A OF COLICODENDRON REVISION (CAPPARACEAE) Hugh XavierCornejo H. litis New The York Botanical Garden Department of Botany and 200th Una Kazimiroff of 430 St. 'Ave. University Wisconsin, New 0458-5 Bronx, York 126, U.S.A. Madison, Wisconsin 53706, t 1 ornejo@nybg.org, xcornejoguay@gmail.com swis@charter.net During our studies of the neotropical species of Capparis (Capparaceae), carried out to produce a generic s.l. realignment extremely polymorphic became of this genus, clear that Colicodendron Mart, a well defined it is South American genus of shrubs and by having stellate trees, characterized a 1-seriate valvate calyx with a dentate or lobed nectary-dish and or nectary-cup, thick-walled, indehiscent amphisarcous fruits. was by Colicodendron established Martius two (1839: 25), for Brazilian species: Colicodendron yco Mart., and the generic type, C. longifolium Mart, (the latter recently placed in Neocalyptrocalyx, as N. [Mart longifolium J & Cornejo H.H. Seeman X. [2008a]). Subsequently, litis (1852: 78) correctly transferred Colicodendron: to Capparis scabrida Kunth, but also the unrelated Capparis Kunth, and avicennifolia Capparis pulcherrima Jacq. nom. (the latter as Colicodendron pulchellum Seem., reject.), as well as the quite unrelated glabrous or simple- haired Capparis Kunth Two subbiloba (=Cynophalla Turczaninow years flexuosa Presl). (1854: [L.] later, J. 327-328) described them synonyms erratically three species in Colicodendron, all of of unrelated species of Capparis a contribution that obfuscated any concept s.l, clear of that genus. was Colicodendron subgenus treated as a of Capparis L. (Eichler 1865: 272), and later as a section of s.l. & Hoffmann Capparis (Pax 1936: 181). In both mentioned infrageneric taxa, the members of Colicodendron were mixed up in heterogeneous assemblage with Mexican Kunth a the Capparis angustifolia (= Quadrella &X. [Kunth] H.H. and South American Kunth angustifolia Cornejo), the and litis C. crotonoides tweediana C. & members Eichler (both H.H. of Capparicordis X. Cornejo, a recently described neotropical litis stellate & genus, Cornejo Most Hutchinson 2007). recently attempted litis (1967: 309), in his generic realignment made of Capparis seven additional, but inappropriate combinations produce s.l, all to a poorly defined Colicodendron. In work, The this a revision of Colicodendron presented. genus validated and amended, mor- is is its phological and limits are defined, the relationships with the morphologically closely related genera are discussed. Journal of the Botanical Research Institute of Texas 2(1) 76 & emend. Cornejo H.H. Colicodendron Mart., Flora 22, Bleibl. 1:25. 1839, X. litis. Type: Colicodendron yco Mart. by trichomes throughout. Leaves simple, short-petiolate Evergreen shrubs or covered stellate to echinate trees, subterminal and/or panicle, whorled. Inflorescences terminal, axilar, a opposite spirally or to sessile, to bracts filiform or Colicodendron raceme or racemose spike, rarely a solitary axillary flower (in yco)\ floral subulate, deciduous. Flowers with calyx 1-seriate, valvate, ± cup-shaped, the 4 sepals free, ± equal, distinct bud from margins and touching each other early until and enclosing the corolla in bud, their tips entirely cup forming hypantium, coated by 4-dentate nectary dish or nectary or near anthesis, the base fused a a at bud, Stamens 14 the filaments 34obed nectary cup within. Petals imbricate or torsivus in sessile. to 60, a 4, on an expanded androgynophore, arranged bud, anthesis