Randia retroflexa (Rubiaceae), a New Species from Southern Mexico

Brittonia, 39(3), 1987, pp. 371-375. 9 1987, by the New York Botanical Garden, Bronx, NY 10458-5126

RANDIA RETROFLEXA (RUBIACEAE), A NEW SPECIES FROM SOUTHERN MEXICO

DAVID H. LORENCE ~ AND MICHAEL NEE Lorence, David H. (Herbario Nacional, Instituto de Biologia, U.N.A.M., Apartado Postal 70-233, 04510 Mrxico, D.F., Mexico) and Michael Nee (New York Botanical Garden, Bronx, NY 10458-5126). Randia retroflexa (Rubiaceae), a new species from southern Mexico. Brittonia 39: 371-375. 1987.--A new species of Rubiaceae, Randia retroflexa, is described and illustrated from Veracruz, Oaxaca, and Chiapas in southern Mexico. An armed canopy vine of lowland and lower montane tropical evergreen forest, it is distinguished by its paired reflexed spines and hirsute pubescence. Its distribution, habitat, and specimen citations are given and affinities are discussed.

Randia L. ex Houst. (Rubiaceae, Gardenieae) is a taxonomically complex genus of shrubs, trees, and lianas formerly considered to comprise some 200 to 300 species in the tropics and subtropics of the world. However, recent studies in Gardenieae based on vegetative characters, floral and fruit morphology, anatomy, and palynology have revealed the Paleotropical members to belong to other genera (Fagerlind, 1943; Keay, 1958; Tirvengadum, 1978; Robbrecht & Puff, 1986). Randia is therefore a Neotropical genus, ranging from the extreme southern United States and the Antilles through Mexico, Central America and South America to Bolivia, with approximately 60 to 70 species by our estimate. The genus is highly diversified in Mexico, with 33 species of which 23 are endemic (Lorence & Rodriguez-Acosta, 1986; Lorence & Dwyer, in press), making it Mexico's third largest rubiaceous genus. Systematic studies on the Rubiaceae of Mexico and Central America and investigations for the Flora de Veracruz and Flora Mesoamericana projects have revealed the presence of a distinctive new species of Randia from Veracruz, Oaxaca and Chiapas which is described below. Randia retroflexa Lorence & M. Nee, sp. nov. (Fig. 1) Liana dioica, spinis geminatis reflexis secus virgas dispersis armata; folia, inflorescentia, calyx, corolla extus trichomatibus flavidis patentibus, 1-2 m m longis hirsuta. Petiolus 1-3 cm longus; lamina elliptica vel obovato-elliptica, 7.5-22.5 cm longa, 3-10 cm lata, margine ciliata. Inflorescentia staminata cymoso-corymbosa, candelabriformis; flores 7-20; pedunculus 3-10 mm, ramis 2-3; pedicellus 3-8 turn longus; hypanthium 2-3 m m longum, 1.5-2 m m latum; calycis cupula 2-2.5 m m longa, 3--4 m m lata, lobi 5 (6), lineares subulati vel oblanceolati, 1-8 (12) m m longi, 0.3-1 m m lati, inaequales; corolla alba, tubo 4-5 cm longo extus patule-hirsutus, lobi 5, anguste elliptici vel ovato-elliptici, 1.54 cm longi, 0.5-1 cm lati; stamina inclusa; antherae sessiles, lineares, 1-1.2 cm longae; stylus glaber, inclusus, 2.5-3.5 cm longus. Inflorescentia pistillata subcapitata, subsessilis; fiores 2--6, sessiles; calycis lobi 12 m m usque longi, 4 m m lati; corollae tubus 3-3.5 cm longus; stylus 2-2.5 cm longus. Fructus solitarius terminalis, subgloboso-obovoideus, 7 cm usque longus, 6 cm diam, umbonatus; pericarpium 2-10 m m crassum; semina discoidea, 8-10 m m longa.

Plants armed, dioecious, woody, sometimes at first shrubby with scrambling branches, eventually vines, high-climbing to the canopy, sometimes forming large tangled masses, 8-13 m or more high, the stems to 7 cm in diam, the bark smooth, greenish, the slash pale, creamy, with little odor; twigs opposite, divaricate, to 25 cm long and 2-5 m m in diameter, the bark thin, exfoliating, reddish- or grayishbrown, lenticellate, the new growth golden- or reddish-hirsute with spreading trichomes 1-2 mm long, the stout spines paired, axillary, recurred and reflexed i Present address: Pacific Tropical Botanical Garden, P.O. Box 340, Lawai, Kauai, HI 96765, USA.

