, in a city softball field, a grass parking lot, and a cemetery. All of the
sites are to some degree disturbed and other vegetation consists primarily
of mixed grasses (predominately Cynodon dactylon (L.) Pers., Paspalum notatum
Fluegge, and Eremochloa ophiuroides (Munro) Hack.) and other bulb species
Qpheion uniflorum (Lindley) Raf , Allium bivalve (L.) Kuntze, and Allium canadense
L,), Copper Lily is so abundant there that in some lawns it forms a com-
plete ground cover when in foliage. The soils are sandy and well drained.
Flowering of Copper Lily is most common in July, August, and September,
when periods of drought are broken by summer thunder showers. The foli-
age appears in late autumn, usually in October, and remains green throughout
the winter. The total area occupied by Copper Lily in Troy is about three to
five acres, and extends in a unbroken population for 276 meters along Three
Notch Street.
In China Grove and in the High Ridge area Copper Lily is found on roadsides
and in lawns. It is associated with many of the same species as the popula-
tion in Troy, with Cynodon dactylon (L.) Pers. and Paspalum notatum Fluegge
Sum 18(1): 353. 1998
354
Sum 18(1)
Fig. 1. Map ot Alabama with locations of Pike and Bullock counties.
predominating. The soils in Cliina Grove and in High Ridge are coarse sands
and the sites tend to be xeric. Undisturbed areas nearby support a dwarf
oak-lichen sandhill community. The population in China Grove extends
for 301 meters along Pike County Highway 37 and the High Ridge popu-
lation extends for 142 meters along Bullock County Highways 7 and l4.
Estimates of total population size are three acres for China Grove and two
acres for High Ridge.
The fourth population is located about eight miles southwest of Troy
near Goshen. It is also found on a roadside and extends for 75 m along Pike
County Highway 25. The soil is sandy and the main associate at this site is
Paspalum notatum Fluegge. This is the smallest population, covering approximately
one-hall acre.
Monitoring of a planted population of copper lily for five years revealed
that the seed generally fell within 15 cm of the parent plant and that seed-
SiDA 18(1): 354. 1998
Notes 355
lings were readily produced. Seedlings generally flowered for the first time
in their third year of growth without special care.
Holmes and Wells (1980) reinforced the suggestion of H.H. Hume (Sealey
1937) that the distribution of Copper Lily in North America was the result
of human activity. They speculated that the distribution of the Copper Lily
in North America was most likely the result of traveling Spanish mission-
aries as all of the Texas and Louisiana populations are near the sites of Span-
ish missions. As there is no record of a Spanish mission ever having been
located in Troy or in Pike county, it is most likely that the species reached
Alabama by other means. Historical records of the Troy-Pike county area
reveal that in 1865 a man by the name of William Murphree left Troy and
moved to Walker County, Texas. In letters to his mother, Murphree states
that he is enclosing seeds for her to plant. In other records Mrs. Murphree
is noted for planting many of the decorative plants in Troy at that time.
The Murphrees lived on Three Notch Street in Troy, the site of the largest
Copper Lily population reported above. Walker County, Texas, where Mr.
Murphree moved is listed by Holmes and Wells as a place where Copper
Lily was collected (1980). We therefore speculate that Copper Lily was in-
troduced into Alabama by the Murphree family from populations in Walker
County, Texas. As all of the sites reported above are old communities, it is
likely that Mrs. Murphree shared the plant with friends or relatives nearby.
Voucher specimens: ALABAMA: Bullock Co.: T13N R21-22E sect. 36-31, intersec-
tion ot Bullock Co. Hwy 7 and 14 and un-numbercd dirt road, high ridge, roadsides and
lawns, hill sun, deep dry sandy soil, abundant, 12 Jul 1996, A.R. Diamond 10413 (AUA).
Pike Co.: TION R21E sect. 29, Troy, U.S. Hwy 29 (Three Notch Street), north of down
town at Knox Field, abundant along sidewalks and road, lawns, and vacant lots, full sun,
sand or clay soil, flowered after shower during drought, no leaves seen, 15 Jul 1988, A.R.
Diamond 5210 (AUA); T12N R21E sect. 9, China Grove, just east of Pike Co, Hwy 7,
roadside, full sun, dry sandy soil, common, 17 Jul 1995, A.R. Diamond 9691 (AUA); T9N
R20E sect. 6, Pike County Hwy 25, 0.2 mi S of U.S. Hwy 29, ca. 75 m along rhe west side
of the road, full sun, flowers yellow, 3 Oct 1997, A.R. Diamond 11131 (AUA).
Duplicates will be distributed at a later date. — Alvin R. Diamond, Jr.,
Charles P. Chapman, and Jim Brummett, Department of Biology, Troy State Uni-
versity, Troy, AL 36082. U.S.A.
REFERENCES
BuRKHALTER, J.R. 1984. Additions to the vascular flora of Florida. Castanea 49:181.
Holmes, W.C. and C.J. Wells. 1980. The distribution oi Hahranthus tubispathus (L'Her.)
Traub in South America and Norrh America- Texas and Louisiana. Sida 8:328-333.
Sealy, J.R. 1937. Zephyranthes, Pyrolirion, Hahranthus and Hippeastrum. J. Royal Hort. Soc.
62:195-209.
SiDA 18(1): 355. 1998
356 SiDA 18(1)
PLANTAGO CORONOPUS (PLANTAGINACEAE) NEW TO TEXAS
Plantago coronopus L,, native to Europe and the Mediterranean area (Chater
& Carrier 1976), was reported as naturalized in California (Dempster 1993).
In the eastern United States it has been variously described as rarely adven-
tive on ballast heaps at seaports (Gleason 1952), occurring sporadically about
ports without persisting (Fernald 1950), and as a repeated introduction in
ballast that does not become established (Gleason & Cronquist 1991)- It
was not reported in standard floras of the southeastern United States (e.g.,
Small 1913; Radford et al. 1968), or in the floras of Missouri (Steyermark
1963), New Mexico (Martin & Hutchins 1981), Great Plains (McGregor &
Brooks 1986), Arkansas (Smith 1994), Oklahoma (Taylor & Taylor 1994),
or Texas (Correll & Johnston 1970). It was also not included in more recent
works on Texas plants such as Hatch et al. (1990) or Jones et al. (1997), nor
was it listed by Rosatti (1984) in his treatment of Plantaginaceae for the
Generic Flora of the Southeastern United States.
As part of the collecting effort for the forthcoming Shinmrs & Mahler's
Illustrated Flora of North Central Texas (Diggs et al., forthcoming), a collec-
tion made in 1998 in Fort Worth (Tarrant County) is apparently the first
documented occurrence of this species for Texas.
Voucher specimen: TEXAS. Tarrant Co.: weedy area in landscape, 301 Cresrwood, Fort
Worth, 24 May 1998, O'Keumm 14221 (BRIT),
At the collection locality numerous individuals were observed as landscape
weeds and other individuals were observed in nearby weedy areas. The mode
of introduction is unclear, but seeds were possibly introduced with landscape
plants. Introduction could also have occurred through spread from intentional
cultivation. Since 1997, seeds oiPlantago coronopus have been offered by Johnny's
Selected Seed Company for the specialty salad green market under the name
"Minutina" or "Erba Stella" and, until recently, seeds were sold by the Orna-
mental Edibles Seed Company as an "edible landscape plant." Other north
central Texas populations have not been found and it is unclear whether the
species will become more widely established. However, some individuals were
very robust, with one having 65 inflorescences. Also, the species is self-com-
patible (Delden et al. 1992) and is tolerant to drought, soil compaction, and
competition (Mook et al. 1992). The seeds are highly viable and germinate
well in soils with low moisture content (Blom 1992).
This species, commonly known as buck-horn plantain, differs from all
other plantains occurring in Texas in having deeply pinnately lobed leaves,
short hairs on the corolla tubes, and 3- or l-locular capsules. Other mem-
bers of the genus occurring in the state have leaves entire or with few teeth
or shallow lobes, corolla tubes glabrous, and 2-locular capsules.
SiDA 18(1): 356. 199<S
Notes
357
Fig. 1. Habit of Piantago corunopus (with permission from Gleason 1952).
Plantago coronopus (Fig. 1) can be recognized by the following descrip-
tion (Butcher 196l;Gleason 1952; Chater& Carrier 1976; Dempster 1993;
Rahn 1996): Pubescent annual or biennial with a taproot and leaves in a
basal rosette; leaves closely spreading on the ground or ascending, linear to
lanceolate in outline, 4—25 cm long, with spreading-ascending, acute, ±
linear lobes, the lobes sometimes toothed or lobed; inflorescences sometimes
numerous, 5—50 cm long including the leafless, densely hairy peduncle terminated
by a narrowly cylindric, dense spike to ca. 12.5 cm long; bracts usually not
surpassing the flowers, broadly scarious-margined at base, the keel prolonged
into an acuminate tip; corolla tubes covered with short hairs; petals acute;
stamens 4; capsules 3- or 4-locular; seeds 3—5 per capsule, dimorphic, one
per locule, ca. 1.5 mm long, and one smaller than the others, ca. 0.5 mm
long, occurring in an upper compartment in the ovary on the abaxial side;
flowering late Apr— Sep.
— Robert J. O'Kennon. Botanical Research Institute of Texas, 509 Pecan Street,
Ft. Worth, TX 76102, U.S.A., bokennon@brit.org; George M. Diggs,Jr., De-
partment of Biology, Austin College, Sherman, Texas 73090, U.S.A. & Botanical
Research Institute of Texas, gdiggs@austinc.edu; and Ronald K. Hoggard, Department
of Biology, University of Central Oklahoma, Edmond, OK 7 5034. rhoggard@ionet.net.
SioA 18(1): 357. 1998
358 Si DA 18(1)
RF.FF.RENCnS
Bi.OM, C.W.P.M. 1992. Ecology oi Plantci^o populations: Germination and establishment.
In: RJ.C;. KtnixT & M. Bos, eds. PLnilago: A Mukidisciplinary Sttidy. Pp. 88-97. Springer-
VerLag, Berlin, Heidelberg, New York.
Bu'rciiiui, R.W. 1961. A new illustrated Britisli flora. Leonard Hill Books Limited, Lon-
don, England, U. K.
C()KK];i.[., D.S. and M.C^. Johnston. 1970. Manual ot the vascular plants ot Texas. Texas
Research Foundation, Rennet.
CiiA'iRR, A.O. and D. Cartif.r. 1976. P/a//tc/go. In: T.G. Turin, V.H. Heywood, N.A. Burges,
D.M. Moore, D.H. Valentine, S.M. Walters, and D.A. Webb, eds. Flora Europaea 4:38-44.
Di-LDiiN, W. Van, 1 1. Van Dijk, and K. WoLif. 1 992. The genetics o^Plantnii^o species. In:
P.J.C. KuiperandM. Bos, eds. PLnitagi): A mukidisciplinary sttidy. Pp. 295-309. Spnnger-
Veriag, Berlin, Heidelberg, New York.
Drmpsthr, L.T. 1993. Plantaginaceae. In: J.C.Hickman, ed. The Jepson manual, higher
|ilants of C^alifornia. Pp. 820-82 I . Univ. of (California Press, Berkeley.
DiGCiS, G.M. Jr., B.L. Lipscomh, and R.J. O'Kt.nnon. Shinners & Mahler's illustrated flora
of North Central Texas. To be published in late 1998 by the Botanical Research Insti-
tute of Texas, Fort Worth.
Ft;RNAi.i), M.L. 1950. Gray's manual ol botany, 8th ed. Reprinteti I 987. Dioscorides Press,
Portland, OR.
Gi.i;ason, H.A. 1952. The new Bntton and Brown illustrated flora ol the northeastern United
States and adjacent Canada, 3 vols. New York Botanical Garden, Bronx.
anti Cronqulsi', A. I 99 1 . Maniial ol the vascular plants of northeastern United
States and adjacent Canada, 2nd ed. Van Nostrand Reinhold Company, New York.
Hatch, S.L., K.N. Gandhi, and L.E. Brown. 1 990. Checklist of the vascular plants of Texas.
Texas Agric. Exp. Sta. Misc. Publ. 1655:1-158.
Jt)Ni;s, S.D., J.K. WiPif, and P.M. Montoomi-rv. 1 997. Vascular plants of Texas: A compre-
hensive checklist including synonymy, bibliography, and index. Univ. of Texas Press, Austin.
Makiin, W.C. and C.R. Hutchins. 1981. A flora oi New Mexico, 2 vols. J. Cramer, Ger-
many.
McCiKHCOR, R.L. and R.E. Brooks. 1986. Plantaginaceae. In: (ireat Plains Elora Associa-
tion. Flora of the Great Plains. Pp. 742-747. Univ. Press ol Kansas, Lawrence.
Mook,J.H.,J. HaI'Ck.J. Vandf.r Took n, and PH. van Tif.ndi;ri.n. 1992. Ecology of /Vrf;//t/,(;«
populations: The tlemographic srructure of iioj-iulations. In: P.J. (7 Kuipcr and M. Bos,
eds. Plantago: A mukidisciplinary study. Pp. 69-87. Springer- Verlag, Berlin, Heidel-
berg, New York.
Radiord, A.E., H.E. Ahlhs, and (7R. Bi-ll. 1968. Manual of the vascular flora of the Caro-
linas. The Ihiiv. of North Carolina Press, Chapel Hill.
Raiin, K. 1996. A phylogenetic study of the Plantaginaceae. J. Linn. Soc, Bot. 1 20: 145-
198.
RosAiii, T J. 198 1. The Plantaginaceae in the soLitheastern United States. J. Arnold Arbor.
65: 533-562.
Small, J.K. 1913- Flora of the sotitheastern United States, l\^^ etl. Published by the au-
thor. New York.
Smiiii, E.B. I99I. Keys to the flora of Arkansas. Univ. of Arkansas Press, Fayetteville.
Sii;vi;RMARK, J.A. 1963. Flora of Missouri. The Iowa State Univ. Press, Ames.
Taylor, R.J. and C.E.S. TAMxm. 1 994. An annotated list of the ferns, fern allies, gymnosperms
and flowering plants of Oklahoma, 3rd ed. Southeastern Oklahoma State Univ., Durant.
SiDA 18(1): 358. 1998
Notes 359
PTERIS VITTATA (PTERIDACEAE), A NEW FERN FOR TEXAS
Pteris vittata L., commonly known as ladder brake, Chinese brake, or Chi-
nese ladder brake, is an Asian native well known as an escape from cultiva-
tion in coastal areas of the southeastern United States. The species was not
included in Small's 1903 or 1913 treatments of the southeastern flora. Small
later (1938) treated the species as Pycnodoria vittata (L.) Small, indicating
that it had been collected in Florida many years previously and that during
the 1930s it was abundant at several Florida localities (e.g.. Everglades).
He also indicated that it was established in Alabama, Louisiana, the West
Indies, and South America. Currently, Pteris vittata is common in southern
Florida where it can be found in almost every habitat with exposed lime-
stone (e.g., pinelands) and on a variety of man-made calcareous substrates
(e.g., sidewalks, buildings, old masonry) (Nauman 1993). Radford et al.
(1968) cited a South Carolina record and noted that it was a rare escape.
Nauman (1993) mapped the species as occurring from southern South Carolina
south to Florida and west to eastern Louisiana; he also showed isolated oc-
currences in southern California and the District of Columbia.
Pteris vittata was not included in the Texas flora by Correll (1956, 1966),
Correll and Johnston (1970), Stanford (1976), Hatch et al. (1990), or Jones
et al. (1997), nor has it been reported from Oklahoma (Taylor & Taylor 1994).