exserted, inserted flat geniculate or spirally in at woody, thick-walled pseudoamphisarcous amphisarcous scabridum), or (in C. anthers basifixed. Fruits mm yellow-embryoned 1-2 cm, cotyledons 1-1.5 wide, and bearing white- or seeds, ca. multilocular, ca. crustaceous. convolute, testa from Due pubescence, valvate calyx with the lobes covering the corolla early the 1-seriate to stellate the bud until near anthesis, and the presence of a nectary cup or nectary dish, Colicodendron is related to & However, H.H. Cornejo. Colicodendron differs described neotropical genus Capparicordis X. recently litis from by having oblong oblanceolate, pinnately nerved leaf blades (vs. cor- the lanceolate, to latter elliptic, and subpalmatly-nerved 4-dentate nectary dish or reniform suborbicular basally leaf blades); a date or to number appendages); higher 3-lobed nectary cup nectary cup with four filiform to triangular-filiform a (vs. amphisarcous with an of stamens, 14 to 60 4 to and larger indehiscent thick-walled fruits, inflexible (vs. 8); cm and with thinly woody 1-2 smaller dehiscent a soft flexible, pericarp bearing larger seeds, ca. (vs. fruits, mm, 5-7 by dispersed which and maturity, bearing smaller seeds, ca. coriaceous pericarp, splits falls off at South American genus Calanthea (DC.) the Cornejo 2002). Colicodendron also related to birds, pers. obs., is with Capparis pulcherrima Miers, as was correctly stated by Martius in the original publication (p. 27, cf. the type of Calanthea [DC] Miers). However, Calanthea mainly differs from Colicodendron by the very Jacq., widely spaced calyx with open aestivation, in which the not valvate linear-ligulate sepals are distinctive from very young bud; and by the seeds with barely folded, from each exposing a valvate corolla the other, mm & 4-7 wide (Cornejo 2008b). very thick cotyledons, litis American-West Indian genus Qua- Central Colicodendron could be also related to the stellate to peltate, hypanthium (DC) However, Colicodendron differs from the latter by having flowers with a coated Presl. drella J. and cup without hypanthium, but with a receptacle four scales by nectary dish or nectary flowers flat a (vs. woody amphisarcous on and globose thick-walled fruits indehiscent, ellipsoid inserted in Quadrella)- to it oblong and few times ovoid, with seeds distinctively with seeds capsular, dehiscent, linear or to larger (vs. row smaller, usually arranged in a single in Quadrella). was known by two widely disjunct endemic species: Colicodendron yco from previously Colicodendron and Ecuador dry western adjacent and scabridum (Kunth) Seem., restricted to the forests of eastern Brazil, C. American endemic both with whorled subsessile leaves Peru. In this work, two additional South species, added and genus. beautiful inflorescences are to this & bahianum nov Colicodendron Cornejo H.H. (Figs. brazil. Amiga rodovia X. sp. 1, 2, 7). Type: Bahia: 1. litis, and Cornejo Revision of Colicodendron litis, Journal of the Botanical Research istitute of Texas 2(1) 78 13-40 3-7(-10) cm, coriaceous narrowly x with blades narrowly oblanceolate to elliptic, stiff, subsessile, sometimes and shortly and acute apiculate chartaceous attenuate revolute the base, to at (dry), (alive), and midvein, nerved glabrous rostrate at the apex, shiny green (alive) with an impressed to often sulcate midvein beneath, prominent and with glabrous or echinate-stellate green strongly above, paler a (alive) on with 25 major upward-arching nerves each side of the margin inconspicuosly