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Bauhinia proboscidea (Fabaceae: Cercidoideae), a new species from Costa Rica and Panama, with notes on B. beguinotii, B. gorgonae and B. pansamalana


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Bauhinia proboscidea, a new species from Costa Rica and Panama, is described and illustrated, and compared to the closely related B. pansamalana of southern Mexico, Guatemala and Honduras. It is also compared with B. beguinotii, with which has been confused in herbaria. Bauhinia gorgonae, endemic to Gorgona Island in Colombia and which has been classified as a variety of B. beguinotii, is here considered a separate species. A key to the Neotropical species of Bauhinia with three fertile stamens is provided. Some observations on the morphology and reproductive biology of B. proboscidea are presented. Global-level assessments of the conservation status according to IUCN Red List criteria indicate that B. proboscidea and B. beguinotii (which was previously assessed using some misidentified records) should both be considered as species of Least Concern (LC), B. gorgonae should be considered as Endangered (EN), and B. pansamalana should be considered Not Threatened (NT). Country-level conservation assessments are also provided for all four species. The homology of the “intrastipular spines” of Bauhinia is briefly discussed.
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Phytotaxa 361 (1): 025–040
Copyright © 2018 Magnolia Press Article PHYTOTAXA
ISSN 1179-3155 (print edition)
ISSN 1179-3163 (online edition)
Accepted by Leonardo Borges: 12 Jun. 2018; published: 18 Jul. 2018
Bauhinia proboscidea (Fabaceae: Cercidoideae), a new species from Costa Rica
and Panama, with notes on B. beguinotii, B. gorgonae and B. pansamalana
1Investigador Asociado, Herbario Nacional de Costa Rica. Departamento de Historia Natural, Museo Nacional de Costa Rica, Apdo.
749-1000, San Joseì, Costa Rica; e-mail:
2Programa de Maestría en Biología Vegetal, Universidad Autónoma de Chiriquí, República de Panamá;
3Jardín Botánico Lankester, Universidad de Costa Rica, Apdo. 302-7050, Cartago, Costa Rica; e-mail:
4Escuela de Biología, Universidad de Costa Rica, Ciudad Universitaria Rodrigo Facio, Apdo. 11501-2060, San José, Costa Rica. OR-
Bauhinia proboscidea, a new species from Costa Rica and Panama, is described and illustrated, and compared to the closely
related B. pansamalana of southern Mexico, Guatemala and Honduras. It is also compared with B. beguinotii, with which
has been confused in herbaria. Bauhinia gorgonae, endemic to Gorgona Island in Colombia and which has been classified
as a variety of B. beguinotii, is here considered a separate species. A key to the Neotropical species of Bauhinia with three
fertile stamens is provided. Some observations on the morphology and reproductive biology of B. proboscidea are presented.
Global-level assessments of the conservation status according to IUCN Red List criteria indicate that B. proboscidea and
B. beguinotii (which was previously assessed using some misidentified records) should both be considered as species of
Least Concern (LC), B. gorgonae should be considered as Endangered (EN), and B. pansamalana should be considered
Not Threatened (NT). Country-level conservation assessments are also provided for all four species. The homology of the
“intrastipular spines” of Bauhinia is briefly discussed.
Se describe e ilustra Bauhinia proboscidea, una nueva especie de Costa Rica y Panamá, y se compara con la cercanamente
emparentada B. pansamalana del sur de México, Guatemala y Honduras. También se compara con B. beguinotii, con la cual
se ha confundido en los herbarios. Bauhinia gorgonae, endémica de la Isla Gorgona en Colombia, se considera aquí como
una especie independiente de B. beguinotii, de la cual se ha clasificado como una variedad. Se brinda una clave para las
especies neotropicales de Bauhinia con tres estambres fértiles. Se presentan algunas observaciones sobre la morfología y
biología reproductiva de B. proboscidea. Evaluaciones a nivel global del estado de conservación de acuerdo con los criterios
de la Lista Roja de UICN, indica que tanto B. proboscidea como B. beguinotii (que había sido previamente evaluada usando
algunos registros mal identificados) deben ser consideradas como especies de Preocupación Menor (LC), B. gorgonae debe
considerarse como En Peligro (EN), y B. pansamalana debe considerarse como No Amenazada (NT). También se presentan
evaluaciones de conservación a nivel de país para las cuatro especies. Se discute brevemente la homología de las “espinas
intraestipulares” de Bauhinia.
Key words: Central America, conservation, intrastipular spines, IUCN Red List categories, taxonomy, reproductive
The delimitation of the genus Bauhinia Linnaeus (1753: 374), the largest in subfamily Cercidoideae Legume Phylogeny
Working Group (2017: 68; formerly tribe Cercideae Bronn 1822: 131), has been controversial (see summaries by
Wunderlin et al. 1981, Lewis & Forest 2005, Sinou et al. 2009). In its widest circumscription (e.g., Wunderlin et al.
1981, 1987, i.e., Bauhinia sensu lato, or s.l.), it contains ca. 350–400 species of pantropical distribution (but absent
26 Phytotaxa 361 (1) © 2018 Magnolia Press
from the south Pacific islands). Numerous generic segregates of Bauhinia s.l. have been proposed. Of these, Barklya
Mueller (1859: 158), Gigasiphon Drake del Castillo (1902: 88), Lasiobema (Korthals 1841: 84) Miquel (1855: 71),
Lysiphyllum (Benth. in Bentham & Hooker 1865: 576) de Wit (1956: 431), Phanera Loureiro (1790: 37), Piliostigma
Hochstetter (1846: 598) and Tylosema (Schweinfurth 1868: 17) Torre & Hillcoat in Exell & Mendonça (1955: 38)
were recognized by Lewis & Forest (2005). Later, Wunderlin (2010) recognized those genera plus Schnella Raddi
(1820: 32) and minus Lasiobema (which he synonymized with Phanera), based on the molecular phylogenetic results
of Sinou et al. (2009). Wunderlin’s (2010) phylogenetic classification has gained general acceptance (e.g., Legume
Phylogeny Working Group 2017). Bauhinia sensu stricto (s.s.), i.e., in Wunderlin’s (2010) circumscription (more or
less equivalent to the “Bauhinia group” in Wunderlin, 1983, and to Bauhinia subgenus Bauhinia in Wunderlin 1987),
is still pantropical, with ca. 160 species of trees, shrubs, or rarely semiscandent plants, with entire or bilobed leaves
(usually interpreted as formed by two partially or completely fused leaflets, see Urban 1885, van der Pijl 1951, Dubey
et al. 1990; truly bifoliolate leaves, i.e., with two free leaflets, are rare in Bauhinia s.s.), often with “intrastipular
spines” (see comments below), devoid of tendrils, and with a calyx that lacks apical nectaries, either spathaceous or
divided into 2–5 lobes at anthesis.
In southern Central America (Costa Rica and Panama), there are 18 documented native species of Bauhinia s.l., 10
of which are members of Bauhinia s.s. (including the one described here). For Panama, Correa et al. (2004) reported
13 native species of Bauhinia s.l., eight of which are members of Bauhinia s.s.; these authors included B. multinervia
(Kunth 1823: 316) de Candolle (1825: 515) as native to Panama (probably based on Caballero 32, F), but that species
occurs naturally only in northeastern South America and the Lesser Antilles (Wunderlin 1983), and is only occasionally
cultivated in Costa Rica and Panama. For Costa Rica, Zamora (2010) reported 13 native species of Bauhinia s.l., seven
of which are members of Bauhinia s.s. Zamora (2010) listed two undescribed species of Bauhinia s.l. for Costa Rica:
the lianescent “Bauhinia sp. A”, later described as Schnella bahiachalensis Zamora (2013: 1–6), and the arborescent
Bauhinia sp. B”, which is formally described below (as B. proboscidea P. Juárez, R. Flores & M.A. Blanco).
Because of their remarkable vegetative similarity and the commonly poor preservation of their delicate flowers
upon pressing, herbarium specimens of the new species described here have been frequently misidentified as Bauhinia
beguinotii Cufodontis (1933: 192). Thus, we provide a detailed comparison between both taxa. Also, we take the
opportunity to compare B. beguinotii with B. gorgonae Killip ex R.S. Cowan (1961: 281–282, f. 2c–d), the latter of
which has been considered a variety of the former (Wunderlin 1973, 1976, 1983, 1986, Quiñones 2005, Castellanos &
Lewis 2012).
Finally, we provide conservation assessments based on the International Union for Conservation of Nature (IUCN)
Red List criteria for B. beguinotii, B. gorgonae, B. pansamalana Donnell Smith (1888: 27–28; a closely-related species
to the one described here as new) and B. proboscidea.