It is now known from Texas based on the following collection:
Voucher collection: Texas. San Saba Co.: ca. 3 mi N of Bend, on the Edwards Plateau,
growing from a limestone boulder with Adiantuni capillns-veneris L. at edge of Rough Creek
(tributary of the Colorado River), 6 Nov 1987 J. W'! Stanford 5 iOS (BRIT, HPC, SPLT).
Subsequent flooding has destroyed the site. The identification was con-
firmed by Jim Blassingame (South Plains College, Levelland, TX).
Of the five Pteris species in the United States (most occurrences are in
the southeastern part of the country), four are introduced and one, P. bahamensis
(J. Agardh) Fee, is native to south Florida (Nauman 1993). Besides P. vittata,
only P. multifida Poir. (spider brake, Chinese brake, Huguenot fern), is known
from Texas. This widely cultivated native of China is naturalized in east
and southeast Texas and the Rio Grande Valley (Correll & Johnston 1970;
Hatch et al. 1990). The deeply palmately 3-divided pinnae (at least some)
of P. multifida are quite distinctive.
Pteris vittata can be recognized and distinguished as follows: stems short-
creeping, stout, densely scaly; leaves clustered, strictly 1 -pinnate (the pin-
nae without lobes or divisions), 0.3-1 m long including petiole; pinnae 12-
20(-30) pairs per leaf, linear-lanceolate to linear-attenuate, 2-18 cm long,
4—9 mm wide, long-attenuate or sharply acute apically, asymmetrically cordate
to widened or truncate basally; serrulate marginally; distal pinnae conspicuously
SiDA 18(1): 359. 1998
360 SiDAl8(l)
longer than proximal pinnae (leaf blades thus oblanceolate in outline) with
the terminal pinna typically longest; petioles and often rachises densely scaly;
sori usually of a continuous narrow band near the margins of the abaxial
surface of the pinnae (thus most of the abaxial blade surface exposed) (Long
& Lakela 1971; Nauman 1993; Smith & Lemieux 1993). The species is a
tetraploid, with 2« =116 (Nauman 1993). An illustration and a detailed
description are available in Small (1938). A recent treatment oi Pteris, in-
cluding a key to taxa occurring in the United States and an illustration of
P. vittata, can be found in Nauman (1993).
— -Jack W. Stanford. Department of Biology. Howard Payne University, Brownwood,
TX 76801. U.S.A.. jstanfor@hputx.edu afui George M. Diggs,Jr, Department
of Biology. Austin College, Sherman, TX 75090. U.S.A. & Botanical Research
Institute of Texas, 509 Pecan Street, Ft. Worth. TX 76102, U.S.A., gdiggs@austinc.edu.
Rni'HRF.NCES
CoRRJiLL, D.S, 1956, Ferns and fern allies of Texas, Texas Research Foundation, Rc-nncr.
1966, Pteridophyta, In: C.L, Lundell, ed. Flora of Texas 1:3—121. Texas
Research Foundation, Renner.
and M.C, John.sr)n, 1970, Manual of the vascular plants of Texas, Texas
Research Foundation, Renner,
Hatch, S,L., K,N. Gandhi, and L.E, Brown, 1 990. Checklist of the vascular plants of Texas.
Texas Agric. Exp, Sta. Misc. Publ, 16^5:1-158,
JoNF.s, S,D,,J,K, Wii'i'i^and P,M, Mon'k.omI'UY. 1997. Vascular plants of Texas: A comprehen-
sive checklist including synonymy, bibliography, and index, Univ. of Texas Press, Austin,
IxjNG, R.W, and O. Lakela. 1971, A flora of tropical Florida, Univ, of Miami Press, Coral
Gables, FL,
Naiiman,C,E, 1993, P(ens. In: Flora of North America Editorial C'ommitree, eds, Fl, North
Amer. 2:1 32-1 35, Oxford Univ, Press, New York and Oxford.
Radford, A,E,, H.E. Aiii.ns, and C.R. Bi;i.i,, 1968, Manual of the vascular flora of the Caro-
linas. The Univ. of North Carolina Press, Chapel Hill.
Small, J. K. 1903- Flora of the southeastern United States. Pubhshed by the author, New
York,
191,3. Flora of the .southeastern United States, 2nd ed. Published by the
author, New York,
1938. Ferns of the southeastern United States. The Science Press Printing
Co., Lancaster, PA,
Smith, A.R. and T, Lpmif.dx, 1993- Pteridaceae. In: J,C, Hickman, ed. Thejep.son manual,
higher plants of California, Pp, 439-1 1 I, Univ, of California Press, Berkeley,
Stanford, J. W. 1976, Keys to the vascular plants of the Texas Edwards Plateau and atlja-
cent areas. Published by the author, Brownwood, TX,
Taylor, R,J, and C,E,S, Taylcjr, 199 i. An annotated list of the ferns, fern allies, gymno-
sperms and flowering plants ot Oklahoma, 3rd ed. Southeastern Oklahoma State Uni-
versity, Durant,
SrDA 18(1): 360, 1998
Notes 361
LYCIANTHES ASA RI FOLIA (SOLANACEAE)
NEW AND WEEDY IN TEXAS
In July of 1997 an unknown plant was collected in Houston and sent to
Texas A&M for identification. It proved to be Lycianthes asarifolia (Kunth
& Bouche) Bitter, a member of the Solanaceae native to South America. This
species has been previously known in North America only from New Or-
leans, Louisiana (Darwin & Feibelman 1991 )• De Rojas and D'Arcy (1997)
incorrectly cited that collection as being from Texas.
This species is recognizable by its prostrate habit, having slender stolons
that root at every node. The leaves are strongly cordate, entire, and long-
petioled. Sometimes a second leaf is produced at non-flowering nodes: if
the primary leaf is relatively large, the second leaf is usually small and of-
ten stipule-like; when the primary leaf is relatively small, the second is of-
ten nearly the same size. The flowers are white, rotate, and ca. 1.5 cm broad.
The yellow anthers are connivent around the style and poricidally dehis-
cent. The fruit is an edible reddish-orange berry ca. 1.3 cm in diameter,
closely subtended by the truncate calyx. Plants are self- infertile (Dean 1997).
A good illustration can be found in the article by De Rojas and D'Arcy and
images of the Houston plants can be seen at Texas A&M's Bioinformatics
Working Group Image Gallery page (http://www.csdl.tamu.edu/FLORA/
imaxxsol.htm). Note that the leaves of the Houston plants are nearly all
rounded apically rather than pointed as in the article illustration.
A visit to the collection site revealed that this plant has overrun several
residential yards in Houston, forming a dense, attractive ground cover in
shaded areas. It is apparently reproducing both vegetatively and sexually,
for numerous flowers and immature fruits were seen in early December 1 997
and mature fruits were collected in March 199H. (Darwin and Feibelman
found no fruit in the Louisiana population and none has since been seen on
plants grown from cuttings taken from that population.) The authors also
suspect that propagules are being carried from yard to yard on the equip-
ment of landscape maintenance companies. The exact time and point of
introduction is not known, but tlie population has been in existence for three
or more years. Residents' attempts to control the plant by hand-pulling,
mowing, herbicide application, and removal of infested sod have proved
unsuccessful. It appears to be tolerant of Houston's winter weather and suffers
only partial dieback during the hottest summer days. Should this plant become
established in nearby Memorial Park (a large, wooded area), it could be nearly
impossible to eradicate. Herbicide trials were begun in one of the Houston
yards in March of 1998. After two rounds of tests, no treatment has pro-
SiDA 18(1): 361. 1998
362 Sum 18(1)
vided complere control, and creacmenrs which have weakened the LyciaJithes
have also damaged the surrounding turf.
It is possible that this plant could become a noxious weed in southern
Texas. The USDA Animal and Plant Health Inspection Service is conduct-
ing a formal risk assessment. (The Louisiana Department of Agriculttn-e
has considered surveying for the plant in the New Orleans area.) The au-
thors encourage identification, documentation, and destruction of any new
colonies subsequently discovered. Should other populations of this plant
be fotmcl, please contact one of the authors.
Voucher specimen: U.S.A. TEXAS. Harris Co.: Houston, shaded residential yard of
502 Lindenwood, with Q^iiercus virginiaiici. Piniis, Stenotaphri/in sec;inclatum , and Trachelosperiiiinn
m moist sandy loam, 6 Dec 1997, Mary Kelchersnl 1 20697 ^A (BH, BRIT, MO, NO, NY,
TACS, TAMU, TEX, US, WAVI).
We thank Michael Nee (NY) for his assistance in identifying this plant.
— Moniq/fe Dubrule Reed, Biology Departvient, Texas A&Al University, College
Station . TX 77843-523S. U.S.A. and Alary Ketchersid, Agricultural and En-
vironmental Safety. Texas Agricultural Extension Service. College Station. TX 77845-
2488. U.S.A.
REFERENCES
Darwin, S.P and T Fiubi'.i.man. 1 99 1 . Lyda>ithes asarifoih/ (Solanaceac) new to North America.
Sida 14:605-606.
Dean, E. 1997. Personal communication. Dr. Dean studies the taxonomy of Lyc/a/zllm at
the University ot California-Davis.
Di- RojAS, C.E. and W.G. D'Arcy. 1997. The genus Lycnnithcs (Solanaceac) in Venezuela.
Ann. Missouri Bot. Ciard. 8-1: 1 67-200.
SiDA 18(1): 362. 1998
Notes 363
FIRST REPORT OF THF GENUS BURMEISTERA
(CAMPANULACEAE) FROM HONDURAS
Burmeistera Triana (Campanulaceae: Lobelioideae) is a genus of 96 species
endemic to the Neotropics (Lammers 1998); it was last monographed by
Wimmer ( 1 943). The genus is closely related to two other Neotropical endemics,
Centropogon C. Presl and Siphocampyhis Pohl (Pepper et al. 1997; Lammers
1998). All are large robust plants (herbaceous or suffruticose perennials,
shrubs, subshrubs, or lianas) with large flowers (corollas averaging 30—60
mm long or more) borne singly in the axils of the upper leaves or aggre-
gated into terminal bracteate or foliose (sometimes corymbiform) racemes;
the tube of the corolla is neither fenestrate nor cleft dorsally and if the lobes
are dimorphic, it is the two dorsal ones that are the larger. Burnmstera is
distinguished from its allies by its combination of usually ebracteolate pedicels,
green or yellow corolla often suffused with maroon or purple, large falcate
or reflexed dorsal corolla lobes, the wide open orifice of the anther tube,
baccate often inflated fruit, and oblong to fusiform seeds much longer than
broad.
Most of the species oi Burmeistera are found in montane areas from Costa
Rica to Ecuador. At the southern limits of this range, two Ecuadorean spe-
cies extend south into Peru (Stein 1987). At the northern end, a single spe-
cies is known to occur north of Costa Rica: Burmeistera virescens (Benth.) Benth.
& Hook, ex Hemsl. It has been reported (McVaugh 1943; Wimmer 1943;
Nash 1976) only from Guatemala, where it occurs in the departments of
Alta Verapaz, Baja Verapaz, Huehuetenango, Quezaltenango, San Marcos,
and Suchitepequez. This is a disjunction from its congeners of nearly 700
km. That gap has been narrowed, however, by the discovery of this same
species in central Honduras. This is the first report of any member o^ Burmeistera
from that nation.
Voucher specimen: HONDURAS. Depto. Olancho: Parque Nacional laMuralla, trail
above Visitors' Center, ca. 14 km above La Union, 1 5°()5'N 86°4()'W, in dense high pri-
mary (2iiercus forest, 1400 m, 27 Oct 1996, P.J.M. Mcun & H. Aiaas 8441 (U).
The new locality in Honduras is approximately 350 km east of the near-
est conspecific populations in Guatemala. The 0//«r//j-dominated primary
forest there was quite rich in epij^hytes, including Pleurothallis tuerckheimii
Schlecht. (Orchidaceae), Columnea rulyrecaulis Standi. (Gesneriaceae), and Peperomia
hoffmannii C. DC. (Piperaceae). Undcrstory shrubs included Besleria solanoides
Kunth (Gesneriaceae), hionnina jerreyrae Taylor (Polygalaceae), Tournefortia
sp. (Boraginaceae), scandent Sphyrospermum majus Griseb. (Ericaceae), and a
treelet species oiClethra (Clethraceae). Species found in the herbaceous layer
SiDA 18(1): 363. 199<S
364 Si DA 18(1)
were Psilochilm macrophyllus (Lindl.) Ames (Orchidaceae), Renealmia mexicana
Klotzsch ex Peterson (Zingiberaceae), Smilacina sp. (Asparagaceae), Spige-
lia sp. (Spigeliaceae), the saprophyte Gymnosiphon s/iaveokns (Karst.) Urb.
(Burmanniaceae), and the root-parasite Monotropa uniflora L. (Monotropaceae).
Adjacent patches of secondary forest contained small trees oiSaurania veraguensis
Seem. (Actinidiaceae), shrubs oiTriumfetta specwsa Seem. (Tiliaceae), and the
herbs Carina tuerckhetnni Kraenzl. (Cannaceae) and Hydrocotyle viexicana Cham.
c& Schlecht. (Apiaceae).
— Thomas G. Lammers, Department oj Botany, Field Aiuse/rm of Natural His-
tory, Chicago, IL 60603-2496, U.S.A.; and P.J. M. Maas. Department of Plant
Ecology and Evolutionary Biology, Herbarium Division. WillemC. van U nnikgebouw,
Heidelberglaan 2, 3584 CS Utrecht. Netherlands.
RFFF.RF.Nr.RS
Lammers, T.G. 1998. Review ot the Neotropical endemics Burmeistem, Centropogon^ and
Siphocampylus (Campanulaceae: Lobelioideae), with description of eighteen new species
and a new section. Brittonia 50:23.^—262.
McVaugh, R. I9'13. Campanulaceae (Lobelioideae). N. Amer. Fl. 32A:1-134.
Nash, D.L. 1976. Campanulaceae. In; Flora of Guatemala, part IX, no. 4. ['ield Mus. Nat.
Hist., Bot. Sen 24:276-431.
Pnppi-R, A.S.-R., M.H.G. Gu.stafsson, and V.A. Albert. 1997. Molecular systematics of
Neotropical Lobelioideae (Campanulaceae), with emphasis on B/innasliri/. Centwpogon.
and SiphiKampyliis. and the utility of friut and floral characters in lobeHoid classification
{abstract], Amer. J, Bot. 84(6, Suppl,):222,
Stein, B.A. 1987. Synopsis of the genus 6«rwe/.f/6'rrf (Campanulaceae: Lobelioideae) in Peru.
Ann. Missouri Bot. Gard. 74:494-496.
WiMMHR, F.E. 1943. Campanulaceae-Lobelioideae, I. Teil. In; R. Mansfeld, ed. Das Pflanzenrcich,
IV.276b. Verlag Wiliielm Engelmann, Leipzig.
SiDA 18(1): 364. 1998
NoTi-s 365
TRIRAPHIS MOLLIS (POACEAE: ARUNDINEAE) A SPECIES
REPORTED NEW TO THE UNITED STATES
Prior to this report the occurrence of Triraphis and T. mollis R. Br. (purple
plumegrass) in naturahzed populations was not documented in the United
States (Chase 1951; Correll & Johnston 1970; Gould 1975; Hatch et al.
1990; Kartesz 1994; Jones et al. 1997). Specimens of this taxon were first
collected in naturalized populations by William Godwin on 15 Mar 1993
in Dimmit County. Since the original U.S. collection, William E. Fox III
collected the species in the same area on 1 May 1996.