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FIG. 1. Randia retroflexa. A. Growth habit. B. Enlargement of stem tip. C. Fruit on leafless stem. D. Mass o f seeds from dried fruit. E. Depauperate staminate inflorescence with two flowers, one in bud, one at anthesis. F. Enlargement of calyx and hypanthium of staminate flower split open to show internal surface. G. Corolla of staminate flower split open to show stamens.

1987]

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or rarely straight, 5-10 mm long, 2-4 m m thick basally, the internodes 1-5 cm long; stipules persistent or eventually deciduous, free or slightly connate basally, broadly triangular to ovate, acuminate, 6-10 mm long, 3-5 m m wide basally, the acumen 1/2the length of or equalling the body, externally densely appressed-hirsute, internally densely white villous, the hairs mixed with slender brown colleters 0.50.7 mm long, the margins scarious. Leaves opposite, those of a pair equal or subequal; petioles (0.3) 1-3 cm long, 1-2 m m thick, densely spreading-hirsute; lamina elliptic to obovate-elliptic, (2) 7.5-22.5 cm long, (1) 3-12.5 cm wide, the apex acute, shortly acuminate or obtuse, the acumen 3-12 m m long, the base narrowly cuneate to obtuse or rounded, the sides often unequal, the costa prominent abaxially, hirsutulous, the secondary veins 6-9 pairs, festoonedbrochidodromous or camptodromous, making a 40-45 ~ angle with the costa, prominent abaxially, the venation visible to tertiary adaxially and to quaternary abaxially, the lamina membranous in shade forms, to slightly fleshy and chartaceous when growing in full sun, slightly discolorous, dark green above, lighter green below, somewhat shiny, drying green, both surfaces appressed-hirsutulous, the trichomes whitish to golden, unicellular, often slightly bulbous basally, denser along the costa and veins, the margin plane, ciliate. Staminate inflorescence terminal, cymose-corymbose, candelabra-like, to 9 cm long and 20 cm wide (including the corollas), (3) 7-20-flowered, terminated by 2-3 short branches each bearing 3-6 flowers, all parts densely spreading-hirsute, the peduncle 3-10 m m long or sessile; staminate flowers with slender pedicels 3-10 m m long, the hypanthium narrowly obconical, 2-3 m m long, 1.5-2 m m wide, the calyx cup 22.5 mm long, 3-4 m m wide, internally sericeous, the lobes 5 (6), filiform, linearsubulate, often unequal, 1-8 (10) m m long, 0.3-1 m m wide; corolla white, the tube 4-5 cm long, 3 m m wide medially, dilated to 4-5 mm wide distally, externally spreading-hirsute, internally glabrous or with scattered hairs distally, the lobes 5, narrowly elliptic or ovate-elliptic, 1.5-4 cm long, 0.5-1 cm wide, acute to acuminate, externally hirsute-ciliate along one side, internally glabrous; stamens 5, included, the anthers sessile, linear, 1-1.2 cm long, affixed 8 m m below the faux; style included, glabrous, 2.5-3.5 cm long including the ellipsoid stigmas 2-3 m m long, the disc fiat, 0.5 m m long. Pistillate inflorescence subcapitate, subsessile, hirsute, the peduncle 3-5 m m long, the flowers 2-6, like the staminate flowers but the hypanthium larger, 3-6 mm long, 2-3 m m wide, the calyx lobes often oblanceolate, foliaceous, to 12 mm long and 4 mm wide, the corolla tube shorter, 3-3.5 cm long, the style shorter, 2-2.5 cm long, the stigma ovoid-capitate, 2 mm long, the ovary unilocular, with 2 intrusive bilobed parietal placentas. Fruit terminal, solitary, subglobose to obovoid-depressed, to 7 cm long and 6 cm in diam, umbonate, the umbo to 10 mm long and 7 m m in diam, the surface at first longhirsute, glabrescent, smooth or slightly reticulate, the pericarp hard, brownish or blackish, 2-10 mm thick; seeds numerous, embedded in a blackish pulp, subcircular-discoid, 8-10 m m in diam, 1.5-3 m m thick, brown. TYPE: MEXICO. VERACRUZ: Municipio de Hidalgotitlfin, Campamento Hermanos Cedillo, a 4 km de Pancho Villa, 150 m, 25 Mar 1975 (fl), B. Orffz & S. J. Martiniano A. 56 (HOLOTYPE:MEXU; ISOTYPES: BM, BR, ENCB, F, MO, NY, UC, XAL). Additional Specimens Examined: MEXICO. CHIAPAS: adelante San Gregorio, E1 Ocote, 2 Abr 1953 (st), Miranda 7733 (MEXU). OAXACA:Mum Sta. Maria Chimalapa, Arroyo Piedra del Tigre, ca 8 km al O de Sta. Maria, 220 m, 8 Sep 1984 (fr), Herndndez G. 417 (CHAPA, MEXU). VERACRUZ:Zapoapan de Cabafias, 1953 (st), Bravo H. s.n. (MEXU); Mun. San Andrrs Tuxtla, Estaci6n de Biologia Tropical "Los Tuxtlas" entre Sontecomapan y Montepio, 150--400 m, 15 Nov 1973 (fr), Calzada 1038 (F, M E X U - - 3 sheets); same locality, 25 Abr 1972 (fl), Cedillo T. 217 (MEXU, NY); same locality, 3 Sep 1970 fir), Herndndez M. & Cedillo T. 745 (F); same locality, 26 Abr 1982 (fl), lbarra M. 89 (MEXU);