Material and Methods
Specimens (or their photographic images) determined as Bauhinia beguinotii, B. gorgonae and B. pansamalana, and
Bauhinia specimens from Mexico and Central America not determined to species level, were studied from the following
herbaria: AAH, BC, BM, COL, CR, CR-INB, EAP, F, GH, K, MEXU, MO, NY, P, PH, PMA, SCZ, TEFH, UCH, US,
USF, USJ and W. Herbarium acronyms cited above and elsewhere in this paper follow Thiers (2018) with the exception
of CR-INB, which denotes specimens deposited in the former INB herbarium, now nominally incorporated in CR but
still housed in the off-site building, formerly of the Instituto Nacional de Biodiversidad, in Santo Domingo de Heredia,
Costa Rica. Measurements of vegetative and reproductive structures were made both from herbarium specimens and
from a living, cultivated plant of B. proboscidea (see below).
A morphological species concept was adopted for practical delimitation, so that apparent gaps in morphological
variation (especially if associated with geographical disjunction) were used to distinguish species (i.e., a phenetic
species concept sensu Judd et al. 2016). Resulting species circumscriptions were compared to type specimens and
to protologue descriptions to verify the proper application of names. The resulting taxa are assumed to represent
independent evolutionary lineages (i.e., evolutionary or phylogenetic species definitions, sensu Judd et al. 2016).
Clarifying comments and a complete list of specimens examined are provided for each taxon. For herbarium
collections that did not include geographical coordinates, these were found or inferred using the Missouri Botanical
Garden´s Gazetteer of Costa Rican Plant Collecting Locales (
shtml#B) or other sources; for those specimens, the corresponding geographical coordinates are indicated in brackets.
For some specimens with unprecise locality data, geographical coordinates could not be inferred with certainty.
BAUHINIA PROBOSCIDEA Phytotaxa 361 (1) © 2018 Magnolia Press 27
Red List conservation assessments were performed for B. beguinotii, B. gorgonae, B. pansamalana and B.
proboscidea, according to the methodology of the International Union for Conservation of Nature (IUCN; IUCN
Standards and Petitions Subcommittee 2017), to determine their corresponding IUCN Red List category of extinction
risk (IUCN Species Survival Commission 2012). We lack precise information on population data and dynamics for
either taxa, and we assume a stable current population trend. The extent of occurrence (EOO) and area of occupancy
(AOO) aspects of the geographic range of the assessed taxa were obtained with the software GeoCAT (Bachman et al.
2011) using the geographical coordinates of herbarium specimens (standardized to decimal degrees). In order to avoid
misidentified records, the specimens cited in this paper (which we were able to study directly or through photographic
images) were used as the only primary occurrence data. We performed both global-level and national-level assessments
for each taxon. These assessments will be submitted to the IUCN Red List Unit upon acceptance of publication of this
Results and Discussion
Bauhinia proboscidea P. Juárez, R. Flores & M.A. Blanco, sp. nov. (Figs. 1, 2)
Similar to Bauhinia pansamalana but differs from that species by its shorter petioles, more elongate leaf blades, inflorescences with 2
(rarely up to 8) flowers, larger flowers, longer fruits and allopatric, more southern geographic distribution.
Type:—COSTA RICA. Prov. Puntarenas: cantón Osa, distrito Drake, poblado Rancho Quemado, cerca de la toma de agua, bosque
primario, ribera de una quebrada, 210 m, 8°40’58.30’’ N, 83°33’36.85’’ W, 4 May 2014 (fr.), Juárez 420 (holotype: USJ!; isotypes:
CR!, MO!).
Shrubs or small trees (occasionally reportedly semiscandent), conforming to Troll’s architectural model, 4–12(–21) m
tall (but see comments) × 10–25 cm at breast height when reproductive, andromonoecious. Stems plagiotropic, with
distichous, alternate phyllotaxy; green, slightly compressed, fractiflex and glabrous when young; becoming cylindrical
with age; bark smooth to striate, lenticelate, light brownish-gray. Leaves simple; petiole bituminous, 10–20 × 1–3
mm; blade chartaceous, entire, ovate, (2.5–) 5–30 × (1.2) 2–9 cm, basally cuneate to rounded, sub-truncate, obtuse, or
slightly acute, apically acuminate or bifid, glabrous on both surfaces, margin entire; venation acrodromous, primary
nerves 5, the 3 central ones reaching the blade apex, secondary nerves irregularly perpendicular to the primaries, tertiary
venation reticulate; stipules ovate, subulate, acute, up to 1 × 0.5 mm, early-caducous; intrastipular spines subconical,
acute, up to 1.5 mm long × 0.4 mm in diameter on the dorsal (upper) side of branches, up to 2.5 mm long × 0.8 mm in
diameter on the ventral (lower) side of branches. Inflorescences both axillary and terminal; terminal ones occasionally
displaced laterally by the axillary branch of the last leaf and thus appearing leaf-opposed, racemose, rarely branched,
sequentially flowered, with 1–2 (–8) flowers, usually only the first flower hermaphroditic, the subsequent flowers
staminate; rachis 1–3 cm long; floral bracts broadly triangular, 1 × 3 mm, minutely strigose; bracteoles ovate, 1.5 × 1.3
mm, minutely strigose, located at the base of the pedicel, adjacent to the floral bract. Flowers zygomorphic to slightly
asymmetric, either hermaphroditic or staminate. Pedicel 3–6 mm long × 1.5–2.5 mm in diameter, minutely strigose.
Flower buds botuliform, up to 5 cm long × 4–5 mm in diameter just before anthesis, pale green. Hypanthium tubular,
8–15 mm long × 4.5–5 mm in diameter externally, 7 mm long × 2 mm in diameter internally. Calyx spathaceous,
formed by 5 connate sepals, 2.5–5.5 × 0.6–1.2 cm, straight or reflex, minutely striate, minutely strigose abaxially,
glabrous adaxially, dehiscing along one or two sepal margins (usually one of the lowermost 3 sepals, and thus opening
obliquely in relation to the rest of the flower), occasionally dehiscing along an additional sepal margin, and frequently
splitting incompletely along the basal margins of other sepals. Petals 5, erect to suberect, subequal or the 2 lowermost
ones smaller, 28–63 × 1.5–5 mm, narrowly spathulate, apically acute, glabrous, white with a green middle vein and
yellowish secondary veins, becoming pink on the second day of anthesis. Stamens 10, heterodynamous, in 2 whorls
of 5 each; the 3 lowermost of the external whorl fertile, the rest staminodial; filaments dark red, glabrous, connate,
forming a staminal tube 15–20 mm long × 3–3.5 mm in diameter; filaments of fertile stamens projecting 30–42 mm
beyond staminal tube, those of hermaphroditic flowers strongly recurved to coiled so that the anthers lie under (or to
the side of) staminal tube, those of staminate flowers straight, slightly incurved apically so that the anthers face the tip
of the pistillode; filaments of staminodes projecting 1–11 mm beyond staminal tube; fertile anthers linear, dorsifixed,
introrsely longicidal, margins minutely villous, grayish pink externally, white internally, 7 × 1 mm each, connate,
forming an oblong pseudosynanther 7 × 3 mm; abortive anthers (of staminodia) narrowly ellipsoid to sagittate or
absent, minutely villous when present, up to 1.5 × 0.5 mm; pollen white. Gynoecium of 1 carpel, stipitate; fertile pistil
28 Phytotaxa 361 (1) © 2018 Magnolia Press
FIGURE 1. Bauhinia proboscidea. A. Fruiting branch. B. Perfect (hermaphrodite) flower; note recurved fertile stamens and incurved
style. C. Staminate flower; note apically incurved fertile stamens. D. Apex of staminodes and bases of free portion of filaments of fertile
stamens in staminate flower. E. Apex of fertile stamen filaments and anthers of staminate flower. F. Floral diagram of perfect flower.
G. Pair of intrastipular spines at a node (leaf scar on opposite side of stem, not visible), with liquid secretion. A based on type specimen
(Juaìrez 420, USJ, CR, MO); B–F based on photographs of Juárez 1241 (USJ). Drawn by P. Juaìrez.
BAUHINIA PROBOSCIDEA Phytotaxa 361 (1) © 2018 Magnolia Press 29
FIGURE 2. Bauhinia proboscidea. A. Staminate flower. B. Perfect (hermaphrodite) flower. C. Apex of fertile stamens of staminate
flower with incurved filaments and open anthers. D. Detail of perfect flower, showing recurved fertile anthers and pistil with incurved
style. E. Detail of staminodes (red and yellow, with white abortive anthers), filaments of fertile stamens (dark red) and pistillode (green) in
staminate flower. F. Immature fruits. G. Two-flowered inflorescence, with one perfect flower and one staminate flower; the perfect flower
(left) has already shed its petals. H. Detail of stem node showing intrastipular spine with liquid secretion. F from Flores et al. 3835 (PMA)
by R. Flores; all other photos from Juárez 1241 (USJ) by P. Juárez.