Cultivated specimens collected from an experiment station in Biloxi,
Mississippi were located in TAES. However, specimens have not been lo-
cated that were from escaped or naturalized populations. TROPICOS has
reported this species as being present in the U.S. because of a listing by the
U.S. Department of Agriculture in their "National List of Scientific Plant
Names" (U.S.D.A. 1982). In that U.S.D.A. publication, the distribution
was not listed because the plant species was not naturalized.
The species was identified to genus using Clayton and Renvoize (1986)
and to species using Simon (1993). The identification was then verified using
the Tracy Herbarium (TAES) plant specimens from Australia and specimens
from U.S. National Herbarium (US). Lazarides (1970) reported the grass
to be grazed in Australia, but stated that it appeared to be less valuable
than many short-lived perennials. Clayton and Renvoize (1986) reported
the genus to be allied with Neyraudia because of gross morphology and the
slender microhairs.
Superficially, T. mollis resembles Aristida. However, upon examination
of the spikelets with a dissecting microscope the following observations become
apparent. Spikelets have several florets with the reduced floret apical. Also
the lemmas are 3-veined, 3-awned. The lemma awns are extensions of the
3-veins of the lemmas. In T. mollis, the central lemma awn originates from
the lemma apex and lateral awns below and on both sides of the central
awn, whereas in Aristida the lemma awns originate from a column or the
lemma apex and branch from a common point of origin.
Triraphis mollis R.Br., Prodr. El. Nov. Holl. 185. 1810. (Fig. 1).
Caespitose perennial. Culms to 50 cm tall, erect to ascending, unbranched;
nodes glabrous, purplish black; internodes solid. Leaves mostly basal; sheaths
glabrous, rounded, margins free; auricles minute; ligules a fringe of hairs
about 1 mm long; blades 6—17 cm long, 1—2 mm wide, flat becoming in-
volute, glabrous. Panicles 7—20 cm long, 8—15 mm wide (excluding awns),
contracted, somewhat interrupted, purplish tinged. Spikelets 4—12 mm long.
Sum 18(1): 365. 1998
366
Si DA 18(1:
Fit;. 1. Triraphis mollis. A. habit; B. spikclet, |;lumes separated trom Horets; C. Horct, palca
view with lemma margin and the associated hairs; and D. ligule.
SiDA 18(1): 366, 1998
Notes 367
crowded, pedicellate, laterally compressed, disarticulation above glumes and
between florets; florets 4—10, callus bearded, reduced florets 1-2 and api-
cal. Glumes subequal (not consistent in which glume is longer), 1 -veined,
hyaline, shorter than spikelets, awnless; first glumes 3.5—5 mm long; sec-
ond glumes 4—5.5 mm long; lowermost lemmas 4—5 mm long, 3-veined
and 3-awned, deeply bifid with teeth to 1 mm long, membranous, median
vein awns 6—9 mm long from between teeth of bifid apex; lateral vein awns
5.5—7 mm long, divergent; lateral veins with long pilose hairs (hairs occa-
sionally on the lemma body); lowermost paleas 2—2.5 mm long, 2-keeled,
hyaline, glabrous to scaberulous. Stamens 3; anthers 0.2—0.3 mm long, yellow.
Collection localities were characterized by deep sandy soils of the Brystal
Fine Sandy Loam (Stevens & Arriaga 1985). Historically, well drained sandy
soils derived from Carrizo Sandstone were called Norfolk Fine Sand (Carter
1931). These soils are derived from parent material in the Carrizo Sands
Formation. Carrizo Sand has been shown to have hygroscopic properties that
ameliorate the effects of a xeric climate and consequently influence its plant
formation (McBryde 1933).
Specimens examined: AUSTRALIA. New South Wales: Sydney; 18??,_/. Maiden s.n.
(TAES). Northern Territory: Simpson Desert, ca 15 km N of Andado Homestead (250
km SE of Alice Sprmgs), f 1 Jul \96^J.Weher 9^« (TAES). Queensland-Warrego Dis-
trict: Charleville, cemetery, Oct-Nov 1945, A'l, Clemens s.n. (TAES).
UNITED STATES. Texas-Dimmit Co.: 14 mi NW of Carrizo Springs on Cometa Road,
near sandy dirt road between Cometa and Hwy 277, 15 Mar 1993 W'^ Godwin s.n. (TAES);
13 mi W of Carrizo Springs, 2 mi W of EM 393 along Hwy 277, then 2 mi N on Cometa
Road, ak)ng W side of road, 1 May 1996 W.E. Fox III. 548A (TAES). Associated with Pappophor/m.
Pennisetuni. Set arid. Acacia and Prosopis.
ACKNOWLEDGMENTS
Financial support was provided in part by Texas Higher Education Co-
ordinating Board— Advanced Research Program and the Texas Agricultural
Experiment Station.
— Stephan L. Hatch, William E. Fox III, and John E. Dawson III, S. M. Tracy
Herbarium. Department of Rangeland Ecology and Managemmt, Texas A&M University,
College Station, TX 77843-2126 U.S.A.; and William B. Godwin, Department
of Entomology, Texas A&M University, College Station, TX 77843-2475 U.S.A.
REFERENCES
Carter, W.T. 1931 . The soils of Texas. Texas Agric. Exp. Sta. Bull. 431 .
Chase, A. 195 1. Hitchcock's manual of the grasses of the United States, 2nd. ed., U.S.D.A.
Misc. Publ. No. 200. U.S. Government Printing Office, Washington, D.C.
SiDA 1H(I): 367. 1998
368 Si DA 18(1)
Clavi'ON, W.D. and S.A. Renvoize. 1986. Gener;i graminum, grasses of thf world. Kew
Bull. Atldit. Scr. XIII. Her Majesty's Stacionary Office, London.
C()Riu:i.i,, D.S. and M.C. Johnston. I9"(). Manual of the vascular planrs of Texas. Texas
Research Foundation, Renner.
Gori.n, F.W. 1975. The grasses of Texas. Texas A&M University Press, College Station.
Haic:ii,S.L., K.N. Gandhi, and L.E. Browx. 1990. Checklist of the vascular i^lants of Texas.
Texas Agric. Exp. Sta. Bull. MP- 165 5.
Jones, S.D., J.K. Wiffe, and P.M. Montgomery. 1997. Vascular plants of Texas, a compre-
hensive checklist including synonymy, bibliograj^hy and index. University ot Texas Press,
Austin.
Karte.sz, J. 1 994. A synonymized checklist of the vascular flora ol the United States, Canada
and Greenland. Univ. of North Carolina Press, Chapel Hill.
Lazaridp.s, M. 1970. The grasses of central Australia. Australian Natl. Univ. Press, Canberra.
McBkydi;, J.B. I9.r3. The vegetation and habitat factors ol tlie (^arrizo Sands. Ecological
Monogr. .r2l7-297.
Simon, B.K. 199.>. A key to Australian grasses. Queensland Department of Primary In-
dustries, Brisbane, Australia.
Stevens, J, W. and D. Akriaga. 1985. Soil survey of Dimmit and Zavala counties, Texas.
U.S.D. A.— S.C'.S and Texas Agricultural Experiment Station.
SiDA 18(1): 368. 1998
PRIORITY OF THE NAME AGALINIS HARPERI
(SCROPHULARIACEAE) OVER THE NAMES AGALINIS
DELICATULA AND AGALINIS PINETORUM
This paper is part of an ongoing study by the author of the vascular plant
genus Agalinis along the Gulf Coastal Plain. The objective of this note is to
clarify one of the many nomenclatural difficulties associated with North
American species of this genus.
Pennell (19 1 3) described three species of Agalinis from Florida: A. delkatula,
A. harperi, and A. pinetorum. These species have recently been shown to be
conspecific (Canne-Hilliker and Kampny 1991). Later, in 1929, Pennell
reduced A. delicatida to a variety of A. pinetorum as A. pinetorum var. delicatula
(Pennell) Pennell, considering it to be a weak variety at best and stating in
his description that it was "Of doubtful rank" (Pennell 1929)- In their study,
Canne-Hilliker and Kampny (1991) concluded that that all three names
represented one species and and chose A. harperi for this complex. Kartesz
( 1 994), although apparently aware of the work of Canne-Hilliker and Kampny,
chose instead to combine the two species under the name A. pinetorum.
Although Pennell later recognized the name Agalims pinetorum var. delicatula
(Pennell) Pennell at the varietal rank, it must still be taken into account for
nomenclatural purposes because the name A. delicatula has equal priority
with A. pinetorum (Pennell 1913)-
The name Agalims harperi and a brief description of the species first ap-
peared in the Elora of Miami, published in 1913 (Small 1913), in which
Pennell prepared the treatment oi Agalims. On 13 August 1913, the August
issue oi Bulletin of the Torrey Botanical Club was published, and in this issue
Pennell published for the first time the description of A. delicatula and A.
pinetorum (Pennell 1913). It is clear that Pennell realized that Small's flora
had appeared (or would appear) before the August 1913 publication of the
Bulletin, this is because he (p. 426) noted that A, harperi was first described
in the Elora of Miami. But due to the fact that Small published the Elora of
Miami himself, I have been unable to determine an exact publication date
for it. However, in accordance with Art. 31-1 of the Code (Greuter et al.
1994), A. harperi was effectively published prior to publication of the 13
August 1913 issue oi Bulletin of the Torrey Botanical Club. The valid publi-
cation date is fulfilled in two ways in accordance with Articles 29-1 and
31.1 of the Code:
1. The preface of Small's Elora of Miami is dated 26 April 1913 (Small
1 9 1 3); this is the only exact date found in the flora. According to Art. 31.1
of the Code, in the absence of any other evidence validating the date of
a work, the earliest date indicating publication is to be accepted as the
SiDA L8(I): 369. 1998
370 SiDA 18(1)
correct publication date.
2. In a 1997 communication with John F. Reed, Director of the LuEsther
T. iVIertz Library at the New York Botanical Garden, I was able to verify
that the publication of Small's Flora of Miaftu did in fact precede the 13
August 1913 publication date of Bulletin of the Torrey Botanical Cl//b-i\ copy
of Small's Flora of Miami at the New York Botanical Garden's library is
date-stamped 21 June 1 91 3. ThLis, publication oi Ai^aTniis harperi in Small's
Flora of Miami obviously preceded the 13 August 1913 publication of
A. delicatula and A. pinetonnn in the Bulletin. The valid name for this
species and its synonyms is as follows:
Agalinis harperi Penned, in Small, Flora of Miami, 176. 1913.
A^cilniis ddicatulci Pennell, BliII. Torrey Bo:. Club 40:425. 1913. Agalinis pinetoriiDi var.
delicatula (Pennell) Pennell, Proc. Acad. Nat. Sci. Philadelphia 81:172-173. 1929.
Gminlia p/dchella \i\v. dcliailnL/{V{:nnv\\) l^ennell, Acad. Nar. Sci. Philadelphia Monogr.
1:441. 1935.
A. pinetorum Pennell, Bull. Torrey Bot. CluLi 40:424-425. 1913.
Gerardia harperi (PenneW) Pennell, Acad. Nat. Sci. Philadelphia Monogr. 1 : 14 1 . 1935.
C. piih-bdla (Pennell) Pennell, Acad. Nac. Sci. Philadelphia Monogr. 1:140. 1935.
ACKNOWLEDGMENTS
I thank John Thieret of Northern Kentucky University for reviewing the
manuscript and John F. Reed of the LuEsther T. Mertz Library at the New
York Botanical Garden for his assistance. I also thank Judith Canne-HiUiker
of the University of Guelph in Ontario, Canada for sharing her unsurpassed
knowledge of Agalinis with me. — -John F. Hays. Ozark Environmental Con-
sulting. 3308 Keenes Edge Drive. Columbia. MO 65201. U.S.A.
REEERENCES
CANNii-HiLLiKER, J.M. and CM. Kamp.ny. 1991. Taxonomic .significance of leaf and stem
anatomy oi Agalinis (Scrophulariaceae) from the U.S.A. and Canada. Canad. J. Bot. 69:1935-
1950.
Greutrr, W., ER. Burdet, W.G. Chaloner, V. Demoulin, D.L. EIawk.swortii, RM. Jorgensen,
D.E^^, NicOLSON, P.C. Lii.vA, P. Treiiane, and J. McNeil. 1994. International code of bo-
tanical nomenclature (Tokyo Code) adopted by the Eifreenth International Botanical Congress,
Yokohama, August-September 1993. Regnum Veg. 131.
Kartesz, J.T. 1994. A synonymized checklist ot the vascular flora of the United States,
Canada, and Greenland, 2d ed. Timber Press, Portland. 2 vols.
Pennei.l, F.W. 1913. Studies in the Agalinanae, a subtribe of the Rhinanthaceae II. Species
of the Atlantic coastal plain. Bull. Torrey Bot. Club 40:401-439.
Pi;nnell, EW. 1929. Agalinis aiul allies in North America-ll. Proc. Acad. Natl. Sci. Phila-
delphia 8 1 : 1 1-249-
Pennell, E.W. 1 935. The Scrophulariaceae of eastern temperate North America. Acad. Nat.
Sci. Philadelphia Monogr. 1:119-476.
Small, J. K. 1913. Elora of Miami. Published by the author. New York.
SiDA 18(1): 370. 1998
HEINZ DIETRICH LUBRECHT (1908-1997),
BOTANICAL BOOKSELLER AND FRIEND
WILLIAM R. BURK
Biology Library
University of North Carolina
Box 5280 Coker Hall
Chapel mil NC 27399-3280 U.S.A.
Heinz (Harry) Dietrich Lubrecht, age 88, well known botanical and natural
history bookseller, antiquarian book expert, appraiser, and former execu-
tive of the publishing company, Stechert-Hafner, died 12 November 1997,
at the Columbia Presbyterian Hospital, New York City, after suffering a
stroke. Harry, as he was affectionately known by friends, colleagues, and
family, devoted nearly 70 years to the scholarly book trade, particularly in
botany and natural history. In 1 974 he established Lubrecht & Cramer, Booksellers
and Publishers, currently located in Port Jervis, NY. His high standards of
business, steadfast honesty, and breadth of knowledge of botanical litera-
ture earned him national and international respect as a bookman. With the
passing of Harry Lubrecht the botanical and bookselling community has
lost a faithful friend and serious scholar.
Born 2 December 1908, in Reutlinger, Germany, Harry was the son of
Adolph August Karl Friederich Wilhelm and Sophie Amalie (Grueninger)
Lubrecht. His father, an architect, was the son of Lutheran Pastor Adolph
Lubrecht and Emma Karoline Kuebel. His mother, well-educated and adept
in three foreign languages, was the daughter of Johann Matthaus Grueninger,
Professor of Mathematics in the Reutlinger Hochschule, and Sophie Kaemerer.
When Harry was five, his parents separated. Subsequently, Harry, his mother,
and two sisters moved to Stuttgart to live in an aunt's apartment home. He
attended Friedrich Eugens Oberrealschule and completed Hochschule in
Stuttgart. Throughout his school years Harry was an avid reader. In order
to ensure enrollment at his school, Harry secured an annual scholarship by
placing in the top five of his class.
Upon graduation at age 17, Harry became an apprentice in Brueninger's
department store as part of his continued schooling. His flare for business
was well recognized by the store's management and in three years he earned
the salary of a regular employee. Refusing to join the Nazi party, he was
compelled to leave Germany. Harry emigrated to the United States where
his Uncle Christian Grueninger, the mayor of Valhalla, NY, would sponsor
him. Arriving in America on 15 May 1928, he soon found a job as a grave
SiDA 18(1): 371. 1998
372 SiDA 18(1)
digger ar the local cemetery in Mt. Kensico, NY. When rhis seasonal work
concluded char fall, Harry looked for employment in New York City. In
October 192S, G.E. Stechert & Co.,^ well-known international book firm,
hired him at a weekly salary of $14.00. The stage was thus set for a distin-
guished, life-long career in the book world, a profession practiced by an-
cestors on two sides of his family. Some of the Kuebels were medieval scribes,
the book "publishers" of their time. In the early nineteenth century two
Lubrecht brothers, Heinrich and August Adolph, separately developed ca-
reers as booksellers in Heilbronn and Blaubeuren, Germany. In the mid-
nineteenth century August's son, Karl Theobold, emigrated from Blaubeuren
to Brooklyn, NY Here he Anglicized his name to Charles T. and founded
with his cousin the publishing company Haasis and Lubrecht.