thickened, 15 to lateral wood, and subterminal racemes, sometimes cauliflorous in older densely terminal midvein. Inflorescences 3-7 same 1-5 cm, bearing 25 flowers the time. Floral bracts linear, axis short, stout, to at stellate-echinate; mm, 1.8-4 cm, the Sepals oblong-elliptic or deciduous, densely echinate-stellate. Pedicels articulate to axis. 12-22 6-8 mm, mainly toward the base, ± acute apex, yellow x abaxially keeled, at lanceolate, ellipsoid to mm 4-6 by hypanthium brown deep, coated a subtet- (when rusty dry), stellate-echinate abaxially; (alive), 2.5-4 x ragonal 4-dentate nectary dish within, nectary appendages broadly deltoid to semiorbicular, 1.5 oblong mm, bud, apparently inserted the edge of the nectary cup, to glandular, Petals torsivae in at stellate. mm, yellow 20-28 7-12 widely obtuse rounded apex, longitudinally nerved, sulfur obovate-oblong, x to at (when on both Stamens 30 rusty-brown sides. to and erect anthesis (Mori, pers. com.), dull dry), stellate at expanded on tomentose torus- 2-3 cm, geniculate in bud, glabrous, inserted a white-stellate- filaments 42, 5-6 mm, Gynophore 1.6-2 cm, glabrous; ovary oblong, white androgynophore, anthers 2 (alive). like ca. mm, with gynophores x 2-3 longitudinally sulcate, glabrous; stigma sessile, capitate. Infructescences ca. 2-4 7-12 cm, subpendulous; Amphisarcous x 2-2.5 x 0.3-0.5 cm, pedicels 3-3.5 cm. oblong-ellipsoid, ca. mm mm 3-7 mature seeds 1.3-17 in diam. wall thick, fruit Colicodendrum bahianum somewhat related to the equally yellow-flowered C. yco, is allopatric in the is more mesophytic bahianum can be distinguished However, easily dry Caatinga the north the C. to (Fig. 7). end branches 1A, sub- by dense whorls arranged toward the of the terminal (Fig. vs. of subsessile leaves, The Colicodendrum more inflorescences of and spaced out along the branches). opposite pedicellate leaves, racemes and paniculate inflorescences, or rarely solitary axillary bahianum are short dense larger longer, (vs. number stamens with sepals anthesis, higher of (30 flowers in yco\ bearing flowers erect reflexed) at C. (vs. mm cm mm), gynophores 4 glabrous (1.6-2 to 42 vs. 14 to 20 in C. yco), smaller anthers (ca. 2 vs. shorter and pubescent) 3.5-5 cm, pubescent) glabrous ovaries. vs. (vs. more Due bahianum seems morphologically closely related whorled Colicodendron the subsessile leaves, to & Bunting H.H. Ruiz differences see G.S. (for herein described Colicodendron valerabellum T. to the also litis, comment under the species). latter November and and August. Phenology.— Collected in flower in February, August, January, fruit in July, hummingbirds The by hermit (Webster 25010). sulfur yellow flowers are visited — Unknown. Cytology. moist wet evergreen Atlantic southern extension of the or Distribution.—Eastern Brazil, restricted to the mm) 2000 Bahia southeastern rain forests into State (Fig. 7). (ca. — wood human and from both har- under pressure agriculture Habitat intense Conservation concern. is comm, Sep bahianum Mori, 1979; vesting, especially for so decorative species Colicodendrum (Scott per. "It ornamental Images potential species are was one of the most beautiful species that I've ever seen"). of this New Web (NYBG, Garden York 2003). Botanical available the site at & 2580 UB, WIS). BRAZIL. Bahia: Mun. Santa Cruz Cabralia, mata costeira, 15 Jul 1966, R.P. Belem R.S. Pinhtiro (F, Est. Paratypes. (CEPEC, DAV, Porto Seguro, Parq. Naca. Monte Pascoal, 15 Jan 1973 T.S. Santos 2692 (CEPEC, NY, WIS); ibidem, trail to WIS)'. (fl.), W (CEPEC, 1971 Santos 1329 Jan T.S. (fl), NY al 2283 (CEPEC, [NY photo in tt hi et (CEPEC, WIS). Pinheiro2202 Enti V- 73, R.S. \A fill "M C [CEPEC photo and fragm. WIS]). at Cornejo and Revision of Colicodendron litis, Itabuno, prominent town CEPEC for a in the Cacao region, where the herbarium mata is located], Cascalheira, pluvial, 13 1 Colicodendron scabridum 2. (Kunth) Seem., Herald Bot. Voy. 78. 