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Hispaniolanthus: A New Genus of Capparaceae Endemic toHispaniolaAuthor(s): Xavier Cornejo and Hugh H. IltisSource: Harvard Papers in Botany, 14(1):9-14. 2009.Published By: Harvard University HerbariaDOI: http://dx.doi.org/10.3100/025.014.0103URL: http://www.bioone.org/doi/full/10.3100/025.014.0103

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TheAmerican species traditionally treated ina very broad sense as Capparis L. s.l. (Cappar-aceae), are a group of ca. 110 species of shrubs,trees, or lianas that range from southern NorthAmerica (Texas and Florida) to northernArgentina and theWest Indies (Cornejo and Iltis,2008a–d). Capparis s.l. exhibits a great diver-sity of calyx types, different calyx and corollaaestivation, different floral nectary structures,and different fruit and embryo types. It wouldbe incorrect to gather all of that morphologicalheterogeneity together under the same genericname. The high morphological diversity of thispolymorphic genus has also been supported bythe clearly demonstrated non-monophyleticcomposition (Hall et al., 2002, 2008). TheNew World species of Capparis were trans-ferred to previously published generic namestraditionally synonymyzed under Capparis(Cynophalla (DC.) J. Presl, Quadrella (DC.)

J. Presl, Capparidastrum Hutch., Neocalyptro-calyx Hutch.), and several new genera wereestablished for the remaining species (Cornejoand Iltis, 2006, 2008a–d; Iltis and Cornejo,2007a–b). While studying the specimens ofCapparis dolichopoda Helwig, it was noticedthat its vegetative parts, in combination with thepeculiar inflorescences and floral morphology,didn’t fit into any of the known genera ofAmerican Capparaceae. Thus, the followingnew genus is herein proposed for this species.

Hispaniolanthus Cornejo & Iltis, gen. nov.TYPE: Capparis dolichopoda Helwig, ArkivBot. 22A (10): 11. 1929. Figs. 1–2.Arbor parva inermis, simplex pilosi,

stipulata, folia spiralata, pedicelata, junioribusconduplicata, plantis juvenilibus cum folialinearia vel linearia-oblonga; inflorescentiasolitaria vel geminata, axillaria et subterminalia,

HISPANIOLANTHUS: A NEW GENUS OF CAPPARACEAEENDEMIC TO HISPANIOLA

XAVIER CORNEJO1 AND HUGH H. ILTIS2

Abstract. Hispaniolanthus (Capparaceae), a new genus segregated from Capparis, is described and illustrated.It is characterized by short simple hairs, minute stipules, spiral phyllotaxy, juvenile individuals with linear leafblades, mature individuals with their young leaves conduplicate, flowers axillary and subterminal, solitary and/orrarely paired, without peduncles and on remarkably long-pedicels. The calyx is decussate with closed aestivation,bearing sepals similar in size to that of the petals, four erect to divergent nectary scales, and ca. 30 to 45 stamenson short filaments, 1.5–2.0 cm long. Hispaniolanthus is a monospecific genus known only from dry forests ofwestern Haiti, facing the Island of Gonaïves. It is represented by the new combination herewith transferred fromCapparis: Hispaniolanthus dolichopodus. Its putative relationships with the morphologically closer genera ofCapparaceae are discussed.

Resumen. Describimos e ilustramos Hispaniolanthus (Capparaceae), un nuevo género segregado de Capparis.Este se caracteriza por presentar pelos simples cortos, estípulas pequeñas, filotaxia en espiral, individuos juvenilescon hojas lineares, individuos maduros con hojas jóvenes conduplicadas, flores axilares y subterminales, solitariasy/o rara vez dispuestas en pares, con distintivos pedicelos muy largos y sin pedúnculos, cálices de estivacióncerrada, con sépalos decusados de tamaño similar al de los pétalos, cuatro escamas nectarias erectas o divergentesy ca. 30 a 45 estambres con filamentos muy cortos, 1.5–2.0 cm. Hispaniolanthus es un género monoespecíficosolamente conocido de los bosques secos del occidente de Haití, frente a la Isla de Gonaïves. Está representadopor la nueva combinación, transferida de Capparis: Hispaniolanthus dolichopodus. Sus relaciones putativas conlos géneros morfológicamente más cercanos son aquí discutidas.