At Stechert Harry advanced from his initial position as billing clerk to
assistant manager of the Antiquarian Department (April 19.^6). He was a
natural for this job as he was familiar with German, French, Latin, and Greek.
His facility for languages assisted him in compiling book catalogs and un-
derstanding the contents of many scientific books, especially those in Ger-
man, acquired by the company. Since his immediate boss lacked signifi-
cant experience in the antiquarian book market, Harry enjoyed the freedom
and breadth to develop the antiquarian department about which topic he
read widely. Under his supervision the process of searching for and supply-
ing out-of-print books was significantly accelerated (Anonymous 1952). Among
Harry's promotions were to: Chief of Reference and Customer Service, G.E.
Stechert & Co. (spring 1945); Division Chief, Antiquarian and Out-of-Print
Department, Stechert-Hafner, Inc. (fall 195 1 ); Vice-President and Editor,
Hafner Publishing Co. (January 1955); and Vice President, Macmillan Publishing
Co.-Hafner Publishing Division (1969).
After conducting a four-year study on subject publishing, Harry con-
cluded that few firms published books on botany. Through Harry's efforts
a new subsidiary called Stechert-Hafner Service Agency was established in
1960. Botanists and natural historians are fortunate to have had Harry's
'a brief chronology of this book Hrm provides an historical perspective. Stechert & Wolff,
founded on 1 September 1872, was renamed Gustav E. Stechert (1876), G.E. Stechert (1897),
and G.E. Stechert & Co. (1904). On 1 October 19^6, the G.E. Stechert partnership was
dissolved and Stechert-Hafner, Inc. (SH) was created with its publishing and reprinting
activities transferred to the newly established Hafner Publishing Co. In 1 96t) Stechert-
Hafner Service Agency was established as a subsidiary of SH. Crowell Collier Jvlacmillan,
Inc. (now Macmillan, Inc.) acquired SH and its subsidiaries (1969). Beginning January
1974, the corporate name, Stechert Macmillan, Inc., became effective. By 1980 Macmillan,
Inc. was divided into three parts of which Macmillan Publishing Co., Inc. contained the
division now called Hafner Press.
SiDA 18(1): 372. 1998
Bdrk, Harry Lubrecht
373
Harry Lubrechr making a presentation at the Botanical Society of America Meeting, His-
torical Section, held at The Ohio State University, Columbus, 1 1 August 1987. Photo Courtesy
Ronald L. Stuckey.
wisdom in developing this subsidiary. This new division served scholarly,
scientific and botanical societies by selling their publications and distrib-
uting scientific and botanical books for European publishers on a large scale.
The Agency distributed publications for the New York Botanical Garden,
Missouri Botanical Garden, Texas Research Foundation, and the Hunt Bo-
tanical Library (Anonymous 1972). During the 1960s and 1970s Harry also
focussed attention on reprinting book classics in botany within the Hafner
Publishing Co. Among the reprint series issued were the Classica Botanica
Americana (CBA) titles by such eighteenth and nineteenth century lumi-
naries as Humphry Marshall, John Torrey & Asa Gray, C. S. Rafinesque,
Stephen Elliott, Thomas Nuttall, and William Darlington. Joseph Ewan,
historian of botany, proposed this series and wrote a bibliographical com-
mentary for each of the reprinted books (Ewan 1967). Ironically, these fac-
simile reprints, as well as many others produced by Hafner, have become as
rare as the originals themselves. The rarity of these volumes was due in part
SiDA 18(1): 373. 1998
374 SioA 18(1)
to their limited printing, high demand by scliolars, and the destruction of
remaining stock when Crowell CoUier Macmillan, Inc. ptirchased Stechert-
Hafner and its subsidiaries in May 1969.
After this corporate change Harry chose to direct the PubHshing Divi-
sion. He could not, however, abandon his interest in and passion for rare
and out-of-print books. Using a substantial part of his lunch hour, he would
work in the Antiquarian Department at noon time. Through his voluntary
work, a veritable labor of love, Harry raised impressive revenues for the firm.
While at Stechert, Harry especially enjoyed appraising private collec-
tions for purchase. He was instrumental in acquiring an outstanding mathematics
library in 1 937, besides many other collections. Representing his company,
Harry participated in the First Antiquarian Book Fair in the United States,
sponsored by the Antiquarian Booksellers' Association of America and held
in New York City (4-9 April 1960). Confident that the fair would be suc-
cessful, he prepared an attractive catalog (#277) of rare books, the first ever
issued from the Stechert-Hafner Antiquarian Department (Anonymous 1 960).
One of his career highlights was the discovery of 1 1 first editions of the
works of Linnaeus in a Paris bookshop. These books had an additional his-
torical value since they were annotated by the French botanist and explorer
Michel Adanson (1 727-1806), noted for his work on the natural classifica-
tion of plants (Lubrecht 1986).
Harry compiled numerotis book catalogs covering general topics, Greek
and Latin authors, and natural history. Although no longer in print, his
catalog (#353), "Early American Botanical Works with a Miscellany of Other
Botanical Rarities" (Lubrecht 1967), provides an historical survey of early
classics in Nortii American botany. This booklet, the result of five years of
searching (Anonymotis 1 967), contains an introduction (pp. 3—5) by Harry,
an annotated list of 217 botanical titles offered for sale, and a section (pp.
55-60), "Reference Tools for Botany," by Joseph Ewan. The Missotiri Bo-
tanical Garden plans to issue a facsimile reprint of this item as a keepsake
of the XVI International Botanical (Congress to be held at the Garden in St.
Louis (August 1999). Through the enthusiastic support of Harry, Hafner
Publishing Co. published and assisted in the production of "A Short His-
tory of Botany in the United States," edited by Joseph Ewan (1969). This
book was distributed at the XI International Botanical Congress in Seattle,
WA, August/September 1969.
Harry retired from Stechert Macmillan on 30 November 1973 and then
briefly compiled catalogs for Jack N. Bartfield, an antiquarian bookseller
of New York City. In December 1 974 Harry founded his own business, Lubrecht
& Cramer, Booksellers and Publishers, which was later incorporated (Au-
SiDA hSd): 371. 199H
BuRK, Harry Lubrecht 375
gust 1 983). Joerg Cramer (deceased 4 June 1 985), a German publisher, specialized
in reprinting classical botanical books, many of which Hafner had distrib-
uted in the United States. Harry's wife Anne, and later his son Charles, assisted
in the business, originally carried out from their country home in Forestburgh,
NY, and dealing predominantly with new, scholarly books, mostly in botany
and natural history. The company is also a distributor for Koeltz Scientific
Books, Gustav Fischer Verlag, and J. Cramer. Although his bookselling company
primarily was concerned with new titles, Harry continued his profound interests
in rare books and the appraisal of libraries. Among the botanical collec-
tions that he appraised were those of Carroll W. Dodge, the New York
Horticultural Society, Emanuel D. and Ann W. Rudolph, William C. Steere,
and R. Gordon Wasson. The remarkable Rudolph Library of 53,()()() vol-
umes of botany and natural history books has since been donated to The
Ohio State University Libraries (Stuessy et al. 1997).
For more than two decades Harry and Anne Lubrecht featured booksale
tables at botanical and scientific meetings and regional mycological forays.
Their service was known not only for the sale of books, typically sold at a
special discount for meeting and foray attendees, but also for their conge-
nial and knowledgeable conversations. Harry captivated customers with his
storehouse of botanical and bibliographical knowledge. He and his wife regularly
attended the annual meetings of the Council on Botanical and Horticul-
tural Libraries, Inc., and national and international book fairs. Concerning
the latter, he attended 49 annual exhibits of the Frankfurt Book Fair in Germany.
Harry was a modest and caring person, yet he forthrightly and candidly ex-
pressed his opinions. I remember him also for many reminiscences of his book
trade career and the botanists with whom he associated. With a rich background
of experience and strong sense of the book market, Harry successfully intro-
duced the quality paperback, also known as the soft bound, to the scholarly book
trade. Among the first in his field, he adopted photo-offset printing as the pro-
cess for producing reprints. Even when this was not widely practiced, Harry
encouraged minority hiring and developed a staff fluent in numerous languages.
He held membership in the American Anriquarian Booksellers' Association,
American Institute of Biological Sciences (AIBS), Council on Botanical and
Horticultural Libraries, Inc., International Association for Plant Taxonomy,
Mycological Society of America, New York Botanical Garden, North Ameri-
can iVLycological Association, and Old Book Table (NY). Harry became a U.S.
citizen on 14 December 1936. As a member of the Lutheran Church, he was a
deacon during the 1960s and early 1970s. Among his honors was a Special Service
Award from the Mycological Society of America presented at the annual meet-
ing of the AIBS, San Antonio, TX, August 1991.
SiDA 18(1): 375. 1998
376 SiDAl8(l)
Harry's successful life was shared with his wife Anne (Johanne Marie Ficke)
whom he married on 2 October 1937. Harry is survived by his wife Anne;
two sons, Peter Thomas Lubrecht, Sr., a retired teacher of Drama and En-
glish and presently Artistic Director of Brundage Park Playhouse, of Randolph,
NJ, and Charles Frederick Lubrecht, co-manager of Lubrecht & Cramer, of
JMontgomery, NY; three grandsons: Peter T. Lubrecht, Jr., Christopher C.
Lubrecht, and Bryan Leone. Harry's two sisters, Use and Ruth Lubrecht,
predeceased him. Funeral services were held 15 November 1997 in St. Peter's
Lutheran Church, Port Jervis and cremation took place at the H. G. Smith
Crematory, Stroudsburg, PA. Interment of his ashes will take place at the
Forestburgh Cemetery. As a tribute to Harry, a memorial program in cel-
ebration of his life will be held at The New York Botanical Garden in the
fall 1998. Memorial contributions may be sent to The LuEsther T. Mertz
Library, New York Botanical Garden, 2()0th Street and Southern Boule-
vard, Bronx, NY 10458.
ACKNOWLEDGMENTS
I am grateful for information on Harry that Anne, Charles, and Peter
Lubrecht provided. Ronald L. Stuckey suggested this biographical essay and
supplied the photograph. The aforementioned individuals and Rogers McVaugh
read the manuscript and made constructive suggestions for changes in the
text. Additional information on Harry was obtained from a feature column
on bookmen (Cfhernofsky] 1986).
REFERENCES
Anonymous. 19')2. Out-of-Print Departmfnt expands. Stechert-I lafner Book News 6(6):H8.
1960. First Antiquarian Book Fair in America. Antiquarian Bookman
25(18):1"'{) 1-1706.
1967. Early American botany catalog. Stcchert-Halner Book News 22(1 ):6.
.. 1972. 100 years [ol} serving libraries. Stechert-Halher Book News 26(5):H— It.
C{iiernotsky], j.L. 1986. Lubrecht spans two eras of bookselling history. AB Bookman's
Weekly 78(5):.i57-358.
Ewan, J. (cd.). 1 967. Ckissica Botanica Americana, ppi. 38-4(), In {H. Lubrecht], Early American
Botanical Works with a Miscellany of Other Botanical Rarities. Stechert-Hafner, Inc.,
New York.
1969. A Short Mistory of Botany in the United States. Hafner Publishing
Co., New York and London.
{Lubrecht, H.]. 1967. Early American Botanical Works with a Miscellany of Other Bo-
tanical Rarities. Stechert-Flafncr, Inc., New York.
1986. Peregrinations of a rare-bookdealer in Europe. AB Bookman's Weekly
78(1 ()):808, 810.
Stuessy, T.E., R.L. Stuckey, W.L. Boomgaarden, and W.R. Burk. 1997. Botanical libraries
and herbaria in North America. 2. The Rudolph natural history library and its acquisi-
tion by The Ohio State University. Taxon 46:643—648.
Sii).-\ 18(1): 376. 1998
First report of the genus Burmeistera (Campanulaceae) from Honduras
Thomas G. Lammers and P.J.M. Maas
363
Triraphis mollis (Poaceae: Arundineae) a species reported new to the United States
Stephan L. Hatch, William E. Fox III, John E. Dawson III, and William B. Godwin
365
Priority of the name Agalinis harperi (Scrophulariaceae) over the names Agalinis delicatula
and Agalinis pinetorum
John F. Hays
369
In Memoriam — Heinz Dietrich Lubrecht (1908-1997), botanical bookseller and friend
William R. Burk
371
Book notices and reviews 192, 236, 240, 246
Index of new names and new combinations in this issue
Agave gracilis Garcia-Mend. & E. Martinez, sp. nov. 227
Corsia purpurata var, wiakabui Takeuchi & Pipoly, var. nov. 164
Crataegus okanagensis J.B. Phipps & O'Kennon, sp. nov. 178
Crataegus okennonii J.B. Phipps, sp. nov. 170
Crataegus series PurpureofructiJ.B. Phipps & O'Kennon, ser. nov. 184
Crataegus phippsii O'Kennon, sp. nov. 185
Cybianthus anthuriophyllus Pipoly, sp. nov, 89
Cybianthus cenepensis Pipoly, sp. nov, 106
Cybianthus comperuvianus Pipoly, sp. nov, 47
Cybianthus croatii Pipoly, sp. nov, 71
Cybianthus flavovirens Pipoly, sp, nov, 134
Cybianthus fosteri Pipoly, sp. nov. 144
Cybianthus grandezii Pipoly, sp. nov. 139
Cybianthus granulosus Pipoly, sp. nov. 133
Cybianthus guyanensis subsp. pseudoicacoreus (Miq. in Mart.) Pipoly, comb, et stat. nov. 49
Cybianthus huampamiensis Pipoly, sp. nov. 130
Cybianthus incognitus Pipoly, sp. nov. 125
Cybianthus jensonii Pipoly, sp. nov. 142
Cybianthus kayapii (Lundell) Pipoly, comb. nov. 84
Cybianthus nestorii Pipoly, sp. nov. 59
Cybianthus pseudolongifolius Pipoly, sp. nov. 101
Cybianthus timanae Pipoly, sp. nov. 53
Cybianthus vasquezii Pipoly, sp. nov. 103
Mandevilla pringleiJ.K. Williams, sp. nov. 231
Mandevilla holosericea (Sesse & Mog.) J.K. Williams, comb. nov. 237
Psychotria osiana Takeuchi & Pipoly, sp. nov. 162
Ruellia jimukensis Villarreal, sp. nov. 223
Sporobolus pinetorum Weakley & P.M. Peterson, sp, nov. 258
Trachypogon mayaensis Wipff & S.D. Jones, sp. nov. 242
Xanthostemon fruticosus Peter G. Wilson & L. Co, sp. nov. 283
Seasonal changes in concentration and distribution ot heavy metals in creosotebush,
Larrea tridentata (Zygophyllaceae), tissues in the El Paso, TX/'Ciudad Juarez, Mexico area
William P. Mackay, Richard Mena, Nicholas E. Pingitorejr., Keith Redelzke.
C. Edward Freeman. Harold Newman. John Gardea. and Hector Navarro
287
New plant records for Dominica, Lesser Antilles
Steven K. Hill and Arlington James
297
A new adder's-tongue (Ophioglossuni: Ophioglossaceae) for North America
James C. Zech, Patricia R. Manning, and Warren Herb Wagner Jr.