30 Phytotaxa 361 (1) © 2018 Magnolia Press
(of hermaphroditic flowers) 30–62 mm long; gynophore ca. 10 mm long, glabrate; ovary narrowly oblong, strigillose,
whitish, 18–20 × 2–3 mm deep × 1 mm thick; style ca. 5 mm, glabrate, incurved; stigma facing the flower base due to
the curvature of the style, oblong, green, ventrally papillose, dorsally glabrous, 6–7 × 2 mm; pistillode (of staminate
flowers) linear, whitish with green margins, 15–30 × 1.5–2 mm. Fruits ensiform to falcate, flattened, basally obtuse
to acute, stipitate (gynophore 2.5–3 cm), apically rostrate with the persistent style, (12–)20–35 × 3 cm, glabrous,
pendulous, green when immature, drying brown to blackish when mature, with 8–12 seeds, dehiscing through both the
ventral and dorsal sutures, valves becoming helically coiled. Seeds sub-elliptic, slightly lenticular, ca. 1.5–2 × 1–1.5
cm, dark brownish, shiny. Seedling erect; cotyledons unknown; eophylls (first leaves after cotyledons) opposite, blade
chartaceous, sub-quadrate, 4.2–4.5 × 5.1–5.3 cm, 5-nerved, each of the 4 lateral nerves terminating in a 2.2–3.5 cm
long acumen extending from the blade, the central nerve terminating in a 1–2 mm long apiculus.
Distribution and habitat:—Bauhinia proboscidea occurs on the Pacific slope of central and southern Costa
Rica and extreme western Panama. It has been collected in many localities in the Costa Rican provinces of Puntarenas
and San José, and from a single locality in the Panamanian province of Chiriquí, in the Burica Peninsula, adjacent
to the Costa Rican border (Figure 3). Unaware of the recent Panamanian collections, Zamora (2010) considered B.
proboscidea (as Bauhinia sp. B) as a Costa Rican endemic. Bauhinia proboscidea grows in primary and secondary
moist forest and rain forest at elevations of 200–1300 m, frequently near streams.
FIGURE 3. Distribution map of Bauhinia beguinotii, B. gorgonae, B. pansamalana and B. proboscidea, according to the localities of
available herbarium collections with geographic coordinates (indicated or inferred, see text).
Phenology:—Flowering has been recorded during both dry and rainy seasons, from January to May and August
to September. Fruiting specimens have been collected from January to August. A small tree now in cultivation at
Lankester Botanic Garden has flowered more or less continuously after reaching a height of ca. 70 cm. Presence of
seedlings has been documented in July (based on the seedling included in Valverde 1060, US).
Observations on reproductive biology:—Anthesis apparently begins during the night or just before dawn. The
petals stay erect and in good condition for ca. 15–20 h (if protected from the sun or heavy rain), and then shrivel,
turning pinkish or magenta before falling (in hermaphroditic flowers). On the second day of anthesis the anthers (in
hermaphrodite flowers) shrivel, but the stigma remains in good condition and apparently receptive. Flowers abscise on
the third day of anthesis.
BAUHINIA PROBOSCIDEA Phytotaxa 361 (1) © 2018 Magnolia Press 31
Most inflorescences in Bauhinia proboscidea are two-flowered. Observations made on numerous inflorescences
produced by the small tree cultivated at Lankester Botanic Garden indicate that most flowers are staminate, and when
hermaphroditic flowers are produced, they are usually the first of the pair to open. Very rarely two hermaphroditic
flowers are produced on the same inflorescence.
Pollinators are unknown, but the floral morphology suggests adaptation to either hawk-moths or hummingbirds.
Flowers of the related Bauhinia pansamalana, which have the same basic morphology, are visited by hummingbirds,
according to Wendt et al. 3399 (MEXU). Bauhinia curvula Bentham in Martius (1870: 194), which has flowers with a
similar morphology, is pollinated by hawk-moths in Brazil (Munin et al. 2008). As in other species of Bauhinia, fruits
of B. proboscidea are explosively dehiscent, and several herbarium specimens show open fruits with coiled valves.
Etymology:—The specific epithet refers to the prominent staminal tube and the projecting pistil (the latter in
hermaphroditic flowers), which together resemble a snout or proboscis of an animal. The recurved fertile stamens
of hermaphroditic flowers also bear a superficial resemblance to the tusks of elephants Loxodonta spp., order
Comments:—Although at least two dozen previous Costa Rican herbarium collections of Bauhinia proboscidea
are known, the earliest dating from 1989, few of them have flowers and these are poorly preserved. During a plant
collecting trip to the Osa Peninsula in 2014, a fruiting specimen of this taxon was collected (P. Juárez 420). One
seed of that specimen was planted and eventually grew; two years later the young plant (vouchered by Juárez 1241)
started blooming, and provided plenty of flower material for a detailed description. This plant is now in cultivation at
Lankester Botanic Garden of the University of Costa Rica.
Bauhinia proboscidea is apparently not rare, judging from the quantity of available herbarium specimens. As
in other arborescent species of Bauhinia, the vegetative morphology and development of B. proboscidea conform to
Troll´s architectural model (Hallé et al. 1978). Valverde 1060 describes the plant as a 21 m tree, which would make
it one of the tallest arborescent Bauhinia in Central America; however, we suspect this could be an error, as all other
specimens of B. proboscidea describe the plant height as 4–12 m. The seedling description is based on the one included
with the specimen Valverde 1060 at US (image available at
The “intrastipular spines” of Bauhinia proboscidea and those of some other species of Bauhinia s.s. are glandular
and secrete tiny drops of nectar (van der Pijl 1951, Dubey et al. 1990, Rezende et al. 1994, P. Juárez & M.A. Blanco,
personal observation), possibly as a reward for ants that defend the plants against herbivores. Marazzi et al. (2012: Fig.
2) documented this phenomenon for B. macranthera Bentham ex Hemsley (1880: 49) but confused the structures with
stipules. In his detailed review of the morphology of Bauhinia s.l., Urban (1885: 82–85) stated that these projections,
seemingly unique among seed plants, originate from the axil of the true stipules (which are often caducous), and
considered them to be derived from the intrastipular trichomes (colleters) present in other species of the genus. However,
González & Marazzi (2018) showed that the corresponding structures (which they call prickles) in B. forficata Link
(1821: 404) subsp. pruinosa (Vogel 1839: 301) Fortunato & Wunderlin (in Fortunato 1986: 550) do not arise from the
axils of the stipules (which are occupied by colleters), but arise from each side of the leaf base instead, just below the
stipules. Thus, the homology of the “intrastipular spines” of Bauhinia remains unsettled. In this regard, it is interesting
to note that similar, paired, spinescent projections borne from either side of the petiole base, adjacent to stipules (and
often confused with them), occur in some unrelated members of Fabaceae (e.g., Prosopis juliflora (Swartz 1788:
85) de Candolle (1825: 447) and most other species of Prosopis section Algarobia de Candolle (1825: 446), Robinia
pseudoacacia Linnaeus (1753: 722), Vachellia seyal (Delile 1813: 286) Hurter (in Mabberley 2008: 1021); Burkart
1937, Bell & Bryan 2008, Sharma & Kumar 2012), and may represent modified short, leafless shoots produced by
accessory, collateral, displaced axillary buds (sensu Bell & Bryan 2008; also Burkart 1937).
As in other species of Bauhinia, B. proboscidea produces both hermaphroditic and staminate flowers (see Wunderlin
1983, Tucker 1988, Torres Colín 1999). Bauhinia s.l. shows a variety of floral morphologies and arrangements of
stamens and staminodes (Urban 1885, Endress 1994); we provide a floral diagram of B. proboscidea (Fig. 1F) for
comparison with Figs. 11–19 in Urban (1885). The petals of B. proboscidea fall easily if the flower-bearing branch
is cut; this probably explains why herbarium collections with complete flowers in good condition are scarce in this
Conservation status:—With an EOO of 8,347.21 km2, an AOO of 52 km2, and known from 11 localities (10 in
Costa Rica, one in Panama), of which four are protected areas (three in Costa Rica, one in Panama), our global-level
assessment for Bauhinia proboscidea shows that this species fits the category of Least Concern (LC) of the IUCN Red
List. Our national-level assessment for Costa Rica also places it in the LC category (EOO= 6,075.06 km2, AOO= 48
32 Phytotaxa 361 (1) © 2018 Magnolia Press
For Panama, however, the national-level assessment categorizes this species as Endangered [EN B2ab (ii, iii, iv)]
because of its restricted AOO (4 km2) and the severe threats facing its only known population. The El Chorogo forest,
which extends for ca. 10 km in a narrow strip on the Panamanian side of the Burica Peninsula along the border with
Costa Rica, is in an isolated hilly area (150–689 m elevation) and contains the largest forest remnant (ca. 800 ha) of
the Pacific slope lowlands in extreme western Panama (Flores et al. 2016). This is the only place in Panama where
12 species of birds globally restricted to the South Central American Pacific Slope Endemic Bird Area (BirdLife
International 2017) occur; thus, it has been designated a National and Global Important Bird Area (Angehr 2003).