1852. (Fig. 7). Basionym: Capparisscc B-W NY!]; 10055-01 [B-W at isolectotype: photo 9478 0! WIS!]). at & & DC, Capparis angulata Ruiz ex Pav. Prodr. 1:253. 1824. Colicodendron angulatum (Ruiz Hutch., Gen. Pav.) 2:309. Fl. PI. 15 PERU: Type: Piura, Paita, Voyage de la Bonite, Jul 1836 M. Gaudichaud B (fl, fr), s.n. (lectotype, designated here: 10 0242739!; isolec- B totypes: 10 0242737! F F G ["misit 1841"]; 876547!, 894168!, F [fragm.] 609130!, [G photo 8472 WIS!], at P!). Nomenclatural note.—The herein selected lectotype of Capparis housed scabrida has with at P, a label a correct origin of the specimen, "in America equatoriale," and the number "19" added. The B isolectotype Cap- of paris scabrida consist of a branch with a terminal inflorescence, with the two one and leaves, inflorescence, floral pieces detached and remounted. That isolectotype, originally only was named identified as Capparis, Capparis Willdenow's nomen non speciosa nudum, (a Capparis = speciosa Griseb. Anisocapparis [1879], speciosa & [Griseb.] X. Cornejo H.H. which litis), as reads the label is at the upper left corner of the photo 9478, now (B-W in a separate sheet 10055 -00 That has label the following handwritten information: 0). "Polyandria subtus glabris tomentosis foliola elliptica. Habitat in America meridionali." The discussed isolectotype later was by correctly identified Eichler himself, in handwritten which a label of determination, reads: "Capparis (Colicodedron) scabrida H.B.K." m cm Multi-branched shrubs and to trees, to 10 25 dbh, low and widely erect to spreading; stem the when gum injured produces a (Litde 6725); terminal branches complanate ± or tetragonal, densely stellate throughout. Leaves mm, few spirally-alternate, times 8-30 opposite-decussate, petioles lacking pulvini; blade coriaceous when (rigidly coriaceous (6-)8-23 2-7 dry), lanceolate oblong, x cm, to acute rounded, to sometimes emarginate, apiculate apex, cuneate rounded, sometimes at to retuse green base, at olive (alive), but drying to a bright sulfur-yellow herbarium when in material, thinly stellate-tomentose young, soon and smooth glabrescent or scabrous above, pale grayish or "ashen" greenish-yellow and densely pale stellate-tomentose with prominent midvein a beneath; (7-)12-23 on nerves each lateral side of the midvein. Inflorescences simple, densely corymbose racemes, and terminal solitary or lateral in the axils of leaves, or branched, compounded terminal racemes complex, into corymbose stout, (sub)erect, panicles, 30 cm, to densely stellate, each raceme with only 1-4 open flowers end one 5-12 at its at time. Floral bracts filiform, mm, soon deciduous. Pedicels 2.5-4 cm, densely brown and stellate sparsely dendroid. Flowers dark brown greenish-brown, to densely Flowers asymmetrical, stellate. in perfect flowers the androgynophore-gynophore off-centered within the calyx cup, arching out and upward. Sepal lobes broadly 10-15 ovate to lanceolate, x mm, 5-8 at anthesis cucullate-ascending, acute the apex, densely brown on both at stellate sides, eventually caducous, broadly inserted on the