307
Notes on the llora of Texas with additions and other significant records
Larr}< E. Brown and Stuart J. Marcus
315
Cyperus fuscus ((^yperaceae), new to Missouri and Nevada, with comments
on its occurrence in North America
PaulM. McKenzie. Brad Jacobs. Charles T. Biyson. Gordon C. Tucker, and Richard Carter
325
The use of animal-dispersed seeds and fruits in forensic botany
Barney L. Lipscomb and George M. Diggs, Jr.
335
NOTES
Glaucium corniculatum (Papaveraceae) in Texas
Zoe Kirkpatnck andJ.K. Williams
yti
Hovenia dulcis (Rhamnaceae) naturalized in central Texas
Douglas H. Goldman
350
Habranthus tubispathus (Liliaceae) new to the flora of Alabama
Alrin R. Diamond, Jr., Charles P. Chapman, and Jim Bmmmett
353
Plantago coronopus (Plantaginaceae) new to Texas
Robert J. O'Kennon. George M Diggs, Jr.. and Ro)iald K. Hoggard
356
Pteris vittata (Pteridaceae), a new fern for Texas
Jack W. Stanford and George M. Diggs, Jr.
359
Lycianthes asarifolia (Solanaceae) new and weedy in Texas
Morn que Dubnile Reed and Mary Ketchersid
361
(continued on inside back cover)
ISSN 0036-1488
CONTENTS
Arnoglossuni album (Asteraccac): New s|X'cies Ironi iiorilicrn Florida
l.iinin C. Aiidei'so}]
377
Sc\-en new nonienciauiral coiiihinatioiis and a new name in Packera fAsieraceae: Senecioneae)
Dchni K. 1 rack and Theodore M. Barkley
Ardisia niambiensis (Mvrsinaeeae). a new species of Ardisia subgenus Ardisia from ihe ('lioeo
I'ioristic i^r()\-ince of (iolonibia
John 1^ Pipoh' 111 (iiid AIniro (J);^olh} P.
389
Clusia niambiensis (Ckisiaceae), a new species from the Choco Florisiic Province of Colonihia ami
Kcuador
John J. Pipoly HI, Alvam Cooallo I' . aiid Maria Sofia Gonzalez
395
New species and nomenclatural notes in Clusia (Clusiaceae) from Andean Colomliia and \'ene/uela
John J. Pipoly HI and Alvaro CogoUuF
401
Dos nuevas adiciones a la orquideoflora iMexicana
Adolfo Kspejo Sema, Ana Rosa Lopez^Femm, Javier Carcia Cruz, Rolando Jimenez Machorroy Luis
Sanchez Saldana
111
Seedling development in species of Chamaesyce (Euphorbun eae) with erect growth habits
U John Hayden and Olga Troyanskaya
4 I')
A revision of the genus Ardisia subgenus Graphardisia (Myrsinaceae)
John J, Pipoly III and Jon M. Ricketson
^ii
Una nueva especie de Polianthes (Agavaceae) del estado de Oaxaca, Mexico
Eloy Solano Camacho andAbisai Carcia-Mendoza
473
Pollen morphology of the genus Echinopcpon (Cuciirbitaceae)
Concepcion Rodriguez J. and Rodolfo Palacios-Chdvez
479
Variation in the Berlandiera pumila (Asteraceae) complex „
G.L. Nesom andB.L. Turner
493
New names and combinations in neotropical Myrsinac eae
J()h)iJ. Pipoly HI and Jon M. Ricketson
S(H
A new species of filmy fern (Hymenophyllaceae: Pteritinphyta) from South India
C. Ahdiil Ilameed and P V. Madhusoodanan
SI9
A second species of Oritropliium fAsteraceae: Astereai) from Mevico
(Iiiy L. Nesom
523
Unic|Lieness of the endangeied [joricia Semaphore Cacuis (Opuntia cofallitola)
Ikniiel F. Austin. Darid .\l. Biiininoer. and Donald J. I'inkava
Ariuinsas Carex (CAperaceae): A lirieflv annoiated list
Philip E. Hyatt
535
(continued on thick corer)
COM'RIBl TIOXS
IX) BOTANY
♦
VOLUMF !H
NuMni'R 2
Dhcfmbhr, 1998
biDA
CONTRIBl'TIONS T(3 BOTANY
I'oi \i)i:i) li^
Lloyd H, Shinnei
19G2
Wm. !■. Mahler
PulilisluT 19^1-1992
— - vf
l^ai"nL'\" I.. Lipscomb
I'dilor
Hoianical Kcscaaii liisiiiuic ot Tcxa.s, Inc.
Sn9 Pecan Slivct
I'ort \\'o\-\h. Texas 7(il02- H)()(), liSA
817 332- Mil sr 332-a 112 FAX
I'.leclfoiiic mail: sida@hril.()i;^
lome page al the IRL: http: \\\\ w.hnl.orj^ siJa
John W, Thieret
Associate hxlitor
Hiol(\L;ical Sciences Dept.
Northern Kenluckv l'ni\ersily
limhlanJ llemhis. Keiitiick\- i KPfi, I'SA
Prof. Dr. Irli.x Llamas
Coiitrihulint^ Sjxmish I-^iilor
Dplo. de Botfmica, L'acullad de Hioloi^i
llni\ersicLid de Leon
L>2 i()~l Leon. Spain
(iiiuk'LiK's lor (."iiiiii"ihiiiiii> aa' avaihibk' upon iwjlk'sI
• ind 1)11 ihi- inside Ixick cover ot the la-it issue ol c.id] xohinie.
SuhstTiption per year: S_!t. iiuli\RliiaL Si^. ISA liisiiuilion.s, J>sl), ( )ulsid(.- kSA;
[lunilxT.s issued twice a \ear
.-WD
•'SlDA. (X)NTK11UTI()NS TO BOTAX^'. \oliinie IS, Xiimher 2, jiai^es r^-(^M)
eopMi^i^hi i')')s
i5olanieal Kese.uxii liisiiiuie ol Te.\as. Iiic
Printed in tin- I nitetl States ol Anierua
ISSX (11)3(1- 1 |S8
ARNOGLOSSUM ALBUM {ASrEKACEAE): NEW
SPECIES FROM NORTHERN FLORIDA
LORAN C. ANDERSON
Department of Biological Science
Florida State University
Tallahassee, FL 32306-4370, U.S.A.
ABSTRACT
The new species Arnoglossum album is described. Its phyllaries have prominently winged
keels — a feature shared with ^. diversifolium, A. floridanum, A. plantagineum, andy4. sulcatum.
Comparisons among members of this closely knit assemblage include morphology, phe-
nology, ecological setting, and range, and a key is provided to distinguish the species.
RESUMEN
Sc describe una especie nueva Arnoglossum album. Sus filarios tienen qtiillas aladas
prominentcs — una caracteristica que comparte con A. diversifolium , A. floridanum, A.
plantagineum, y A. sulcatum. Las comparaciones entre miembros de este conjunto fuertemente
reticulado incluye morfologia, fenologia, caracteristicas ecologicas y areal. Se ofrece una
clave para diferenciar las especies.
Species of the x.w?dA?i^\wd\di Arnoglossum, commonly called "Indian Plantains,"
were formerly placed in the heterogeneous '^Cacalia" (Krai & Godfrey 1958;
Cronquist 1978; Pippen 1978), but, because Cacalia is now typified by a
species of the senecionoid European genus '' Adenostyles' and because ge-
neric circumscriptions are now more narrowly drawn, Arnoglossum is con-
sidered distinct by many recent authors (Robinson 1974, 1980; Jeffrey 1979,
1992; Godfrey &Wooten 1981; Wetter 1983; Funk 1985; Brummitt 1992).
Arnoglossum has three species with extensive ranges in the eastern United
States \A. atrip licifolium (L.) H. Robins, andyl. reniforme {Wook.) H. Robins.]
or southeastern United States [A. ovatum (Walt.) H. Robins.] and one from
the central part of the country \A. plantagineum Raf ] . The remaining three
species have more limited ranges, one as a Florida endemic [A. floridanum
(A. Gray) H. Robins.] and the others with ranges barely extending from
Florida into neighboring states [A. diversifolium (Torn & A. Gray) H. Robins,
andyl. sulcatum (Fern.) Fi. Robins.] . The latter four species {i.e., A. plantagineum
and "the Florida group") are similar in having phyllaries with keels that are
strongly winged. The new species described below is now added to this assemblage
with winged phyllaries. . ■ - • -:
SiDA 18(2): 377-384. 1998
378 SiDA 18(2)
MATERIALS AND METHODS
Fresh and dried materials were processed as described by Anderson (1994).
Heads from at least five different collections per species were dissected and
measured for involucral and floral quantitative data. Living plants of the
five species under study were pro]:)agated from seed or as transplants in a
greenhouse or common garden in Tallahassee, Florida. Vouchers of seed-
lings, plants at various stages of development, and all cited specimens are
deposited at FSU unless noted otherwise.
SPECIES DESCRIPTION
Arnoglossum album L.C. Anderson, sp. nov. (Figs. 1, 6-9). Tyei,: U.S.A.
FLORIDA. Bay Co.: locally common in acidic, poorly drained .sandy soil of open,
wet savanna with Sarracenia flava, Rhynchospord, Rhtxici, and Polygala spp. just E of
Rte 77 on N side of Southport, T2S, R14W, NElAl of SEl/4 Sec 21, 6 Jun 1995,
L.C. Amkrmi 75555 (HOLOTYPE: NY!; ISOTYPES: BRIT! FSU! MO! US!).
Ilerbac perennes plertimqLie 8—1 dm aka. Folia basalia 20—63 cm lon^'a, 5—16 cm lata,
longe petiolata, laminis late ovatis vel anguste oblongo-lanceolatis, verms lateralibus principalibus
per 2—4 cm supra basin laminae ad venam mediam arete parallelis dein divergentibus. Folia
caulina pauca, 7-26 cm longa, petiolata, serrata, foliis superioribus reductis sessilibusque.
C^apitula in cymis corymbosis disposita, involucris 10—1 3.4 mm longis, piiyllariis albis carinis
promincnter alatis. CoroUae albae, 9—10.5 mm longae.
Essentially glabrous cespitose herbs. Stems (5-)8-l()(-l 0.5) dm tall, light
green, strongly ridged. Basal (radical) leaves (2()-)3()-55(-63) cm long overall,
5-10(-l6) cm wide, long petiolate, (1()-)1 5-18(-3l) cm, blades broadly
ovate to narrowly oblong-lanceolate, margins entire, shallowly sinuate, or
rarely serrulate-denticulate, bases attenuate, apices rounded, mucronulate,
basally disposed lateral veins appressed to the midrib 2-4 cm then abruptly
extending toward the leaf margins; cauline leaves few, blades (7-)12— 1 8(—
26) cm long overall, (2. 5-)4-5(-6) cm wide, petioles (I-)4-5(-l 2) cm long,
ovate, bases cuneate, apices acuminate, serrately toothed, upper leaves re-
duced in size and sessile. Inflorescence compound, freely branched, ± flat-
topped, with heads crowded in corymbose cymes. Heads 5 -flowered; in-
volucres 1 {)— 1 2.5(— 1 3.4) mm long, cylindric, phyllaries 5, chalky white,
margins hyaline, keels prominently winged (the wings rising 1 — 1.5(— 2.0)
mm from the phyllaries with margins sinuate to erose and reduced apically).
Receptacles with short central cusp. Corollas white, rarely tinged with pink,
9—10.3 mm long, veins 10, tube 5-6.4 mm, throat 0.2—1 .0 mm, lobes 2.9-
3.8 mm, lanceolate; anthers 2.5—2.6 mm long, collars cylindrical; styles 1 1 .8—
12.5 mm long, style branches 1.5-1.7 mm long, stylopodium immersed
in nectary; achenes glabrous, 4.2—5 mm long, cylindric to narrowly clav-
ate, carpopodium cells erect, pappus white, 6-7 mm long, n = 25.
Di.strihution. — Endemic to Bay and Gulf counties of Florida in poorly
Andrrson, a new species of Arnoglossum
379
Figs. 1—24. Representative leaves of selected Arnoglossum species, all at 1/6 full size. Figs.
1-5. Radical leaves showing major venation. Fig. \ . A. album with appresscd lateral veins,
Anderson 15600, Bay Co., FL. Fig. 2. A. divcrsifoliunu Anderson 13414, Levy Co., FL. Fig.
3. A. flortdanurn, Anderson 72/65, Taylor Co., FL. Fig. A. A. plantagineum, Anderson 12166,
Riley Co., KS. Fig. 5. A. sulcatum, Anderson 16035, Santa Rosa Co., FL. Figs. 6-24. Out-
lines of cauline leaves (major venation similar to respective radical leaves). Figs. 6—9. A.
album. Fig. 6. Anderson 12137, Cuh Co., FL. Figs. 7-8. Anderson 15600, Bay Co., FL.
Fig. ^. Anderson 13432, Gulf Co., FL. Figs. \{)-\5-A. diversifolium. Fig. \'^. Anderson 13475,
Levy Co., FL. Figs. 11, 13. Anderson 13478, Putnam Co., FL. Fig'^ 12. McDamel 9059,
Houston Co., AL. Figs. 14—15. A. floridanum. Fig. 14. Anderson 12160, Taylor Co., FL.
Figs. I'b-lG . Anderson 12077, Clay Co., FL. Figs. 17-20. A. plantagineum. Fig. 17. Mehrhoff
12299 (CONN), Bruce Co., Ontario Prov., Canada. Fig. 18. Anderson 11991, Loundes
Co., AL. Fig. 1 9. Bryson 9861, Oktibbeha Co., MS. Fig. 20. Anderson 11968, Tangipohoa
Par., LA. Figs. 1\ -24. A. sulcatum. Fig. 2\. McDamel 7978, Geneva Co., AL. Fig. 22. Anderson
12342, Leon Co., FL. Fig. 23. Anderson 16058, Walton Co., FL. Fig. 24. Anderson 16057,
Walton Co., FL.
380 SinAl8(2)
drained, acidic (pH 5-6), loamy sands (with 0.35-1.7% organic matter)
of wet savannas, open pinewoods, and most frequently the ecotone between
the two, often in close association with several of the following: Aletris liitea,
Aristida heyrichiana, Asclepias longifolia, A. michauxii, A. viridula. Aster
eryngiifolius, Cleistes bifaria, Drosera capillaris, Eriocaidon compressum, Hy-
pericum gymnanthum, Lachnanthes caroliniana, Lachnocauion anceps, Lo-
heliaflorid^ina, Ludwigia linifolia, L. microcarpa, L. virgata, Murshallia tenuijolia,
Oxypolis greenmanii, Panicum rigididum, Physostegia godfreyi, Platanthera
nivea, Polygala balduinii, P. cruciata, P. ramosa, Rhexia lutea, Rhynchospora
ciliaris, R. curtisii, R. fascicularis^ R. filifolia, R. globularis, R. oligantha, R.
pusilla, R. rari flora, R. wrightiana., Rudbeckia graminifolia, Ruellia noctiflora,
Sabatia bartramii, S. campanidata^ Sarracenia flava, Scutellaria floridana,
Tofieldiaracemosa, Verbesina chapmanii, Xyris ambigua, X. baldwiniana, and
X. elliotii. The woody associates (less frec[uent) include: Cyrilla racemiflora,
Hypericum cistifolium, H. exile, H. reductum. Ilex coriacea, I. vomitoria. Magnolia
virginiana, Myrica cerifera, Nyssa ursimi, and Pinus elliottii. Flowers June to
mid-July.