It also contains a population of the similarly restricted and globally endangered Red-Backed (or Central American)
Squirrel Monkey (Angehr 2003). The only part of this forest under official protection is that within the Audubon-El
Chorogo (178 ha) and San Bartolo (105 ha) Private Nature Reserves. The remaining forest is privately owned by
various farmers, has no formal protection and is in danger of being deforested for cattle ranching and other agricultural
pursuits, or for selective logging.
Additional specimens examined (paratypes):—COSTA RICA. Cultivated: [Germinated from seeds of the type
collection; cultivated at San Rafael de Heredia, later planted at Lankester Botanic Garden], 1 July 2016, Juárez 1241
(USJ-spirit). Puntarenas: Buenos Aires, Cuenca Térraba-Sierpe, Potrero Grande, La Lucha, alrededores de Rancho
Amuo, 1000–1100 m, 9º06’ N, 83º06’ W, 18 February 2008 (fr.), Santamaría 6989 (CR, CR-INB, PMA); same locality,
3 November 2010 (sterile), González et al. 1862 (CR-INB); same locality, 3 November 2010 (sterile), González et al.
1863 (CR-INB); same locality, 3 November 2010 (sterile), González et al. 1864 (CR-INB); same locality, 3 November
2010 (sterile), González et al. 1865 (CR-INB); Bueno Aires, Cuenca Térraba-Sierpe. Potrero Grande, Lodge Monte
Amou, La Lucha, Sendero Quiero a 2.5 km de La Lucha, 1100 m, 9º06’ N, 83º06’ W, 22 Febrary 2000 (fl., fr.), Aguilar
5989 (CR, CR-INB, MO); Golfito, Dos Brazos de Río Tigre, Jiménez, Cerro Müller, bajando por la fila a los afluentes
de Río Niño, 400–744 m, 8º30’ N, 83º28’ W, 30 August 1990 (fr.), Herrera & Fallas 4173 (CR, CR-INB, MO); Osa,
Parque Nacional Corcovado, Sirena, Los Patos, Mirador Trail, 200–300 m, 8º41’40” N, 83º34’48” W, 26 October 1989
(fl.), Kernan 1303 (CR-INB, K); Osa, Aguabuena, cuenca superior de la Quebrada El Campo, Rincón, 300 m, 8º42’35”
N, 83º31’50” W, 1 October 1990 (fl., fr.), Herrera 4421 (CR, K, USJ); Osa, Rancho Quemado, Sierpe. siguiendo el
camino a Drake, cabeceras de Río Drake, 400 m, 8º41’ N, 83º35’ W, 18 June 1990 (fr.), Herrera 4204 (CR, CR-INB,
MEXU, USJ); Osa, Reserva Forestal Golfo Dulce, Rancho Quemado centro, 300 m, 8º41’40” N, 83º33’35” W, 15
December 1991 (fr.), Aguilar 751 (CR, CR-INB, MO); Osa, Sierpe, Agua Buena, bosques al Noreste de la Fundación
Neotrópica, 290 m, 8º42’36.31” N, 83º31’21.30” W, 15 September 2013 (sterile), Jiménez & Juárez 1733 (USJ); Osa,
Sierpe, Campo de Aguabuena, Parcela Proyecto BDEF, 95 m, 8º42’11” N, 83º31’05” W, 30 January 2013 (sterile),
Chacón et al. 1613 (USJ); Osa, Palmar Norte, bosque aledaño a la toma de agua potable, faldas de Fila Retinto, 700 m,
08°57’36”N, 83°27’36”W, 13 May 1992 (fl., fr.), Jiménez 1077 (CR, CR-INB, MEXU, MO, USJ). San José: Acosta,
cuenca del Pirrís-Damas, Fila Ayarales, falda NE, Quebrada Ayarales, camino al Cornelio, 1250–1300 m, 9º43’ N,
84º12’ W, 13 May 2001 (fl.), Morales 8050 (CR-INB, K, MO, USJ); Acosta, Fila Bustamante, 3 km NO Hacienda
Tiquires, Cerro El Cornelio, faldas Fila Aguabuena, 1300 m, 9º42’ N, 84º13’ W, 16 December 1994 (sterile), Jiménez
1699 (CR, CR-INB); Acosta, Fila Bustamante, Tiquires, bosque primario en el camino a Zoncuaco, cerca del paso
sobre la quebrada El Ayaral, hasta cerca del cruce, 1100–1250 m, [9°43’N 84°12’W], 4 June 1995 (sterile), Morales
4343 (CR-INB, MO); Aguirre, Londres, estribaciones de Cerro Nara, 800 m, 9º29’30” N, 84º01’30” W, 13 May 1994
(fr.), Herrera 7030 (CR); Aserrí, cuenca del Pirrís-Damas, Fila Agua Buena, Quebrada Laja, 1100–1200 m, 9º40’ N,
84º11’ W, 23 January 2003 (fr.), Morales & Hammel 9060 (CR-INB); Aserrí, cuenca del Pirrís-Damas, entre Escuadra
y Zoncuano por Hacienda Tiquires, 1000–1500 m, 9º40’ N, 84º11’ W, 28 June 1995 (fr.), Hammel 19896 (CR, CR-
INB, K, MO); Dota, San Isidro, Zona Protectora Cerros Nara, 900–1000 m, 9º29’40” N, 84º00’50” W, 21 July 1998
(fl., fr.), Valverde 1060 (CR, US, USJ); Dota, Copey, área no protegida, La Chaqueta, márgenes de Río Savegre, 750
m, 9º29’55” N, 83º54’50” W, 16 May 2002 (fr.), Estrada & Bustamante 3271 (CR, K, MO, US); Tarrazú, San Lorenzo,
área no protegida, Cerro Diamante, 600–900 m, 9º32’40” N, 82º02’30” W [erroneous coordinates; actual approximate
coordinates: 9.554985 -84.043280], 9 February 1992 (fr.), Estrada 2097 (CR); Tarrazú, cuenca del Naranjo y Paquita,
2.5 Sur de Napoles, sobre la carretera de San Marcos de Quepos, 1300 m, 9º35’ N, 84º04’ W, 17 February 1998 (fl.,
fr.), Rodríguez et al. 3039 (CR, CR-INB, MO). PANAMA. Chiriquí: Distrito de Barú, Corregimiento de Puerto
Armuelles, Reserva Natural Privada Audubon-El Chorogo, 547 m, 8°17’26.4” N, 82°59’52.4” W, 1 April 2016 (fl,
fr) Flores et al. 3835 (PMA, SCZ, UCH); same locality, 1 April 2016 (sterile), Flores et al. 3836 (PMA, SCZ); Punta
Burica, El Chorogo, desde la finca de Fernando Chavarría hacia la trocha en límite fronterizo, altos de la Laguna, Finca
Audubon, 16 May 2007 (fr.), Aranda et al. 3921 (SCZ).
BAUHINIA PROBOSCIDEA Phytotaxa 361 (1) © 2018 Magnolia Press 33
Bauhinia pansamalana Donnell Smith (1888: 27–28). Casparia pansamalana (Donn. Sm.) Britton & Rose (1930:
Type:—GUATEMALA. [Alta Verapaz]: River banks of Pansamalá and Sacolol, Pansamalá, [ca. 15º28’26” N, 90º18’40” W], alt. 37–3800
ft, June 1885 (fl., fr.), H. von Tuerckheim 681 (holotype: US!, isotypes: GH!, K!, NY!, P!, US!).
Distribution and habitat:—Bauhinia pansamalana occurs on the Caribbean slopes of southern Mexico (Chiapas,
Tabasco and Veracruz states), Guatemala and western Honduras, at elevations of 30–1600 m (see also Standley &
Steyermark 1946, Wunderlin 1986, Nelson-Sutherland 2008; Fig. 3). Available specimens consistently describe the
habitat as evergreen montane forest (selva alta perennifolia), and occasionally on limestone-based, rocky soils.
Comments:—Bauhinia pansamalana and B. proboscidea appear to be sister species. Bauhinia proboscidea
differs from B. pansamalana in its shorter petioles (1–2 vs. 3–6 cm, respectively), more elongate leaf blades (length-
to-width ratio 2.5–3.3 vs. 1.3–2.8), inflorescences with fewer flowers (1–8 vs. 20–40), longer petals (28–63 vs. 20–25
mm), longer fertile stamen filaments (45–62 vs. 20–25 mm), longer anthers (7 vs. 4–5 mm) and somewhat longer fruits
(20–35 vs. 10.7–29 cm). The leaves of B. pansamalana are occasionally bifid to markedly bilobed apically (even on
the same plant, as in Pittier 333 US no. 292494), while those of B. proboscidea are either entire or only slightly bifid
apically. Also, both species are markedly allopatric.