Keywords: Capparaceae, Hispaniolanthus, Haiti, Hispaniola, endemic

The authors are indebted to the herbaria B, C, S, and US, for the loans of Capparis dolichopoda. The plate was preparedby Kandis Elliot, artist in residence at the UW Botany Department. Special thanks to Thirza Ruiz-Zapata (MY) and ananonymous reviewer for their critical review of the manuscript.

1The New York Botanical Garden, 200th Street and Kazimiroff Avenue, Bronx, New York 10458-5126, U.S.A.Email: xcornejo@nybg.org, xcornejoguay@gmail.com. Author for correspondence.

2Department of Botany, University of Wisconsin, 430 Lincoln Drive, Madison, Wisconsin 53706, U.S.A.Email: swis@charter.net

Harvard Papers in Botany, Vol. 14, No. 1, 2009, pp. 9–14.© President and Fellows of Harvard College, 2009.

10 HARVARD PAPERS IN BOTANY Vol. 14, No. 1

FIGURE 1. Hispaniolanthus Cornejo & Iltis. A, terminal leafy branches bearing axillary inflorescences withlong-pedicellate flowers; B, close-up of flower bud just before anthesis; C, flowers at postanthesis; D, veryyoung infructescence with divergent nectary scales; E, drawing of longitudinal section of immature fruit; F,mature leaves; G, leaves from a juvenile individual; H, flower at anthesis. A–D, Ekman 9479 (C); E, Ekman9333 (B); F, Ekman 5398, (B, an isotype); G, Ekman 9333 (US); H, Ekman 9479 (S).

longissima pedicellata, pedunculis absens;alabastra globosa, cum calyce aestivatiodecussata, clausa; floris regularis, sepalis etpetalis similis longis; stamina 30 usque 45;esquama nectaria 4, erecta vel divergentia.Unarmed trees, glabrous or covered by sim-

ple short trichomes. Stipules present, ± triangu-lar, minute. Leaves spiral, the petioles of evenlengths; blades simple, conduplicate when veryyoung; juvenile individuals with linear to linear-oblong blades. Flowers axillary and subtermi-nal, solitary or in pairs and long-pedicellate,without peduncles, actinomorphic; flower budsglobose, the calyx decussate, with closed aesti-vation nearly to anthesis. Sepals 4, ovate, all ofsimilar shape and size. Petals 4, sessile, ovate-orbicular, of size similar to those of the sepals,inserted on a flat receptacle. Nectary scales 4,erect to divergent, deltoid to triangular. Andro-phore conical; stamens 30 to 45, inserted in 3 to 4alternate whorls, the filaments short, 1.5–2.0 cmlong. Gynophore 9–12 mm long, glabrous; ovarydensely verruculose. Fruits presumably pepos(see below), the immature ones 2–4 cm long,with persistent stigma; seeds ca. 20, cochleate,ca. 4–5 × 4 mm.

Etymology: named after the West IndianIsland of Hispaniola, of which the eastern halfconstitutes the Dominican Republic, and thewestern half the Republic of Haiti.Distribution: a monospecific genus, only

known along the coast in south- and central-west-ern Haiti, facing the Island of Gonaïves (Fig. 2).Helwig described Capparis dolichopoda, the

generic type of Hispaniolanthus, from a collec-tion with immature flowers as having pedicels3.0–5.5 cm long (see protologue). But E. L.Ekman 9479, a collection of the same species(not cited in the original publication but laterdetermined by Helwig himself as Capparisdolichopoda; see additional cited specimensbelow) has mature flowers on pedicels 10–15cm long. We have not seen mature fruits of thisgenus, but judging by the immature fruits ofE. L. Ekman 9633, and the vernacular names“Avocatier marron, L’avocat marron” (French,the brown avocado), which are written at the endof the description in the protologue of Capparisdolichopoda, we suspect that the fruits ofHispan-iolanthus presumably are large, brown pepos.

2009 CORNEJOAND ILTIS, HISPANIOLANTHUS,ANEW GENUS OF CAPPARACEAE 11

FIGURE 2. Map of Haiti showing the occurrence of Hispaniolanthus dolichopodus (Helwig) Cornejo & Iltis(hollow circles).