rim of a wide bowl-shaped calyx cup, 9-13 x 6-8 mm, with 3-lobed a nectary cup hypanthium coating the within. narrowly Petals broadly oblong 15-23 to ovate, or x elliptic, mm, 10-15 and reflexed outrolled at the tip at anthesis, widely cuneate to truncate at the sessile base, obtuse rounded to at the apex, white to cream or yellowish, sometimes greenish, on both Stamens stellate sides. mm 24 borne on to 35, in a ring top of the short 2-3 androgynophore, filaments (3.5-)5-7(-8) cm, stellate mm and dendroid, adnate each other 0.5-2 3-4 mm, to for the base; anthers white Gynophore at (alive). 5-8 cm, creamish-white, maturing purple 4-6 to sparsely ovary ovoid 3-5 (alive), stellate; to ellipsoid, x mm, densely stigma stellate; truncate, Infructescences with gynophores 7-10 sessile. x 0.4-0.6 cm, pedicels 2.5-3.5(-4) cm. Pseudoamphisarcum oblongoid, ovoid, or 8-15 5-9 asymmetrical, ellipsoid, slightly x Research of Texas Journal of the Botanical Institute 2(1) 80 when mature exhibiting 8 longitudinal ± sulcate umbonate apex, densely cm, pendulous, ± the stellate, at mm 6-10 pulp bright orange wall thick, 4 major, indicating the valves); fruit fibrous, (dried) lines (usually cm, 1-2 x 1-1.5 sur- subspherical-reniform, often strongly beveled, ca. seeds 15 maturity, insipid; to 60, at many reddish-brown by from the crustaceous testa; rounded by orange hairs densely sarcotesta infiltrated a embryo yellow. woody subwoody with fruit-wall, The amphisarcum indehiscent provided a thick, to a simple, fruit, is maturity 1994: upon removal and never in valves after (Spjut which shape of sarcocarp, splits the retains decompose and on ground eventually maturity the to In Colicodendron scabridum, the fruits at fall 23, 37). convex, unattached, persistent thickly-subwoody, abaxially the seeds, leaving usually four elliptical, release known which any the types (Spjut doesn't in of fruit That type Colicodendron scabridum, of fit valves. fruit which amphisarcum) Cornejo (mod. nov), Pseudoamphisarcum (pseudo + X. proposed herein as 1994), is campo Van den Eynden ECUADOR: names.— sapote de (Spanish, et Sapote (Spanish, 6725), Local Little & gomoso Steyermark (Spanish, 54842)] sapote 1999: sapote de perro (Spanish, Cornejo Bonifaz 7583), 42), al. Madsen 8521 64103). Zapote de perro (Spanish, Acosta-Solis Mille 1940, Brandbyge Zapotillo (Spanish, 42789); & Zapote (Spanish, Woytkowski de perro (Spanish, 5672). PERU: Diaz Baldeon sapote Sapote (Spanish, 2374), & Simpson Schunke (Spanish, 567). Vargas Zapotillo 42), The mature pulp Colicodendron scabridum strongly resembles the color bright orange color of the fruit of known [Bonpl] {Quararibea cordata "Sapote" pulp Ecuador) well edible fruit called of the of a locally (in names and Colicodendron similar derived local of Bombacaceae). This the origin the several for Vischer, is Thomas. This an endemic one names, Pseudalopex sechurae is "Sapote de of the local refers to perro," known has distributional pat- de monte" (Spanish) or "Sechura fox" (English), that a canid as "Perro locally When Ecuador and the food scabridum in the dry coastal areas of Peru. Colicodendron tern similar to that of (Bruning scabridum Colicodendron the Pseudalopex sechurae facultatively eats the fruits of really scarce, is Cornejo 2005). 