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same locality, 4 Jun 1982 (fr), Ibarra M. 154 (MEXU); same locality, 13 Ene 1983 fir), Ibarra M. 388 (MEXU); same locality, 13 Jul 1984 (fr), Ibarra M. 1904 (MEXU); same locality, 20 Ene 1985 (ft), Ibarra M. 2239 (MEXU); same locality, 22 Abr 1985 (fl), lbarra M. 2390 (MEXU); same locality, 21 Mar 1986 (st), Lorence 4957 (MEXU); Mun. Minatitl~n, 13.7 k m a l E de La Laguna, luego 5-7 km al N, 130 m, 29 Mar 1982 (fl), Lorence et al. 3979 (CHAPA, MEXU); Mun. Jesfis Carranza, 13 km al NNO de Poblado 2, 120 m, 30 Mar 1982 (fl), Lorence et al. 3989 (CHAPA, MEXU); Mun. Soteapan, camino de Tebanca a Bastonal, a 3--4 km al SE de Tebanca, 800-850 m, 25 Abr 1982 (fl), Lorence et al. 4135 (MEXU); Mun. Catemaco, Cumbres de Bastonal, a 11 k m a l S de Tebanca, 26 Abr 1982 (fl), Ibarra M. 89 (MEXU); Mun. Catemaco, highest point on road from Catemaco to Sontecomapan, 8 km NE of Calemaco, 500-550 m, 31 Oct 1981 (st), Nee & Calzada 22529 (BM, F, MEXU, MO, NY, XAL); Mun. Catemaco, Playa Escondida, 10 km N of Catemaco, 0-150 m, 31 Oct 1981 (st), Nee 22547 (F, NY, XAL); same locality, same vine, 6 Apr 1983 (tt), Nee 26498 (BM, CAS, CHAPA, ENCB, F, G, GH, MEXU, MICH, MO, NY, US, XAL); Mun. Catemaco, road from Catemaco to Coyame, 6 km NE of Catemaco, 350 m, 20 Apr 1983 (st), Nee et al. 26673 (CAS, CHAPA, F, MEXU, MO, NY, P, TEX, VEN, XAL); 8.2 km N sobre la carretera La Laguna-Rio Grande, luego sobre terraceria a Ejido Belisario Domlnguez, luego al N, 130 m, 16 Jul 1980 fir), Wendt et al. 2598 (CHAPA). Distribution: Known only from southern Mexico in the states of Veracruz, O a x a c a , a n d C h i a p a s (Fig. 2). E c o l o g y : R a n d i a r e t r o f l e x a is a c a n o p y v i n e in l o w l a n d a n d l o w e r m o n t a n e t r o p i c a l r a i n f o r e s t f r o m n e a r sea l e v e l t o a b o u t 850 m e l e v a t i o n . T h e n e w s p e c i e s is i m m e d i a t e l y r e f e r a b l e to R a n d i a sens. str. b e c a u s e it p o s s e s s e s the following combination of characters: unilocular ovary, parietal placentation with two massive, intrusive bilobed placentas, numerous discoid seeds embedded in a b r o w n - b l a c k p u l p , a n d p o l l e n s h e d in p e r m a n e n t t e t r a d s . B e c a u s e o f its large flowers a n d fruits, R a n d i a r e t r o f l e x a s e e m s b e s t p l a c e d in t h e s u b g e n u s B a s a n a c a n t h a ( H o o k . f.) L. O. W i l l i a m s ( W i l l i a m s , 1972), e v e n t h o u g h m o s t o t h e r m e m b e r s o f t h i s g r o u p h a v e straight, t e r m i n a l o r s u b t e r m i n a l s p i n e s a r r a n g e d in g r o u p s o f t h r e e o r four. T h e s p i n e s a r e s t r a i g h t in N e e e t al. 2 6 6 7 3 , h o w e v e r , suggesting