Representative specimens examined. U.S.A. FLORIDA. Bay Co.: rvpc locality, /..C/l;/('/i:';w;^
/ 552.9 (FSU), 20 Jim 1995, L.C. Anderson 15600 (I'SU, MO, NY), 20 (un 1995, L.C.
Anderson 1 5601 (FSU), L. C. Anderson 15708 (FSU); S side County Rd 2300, 1 .9 mi W of
Rte 77. 1 .5 air mi NW of Southport, 1 3 Jim 1 995, L.C. Anderson /5577(FSU); 0.5 mi E
of Butnt Mills C;rcek bridge on S side of Rte 388, 1 .9 mi W of Rte 77, 1 .5 air mi WNW
ofSouthporr, 13 jun 1995\ L.C. Anderson I558I (FSU). Gulf Co.: 5.3 miSofRte22on
E side of Daniels Rd near Wetappo Creek, ca. 5.5 air mi WSW of Wewa- hitchka, L.C.
Anderson 1 1642 (FSU), L. C. Anderson 1 167.3 (2), L.C. Anderson 1 1934 (FSU), L.C. Anderson
/2/37(BRi'F FSU, CiH, MO, NY, US), L.C. Anderson 121.38 (FSU), L.C. Anderson 1.3421
(FSU), 5.2 mi S of Rte 22 on W side Daniels Rd, L.C. Anderson 11950 (FSU), L. C Anderson
12171 (FSU).
niSCUSSiON
The Indian plantains with winged phyllaries form a closely related alli-
ance of species. They differ from each other in several floral features as summarized
in Table 1 . The longest involucres occur in A. floridanum, and the shortest
in ^. sulcatum.. Phyllaries mA. album have prominent wings proximally
that become highly reduced apically, whereas in all other species the wings
are shorter in height but ± equally cieveloped for the length of the bract
(often broadest apically). Arnoglossum album also has the whitest phyllar-
ies; in the other species they are more greenish-white.
Corollas oiA. album are white, rarely tinged with pink; in the other species
they are white, light lavender, or ochroleucous (more greenish-white or yellowish-
white in A. floridanum and sometimes more pinkish in A. diversifolium) .
Corollas are generally longest in A. floridanum and shortest in A. sulcatum,
but not as short as reported by Pippen (1978) and Cronquist (1980). In all
species the corolla tubes are relativelv long and the throats below the lobes
Anderson, A new species of Arnoglossum 381
Taiu I 1 . Size floral features (in mm) of "winded" Arnoglossum species.
Taxon
Involucre length
C^orolla length
Tube width
Lobe length
A. album
10.0-13.4
9.0-10.5
0.9-1.0
2.9-3.8
A. diversifolium
A. floridanum
10.2-11.0
11.0-14.8
9.1-10.2
9.8-12.2
0.8-0.9
0.8-0.9
3.1-3.5
2.8-4.1
A. plantagmeum
A. sulcatum
9.8-12.8
8.5-9.9
9.2-11.4
7.8-9.6
0.7-0.8
0.6-0.7
3.1-3.6
2.7-3.4
very short. The tube widths are greatest in A. alburn and most slender in A.
sulcatum. Carpopodial cells of the achenes in A. album are generally erect,
whereas they are procumbent in A. plantagineum. Cell shape varies some-
what within a given population because of differing stages of achene matu-
rity. Robinson and Brettell (1973) suggested carpopodial structure was an
excellent character for distinguishing genera in the Senecioneae; they re-
corded procumbent carpopodial cells iox Mesadenia (i.e., Arnoglossum) .^&\ter
( 1 983) found considerable variation in this and other micromorphological
characters and questioned their value as generic markers.
Vegetatively, yi . album looks most X'^lq A. plantagineum, and that species
is probably its closest relative. The two differ in seedling morphology. Cotyledons
ofyl. album are 1-1.5 cm long and 9-12 mm wide (± orbicular), whereas
they are 3—4.5 cm long and 8—9 mm wide (spatulate-oblanceolate) in ^.
plantagineum.
The five species differ more in leaf morphology than they do in floral
features (Figs. 1—24). Basal leaves oi A. album have distinctive venation.
The principal lateral veins are appressed and closely parallel the midvein
into the blade for a few centimeters and then spread abruptly toward the
leaf margins (Fig. 1). This feature of appressed lateral veins is also found in
the related Yermo xanthocephalus ofWyoming. In the oxhei Arnoglossum species,
the lateral veins spread immediately at the base of the blade (Figs. 2-5).
Relative length of the petiole on radical leaves is often used in keys to
the species (Krai & Godfrey 1958; Pippen 1978; Cronquist 1980). This
feature is correlated to habitat preferences of die species. Generally, A. diversifolium
andy4. sulcatum grow in deep shade, and their radical leaves have long peti-
oles (Figs. 2, 5). Arnoglossum floridanum usually grows in full sun, and its
radical leaves have short petioles (Fig. 3). Petiole lengths are more variable
in radical leaves of /I. album ^ind A. plantagineum. These two species often
grow in savannas or prairies. Their petiole lengths are related to the density
of grass and other vegetation near the plants; plants from the same seed
source that were grown in different shade or vegetation densities in my garden
produced short-petioled leaves in sunnier or less crowded situations and
longer petioles in shaded or crowded conditions.
382 SmA 18(2)
Cauline leaves of A. diversifolium (Figs. 1 0-1 3) are periolate and basally
truncated or cordate; they are petiolate and rounded to broadly cuneate in
A. album (Figs. 6—9), A. floridanum (Figs. 14—16), -^nd A. piantagineum
(Figs. 17-20), whereas they are ± sessile and more narrowly cuneate in A.
sulcatum (Figs. 21-24). Cauline leaves of^. diversifolium are deltoid-hastate
in oudine and dentately toothed. They tend to be ovate, serrately lobed,
and acute in A. album; ovate-oblong, crenulate, and obtuse or rouncied in
A. floridanum\ ovate to lanceolate or narrowly spatulate, entire or serru-
late, and acute in^l. plantagineum\ and ovate to rhombic or narrowly lan-
ceolate, sparsely serrate-dentate, and acute to obtuse in A. sulcatum.
Arnoglossum plantagineum has the greatest geographical range among the
five species (east Texas, Louisiana, and the black soil belt of Alabama north
through Kansas and Ohio to southern Minnesota, Michigan, and Ontario,
Canada); it occurs mainly in calcareous, tall-grass prairie (Pippen & Chapman
1986), but minor habitats include glades, fens, and pine-oak woodlands.
Arnoglossum floridanum occurs in well-drained sands in open pine-scrub
oak, dry flatwoods, and old fielcis from Duval and Madison through pen-
insular Florida south to Highlands and Manatee counties.
The other three species occur mainly in the Florida panhandle. Arnoglossum
diversifolium occurs in river swamps and wet hammocks and is the most
wide-ranging of these three; it has a few populations in southern Alabama
and southwestern Ceorgia, ranges fi-om Walton County to Leon County
in the panhandle, and has disjunct populations in Putnam, Volusia, and
Levy counties, Florida. Arnoglossum sulcatum occupies shaded acid bogs or
swamps; it occurs sporadically in extreme southern Alabama and south-
western Georgia and from F^scambia to Leon Counties of the Florida pan-
handle. Arnoglossum album is restricted to Bay and Gulf Counties of the
Florida panhandle; the only other species of the group that occurs in those
counties is A. sulcatum, but it has different habitat preferences and
phenology.
Flowering time in Arnoglossum plantagineum is clinal from south to north;
the plants blooming in late April in southern Louisiana to early August in
Minnesota and Michigan (l^ippen & Chapman 1986). A reverse dine ex-
ists for A. diversifolium; it blooms in the Florida panhandle from May to
early July (same period fox A. floridanum) , whereas, to the south, in Levy
County (where it is sympatric with Hasteola robertiorum) A. diversifolium
blooms from mid-August through September. Arnoglossum album blooms
from June to mid-July, and A. sulcatum is the latest to bloom (September
to October).
The new species, A. album, is distinguished from all of its immediate
relatives by the following features: it is geographically isolated from all other
species except A. sulcatum, and it is totally separated from A. sulcatum by
Andkrson, a new species of Arnoglossum 383
habitat and phenology; basal leaf venation is distinctive; cauline leaves are
most similar to those of /}. sulcatum, but they are generally narrower and
sessile in A. sulcatum; its involucres and corollas appear whiter than those
of the other species; the wings of the phyllaries are higher proximally and
attenuated and distally, whereas wings are lower in height overall and evenly
raised along the keel or somewhat higher distally in the others; the phyl-
lary wings have erose margins distally in A. album, and wing margins are
entire or sinuate in the other species; and the corolla tubes are wider than
those in any other species in the group.
KEY TO THE FIVE SPECIES Ol- ARNOGLOSSUM WITH WINGED PHYEEARIES
Most keys to species of Arnoglossum include significant habitat and phe-
nological data. The following key uses only morphological features.
1. Blades ot radical leaves truncate, ovate to cordate-ovate, sparsely den-
ticulate; lower cauline leaves deltoid-hastate, dentately toothed A. diversifolium
1. Blades of radical leaves ovate to ovate-oblong, entire, sinuate or crenate;
lower cauline leaves ovate, entire, crenulate or serratcly toothed.
2. Phyllaries with prominently winged keels, wings highest at base, erose;
radical leaves with lateral veins appressed to midvcins lor 2—4 cm
then spreading A. album
2. Phyllaries with keels + evenly winged throughout or highest apically,
entire or sinuate; radical leaves with lateral veins diverging from base
of blade.
3. Involucres and corollas mostly over 1 1 mm and 10 mm long, re-
spectively; cauline leaves crenulate A. floridanum
3. Involucres and corollas mostly shorter; leaves entire or sparsely serrulate.
4. Involucres mostly over 1 mm long, corollas over 9 mm; cauline
leaves at midstem rounded to petiokue base A. plantagineum
4. involucres less than 10 mm long, corollas 8-9(-9.6) mm; cauline
leaves at midstem broadly cuneate and sessile A. sulcatum
acknoweedc;m ents
This study was supported in part by a grant from the Council on Re-
search and Creativity of Florida State University. Richard Pippen graciously
shared his knowledge of the group. Mark Garland assisted with the Latin
diagnosis, and Ken Womble helped with the illustrations. Ted M. Barkley,
R.R. Kowal and A.B. Thistle are thanked for providing critical reviews of
the manuscript.
references
AndI'RSON, L.C. 1994. A revision of Hasteola (Asteraceae) in the New World. Syst. Bot.
19:211-219.
Brummiti , R.K. 1992. Vascular plant families and genera. Kew: Royal Botanic Gardens.
Cronqulsi , A. 1978. Compositae, tribe Senecioneae. N. Amer. Flora, sen II, 10:14-16.
. 1980. Vascular plants of the southeastern United States. Vol. 1 . Asteraceae. Univ.
384 SinAl8(2)
of North Carolina Press, Chapel Hill.
Funk, V.A. 1985. Cladistics and generic concepts in the Compositae. Taxon 34:72-80.
GoDi-RKY, R.K., and J.W. WocriEN. I 98 1 . Aqtuuic and wetland plants of the southeastern
United States. Dicotyledons. Univ. of^ Georgia l^'ress, Athens.
Jl.i FRiv, C. 1979. Generic and sectional limits in Senecio (C'oniposltae): II. F.vahiation of
some recent studies. Kew Bull. 34:49-58.
. 1992. 1 he tribe Senecioneac (Compositae) in the Mascarene Islands with an an-
notated world check-list of the genera of the tribe. Kew Bull. 74:49-109.
Kral, R., and R.K. GoDFRl-Y. 1958. Synopsis ol the Florida species of ('iicnliii (Compositae).
Quart. J. Florida Acad. Sci. 21:193-206.
PiPPHN, R.W. 1978. Cnailia. N. Amer. Flora, series 11, 10:151-159.
and K.A. CjiaI'MAN. 1986. Comparison oi morphological characters between re-
gional habitats of Cacalia plantaginea (Asteraceae). In: G.K. Clanby and P.N. Pemblc,
eds. Proceedings of the 9rh North American Prairie C'onference. IVi-college University
C^enter for Environmental Studies, Faijjo, ND.
Robinson, F1. 1974. Studies in the Senecioneae (Asteraceae). VI. Fhe genus Arnoglossiim.
Phyrologia 28:294-295.
. 1980. Arnoglossiim rcniforme (Hook.) li. Robins., comb. nov. Phytologia 46:441.
and R.D. Brettcll. 1973. Studies in the Senecioneae (Asteraceae). IV. The genera
Mcsiidoi'uL Syiieik'sis, Miricncalicu Koyamacalia and Sirniciiliti. Phytologia 27:265-276.
Wlil'll.R, M.A. 1983. Micromorphological characters and generic delimitation of some
New World Senecioneae (Asteraceae). Brittonia 35:1-22.
SEVEN NEW NOMENCLATURAL
COMBINATIONS AND A NEW NAME IN
PACKERA (ASTERACEAE: SENECIONEAE)
DEBRA K. TROCK and THEODORE M. BARKLEYi
Herbarium, Division of Biology
Kansas State University
Manhattan, KS 66506, U.S.A.
dkwel@ksii. edu
ABSTR/Uri'
The following new combinations and new name arc proposed: Packera boLinderi var.
harfordii (Greenm.) D.K. Trock & T.M. Barkley, comb, nov.; Packera dimorphophylla var.
intermedia (T.M. Barkley) D.K. Trock & T.M. Barkley, comb, nov.; Packera dimorphophylla
var. paysoni (T.M. Barkley) D.K. Trock & T.M. Barkley, comb, nov.; Packera pseuda urea
Nss.flavula (Greene) D.K. Trock & T.M. Barkley, comb, nov.; V-A.cks.x-3,pseudmirea\'\\x. semicordata
(Mack. & Bush) D.K. Trock &T.M. Barkley, comb, nov.; Packera neomexicanavix. tonmeyi
(Greene) D.K. Trock & T.M. Barkley, comb, nov.; Packera neomexicana var. mercalfei (Greene)
D.K. Trock & T.M. Barkley, comb, nov.; and Packera buekii D.K. Trock & T.M. Barkley,
nom. nov.
RESUMEN
Sc proponen las siguientes nuevas combinaciones y nuevo nombrc: Packera bolanderi
var. harfordii (Greenm.) D.K. Trock &T.M. Barkley, comb, nov.; Packera dimorphophylla
var. intermedia (T.M. Barkley) D.K. Trock & T.M. Barkley, comb, nov.; Packera dimorphophylla
var. paysoni (T.M. Barkley) D.K. Trock & T.M. Barkley, comb, nov.; Packera pseudaurea
\''ix. fldvula (C irccne) D.K. Irock & T.M. Barkley, comb, nov.; Packera /^ifwd^z/rcd var. semicordata
(Mack. & Bush) D.K. Trock & T.M. Barkley, comb, nov.; Packera neomexicana var. toumeyi
(Greene) D.K. Trock & T.M. Barkley, comb, nov.; Packera neomexicana var. mercalfei (Greene)
D.K. Trock & T.M. Barkley, comb, nov.; y Packera buekii D.K. Trock & T.M. Barkley,
nom. nov.
The genus Packera (Asteraceae: Senecioneae) has been known widely as
the Aureoid group oi Senecio, and the recognition o^ Packera necessitates
many nomenclatural transfers. Preparation of the treatment o{ Packera for
the Flora of North America project has drawn our attention to the trans-
fers proposed here. Type specimens for the basionyms are provided by Barkley
( 1 978) . The combinations P. pseudaurea var. semicordata and P. dimorphophylla
var. intermedia are proposed simply for consistency in the use of "varietas"
'Present address: Botanical Research Institute ofTexas, 509 Pecan Street, Fort Worth, TX
76102-4060, U.S.A, e-mail: barkley@brit.org
SiDA 18(2): 385-387. 1998
386 SiDA 18(2)
as the infraspecific rank of choice in Packera The two epithets are other-
wise available as subspecies.