1985; Southwestern Ecuador, Zapotillo (Spanish), a town located in a dry forest at the Province of Loja in known name name Colicodendron scabridum. owes the local of the locally well to its — September December. mostly Flowering mostly between February October, fruiting to to Phenology. m A 4-6 60 per Within the nectary cup, three produces 20 flowers anthesis night. single tree high, to at By be produced 19:00 each droplet from below of each nectary lobe. 19:30 h, nectar droplets of start to at h., anymore, morning have nectar the nectary cup of nectar. But by the next at 6:00 h, the flowers don't full is The during one night per flower during nectar secreted perhaps due the intense bat activity the night. is to Bahia de Caraquez, The one night only and don't produce any scent (Cornejo field obs. in flowers only. live Ecuador, July 2004). ECUADOR. WIS, unpublished n = Manabi: and 243 ms). Cytology.— litis 8 [!Pazy, litis litis element dry very dry or xerophytic of to Distribution.— Colicodendron scabridum a characteristic floristic is Ecuador from Ecuador and where dune former. This species ranges sea level in western Peru, a areas of is m and Peru south 2500 mountains Province in to from Province Manabi, in the of the Loja, the of to ca. Ancash Dept. (Fig. 7). by scabridum are visited the Colicodendron and observations.—-The flowers of Ecological interactions field when common Dozens buzzing bees can be heard from under a tree honey of domestic bee, Apis mellijera L. and around The honey peaks in the flowers of the nocturnal flowers are just about to open. bee's activity The domestic bees re-visiting the flowers the between 18:30 19:30 re-start Colicodendron scabridum to h. is such presumably pollen predator bees as Halictidae morning, between 6:00 8:00 Nocturnal, next as well to h. and While both bees touch the anthers the ovaries, 19:00 species of the flowers h. (Megaloptal), also visit at many From small bats frequent the 20:00 5:50 seems however that the true pollinators are bats. to h, it The which exposed on long peduncles high above the foliage. bats flowers of Colicodendron scabridum, are from during than one second, wings hovering while licking the nectar each flower less beat their in the air and Cornejo Revision of Colicodendron litis, 81 hummingbirds The as are apt to do. bats do not hold themselves -om the flowers as happens the African to Bignoniaceae, same Kigelia ajricana [Lam.] Benth. (seen in Bahia de Caraquez, Ecuador, the where locality When was by Colicodendron scabridum studied the senior author). leaving the flower, the bat shakes the inflo- presumably rescence, helping pollination. interesting to note that only the individuals of Colicodendron It is growing scabridum darkness by These in total are visited bats. conspicuously have a higher productiv- fruit ity per tree than the remaining individuals located around the nocturnal lights along the roadsides in the same locality. Additionally, the flowers of Colicodendron scabridum are also visited by Amazilia amazilia, a who hummingbird new visits the old (previous night's) flowers between 7:00 to 8:00 h, and the flowers just open about commonly to at ca. 18:00 h. Finally, the flowers of Colicodendron scabridum are visited by wasps, and coleoptera several species of ants (Cornejo field obs. in Bahia de Caraquez, Ecuador, July 2004). — The have been humans and Uses. fruits reported as edible for as medicinal against colds and coughs. commonly They are eaten by and donkeys Ceron Van den Eynden cattle 22424, 1999: (C. et et 42, al. al. & wood make Cornejo while the used and Van field obs.), is for fuel to handicrafts {Cornejo Bonijaz 7583, den Eynden et al. 1999: 42). Finally, a "glue" for paper can be obtained from the cut, gum-oozing ends of the stems (Ceron 11876). — A common Conservation biology. characteristic species in the dry landscape, does not appear be to it danger in of extinction. & m,7Jan IK7 QCNE); 1994, Cornejo (GUAY, X. Bonijaz ibidem, C i ;. QCNE, (MO, 11671 WIS); al 22424 (QAP);JoahaciaMer 420 Sep 111,6 1991, C.Jossc 638 QCNE & ,ca.2m,22 Los Playa Frailes Jan 1991, A. Gentry [2]); C. 1 , ^! 