In his unpublished revision of Bauhinia s.l. for Mesoamerica, Torres Colín (1999) reported the occurrence of B.
pansamalana in Costa Rica based on three collections of B. proboscidea (Aguilar 751, Herrera & Fallas 4173 [as
Herrera 4173] and Herrera 4204); even his illustration of B. pansamalana was based on a collection of B. proboscidea
(Herrera 4204). Because the description of B. pansamalana by Torres Colín (1999) represents an amalgamation of
both species, the dimensions reported here for both species are taken directly from herbarium specimens or their
According to the classification of tribe Cercideae of Wunderlin et al. (1987), both Bauhinia pansamalana and B.
proboscidea belong in series Triandrae Wunderlin, Larsen & Larsen (1987: 14, characterized by three fertile stamens
separated from each other by a staminode) of subgenus Bauhinia section Amaria (S. Mutis ex Caldas 1810: 25)
Endlicher (1840: 1318). More recently, Wunderlin (2006: 97) synonymized series Triandrae with series Remotae
Wunderlin, Larsen & Larsen (1987: 12) of section Bauhinia.
Conservation status:—The conservation status of Bauhinia pansamalana was previously assessed by Walter &
Gillett (1998: 349), who placed it, without an explicit rationale, in the Rare (R) category (based on herbarium records
from Guatemala and Mexico only). With an EOO of 43,191.97 km2, an AOO of 40 km2, and known from at least 15
localities (10 in Mexico, four in Guatemala, one in Honduras), of which at least two are protected areas (Reserva de la
Biosfera Montes Azules in Chiapas, Mexico, and Parque Nacional Cerro Azul, Honduras), our global-level assessment
places this species in the Not Threatened (NT) category.
Our national-level assessment for Mexico places Bauhinia pansamalana in the Least Concern (LC) category
(EOO= 23,643.92 km2, AOO= 28 km2), also considering its occurrence in one protected area. Our national-level
assessment for Guatemala places this species in the Endangered (EN B2ab (ii, iii, iv)) category (AOO= 8 km2);
however, we had only two collections with inferred geographical coordinates (we were unable to infer with certainty
the precise locality and corresponding geographical coordinates of four other herbarium collections), which made it
impossible to calculate the EOO and likely underestimated the AOO. Bauhinia pansamalana has not been collected
in any protected area in Guatemala. Finally, our national-level assessment for Honduras places this species in the
Vulnerable [VU B2ab (ii, iii, iv)] category (AOO= 4 km2), considering its occurrence in a protected area.
Additional specimens examined:—GUATEMALA. Alta Verapaz: Between Sepacuité and Secoyocté, 1100
m, [ca. 15º25’59” N, 89º45’07” W], 24 May 1905 (fl., fr.), Pittier 333 (MO, US [2 sheets]); Near Chirriacté, on the
Petén highway, 900 m, 9 April 1941 (fl.?), Standley 91659 (US); Near the finca Sepacuite, April 1902 (fl., fr.), Cook
& Greggs 581 (US [2 sheets]); Pansamalá, alt. 2800 pp, April 1889 (fl., fr.), Donnell Smith 1631 (PH, US [2 sheets]).
Izabal: Along Río Bonita, 30–150 m, 21 December 1941 (st.), Steyermark 41738 (AAH). HONDURAS. Copán: La
Meseta, Lado Sur de la montaña alrededor de la meseta, 12 km al Noroeste de Florida, Parque Nacional Cerro Azul,
1600 m, 15º06’N, 88º55’W, 9 February 1992, Mejía & Hawkins 32 (EAP, MO). MEXICO. Chiapas: Finca Irlanda,
June 1914 (fl.?), Purpus 7302 (GH [2 sheets, as “7203”], US) Municipio Berriozábal, La Pera, 1200 m, 16º52’01.5”N,
93º19’39.7”W, no date (fl., fr.), Ortiz Rodríguez 051 (MEXU); Loc. 25 km al E de las Lagunas de Monte Bello, 1
March 1982 (fr.), Cabrera et al. 1826 (MEXU [2 sheets]); Loc. 9.6 km al E de Lago Tziscao y 25 km al E de lagunas
de Montebello, 1140 m, 19 February 1983, Torres et al. 2324 (GH, MEXU, MO); Municipio Ocosingo, alrededor del
ex-poblado del Charrial, 800 m, 3 December 1976 (fl., fr.), Valdivia 2404 (MEXU); Municipio Ocosingo, Naha, Selva
Lacandona, 900 m, 16º56’N, 91º33’W, 30 December 1994 (fl.), Durán 260 (MEXU); Municipio Ocosingo, zona de
34 Phytotaxa 361 (1) © 2018 Magnolia Press
trabajaderos del viejo poblado Charical, aprox. 10 km desde el puente de Nueva Palestina, Com. Lacandona, 800–500
m, 16º20’N, 91º15’W, 8 April 2000 (fr.), Paladino & Sánchez 425 (MEXU [2 sheets]); Municipio Ocosingo, Reserva
de la Biosfera Montes Azules, callejón de amejoramiento de la zona de trabajaderos del viejo poblado de San Pablo,
Nueva Palestina, Comunidad Lacandona, 700 m, 16º50’N, 91º15’W, 2 October 2000 (fr.), Paladino & Sánchez 463
(MEXU); Municipio La Trinitaria, 15 km east northeast of Dos Lagos above Santa Elena, 1000 m, [ca. 16º05’70” N,
90º28’07” W], 29 December 1981 (fr.), Breedlove 56556 (MEXU [2 sheets], MO). Tabasco: San José Puyacatengo, a
orillas del arroyo Ogoiba a 3 km a la der. del CRUSE-UACh, 60 m, 17º33’N, 92º57’W, June 1996 (fl.), Guadarrama
s.n. (MEXU no. 1325866). Veracruz: Municipio Hidalgotit, Río Soloxóxhil entre Hnos. Cedillo y La Escuadra, 150
m, 17º17’N, 94º38’W, 3 April 1974 (fl., fr.), Vásquez et al. V-338 (MEXU); Municipio Minatitlán, lomas al S del
Poblado 11, ± 27 km al E de La Laguna, 200 m, 17º14’N, 94º17’30”W, 3 June 1981 (fl.), Wendt et al. 3399 (MEXU).
Bauhinia beguinotii Cufodontis (1933: 192).
Type:—COSTA RICA. [Limón]: In regione Atlantica, “Waldeck”, 28 milia a Puerto Limon, ad silvarum margines versus Río Barbilla,
[10°06’N, 83°23’W], 40 m, 12 May 1930 (fl.), Cufodontis 664 (holotype: W! [photos F!, MEXU!, MO!]).
Distribution and habitat:—Bauhinia beguinotii occurs on the Caribbean slope of Costa Rica and Panama, at
elevations of 50–700 m (Fig. 3). See additional comments below. Available specimens describe the habitat as primary
and secondary tropical wet forest, frequently in the understory.
Comments:—In herbaria, Bauhinia proboscidea has often been confused with B. beguinotii because of its
arborescent habit, shortly petiolate, entire to apically bifid leaves of similar shape and occurrence in both Costa Rica
and Panama. However, B. proboscidea can be distinguished from B. beguinotii by its more elongate leaf blades (length-
to-width ratio 2.5–3.3 vs. 1.5–2.3), the inner pair of leaf blade nerves more or less equidistant between those of the
outer pair and the midrib (vs. much closer to the midrib), linear (vs. oblanceolate) petals, 3 (vs. 10) fertile stamens
and longer fruits (20–35 vs. 12–18 cm). Zamora (2010) mentioned the number of leaf blade nerves as a distinguishing
character between B. beguinotii and B. proboscidea (as Bauhinia sp. B”; 7 vs. 5, respectively), but an exhaustive
examination of herbarium specimens shows that this is not a reliable difference. Also, while B. proboscidea occurs
on the Pacific slope of central Costa Rica and extreme western Panama at 200–1300 m, B. beguinotii is naturally
distributed on the Caribbean slope of Costa Rica to central Panama at 50–700 m (Fig. 3). Thus, although occurring
in the same two countries, both species are allopatric. Because of misidentified specimens of B. proboscidea and
B. gorgonae, some studies based on determinations mined from herbarium databases have erroneously reported the
occurrence of B. beguinotii on the Pacific slope of Costa Rica and/or Panama, and also in Colombia and/or Venezuela
(e.g., Contu 2012, Cornejo et al. 2012, Ulloa Ulloa et al. 2017; see further comments in the following paragraphs and
under B. gorgonae below).
A sterile specimen from 800–1300 m in Darien Province in Panama, identified as Bauhinia beguinotii (Gentry
& Mori 13607, MO), has exceedingly long petioles (3.5–7 vs. 1.5–3.5 cm, the range otherwise known from the
species even in the juvenile condition) and long-acuminate to caudate apices of the leaf lobes (up to 4 cm long, a
condition never seen in B. beguinotii; see below). It is apparent that this specimen is misidentified. Another specimen
misidentified as B. beguinotii from Cerro Obu in the Panamanian province of San Blas (de Nevers et al. 8099, MO),
described as having white petals but lacking any flowers on the sheet, has large, deeply divided leaf blades with a
ferrugineous abaxial surface (a condition never present in that species, see below).