Hispaniolanthus is characterized by the pres-ence of simple hairs or a lack of any pubescence,a pair of minute, ± triangular stipules, axillaryexceptionally long-pedicellate, mostly solitaryflowers, without peduncles, these with decus-sate calyces with closed aestivation, and sepalsand petals of similar size.Axillary, erect, solitary flowers are also pres-

ent in Capparis s. str. (e.g., Capparis spinosa L.,the generic type), but the latter genus sharplydiffers from Hispaniolanthus by the scrambling(vs. erect) habit, the presence (vs. absence) ofstipular retrorse spines, flowers with one galeate(helmet-shape) sepal (vs. all sepals similar,never with a helmet-shaped one), the presenceof one nectary gland with the apex directedtoward the interior of the flower (vs. four nec-tary scales, erect to divergent), and two of thefour petals irregular, with their asymmetricalbases laterally fused to each other and folded,forming a petaloid, hood-like structure that par-tially envelops and protects the nectary gland(vs. the petal bases always free, ± symmetric,and never forming any structure that envelopsthe nectary glands). With the exception of thehabit, none of theAmerican species traditionallyidentified as Capparis share the remaining char-acters of C. spinosa (mentioned above). In addi-tion, a recent molecular study of Capparaceae(Hall, 2008), demonstrated that the Old WorldCapparis s. str. (including Capparis spinosa)belong to a separate lineage that is not the clos-est relative of the NewWorld Capparaceae.Among the neotropical genera of Cappar-

aceae, axillary, erect, solitary flowers are presentin Mesocapparis (Eichl.) Cornejo & Iltis, aSouth American, monospecific genus, recentlysegregated from Capparis s.l. (Cornejo and Iltis,2008b). The similar scrambling habit and thepresence of solitary, axillary flowers, with 2-seriate calyces of Mesocapparis, somewhatreminiscent of Capparis spinosa and its allies,seems to be the reason why Eichler consideredit to be intermediate between the, at that time,so-called New World and Old World Capparis(Eichler, 1865 [e.g., as Capparis subgen.Mesocapparis Eichler]). However, Capparisdolichopoda cannot be placed within Meso-capparis because the former genus/species differsfrom the latter by the simple hairs or lack ofpubescence (vs. densely stellate-multiangulatetrichomes), spiral (vs. alternate) phyllotaxy, thepresence (vs. absence) of stipules, flowers withremarkably long-pedicels (vs. flowers of shorter,medium-size pedicels), and petals of similar size

to those of the sepals (vs. sepals and petalsremarkably differentiated).Solitary, axillary flowers are also found scat-

tered in a few species of neotropical genera ofCapparaceae that always bear terminal, peduncu-late, racemose inflorescences (e.g., Capparidas-trum Hutch., Caphexandra Iltis & Cornejo, andNeocalyptrocalyx Hutch.). However, in thosecases, the axillary flowers always have pedicelsof similar size to those of the flowers from thepedunculate inflorescences.The simple pubescence, spiral phyllotaxy,

presence of minute stipules, and stamens insertedin several rows, ofHispaniolanthus are reminis-cent of Capparidastrum, a genus distributedfrom Mexico to northern Argentina and in theWest Indies (Cornejo and Iltis, 2008d). However,Hispaniolanthus differs from the latter mainly bythe presence of flowers long-pedicellate, axillaryand/or subterminal, solitary or rarely paired (vs.inflorescences terminal, racemose, with the flow-ers always on medium-sized pedicels), calycesdecussate with closed aestivation in bud (vs.calyces valvate to imbricate, with open aestivationin bud), the petals of similar size to that of thesepals (vs. the petals two to four times larger thanthe sepals) and the stamens not much exserted(vs. the stamens remarkably exserted). In addi-tion, the juvenile individuals ofHispaniolanthushave linear to linear-oblong leaf blades and themature individuals present very young condu-plicate leaves; these types of leaves are neverpresent or at least are unknown for the species ofCapparidastrum.Hispaniolanthus is representedby the following combination.

Hispaniolanthus dolichopodus (Helwig)Cornejo & Iltis, comb. nov.Basionym:Capparis dolichopodaHelwig,Arkiv

Bot. 22A (10): 11. 1929. TYPE: HAITI.Massif de la Hotte, western group, Anse àVeau, on the coral reef west of town, ca.18˚30'N, 73˚20'W, 11 January 1926 (fl),E. L. Ekman 5398 (Holotype: S [not seen];Isotypes: B [photo at NY,WIS], EHH [notseen], US [1410717]).