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LORENCE & NEE" RANDIA

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this c h a r a c t e r m a y b e s o m e w h a t v a r i a b l e w i t h i n the species. C e r t a i n l y m u c h m o r e w o r k r e m a i n s to be d o n e o n d e l i m i t i n g s u b g e n e r i c categories i n R a n d i a sens. str. T h e long, s p r e a d i n g h i r s u t e p u b e s c e n c e a n d p e c u l i a r paired, reflexed s p i n e s o f R a n d i a r e t r o f l e x a are d i s t i n c t i v e , a n d t h e species a p p e a r s to h a v e n o close relatives i n the genus. It is o n e o f o n l y three l i a n e s c e n t species o f R a n d i a k n o w n f r o m M e x i c o , a key to w h i c h is p r e s e n t e d below. T h e specific e p i t h e t refers t o the p a i r e d , reflexed spines, as d o e s its v e r n a c u l a r n a m e i n Veracruz, "'ufia de gato'" (cat's claw), w h i c h is also a p p l i e d to v a r i o u s o t h e r p l a n t s a r m e d w i t h c u r v e d spines. K e y to the l i a n e s c e n t species o f R a n d i a i n M e x i c o 1 Leaves strongly bullate; coastal dunes and deciduous or semideciduous dry forest; Guerrero ........................................................................................................................................... R. guerrerensis Lorence & Rodriguez-Acosta 1 Leaves not bullate; lowland or lower montane tropical evergreen forest; Chiapas, Oaxaca, and Veracruz. 2 Spines paired, reflexed, not terminal; leaves hirsutulous on both surfaces; flowers and inflorescence densely spreading-hirsute ........................................................... R. retroflexa Lorence & M. Nee 2 Spines in groups of 3 or 4, ascendant, terminal or subterminal; leaves strigose primarily on costa, veins, and margin; flowers and inflorescence strigillose or strigose ............R. sp. nov.2 Acknowledgments

W e w o u l d like to t h a n k Ms. Sheila S e y b o l t for t h e i l l u s t r a t i o n . Literature Cited

Fagerlind, F. 1943. Die sprossfolge in der Gattung Randia und Ihre Bedentung ffir die Revision der Gattung. Ark. Bot. 30A(7): 1-57. Keay, R. W.J. 1958. Randia and Gardenia in West Africa. Bull. Jard. Bot. l~tat 28: 15-72. Lorence, D. H. & J. D. Dwyer. In press. New taxa and a new name in Mexican and Central American Randia (Rubiaceae, Gardenieae). Bol. Soc. Bot. Mrxico 47. - & M. Rodriguez-Acosta. 1986. Randia guerrerensis, una nueva especie de Rubiaceae de Mrxico. Biotica 11(3): 195-199. Robbrecht, E. & C. Puff. 1986. A survey of the Gardenieae and related tribes (Rubiaceae). Bot. Jahrb. Syst. 108: 63-137. Tirvengadum, D. D. 1978. A synopsis of the Rubiaceae--Gardenieae of Ceylon (Sri Lanka). Bull. Mus. Hist. Nat. (Paris), srr. 3, no. 521, Botanique 35: 3-33. Williams, L. O. 1972. Randias from Central America. Phytologia 24:159-163. 2 Species epithet will be published by Lorence and Dwyer (In press), honoring Mario V~zquez.

BOOKS RECEIVED Biology. By Neil A. Campbell. Benjamin/CummingsPublishingCompany, 2727 Sand Hill Rd., Menlo

Park, CA 94025. ISBN 0-8053-1840-2. 1987. 1101 pp. $39.95 (cloth). Biotcchnology of Plants and Microorganisms. Edited by O. J. Crocomo, W. R. Sharp, D. A. Evans,

J. E. Bravo, F. C. A. Tavares, and E. F. Paddock. Ohio State University Press, 1050 Carmack Rd., Columbus, OH 43210-1002. ISBN 0-8142-0375-2. 1986. 473 pp. $60 (cloth). Leaf Protein and Its By-Products in Human and Animal Nutrition. Second edition. By N. W. Pirie. Cambridge University Press, 32 East 57th Street, New York, NY 10022. ISBN 0-521-33030-0. 1987. 209 pp. $37.50 (cloth). Hendrik Adriaan van Reede tot Drakenstein (1636-1691) and Hortus Malabaricus. A Contribution to the History of Dutch Colonial Botany. By J. Heniger. A. A. Balkema Publishers, P.O. Box 230,

Accord, MA 02018. ISBN 90-6191-681-X. 1986. 295 pp. $75 (cloth).