Bauhinia beguinotii has been erroneously reported from Colombia (Ruiz et al. 2016, Ulloa Ulloa et al. 2017)
based both on misidentified specimens of Schnella (Forero 1499, Romero 10081 and Romero 10087, all at COL, and
Gentry & Cuadros 63779, MO, which are lianescent and have visible tendrils), and on B. gorgonae; see comments
under that species below).
The collection Kriebel 1613 indicates that the white flowers of Bauhinia beguinotii stay open at night.
Conservation status:—Contu (2012) provided a Red List conservation assessment of Bauhinia beguinotii
according to the methods and criteria of the IUCN Species Survival Commission (2012), resulting in the placement
of this species in the Least Concern (LC) category. However, Contu cited, as sites of occurrence of B. beguinotii,
several protected areas on the Pacific slope of Costa Rica where B. proboscidea is known to occur (Reserva Forestal
Golfo Dulce, Parque Nacional Corcovado–Estación Sirena and Parque Nacional Barbilla; see specimens cited for
that species), and others in Colombia (Isla Gorgona and Paramillo National Park on the mainland; localities for B.
gorgonae and for Gentry & Cuadros 63779 [Schnella sp., cited above], respectively). Contu (2012) also listed the
upper elevation limit of B. beguinotii as 1300 m, which agrees with the corresponding value for B. proboscidea but
not for that of B. beguinotii (700 m). This, together with B. beguinotii (as circumscribed here) being confined to the
BAUHINIA PROBOSCIDEA Phytotaxa 361 (1) © 2018 Magnolia Press 35
Caribbean slope of Costa Rica and Panama, shows that Contu (2012) included specimens of B. gorgonae and also
misidentified specimens of both B. proboscidea and of the genus Schnella in her calculation of the EOO and AOO for
B. beguinotii, possibly underestimating the extinction risk of the latter.
With an EOO of 45,152.88 km2, an AOO of 56 km2, and known from nine localities (seven in Costa Rica, two in
Panama), our global-level assessment for Bauhinia beguinotii shows that this species fits the category of Endangered
(EN). However, because of its presence in at least four protected areas (three in Costa Rica, one in Panama), it can be
safely placed in the LC category. Thus, even when the EOO calculated by us with curated data is much smaller than
that calculated by Contu (2012, 270,000 km2), this species maintains the same Red List category.
Our national-level assessment for Costa Rica also places Bauhinia beguinotii in the LC category (EOO= 7,217.66
km2, AOO= 32 km2). For Panama, our national-level assessment places this species in the Vulnerable category (VU
B1ab(ii,ii,iv)+2ab(ii,iii,iv), EOO= 3,263.06 km2, AOO= 24 km2).
Additional specimens examined:—COSTA RICA. Alajuela: San Ramón, distrito Peñas Blancas, Reserva
Biológica Soltis Center, sendero a la catarata, 500 m, 10º23’14” N, 83º37’19” W, 29 August 2016 (fl., fr.), Cedeño-
Fonseca et al. 935 (USJ [2 sheets]); Upala, Volcán Tenorio, Pilón, [ca. 10°40’N 85°01’W], 19 November 1987
(sterile), Sánchez & Poveda 1285 (CR, MO); Upala, área no protegida, cuenca del Zapote, entrada la Carmela a salir
a la estación, 600–700 m, 10º43’15” N, 84º59’45” W, 19 May 2004 (fl.), Kriebel 4629 (CR, CR-INB, K, MO); Upala,
Bijagua, El Pilón, Cabeceras del Río Celeste, 700 m, 10º49’ N, 84º27’ W, 14 November 1987 (fl., fr.), Herrera 1275
(CR, F, MO [2 sheets]). Cartago: Turrialba, R.B. Barbilla, cuenca del Matina, sendero que lleva al Río Danta, 500
m, 9º58’24” N, 83º27’02” W, 16 July 1998 (fr.), Rodríguez & Vargas 3822 (CR, CR-INB); Turrialba, P.N. Barbilla,
cuenca del Matina, márgen Sur del Río Danta, 400 m, 9º58’19” N, 83º27’39” W, 16 May 2001 (fl.), Acosta et al. 3078
(CR, CR-INB, K, MO, USJ). Heredia: Sarapiquí, Rara Avis, ca. 15 km al suroeste de Horquetas, sendero Atajo, 700
m, 10º17’ N, 84º02’ W, 24 June 1989 (fl., fr.), Vargas 1 (CR, CR-INB, MO [2 sheets], USJ); Sarapiquí, Rara Avis,
ca. 15 km al suroeste de Horquetas, siguiendo los caminos Real y Atajo, hasta la Catarata Grande, en su base, 400 m,
10º17’ N, 84º02’ W, 10 October 1989 (fr.), Vargas 213 (CR, CR-INB, MEXU, USJ); Sarapiquí, Rara Avis, ca. 15 km
al suroeste de Horquetas, 400–600 m, 10º17’ N, 84º02’ W, 23 November 1989 (fr.), Vargas 321 (CR, CR-INB, MEXU,
MO); Sarapiquí, Horquetas, El Plástico, sendero El Tigre y Atajo, 700 m, 10º17’03” N, 84º02’47” W, 11 January 1994
(fl., fr.), Vargas et al. 1668 (CR, K, MO, USJ). Limón: Drenaje de Rio Valle Estrella, Montana de Andromeda, [ca.
9°40’N 83°02’W], 150 m, 26–28 October 1951 (sterile?), Shank & Molina 4466 (EAP, US); Pococí, área no protegida,
cuenca Tortuguero-Sierpe, Estación Biológica La Suerte, Primavera, 100 m, 10º26’30” N, 83º47’20” W, 2 November
2002 (fl.), Kriebel 1613 (CR, CR-INB, MO); Pococí, Llanura de Santa Clara, Finca La Suerte, 50 m, 10º26’30” N,
83º47’20” W, 8 July 1995 (fr.), Aguilar et al. 4163 (CR, CR-INB, MEXU, MO). PANAMÁ. Bocas del Toro: Al N.O.
del campamento Changuinola 1 de Corriente Grande, Cerro Bracha, [ca. 9º17’58” N, 82º32’04” W], 18 January 1980
(fl.), Correa et al. 3227 (PMA, USF); Al SE y NE del campamento Changuinola 1 del IRHE, 19 January 1980 (fl., fr.),
Correa et al. 3316 (PMA, USF). Panama: Ca. 10 km N of Pan American Highway on El Llano-Carti Road, 500 m,
12 December 1973 (st.), Gentry et al. 8881 (MO); Cartí Rd., 7 ml from turnoff at El Llano, 320 m, 9º15’ N, 78º58’ W,
16–17 April 1985 (fl.), Hammel 13547 (MO, USF); El Llano-Carti Road, 10–12 km from junction with Inter-American
Hwy, 410 m, 30 October 1974 (fl.), Mori & Kallunki 2847 (MO); El-Llano-Carti-Tupile road, 16 km N of Pan-Am
Highway at El Llano, 400–500 m, 16 January 1974 (fl.), Nee & Dressler 9362 (MO [2 sheets]); Panama/San Blas,
El Llano-Cartí road, 300–400 m, 9º20’ N, 79º00’ W, 28 August 1982 (fr.), Hamilton & Stockwell 1089 (MO, USF);
Road from El Llano to Carti, 14.8 km N of the Panamerican Highway, 300–500 m, 3 September 1977 (fl.), Folsom &
Maas 5202 (MO). San Blas [now Guna Yala]: El Llano to Carti road, near Nusgandi, along Sendero Nusgandi, west
of buildings, 250–300 m, 9º15’ N, 79º00’ W, 16 June 1987 (fl.), McPherson 11033 (MEXU, MO [2 sheets], USF); El
Llano-Carti road, near Nusagandi, along sendero Nusagandi, ca. 11 road-miles from Pan-American Highway, 300 m,
9º15’ N, 79º00’ W, 12 July 1988 (fl., fr.), McPherson 12734A (MO); El Llano-Carti Road, km 19.1, 350 m, 9º19’ N,
78º55’ W, 11 March 1986 (fr.), de Nevers et al. 7326 (MO); El Llano-Carti Road, 19.1 km from Interamerican Hwy,
350 m, 9º19’ N, 78º55’ W, 5 March 1985 (fl., fr.), de Nevers et al. 4957 (MEXU, MO, PMA, USF); El-Llano-Carti
Rd. Km 19.1, 350 m, 9º19’ N, 78º55’ W, 1 July 1985 (fl., fr.), de Nevers 5946 (MEXU, MO, PMA, USF); Nusagandi,
along continental divide on El Llano-Carti Road, headwaters of Atlantic draining creeks, 320 m, 9º19’ N, 78º15’ W, 12
August 1984 (fl.), de Nevers & Pérez 3701 (MO, PMA, USF); Nusagandi, along the continental divide on the El Llano-
Carti Road, 350 m, 9º19’ N, 78º15’ W, 30 July 1984 (fl.), de Nevers & Pérez 3582 (MO, USF); Nusgandi ridge running
NW from Punta Mamá, 300–350 m, 9º19’ N, 78º15’ W, 13 August 1984 (fl., fr.), de Nevers et al. 3728 (MEXU, MO
[2 sheets], USF [2 sheets]); Pemasky, sendero Nusagandi, Km 18 de la carretera Llano-Cartí, 350 m, 9º20’ N, 78º58’
W, 4 July 1994 (fl., fr.), Galdames et al. 1306 (CR-INB, PMA, SCZ); Trail along continental divide, 400 m, 9º20’ N,
78º56’ W, 23 July 1986 (fl.), McDonagh et al. 287 (MO).