9 ^jr™^ L^7nmZ^Sl 1 Aug-8 Sep 1836 [voyage of H.M.S. Sulphur], G. W. Barclay 422 (BM I Pum H. Eggers 14735 (GH, K, LE, M, US); ibidem, 1852, Andersson 12, Isla s.n. (S); 50 m, 4 Nov J. Madsen 64103 (AAU); path Rio Hondo Cerro Yanzu >9'S, ca. 1987 to V QCA, (AAU[2], QCNE). El Oro: Parroquia San Antonio, Hcda. Montecarlo, 2: 2 Journal of the Botanical Research Institute of Texas 2(1) W3 o o S,5m,7 Apr Holm-Nielsen WI! Huaq t,80 13 27 1980, Anderssc (GB, Lorn:scrub fores , , L. i i); QCNE, W] (MO, QAP, 1876 Loja: I L^^sTS IB^!^^^X eM^ Hl'S V, V^T.'. mT3°F QCA) 5^400 *eenLoja (GB, }] S QCNE); Catama above Van den Eynden 615 (LOJA, 1996, et a\. :b m ndU NY, between mbo and La Toma, 1000-22 >i.o,^ cry dry slope, 1650 9 Oct .955, E. Asplu J077(B,K, LD, S); El Ta: c a. ] , G.HarUng 6065 (MO.QCNE.WIS; (S);Cata >59, Sep 199' etai: 6 a, Little Apr ma,ca. 1500 m, 24 19 3 (GH 54842 Steycrmflrfc 43, J. Gua -651 (NY, WIS[2]); Rio :?H >S L°r229^^^ E? NWof rB il m, 537 Knight 609 0* .400 R. Esplrwzfl CF QCA, QCNE, V 6584 (NY, Valdespino SoalZs^i 10 Jul 1 i, X\^' T^l 1 ckTwkhcaa i^79< 'roads) Quebrada Las Penas, betw. San Juan de la Virgen and Lam- >374 (MO). La Wright 3518 (MO); ison &J. Maranon Celendin-Balsas 25 kn Valley, rd. & Apr Baldedn 2784 (MO); Ce Libcrtad: 15 Prov. Trujillo, Trujillo, 1 iz S. A (MO, band NY); 10965 (MO); ibidem, 100 m, 14 Jul 1973, Araujo 7978 :. *s™^ 7'maZ GH, tm&J. Wright 3352 G, (F, & 2-7). Colicodendron valerabellum H.H. Ruiz Bunting, nov. (Figs. G.S. sp. 3. T. i litis, damaged with one two unbranched stems from near the Evergreen palmoid in plants or additional treelets [or and sometimes flower-bearing some decumbent and sending lateral- base, of these forth 1 to 3, erect, leaf- m cm 3-5 dbh crowded young branches) dark-brown and long often (colonal?); horizontal 5 shoots], to grouped cm, in echinate or echinate-stellate; stipules absent. Leaves subsessile to short-petiolate, to 1.5 nodes crowded whorls the end of the terminal branches exposing mostly leafless internodes, of leaf inser- at 4-9 cm, 3-6 narrowly (16-)20-50 x coriaceous cm, blades oblanceolate stiffly tions swollen, to elliptic, ca. and Cornejo Revision of Colicodendron litis, and hard but somewhat flexible chartaceous attenuate but rounded-retuse subcordate the (alive), (dry), to at base, acuminate or acute to widely obtuse the apex, margin inconspicuosly thickened and at revolute, lustrous green (alive) with a impressed to sulcate midvein, prominently and coarsely rugose with the bullate), (i.e., blade surface between the deeply impressed midrib and main nerves prominently convex and smooth, ap- Journal of the Botanical Research Institute of Texas 2(1) H.H. Ruiz & G.S. Bunting. A. Habit. B. A leaf blade. C. An inflorescence with three hermaphrodite flowers in T. litis, i An male above them bloom and three beginning to wilt; and, above these, some 50 male flower buds. D. postanthesis at base; four flowers in full & & inflorescence bearing (sub)sessil flowerbuds. E-F. Immature fruit. (A. After photo of Bunting Trujillo 13120. B-D. litis etal. 30547, WIS. E-F. Ruiz fane 4764, MY). A-E, ni; by T. Ruiz Z. Villa F,