Packera bolanderi var. harfordii (Greenm.) D.K. Trock &T.M. Barkley,
Comb.nov. BASIONYM: Senecio Ijarfordii Gveenm. in PipeT, CA)nrr. U.S. N;itl. Herb.
1 1 :567. 1906. St'iu'do hoLiiideri var. hiirforflii {(;recnm.) I'.M. Barkley
Packera dimorphophylla var. intermedia (T.M. Barkley) D.K. Irock & T.M.
Barkley, comb. nov. By\.SIC)NYi\l: Senecw dimoypliophyllmxAX. intcTiiiedius'Y.M..
Barklev. Trans. Kansas Acad. Sci. 65:362. 1 963. Packera dimorphophylla subsp. intermedia
(T.M.'Barkky) Weber & Love.
Packera dimorphophylla var. paysoni ( l.M. Barkley) D.K. Trock & l.M.
Barkley, comb. nov. BA.SIONYM: Seuecio dimorphophyllm\':\\. payu^ii^WW . Barkley,
Trans. Kansas Acad. Sci. 65:362. 1963.
Packera pseudaurea var. flavula (Greene) D.K. Trock &T.M. Barkley, comb,
nov. BaSIONYM: Senecio flamdus i^\\xi:nc, Pittonia 4: 108. 1900. Setiecio pseudaureus
var. flavula (C^reene) Greenm. Packera pseudaurea suhsp. [lavula (Cirecne) Weber &
L.ove.
Packera pseudaurea var. semicordata (Mack &C Bush) D.K. Trock & T.M.
Barkley, comb. nov. BaSIONYM: Senecio semieordatus Mack. & Bush, Rep.
Missouri Bot. Card. 16:107. 1905. Senecio aureus var. semieordatus (Mack. & Bush)
(u'eenm. Soiecio pseudaureus viw se)nicordatus (Mack. & Bush) F.M. Barkle\'.
Packera neomexicana var. toumeyi (Greene) D.K. Trock & f.M. Barkley,
comb. nov. ^i\S\OW\\\: Senecio toumeyi Gvciinc, Pittonia 3:349. \89S. Senecio neo-
mexicanus \'AV. /o //;?«')'/ (Greene) l.M. Barkley.
Packera neomexicana var. metcaltei (Cireene) D.K. Trock & F.M. Barkley,
comb. nov. BASIONYM: Senecio ?netcalfei CiKcne ex Wooron & Standi., (lonlr. U.S.
Natl. I lerb. 16:193. 1913. Senecio ueowe.\ica>/us v.w. metcalfei [Greene) I'.M. Barkley.
The situation with the epithet '\ymbaiarioiJef is complicated. A single-
headed plant of wet, open sites in western U.S. and adjacent Canada was
long known as Senecio subnudus DC., a name that was introduced by A. P.
de Candolle in 1 837. Unfortunately, de Candolle used the epithet "subnudus"
twice; the other time For an African plant. Heinrich W. Buck (1840) rec-
ognized the error in his index to the Prodromus and he chose the African
plant to be the correct Senecio subnudus. He then called the American plant
Senecio cymbalnrioidesW. Buck, and unfortunately, this name remained forgotten
until the 1960s (Barkley 1962). In the meantime, a widespread plant of
western North America was named Senecio cynibalarioides byThomas Ntittall.
It was necessary to change the name of the North American S. subnudus
DC. to .V. cynibalarioides H. Buck, and S. cymbalarioides Nutt. then be-
came S. streptanthifolius Greene, which is the next oldest available name
for that entity (Barkley 1962). Recognition of the gentis Packera retjuired
Trock and Barkley, New combinations and a new name in Pacltera 387
the transfer of many epithets into that genus and, unfortunately, Senecio
cymbalarioidesNutt. was transferred into Packera where it became /? cymbalarioides
(Nutt.) W.A. Weber & A. Love, (Weber & Love 1981). Shortly thereafter,
Weber and Love recognized that it was S. cymbalarioides H. Buek that should
have been transferred to Packera, so they made the combination: P cyimbalaroides
(H. Buek) WA. Weber & A. Love (Weber 1 984). The transfer of the Nuttall
epithet into Packera preoccupies that name in Packera so Packera cymbalioides
(H. Buek) W.A. Weber & A. Love is illegitimate and must be rejected ac-
cording the International Code of Botanical Nomenclature (Greuter et al.
1994). Thus, the American plant first known as Senecio subnudus DC. and
then as Senecio cymblalrioides Buek must have a new name. The new epi-
thet that we chose for this distinctive plant commemorates Heinrich Wilhelm
Buek, the 19th Century German botanist whose carefully prepared index
to de Candolle's Prodromus gave rise to the need for this new name.
Packera buekii D.K. Trock &T.M. Barkley, nom. nov. Basionvm: Sawcio suhimdus
DC]., Prodr. 6:37. 1 837 (non S. subnudus DC, Proclr. 6:405. 1 837). Senecio cytnbaLnioides
Buck, Gen. Sp. Synon. Cand. 2:6. 1840 (non .V. cynd)alarioides Nutt., Trans. Amer.
Phil. Soc. 11. 7:412.1841.).
ac:knowledgmf.nts
We thank John Bain, John Pipoly, Alan Whittemore, John Strother, K.
Gandhi, and D.H. Nicolson for their advice and editorial wisdom in the
preparation of this work. This is contribution no: 99'144-J [xom the Kan-
sas Agricultural Experiment Station, Manhattan.
RKFERHNCES
Barkley, T.M. 1962. A revision oi Senecio aureus Linn, and allied species. Trans. Kansas
Acad. Sci. 65:318-408.
. 1978. Senecio. In: North American flora 11. 10:50-137.
Buek, H.W. 1840. Genera, species, et synonyma Candolleana. 2.
Candoeie, A.R de, 1837. Prodromus systematis.. . 6:405.
Gri^uter, W., F.R. Barrie, H.M. Bure^et, W.G. Chalonir, V. Demoulin, D.L. Hawksworth,
RM. J()erc;en.sen, D.H. Nicoe.son, PC. Sieva, R Tehani;, & J.M. McNeii e, eds. Inter-
national code of botanical nomenclature (Tokyo Code). Regnum. Vcg. 131:1-389.
Weber, W.A. 1984. New names and combinations, principally in the Rocky IVIountain
flora-IV. Phytologia 55:1-1 1.
Weber, W.A. and A. Lc)VE. 1981. New combinations in the genus Packera (Asteraccae).
Phytologia 49:44-50.
388 SinAl8(2)
BOOK REVIEWS
A. Esi'Ejc) and A.R. Lc'^pez-Feriuri. 1 997. Las Monocotiledoneas Mexicanas.
Una Sinopsis Floristica 1 . Lista de Referenda PARTE VIII. Orchidaceae
1. (ISBN 968-6144-10-2; 968-6 144-20-X pbk). Consejo Nacional de
la Flora dc Mexico, Apartado Postal 70-261, Delegacion Coyoacan,
04510 Mexico, D.E iind Universidad AutonomaMetropolitana-Iztapalapa,
Av Michoacan y La Purisima, Col. Vicentina, Delegacion Iztapalpa,
09340 Mexico, D.E No price given. 90 pp. Color cover.
A. EsPF.jO and A.R. Loi'hy.-Fp.RRARi. 1997. 1998. Las Monocotiledoneas
Mexicanas. Una Sinopsis Floristica 1. Lista de Referencia PARTE
VIII. Orchidaceae 2. (ISBN 968-6144-10-2; 968-6144-21-8 pbk Consejo
Nacional de la Flora de Mexico, Apartado Postal 70-261 , Delegacion
Coyoacan, 045 1 Mexico, D.E and Universidad Autonoma Metropolitana-
Iztapalapa, Av Michoacan y La Pinisima, Col. Vicentina, Delegacion
Iztapalpa, 09340 Mexico, D.E No price given. 1 15 pp. Color cover.
'Hicsc two vdliiiiics list the species ol Orchidaceae known from Mexico, in alphabetical order by ge-
ntis, then by species, from Aciucta to Lnclia (Orchidaceae 1) and hom l.cnihoglosstDn to Xylohiunt
(Orchidaceae 2). There arc also errata and additions to the first six parts (Agavaceae throiit^h Nolinaceac).
l-'or each species, there is a complete literature citation, followed b\' the basionvm, along with all taxonomic
and nomenclarin-al synonyms and respective t\'pe citations. It is clear that the authors have seen the
vast majority of the type specimens involved. There are no lectotypcs chosen in the work, but lecto-
types chosen b\' othei^ ainhors are cited clearK'. I\'pe specimens seen have the traditional"!" marking;
otherwise, location tjf'the holotype is cited as it appears in the |iiotologtie. Each species' distribution
within Mexico is also given.
Tiie list is a very handy reference for anyone interested in the Orchid flora of Mexico, t)r for that
matter, Mesoamerica. It is printed on high quality paper and is clear anti easy to read. The author
abbreviations correspond to Authors of Plant Names, published by the Royal Botanic Gardens, Kew,
and the literature citations correspond to the B'T-EI and TL-2. 1 have seen no notable t\'pographital
errors.
Annotated checklists such as these are extremeh' useful, especiall)- gi\-en the complete index to ac-
cepted names and synomyms in the back. This series is absolutely necessary for any curator dealing
with Mexican monocotyledonous plants. We anxiously await the arrival of the other volumes, and
unhesitatingly recommend them for swm'oncl-Johri J. Pipoly ill
SioA 18(2): 388. 1998
ARDISIA NIAMBIENSIS (MYRSINACEAE),
A NEW SPECIES OF ARDISIA SUBGENUS
ARDISIA FROM THE CHOCO FLORISTIC
PROVINCE OF COLOMBIA
JOHNJ. PIPOLYIII
Botanical Research Institute of Texas
509 Pecan Street, Fort Worth, TX 76102-4060 U.S.A.
jpipoly@brit. org; clusia@latinmail. com
ALVARO COGOLLO P.
Fundacion Jardin Botdnico, Joaquin Antonio Uribe
Apartado Aereo 51407, Medellin, Antioquia, COLOMBIA
acogollo @latinmaiL com; jardinbo @epm. net. co
ABSTRACT
Documentation oFthe flora of the Natural Reserve Ri'o Nanibf has resuhed in the dis-
covery of a heretofore undescribed species, Ardisia niambiensis. The species is described,
illustrated and its salient morphological features are elucidated. Ardisia niambiensis is best
placed in subgenus Ardisia because of its terminal, glandular-papillate, paniculate inflo-
rescences with tripinnate branches, and the evenly divided, 5-lobed calyx, whose symmet-
ric lobes have glandular-ciliolate margins. It is most closely related io Ardisia premontana,
a taxon endemic to the eastern slopes of the Eastern Andean Cordillera of southern Ecua-
dor and Peru.
RESUMF.N
Al documentar la flora de la Reserva Natural Rio Nambi, se encontro una nueva especie,
Ardisia niambiensis. Se describe, se ilustra yse discuten sus caracteres principales y sobresalientes.
Ardisia niambiensis sc ubica mejordentro del subgenero^ri?/zi-Mdebido asus pani'culas terminales
y glandular-papillosas con ramificacion tripinnada, su caliz simetricamente dividido con
lobulos simetricos y glandular-ciliolados a lo largo de los margenes. La nueva especie es
muy afi'n a Ardisia premontana, una especie endemica a la Hilda oriental de la Cordillera
Oriental del Ecuador y Perti.
INTRODUCTION
The pantropical ^enus Ardisia Swartz contains approximately 400-500
species (Chen Cheih & Pipoly 1996) with centers of high species diversity
in Malesia and the Choco Floristic Province from Panama through Co-
lombia to Ecuador (Pipoly 1991, 1992, 1994, 1995, 1996). Within the
Neotropics, approximately 300 species have been described, the majority
SiDA 18(2): 389-393. 1998
390 Sum 18(2)
from Mesoamerica. There are relatively few South American taxa, perhaps
not passing 40 total. During the course of an analysis of the vegetation of
the Ri'o Nambi Natural Reserve, located in western portion of Nariiio cor-
responding to the Choco Floristic Province, a new species of Ardisia was
discovered, which is described herewith.
Ardisia niambiensis Pipoly & CogoUo, sp. nov. (Fig. 1). Typi : COLOMBIA.
NarINO: Mpio. Barbacoas; Corrcgimicntos Ortiz y Zamora; Vcicda El Barro; Rescrva
NacLiral Ri'o Nambi, ca. 5 dm al W dc Altaquer, Faldas Occidcntalcs de la Cordillera
Occidenral, 01° 15'N, 78° 08'W, 1,250-1,350 m, 9 Sep 1997 (stam. Fl),/ Ptpoly,
A. Corolla, M. Lopez&M. Rodriguez 2 1 572 {HOl.OJWY^ l\SO; ISOlYi'F.S: BRIT, COL,
FMb', GH, HUA, JAUM, K, mo, NY, 'FULV).
Ob infloresccnriam tcrminalcni iripinnacipaiiicularam, calyceni aequalitcr divisum, necnon
lobos calycinos simetricos glandulari-ciliolaribusquc subgencri Ardisiac pertinet. Propter
laminas ad apicc breviacuminatas, rhachides inflorescentiares angulatos, margines scpalinos
hyalinos carinatos subintegros erososqtic/^. p)~ernonM)!oy^\dt: arete afhnis, sed ab ea raniulis
glabris (non fufuracco-lepidotis), laminis coriaeeis (non charraceis), subtcr glabris dense
punctatis punctato-lincatisque (nee furfuraeeo-lepidotis), petiolis 7-10 (non 15-25) mm
longis, floribus pedicellatis (non sessilis), denique lobis calycinis deltatis vel late triangularibus
(non ovatis) 2-2.2 (nee 1-1.2) mm longis statim distinguitiir.
Tree to H) m X 15 cm DBH. Bmnchlets terete, 4-8 mm glabrous. Leaves
alternate; blades coriaceous, oblanceolate to obovate, { 1 4. 1-) 19-25.4 cm
long, (6.8-)7.5-9.4 cm wide, apically abruptly acuminate, the acumen 5-
8 mm long, basally acute, decurrent to petiole base, midrib canaliculate
and prominently raised above, prominently raised and densely black punctate-
lineate below, the secondary veins numerous, .39-47 pairs, prominulous
above and below, drying dark above, pallid and densely perpuncticulose
below, the punctations in the form of short lines, the margin slightly inrolled
except revolute basally; petioles marginate to base, 7-10 mm long, glabrous.
Inflorescence iQUTiinA, pyramidal, tripinnately paniculate, 12.8—22.5 cm long,
1 1 .5-33 cm wide, peduncle 1-2.5 cm long, the rachis prominently angulate,
the secondary branches with smaller panicular units 1 2-16 cm long, 6-10
cm wide, the ultimate branches racemose, moderately rufous glandular-
granulose and prominently and densely black punctate-lineate; floral bracts
caducous, unknown; pedicels cylindrical, (1.0— )1 .5-2.5(-3.0) mm long.
Flowers unknown, but fruiting calyx 3-3.5 mm long, rufous glandular papillate
and darkened at junction of lobe and tube, the lobes deltate to widely tri-
angular, 2—2.2 mm long, 2.2—2.5 mm wide, apex acute to a minutely acuminate
apex, minutely rufous-lepidote and carinate medially without, prominently
punctate and punctate-lineate, the margin hyaline, somewhat erose and sparsely
glandular-ciliolate apically. frw/V globose, 5-8(-l 0) mm long and wide when
dried; when fresh, exocarp fleshy, bitter tasting.