36 Phytotaxa 361 (1) © 2018 Magnolia Press
Bauhinia gorgonae Killip ex R.S. Cowan (1961: 281–282, f. 2c–d). Bauhinia beguinotii var. gorgonae (Killip ex R.S.
Cowan) Wunderlin (1973: 570).
Type:—COLOMBIA. Nariño [actually Valle del Cauca]: dense forest along stream, east side of Isla Gorgona, 50–100 m, 11 February
1939 (fl.), Killip & Garcia-Barriga 33170 (holotype: US!; isotypes: BC!, BM!, COL!, US!).
Distribution and habitat:—Bauhinia gorgonae is endemic to Gorgona Island (Isla Gorgona) of the Department of
Nariño, Colombia, located 35 km off the Colombian Pacific coast (Fig. 3). The few available herbarium specimens
indicate elevations of 50–305 m (the highest point in the island reaches 338 m). The island is covered by tropical wet
forest without a dry season, with an average annual precipitation of 6891 mm. See additional comments below.
Comments:—Bauhinia gorgonae has been considered as a variety of B. beguinotii (B. beguinotii var. gorgonae
Wunderlin 1973; see also Wunderlin 1976, 1983, 1986, Quiñones 2005, Castellanos & Lewis 2012). However, the
leaves of B. gorgonae are consistently divided for more than one third of their length, sometimes even completely
divided in two separate leaflets, while leaves in B. beguinotii are entire or only slightly bifid apically (sinus less than
one sixth the length of the leaf blade). The flowers of both species are very similar; however, five of the ten stamens of
B. beguinotii (the shorter ones) are connate for ca. three fourths of their length, forming a prominent “staminate ligule”
(Wunderlin 1983, Fig. 1E in Torres Colín 1999), but in B. gorgonae the shorter stamens appear to be connate for less
than a fourth of their length (Fig. 2d in Cowan 1961). The fruits of B. gorgonae have not been documented.
Wunderlin (1976, 1983, 1986) stated that B. gorgonae and B. beguinotii are distinguished only in the degree of
leaf lobing, and that further work would likely demonstrate the intraspecific variability of this character. We agree that
the degree of leaf lobing is variable in many species of Bauhinia s.l. (especially in lianescent Schnella, even in the
same plant); however, the available herbarium specimens and our own observations in the field and from cultivated
plants indicate that the leaf shape of B. beguinotii in Costa Rica and Panama described above is fairly constant, and
that markedly divided leaves are never produced in any developmental stage in this species.
Given the consistent difference in leaf shape, the apparent difference in the degree of connation of the shorter
stamens, and their allopatric distribution (with resulting reproductive isolation), we consider B. gorgonae and B.
beguinotii as different species.
Two sterile specimens from Gorgona Island, misidentified as B. gorgonae (Fernández Alonso 7393 and Lozano
5189, both at COL), are lianescent and represent species of Schnella.
Bauhinia beguinotii var. gorgonae was reported for Venezuela by Stergios et al. (2008), unvouchered and as
with “unknown geographical distribution”. This erroneous report is likely based on the original description of B.
gorgonae in a Venezuelan journal (Cowan 1961), even when the protologue only cited specimens from Isla Gorgona
in Colombia. This also led to the erroneous report of B. beguinotii from Venezuela by Ulloa Ulloa et al. (2017).
According to the classification of tribe Cercideae of Wunderlin et al. (1987), both Bauhinia beguinotii and B.
gorgonae belong in series Decandrae Wunderlin, Larsen & Larsen (1987: 14, characterized by 10 fertile stamens), of
subgenus Bauhinia section Amaria.
Conservation status:—Because of its restricted AOO (estimated at 4 km2), Bauhinia gorgonae fits in the category
of Endangered [EN B2ab (ii, iv)]. For this placement we have taken into account that the only known locality where
this species occurs is currently a protected area, the Gorgona Island Natural National Park of Colombia. We used
the same inferred geographical coordinates for all four available collections (indicated for St. George Exped. 360,
Additional specimens examined:—COLOMBIA. Valle del Cauca: Gorgona Island, 800 ft, [2º58’06” N,
78º11’04” W], 7 July 1924 (fl.), St. George Exped. 360 (Kelsall s.n.) (US); Gorgona Island, 1000 ft, 10 July 1924 (fl.),
St. George Exped. 380 (Cheeseman s.n.) (US); Gorgona Island, sea level, 20 October 1924 (fl.), St. George Exped. 628
(Collenette s.n.) (K, US).
Notes on the Neotropical triandrous species of Bauhinia:—Including the species described here, only four
Neotropical species of Bauhinia with three fertile stamens are known. Bauhinia proboscidea and B. pansamalana are
quite similar and differ mostly in their leaf proportions and the dimensions of their reproductive characters (Table 1).
The other triandrous neotropical Bauhinia species are: B. coulteri Macbride (1919: 59), a species of open oak-pine,
juniper, or deciduous forest of central Mexico with shortly bilobate leaves, pink (rarely white), sessile, narrowly
elliptic petals, and 2–10-flowered racemose inflorescences, and B. pinheiroi Wunderlin (1987: 62), a rare species found
exclusively in eastern Atlantic Brazilian coastal forest (Vaz et al. 2010) with divaricate bilobed leaves, yellowish-
white, shortly clawed petals, and 24–30-flowered racemose or paniculate inflorescences. These species are related
to B. pansamalana (and B. proboscidea) because of their three fertile stamens and their pollen morphology (Torres
Colín 1999). According to Wunderlin et al.´s (1987) classification, modified by Wunderlin (2006), B. pansamalana, B.
BAUHINIA PROBOSCIDEA Phytotaxa 361 (1) © 2018 Magnolia Press 37
pinheroi and B. proboscidea belong in Bauhinia series Remotae, but B. coulteri belongs in its own monotypic section
Coulterae Wunderlin, Larsen & Larsen (1987: 12), also of section Bauhinia. A key to the triandrous Neotropical
Bauhinia species is provided below.
TABLE 1. Comparison between Bauhinia pansamalana and B. proboscidea.