Additional specimens examined: HAITI.Massif desMatheux, l’Arcahaie, Lully, in forests,common, 11 Jan 1928 (fl, y fr), E. L. Ekman9479 (C, EHH [not seen], MO [not seen], S[2],US); ibidem, 14 Nov 1927 (juvenile and maturesterile leaves), E. L. Ekman 9333 (C, EHH [notseen], S, US); 29 Feb 1928 (y fr), E. L. Ekman9633 (B, S); Williamson, ca. 18˚51'N, 72˚35'W,18Apr 1930 (st), A. Dèsert 203-N (WIS [MAD]).

12 HARVARD PAPERS IN BOTANY Vol. 14, No. 1

Local names: avocatier marron, L’avocatmarron (French, the brown avocado).Habitat and distribution: dry coastal forests

of western Haiti.Phenology and biological interactions: as in

several other bat-pollinated species of neotropicalCapparaceae (e.g., Colicodendrum scabridum(Kunth) Seem. [Cornejo and Iltis, 2008e]), thenot fully opened flowers in the herbarium mate-rial studied lead us to think that the flowers ofHispaniolanthus dolichopodus are nocturnal.These presumably nocturnal flowers, correlatedwith the presence of extremely long-pedicelsthat subtend them above the foliage (hence the

specific epithet dolichopoda of the generictype), it seems are a replacement for the absenceof peduncles and thus are adaptations that fitwithin the bat pollination syndrome.Conservation: known from five collections:

the type plus the four additional specimens citedabove, all of them were gathered in Haitibetween 1926 to 1930. The status of the existingpopulations and the biology of this species isunknown.We encourage local botanists to redis-cover this interesting, isolated genus andspecies, previously recorded as locally common(E. L. Ekman 9479), and include it in programsof conservation and propagation.

2009 CORNEJOAND ILTIS, HISPANIOLANTHUS,ANEW GENUS OF CAPPARACEAE 13

KEY TO THE GLABROUS OR SIMPLE-PUBESCENT AMERICAN GENERA OFCAPPARACEAE WITH SIMPLE LEAVES

1a. Inflorescences axillary, without peduncles, flowers often solitary or in pairs, with pedicels elongate (10–15 cmlong); calyx with closed aestivation nearly to anthesis, decussate and isosepalous; sepals and petals of similarsize . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Hispaniolanthus Cornejo & Iltis

1b. Inflorescences always terminal and pedunculate (sometimes also axillary in Anisocapparis Cornejo & Iltis),flowers with pedicels short to medium size (1–5 cm long); calyx with open aestivation in buds, valvate to imbri-cate and isosepalous or decussate and anisosepalous (only in Anisocapparis); petals 2 to 3 times longer than thesepals . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2

2a. Terminal branchlets with extrafloral, supra-axillary nectaries and distichous leaves (spiral leaves only inCynophalla sclerophylla (Iltis & Cornejo) Iltis & Cornejo), petioles of similar length, generally short, pulviniabsent; calyx decussate, anisosepalous, the outer pair of sepals smaller; floral nectaries ± horizontally disposedon the receptacle, flat to somewhat concave, or sometimes nectary scales rudimentary and widely divergent;pollen tectate-spinulose; fruits capsular, dehiscent; seeds with green embryo. . . . . . . . Cynophalla (DC.) J. Presl

2b. Terminal branchlets without extrafloral, supra-axillary nectaries and spiral, subopposite and/or 3-verticillateleaves, the larger ones often with petioles of different length and pulvinate (often not obvious in herbariummaterial); calyx valvate to imbricate, isosepalous (decussate and anisosepalous only in Anisocapparis); floralnectaries fleshy, usually rounded and arranged on a flat receptacle, or erect nectary scales or cupular or disci-form floral nectaries within a small hypanthium; pollen tectate-spinulose, tectate-perforate, striato-reticulate orreticulate; fruit capsular or/to pepo or amphisarcous; seeds usually with a white or cream embryo, very rarelyyellow or green . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3