Distribution. — Known only from Reserva Natural Rio Nambi, Nariho,
Colombia, at 1,250-1,350 m elevation.
PiPOLY AND COGOi.LO P., A new species of Ardisia
391
Fig. 1 . Ardisia niambiensis Pipoly, showing the pyramidal tripinnate inflorescence, densely
and prominently black punctate leaf surfaces, short petioles marginate to base, and pedi-
cellate flowers. Drawn from the holotype.
392 SiDAl8(2)
Ecology. — Ardisia niambiemis occurs in premontane pluvial forest, receiving
approximately 7,800 to 8,200 mm rain per year. The species occurs in relatively
low densities, in at the forest margin on ridgetops.
Etymology. — Ardisia niambiensis is named phoenetically for the Reserva
Natural Rio Nambi, a private, nonprofit, integrated conservation and de-
velopment area in the state of Narino, Colombia, operated by FELCA, the
Ecological Foundation of the Hummingbirds of Altaquer (Fundacion Ecologica
de los Colibris de Altaquer).
Parai'vpi:. COLOMBIA. Narino: Mpio. Rarbacoas; C "orregimienros Ortiz Zamora; Vereda
El Rarro, Rcscrva Natural Ri'o Nainbf, ca. S km al W de Altaquer, Faldas Occidentales de
la Cordillera Occidental, OT 15' N, 78° 08' W; 1,350-1,450 m, 3 Sep 1997 (ster.), /.
Pipoly, A Cogolio. M. Lopez & M. Rodr!f(uez 21227 {BRIT, FMR, JAUM, PSO).
On account of the shortly acuminate leaf apices, angulate inflorescence
rachises, carinate sepals with hyaline and subentire to erose margins, Ardisia
niambiensis is most closely related to Ardisia premontana Pipoly. It is sepa-
rated from /I, premontana by its glabrous (not ftirftiraceous-lepidote) branchlets,
the coriaceous (not chartaceous) leaf blades that are densely black punctate
and punctate lineate (and not furfuraceous-lepidote) below, petioles 7-10
(not 1 5-25) mm long, pedicellate (not sessile) flowers and deltate or widely
triangular (not ovate) sepals that are 2.0-2.2 (and not 1.0-1.2) mm long.
Ardisia subgenus Ardisia, to which A. niambiensis belongs, is defined by
the well-developed but early caducous floral and inflorescence bracts, race-
mose or rarely spicate branchlets of the inflorescence, the anthers dehisc-
ing by subapical pores opening into wide longitudinal slits, and symmetric
sepals. Within that subgenus, the other species known from the Choco include
A. colombiana Lundell, A. granatensis Mez, A. opaca Lundell, A. valida Mez
(= A. perinsignis Lundell syn. nov.), A. monsalveae Pipoly, and A. cabrerae
Pipoly (Forero & Gentry 1990; Pipoly 1991, 1995) Among these species,
the only ones with inflorescence shape, pedicellate flowers and sepals simi-
lar to those o^ Ardisia niambiensis are A. cabrerae and A. monsalveae. How-
ever, the three species may be easily separated by the following key:
1. Calyx 2.4-2.9 mm long, apically ohurse, the margin.s glabrous, entire to
subentire.
2. Leaf blades coriaceous, elliptic; sepals 3.4-3.3 mm wide, glabrous A. cabrerae
2. Leal blades chartaceous, widely oblanceolate to obovate; sepals 0.5—0.6
mm wide, densely and minutely rufo-puberulcnt A. monsalveae
L C'alyx 3—3.5 mm long, apically acute to a mintitcly acuminate apex, min-
utely rufous-lepidote and carinate, the margin spaisely ciliolate A. niambiensis
ACKNtWIJ'.D(;MK.Nr,S
Support for our studies in plant diversity at the Rio Nambi Natural Reserve
is provided by a generous grant from the National Geographic Society, No.
PiPOLY AND Coc.OLLO P., A new species of Ardisia 393
5575-95. We thank the collaboration of the herbarium of the Universidad
del Narino (PSO), especially the Director, Biol. Marta Sofia Gonzalez Insuasty,
and the herbarium technician, Bernardo Ramirez. We gratefully acknowl-
edge the logistical collaboration of FELCA (Fundacion Ecologica de los
Colibris de Altaquer), especially that of Don Marcial Bisbicuz and Mauricio
Flores. We also thank Juan Guillermo Ramirez Arango and Adriana Gomez
for technical and logistical help. We are most grateful to Dr. Ramiro Fonnegra,
director of HUA, for making the services of Gloria Mora available to us,
who skillfully prepared the illustration.
REFERENCES
Chkn, CiiiEH andj. Pii'OLY. 1996. Myrsinaceae. Pp. 1-38. In: Wu Zheng Yi and PH. Raven,
eds. Flora of China. Vol 15. Science Press, Beijing and Missouri Botanical Garden, St.
Louis, Missouri.
FoRiiRO G., E. and A.H. Gi:ntry. 1989. Lista Anotada de las Plantas del Departamento
del Choco, Colombia. Biblioteca Jose Jeronimo Triana, No. 10. Instituto de Ciencias
Naturales, Museo de Historia Natural, Bogota, D.E.
PiPOLY, J. 1991. Noras sobre el generoylrf/wz^Swartz (Myrsinaceae) en Colombia. Caldasia
l6(78):277-284.
. 1992. Ardisia callejasii (Myrsinaceae): a new species from the Antioquian Choco
of Colombia. Novon 2:389-391.
. 1 994. New species oi Ardisia (Myrsinaceae) from the Corciillera Occidental of Colombia
and Ecuador. Noyon A-3%-AA.
_. 1 995. Dos nuevas especies del ^tnero Ardisia (Myrsinaceae) de la provincia flon'stica
chocoana de Colombia. Caldasia 17(82-85):4l9-424.
1996. New species oi Ardisia (Myrsinaceae) from Ecuador and Peru. Sida 17:445-
458.
394 Sum 18(2)
BOOK REVIEW
TnoM/\s LuMFKiN and Dhan Mc:Claky. 1994. Azuki Bean: Botany, Pro-
duction and Uses. (ISBN 85918 765 6, hbk). CAB International,
Oxford University Press, 198 Madison Ave., New York, NY 10016.
800-451-7556. $70.00. 268 Pp.
This book purports to be a monograph, inckiding both the basic botanical and apphcd
agronomic aspects among the "Azuki" members of Vigna subgenus Cemtotropii. The book
is divided into nine chapters, inckiding an introduction, botany, physiok-)gical character-
istics, production, insects and nematodes, diseases, breeding characteristics, food chem-
istry and processing, and uses and marketing. The agronomic chapters contain a thor-
ough review of the literature, but much of it refers only to the A/uki beans in general, and
not to a specific taxon. As a plant systematist, I am most disappointed with Chapter 2,
the Botany of Azuki. I find it superficial, and not useful for understanding the differences
among the species, or the infraspecific raxa (subspecies, varieties and cultivars). A synop-
sis or taxonomic revision for this relatively small group could have been contributed by
any t:ine of a number of qualified taxonomists, but the authors chose to avoid a formal
treatment, f hey state that in their Table 2. 1 , "No attempt has been made to correct the
confusion ol different classifications in the literature." That statement amounts to a fatal
flaw for the work, because many of the subsequent chapters make frequent reference to
various cultivar names, witkout link to a taxon described in the Botany chapter, for ex-
ample, in the discussion of starch granules and processing (Chapter 8), reference is made
to "azuki cv lakara,'' yet there is no mention of that or any other cultivar in the taxonomy
chapter. Because the reader is most often unsure of the exact entity I'eferred to, the rest of
the book loses much of its usefulness, except in broad terms. I find the book hardly more
than a literature review, and not a definitive monograph. Its extensive bibliography may
be tiseftil for agronomic investigators, but hardly justifies its hefty price. I would ha\'e
preferred to have seen a comprehensive, well-executed systematic, agronomic and agri-
cultural treatment, like the cla,ssic work off. van der Maesen for Cicer arietinu7n, pub-
lished in Medcd. Landbouwhogeschool 72-10:1-342. 1972. 1 highly recommend that
Maesen's monograph be used as a model (with phylogenetic systcmatics added) for eco-
nomically critical and highly domesticated plant taxa. By following Maesen's model, we
may better take advantage of wild relatives of ctirrently cultivated plants, to bring useful
characteristics into the cultivated from the wild. Fhe book should be part of any major
agi'icukural library, as a literature review souvcc— Job// Pipoly, III
SiDA 18(2): 394. 1998
CLUSIA NIAMBIENSIS (CLUSIACEAE),
A NEW SPECIES FROM THE CHOCO FLORISTIC
PROVINCE OF COLOMBIA AND ECUADOR
JOHNJ. PIPOLYIII
Botanical Research Institute of Texas
509 Pecan Street
FortWorth, TX 76102-4060, U.S.A.
jpipoly@brit.org; clusia@latinmail. com
ALVARO COGOLLO P.
Fundacion Jardin Botdnico, Joaquin Antonio Uribe
Apartado Aereo 51407, Medellin, Antioquia, COLOMBIA
acogollo @latinmail. com; jardinbo @epm. net. co
MARTA SOFIA GONZALEZ
Departamento de Biologia, Universidad de Nariho
Apartado Aereo 1176, Pasto, Narino, COLOMBIA
ABSTJIACT
Docuniencation of the flora of the Natural Reserve Rio Nambf has resulted in the dis-
covery of a heretofore undescribed species, China niambiensis. The species is described,
ilhistrated and its sahent morphological features are elucidated. While its vegetative mor-
phology is reminiscent of both Clusia laurifolia and C. venusta, the free, numerous, and
subulate stamens with a central mass of staniinodia producing a waxy resin indicate it is a
member of section Chlamydoclusia rather than section Retinostemon, where the aforemen-
tioned congeners belong. At present, the relationships of Clusia niambiensis within sec-
tion Chlamydoclusia, are unknown. It is anticipated that continuing studies of the genus,
leading to a treatment of the family for Flora de Colombia, will resolve this problem.
RHSUMRN
Al documentar la flora dc la Reserva Natural Rio Nambf, sc encontro una nucva especie,
Clusia niambiensis. Se describe, se ilustray se discuten sus caracteres principales y sobresalientes.
Mientras que sus caracteres vegetativos son muy semejantes a los de Clusia laurifolia y C.
venusta, sus estambres libres, numerosos y subulados indican claramentc que C niambiensis
pertenece a la seccion Chlamydoclusia y no a la seccion Retinostemon, donde se ubican las
otras especics congenericas susodichas. En este momento, no se sabe exactamente la ubicacion
taxonomica de Clusia niambiensis dentro de la seccion Chlamydoclusia. Esperamos que a
travcs de los estudios en proceso para el tratamiento taxonomico de la familia para Mora
de Colombia, se pueda resolver este problema.
Recent exploration of the Reserva Natural Rio Nambi, located on the
SiDA 18(2): 395-399. 1998
396 SiDAl8(2)
western slopes of the Western Cordillera of the Colombian Andes, has re-
sulted in the discovery of this spectacular new species, described herewith.
Clusia (§ Chlamydoclusia) niambiensis, Pipoly, Cogollo et Gonzalez, sp.
nov. (Fig. 1). Typi;: COLOMBIA. NarinO: Mpio. Barbacoas; C^orrcgimientos Ortiz
y Zamora; Ve rcda HI Barro; Reserva Natural Rio Nambi, ca. 5 km al W dc Aliaqucr,
Faldas Occidcncales de la Cordillera Occidental, 01° 1 5' N, 78° 08' W; 1 ,450-1 ,500
m, 8 Sep 1997 (stam. Fl), /, Pipok A. Cogollo, M. Lopez 6- M. Rodriguez 2 1489 {UO-
LOTYI'E: l\SO; ISOTYPl'S: BRIT, COL, FMB, jAUM, K. MO).
Qtioad lamina biiHata nervatiosque brochidodromos, resinam albam, C laurifoliarn 3ixae
•similans sed ab ea petiolis profundc canalictilati.sqtic marginafis (non aliqiiantam canalictdatis),
ad bases laminas aciitas decurrentcsqtie (non obrii.sis baud deciirrentes), deniqiie fructibiis
ovoideis vcl ellipsoideis (non subglobosis) pracclarc distat. Propter stamina inter se libera
stibulataque centraliter resinifera, stigmata connivcntaque scctio Clilamydoclusia pcrtinet,
interspecies aliis petiolis profunde canalictilatis marginatisque, lamini.s bullads nervatiosque
brocbidodromis, Iructibus iongirudinaliter costatis starim distinguitur.
Glabrous, dioecious, shrubs or treclets to 2-3(-5) m tall, growth dynam-
ics corresponding to Scarrone's Model; latex white. Branchlets terete, ap-
pearing tetragonal when dried, 8-10(— 15) mm diam., the bark reddish-
brown, exfoliating transversely in the upper nodes, appearing furfuraceous
at times. Leaves decussate; blades chartaceous, obovate, (19— )27-38.5 cm
long, (1 4.5-) 15.2—23 cm wide, apically rounded to obtuse, basally acute,
decurrent to the petiole base, bullate, the midrib prominently raised or raised
within a depression above, prominently raised below, the secondary veins
brochidodromous, 28-46 pairs, ca. 7—10 mm apart, alternating so every
other vein has twice the diameter of the others, deeply impressed above,
prominently raised below, the submarginal connecting vein ca. 2-4 mm
from margin, glabrous above and below, at times when dried with small
linear latex canals oxidizing red, the margin scarious, entire; petioles deeply
canaliculate and marginate, (1.0-)2.5— 3.5 cm long, without adaxial mar-
gined pit. Staminate inflorescence erect in bud, pendulous at maturity, a compound
cyme, 20—25 cm long, 1 0—20 cm wide; pedtmclc appearing tetragonal when
dried, 10.5—17 cm long, the bark transversely checked and exfoliating; primary
inflorescence bract coriaceous, lanceolate, 1 8—21 mm long, 4-6 mm wide,
apically acute, the midrib prominently raised above anci below, the margin
scarious, entire, glabrous; secondary inflorescence bracts cartilaginous, sub-
orbicular, 3.5-4.0 mm long, 3.8-4.2 mm wide, apically broadly rounded,
medially carinate, the margin scarious, entire; tertiary and floral bracts as
in secondary but smaller acroperally, to the smallest that are rhomboid, 3
mm long and wide, apically acute; bracteoles 2, cartilaginous, ovate, .3.8-
4.0 mm long, 2.5-3.0 mm wide, apically rounded, cucuUate, the margin
hyaline, not scarious, entire; pedicels 1-2. 5 (—5) mm long. Staminate flow-
ers sepals 8, the lower 4 opposite, the upper 4 contorted, coriaceous, aero-
PiPOLY AND CoGOi.i.o P., A new species of Clusia
397
I cm
Fk;. 1 . Clusia niambiensis Pipoly, C^ogollo & Gonzalez. A. Habit. B. Opened pistillate flower,
showing staminodes and stigmatic area. C. Opened staminate flower, showing stamens and
central staminodial mass. D. Fruit, showing costae and mature stigmas. Figs. A— D, drawn
from holotype.
398 SinAl8(2)
petally larger, the lowermost oblate, 4.4-5.5 mm long, 5.5-6.5 mm wide,
apically broadly rounded, translucent toward the margins, the linear latex
canals black, conspicuous, the margins flat, scarious, irregularly notched,
at times also appearing erose; uppermost sepals as in lowermost but ob-
long, 10-12 mm long, 7.5-8 mm wide, petals 12-16, contorted, mem-
bran 
TVNews
OpenLibrary