Character B. pansamalana B. proboscidea
Petioles 3–6 cm long 1–2 cm long
Inflorescence 20–40 flowered 1–2(–8) flowered
Petals Narrowly elliptic, 20–25 mm long Linear, 28–63 mm long
Stamen filaments 20–25 mm long 50–55 mm long
Anthers 4‒5 mm long 7 mm long
Gynoecium 20‒25 mm long 50–55 mm long
Fruit 10.7–29 cm long 20–35 cm long
Distribution Caribbean slope of southern Mexico, Guatemala and
western Honduras
Pacific slope of central and southern Costa Rica
and extreme western Panama
Key to the Neotropical species of Bauhinia with three fertile stamens
1. Leaves blades entire, sometimes apically bifid (sinus < ¼ of the blade length) ................................................................................2
- Leaves blades bilobed (sinus > ¼ of the blade length).......................................................................................................................3
2. Inflorescences 1–8-flowered; petals narrowly linear; inhabiting evergreen forest of the Pacific slope of Costa Rica and extreme
western Panama ............................................................................................................................................................B. proboscidea
- Inflorescences 20–40-flowered; petals narrowly elliptic; inhabiting evergreen or deciduous forests of the Caribbean slope of
southern Mexico, Guatemala and western Honduras .................................................................................................B. pansamalana
3. Leaf blades divided for one half to two thirds of their length; from Brazil ......................................................................B. pinheiroi
- Leaf blades divided for less than one third of their length; from Mexico, Guatemala and/or Honduras ...........................................4
4. Leaf lobes apically rounded; petals sessile, white to pink; inhabiting open oak-pine, juniper, or deciduous forest of central
Mexico .................................................................................................................................................................................B. coulteri
- Leaf lobes acuminate; petals shortly clawed, white; inhabiting moist deciduous or tropical rainforest of southern Mexico,
Guatemala and western Honduras ..............................................................................................................................B. pansamalana
We thank Gustavo Romero (AMES), Ivón Ramírez (CICY), Eliana Noguera Savelli and María Alejandra Guillén
(Venezuelan National Herbarium, VEN) for providing access to critical literature, and Marco V. Cedeño and Michael
Grayum, José Esteban Jiménez, Rina Díaz, Lilian Ferrufino and Anthony Brach for providing specimen images from the
Missouri Botanical Garden Herbarium (MO), Mexican National Herbarium (MEXU), Escuela Agrícola Panamericana
Herbarium (EAP), Universidad Nacional Autónoma de Honduras Herbarium (TEFH), and the Harvard University
Herbaria, respectively. Suggestions by Leonardo Borges (SPSC) and two anonymous reviewers helped to improve
the manuscript. Pedro Juárez thanks Albert Morera Beita and the Laboratorio de Ecología Tropical of Universidad
Nacional de Costa Rica (UNA) for logistical support for field work. Rodolfo Flores thanks the Smithsonian Tropical
Research Institute (STRI) for logistical support for fieldwork, the Sociedad Audubon de Panamá/Panama Audubon
Society for facilitating his visit to the Reserva Natural Privada Audubon-El Chorogo and for logistical support, the
Panamanian Ministry of Environment for issuing collection permits, the Universidad de Panamá Herbarium (PMA)
for assistance in processing specimens, and Armando Quiróz, José Caballero, and Eliécer Caballero for support during
field work. Mario A. Blanco is grateful to the Universidad de Costa Rica for providing financial support to visit
Panama in 2016. This is a contribution to the Global Tree Assessment initiative (, led
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... While not stipular in origin, their homologies are unclear. They appear to be similar to intrastipular spines described in some legumes and suggested to be either modified short shoots (Sharma and Kumar 2012;Juárez et al. 2018) or emergences (Bell 2008). ...
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Bahiana is expanded from 1 to 2 species with the description of B. occidentalis K. Wurdack, sp. nov. as a new endemic of the seasonally dry tropical forests (SDTFs) of Peru. The disjunct distribution of Bahiana with populations of B. occidentalis on opposite sides of the Andes in northwestern Peru (Tumbes, San Martín) and B. pyriformis in eastern Brazil (Bahia) adds to the phytogeographic links among the widely scattered New World SDTFs. Although B. occidentalis remains imperfectly known due to the lack of flowering collections, molecular phylogenetic results from four loci (plastid matK , rbcL , and trnL-F ; and nuclear ITS) unite the two species as does gross vegetative morphology, notably their spinose stipules, and androecial structure. Spinescence in Euphorbiaceae was surveyed and found on vegetative organs in 25 genera, which mostly have modified sharp branch tips. Among New World taxa, spines that originate from stipule modifications only occur in Bahiana and Acidocroton , while the intrastipular spines of Philyra are of uncertain homologies.
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Los ejemplares tipo son especímenes de herbario que se usan para hacer la descripción de una nueva especie de planta válidamente publicada. En este estudio se revisaron las colecciones de especies tipo de Leguminosae, subfamilia Cercidoideae procedentes de Colombia depositadas en herbarios colombianos y de otros países. La subfamilia Cercidoideae está compuesta por 14 géneros, de los cuales dos se encuentran en Colombia (Bauhinia y Schnella). Doce especies y dos variedades de estos géneros se han descrito usando colecciones originarias de Colombia. Los especímenes que las conforman se han distribuido en 19 herbarios a nivel mundial, en tanto que en herbarios de Colombia solamente existen un holotipo y un isotipo. Se presentan datos adicionales sobre las especies analizadas en cuanto a endemismos, hábito, distribución geográfica y estado de conservación. Se incluye información complementaria sobre el estado actual del conocimiento de los dos géneros de la subfamilia en el país.
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A newsletter to promote communication among research scientists concerned with the systematics of the Leguminosae/Fabaceae Issue 65, Year 2018 CONTENT Page
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The classification of the legume family proposed here addresses the long-known non-monophyly of the traditionally recognised subfamily Caesalpinioideae, by recognising six robustly supported monophyletic subfamilies. This new classification uses as its framework the most comprehensive phylogenetic analyses of legumes to date, based on plastid matK gene sequences, and including near-complete sampling of genera (698 of the currently recognised 765 genera) and ca. 20% (3696) of known species. The matK gene region has been the most widely sequenced across the legumes, and in most legume lineages, this gene region is sufficiently variable to yield well-supported clades. This analysis resolves the same major clades as in other phylogenies of whole plastid and nuclear gene sets (with much sparser taxon sampling). Our analysis improves upon previous studies that have used large phylogenies of the Leguminosae for addressing evolutionary questions, because it maximises generic sampling and provides a phylogenetic tree that is based on a fully curated set of sequences that are vouchered and taxonomically validated. The phylogenetic trees obtained and the underlying data are available to browse and download, facilitating subsequent analyses that require evolutionary trees. Here we propose a new community-endorsed classification of the family that reflects the phylogenetic structure that is consistently resolved and recognises six subfamilies in Leguminosae: a recircumscribed Caesalpinioideae DC., Cercidoideae Legume Phylogeny Working Group (stat. nov.), Detarioideae Burmeist., Dialioideae Legume Phylogeny Working Group (stat. nov.), Duparquetioideae Legume Phylogeny Working Group (stat. nov.), and Papilionoideae DC. The traditionally recognised subfamily Mimosoideae is a distinct clade nested within the recircumscribed Caesalpinioideae and is referred to informally as the mimosoid clade pending a forthcoming formal tribal and/or cladebased classification of the new Caesalpinioideae. We provide a key for subfamily identification, descriptions with diagnostic charactertistics for the subfamilies, figures illustrating their floral and fruit diversity, and lists of genera by subfamily. This new classification of Leguminosae represents a consensus view of the international legume systematics community; it invokes both compromise and practicality of use.
Extrafloral nectaries (EFNs) are particularly common in Fabaceae, and their morphological diversity has proved taxonomically useful. However, their anatomy remains poorly investigated. In this study we analysed EFN anatomy in 12 species from ten genera of Fabaceae, using scanning electron microscopy and light microscopy. We found three main structural organizations of EFNs. (1) The ‘embedded’ EFNs (cryptic and externally non-visible) in Bauhinia forficata are secretory prickles in which nectar-secreting tissue is completely embedded within the prickle, representing a new kind of EFNs in plants. Among externally visible EFNs, (2) ‘elevated’ EFNs were found in seven species, including a Mimosa sp. belonging to a clade considered to lack EFNs, and (3) ‘swollen-scar’ EFNs derived from the abscission site of aborted buds were found in four species, including a new report in Galactia. In three species, we also found colleters anatomically similar to other colleters reported for Fabaceae. We discuss terminological issues in the (scarce) literature on EFN anatomy in Fabaceae, mainly due to incorrect interpretation of structures or misuse of existing terms.
Bauhinia malabarica and B. divaricata have both been reported to have dimorphic flowers; floral development of these species has been investigated and compared using SEM. B. malabarica is subdioecious, with three types of flowers: perfect, staminate, and carpellate. Individual trees usually have only one type of flower. Perfect and carpellate flowers have similar initiation of floral organs; each has five sepals, five petals, two whorls of five stamen primordia and a carpel primordium. The carpels of carpellate flowers do not differ from those of perfect flowers throughout development. Both have a gynophore or stipe and a cuplike hypanthium. Stamen development diverges markedly after mid-development: the perfect flowers have ten stamens in two whorls, the outer with longer filaments than the inner. All stamens have anthers, which are covered abaxially with abundant inflated trichomes. Carpellate flowers have a circle of short cylindrical staminodia, each bearing a few hairs, about the base of the carpel on the rim of the hypanthium. Heteromorphy in B. malabarica is effected by suppression of stamen development, even though the usual number of stamen primordia is initiated. Suppression of stamens occurs at midstage in development in carpellate flowers of B. malabarica, and is complete. In B. divaricata nine stamen primordia are released from suppression in late stage, undergo intercalary growth and form a staminodial tube around the carpel stipe. The dimorphy in B. divaricata is expressed late in bud enlargement as divergent rates of growth in the carpel in the two morphs.
A taxonomic treatment of Bauhinia subgenus Bauhinia section Amaria is presented. Fifteen species are recognized in series Decandrae and one species is recognized in series Stenanthae. Bauhinia amatlana Wunderlin from southern Mexico, Bauhinia arborea Wunderlin from Ecuador, Bauhinia ayabacensis Wunderlin from Ecuador and Peru, and Bauhinia geniculata Wunderlin from Colombia are described and the combination Bauhinia petiolata (DC.) Triana ex Hooker f. var. caudigera (S.F. Blake) Wunderlin is proposed. Keys, descriptions, and distribution data are provided.