3a. Calyx decussate, anisosepalous, the outer pair of sepals half of the length of the larger inner pair; floralnectaries of 4 scales, strongly dimorphic (2 major and 2 minor); pollen finely reticulate, seeds ± subglobose,embryo highly anisocotylar, the major cotyledon subglobose, massive, compact, specialized for storagestarches, and the minor cotyledon rudimentary or absent; cataphylls present in the earlier stages ofgermination . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Anisocapparis Cornejo & Iltis

3b. Calyx valvate to imbricate, isosepalous; floral nectaries of 4 isomorphic usually rounded fleshy glands, or4 isomorphic scales, or cupular to disciform floral nectary within a small hypanthium; pollen tectate-spinulose,tectate perforate, or striato-reticulate; seeds cochleate to ellipsoid, embryo convolute, thin and flexible(when fresh), cotyledons of similar sizes to somewhat asymmetric; cataphylls absent in the earlier stagesof germination. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4

4a. Bracts, if present, often triangular and usually less than 2 mm, arranged at and near the base of the inflorescence’speduncle; floral nectaries of 4 fleshy glands, usually rounded and inserted on a flat receptacle and/or sometimesfused to the base of the androgynophore; petals inserted on a ± flat receptacle; stamen filaments inserted on sev-eral alternate whorls on a hemispherical convex surface at the upper part of the androphore (seen at anthesis infresh material), the pollen tectate-spinulose or tectate-perforate; fruit capsules or/to pepos or amphisarca, usuallyoblong to elliptic or globose (± oblong-torulose in C. frondosum), the apex ± rounded, few times acute (whenfresh), pulp ± present (sometimes scarce when fresh and disintegrating when drying); seeds cochleate or reniformarranged in 2 or more rows, the cotyledons symmetric. . . . . . . . . . . . . . . . . . . . . . Capparidastrum (DC.) Hutch.

4b. Bracts linear, to 13 mm, present at and near the base of the inflorescence’s peduncle; floral nectaries cupularor disciform within a small hypanthium; petals fused at the edge or at the outer edge of a cupular or disciformfloral nectary; stamen filaments inserted on 2 alternate whorls on a slightly swollen androgynophore, thepollen striato-reticulate; fruits moniliform or linear-torulose, the apex narrowly-attenuate (when fresh),essentially without pulp; seeds ellipsoid arranged in 1 row, the cotyledons jackknifed, somewhatasymmetric . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .Monilicarpa Cornejo & Iltis

14 HARVARD PAPERS IN BOTANY Vol. 14, No. 1

LITERATURE CITED

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–––––––. 2008a. New combinations in SouthAmericanCapparaceae. Harvard Pap. Bot. 13(1): 117–120.

–––––––. 2008b. Two new genera of Capparaceae:Sarcotoxicum andMesocapparis stat. nov., and thereinstatement of Neocalyptrocalyx. Harvard Pap.Bot. 13(1): 103–116.

–––––––. 2008c. Anisocapparis y Monilicarpa: dosnuevos géneros de Capparaceae de América delSur. J. Bot. Res. Inst. Texas 2(1): 61–74.

–––––––. 2008d. The reinstatement of Capparidastrum.Harvard Pap. Bot. 13(2): 229–236.

–––––––. 2008e. A revision of Colicodendron Mart.(Capparaceae s.s.). J. Bot. Res. Inst. Texas 2(1):75–93.

EICHLER,A.W. 1865. Capparideae. Pages 237–292 inC. F. P. VONMARTIUS, ED., Flora Brasiliensis 13(I).

HALL, J. C. 2008. Systematics of Capparaceae andCleomaceae: an evaluation of the generic delimita-tions of Capparis and Cleome using plastid DNAsequence data. Botany 86: 682–696.

HALL, J. C., K. J. SYTSMA, AND H. H. ILTIS. 2002.Phylogeny of Capparaceae and Brassicaceae basedon chloroplast sequence data. Amer. J. Bot. 89:1826–1842.

ILTIS, H. H., AND X. CORNEJO. 2007a. Studies in theCapparaceae XXX: Capparicordis, a new genusfrom the neotropics. Brittonia 59(3): 246–254.

–––––––. 2007b. Studies in the Capparaceae XXXII.A new combination in Quadrella. Novon 17:452–453.