Selbyana 11: 26-34 REVISION OF SUBG. RAVNIA (: CINCHONIOIDEAE)

CHARLOTTE M. TAYLOR Department of Biology, University of Puerto Rico, Rio Piedras, Puerto Rico 00931

ABsTRACT. The characteristics used previously to separate Ravnia Oersted from Hillia Jacq. can now be considered mistaken or inconsistent. The subgenus Ravnia (Oersted) c. M. Taylor is erected in Hillia for three , H. triflora (Oersted) C. M. Taylor, H. allenii C. M. Taylor (nom. nov.) and H. longiftl­ amentosa (Steyerm.) C. M. Taylor, and a new variety, H. triflora var. pittieri (Standley) C. M. Taylor, from Central America.

Oersted (1852) erected Ravnia for a distinctive ment was made by Standley, although he had epiphytic shrub he found during his 1846-1848 seen more specimens (Standley, 1921) and first explorations of Costa Rica. In addition to its described the fruits (Standley, 1938). Steyermark succulent leaves and stems, R. triflora Oersted (1952) suggested that Ravnia was closely allied has showy bright red flowers 3-7 cm long with to Coutarea Aublet, but presented only the a curiously inflated corolla tube, which is swollen exserted stamens ofR. longifilamentosa Steyerm. in the middle portion but constricted at either as evidence for this. Ravnia has not been men­ end (FIGURE la, b). These flowers are similar to tioned in subsequent discussions of subfamilial those of some Gesneriaceae, such as species of classification in the Rubiaceae, although both Besleria, and in general aspect of Ravnia Verdcourt (1958) and Bremekamp (1966) have are easily mistaken for gesneriads. Oersted did suggested that Hillia deserves it own based not see them, but the fruits of Ravnia are cylin­ on its plumose seeds. drical septicidal capsules with numerous small Determination of the relationships of Ravnia winged seeds, each bearing a tuft of long brown within the Rubiaceae has been complicated by trichomes at one end (FIGURE 1t). Oersted named a poor understanding of the itself. Al­ the to honor the Danish naturalist Peter though additional species have been published Ravn (1783-1839), who worked for the Botan­ (Standley, 1921; Steyermark, 1952), most tax­ ical Museum in Copenhagen and later collected onomists who have turned their attention to Rav­ specimens in Denmark's Caribbean possessions nia have finally combined all of the published (now the U.S. Virgin Islands) (Oersted, 1852; names, either explicitly (Dwyer, 1980) or in prac­ Vegter, 1983). Several species of Ravnia have tice (Standley, 1938: 1368). The resulting "R. been described since, all from Costa Rica and triflora" has become a variable taxon with a "re­ Panama. markable" (Dwyer, 1980) range ofleaf size, leaf Relatively few flowering specimens and no shape, anisophylly, corolla form and color, and fruiting material of Ravnia were collected in the sizes of floral parts. succeeding 80 years. Consequently published de­ Ravnia received monographic attention only scriptions of Ravnia remained incomplete, and from Standley (1921: 114). He recognized only its relationships within the Rubiaceae obscure. R. triflora and a new species, R. pittieri Standley With his original description Oersted also pre­ from Costa Rica. Later, however (1938: 1368), sented a diagnosis that separated Ravnia from he suggested that his new species might be "only Hillia Jacq., which he thought it strongly resem­ a narrow-leaved variety of R. triflora." Steyer­ bled. Schumann (1891: 51) later suggested in his mark (1952) described two more species from monograph of the world's Rubiaceae that from Panama, R. panamensis Steyerm. and R. lon­ Oersted's description, R. triflora was probably gifilamentosa Steyerm., but did not survey the allied to either Hillia or Ruiz & genus. In the most recent consideration of the Pavon. No elaboration of this suggested place- genus, Dwyer (1980) combined all of the species

---+ FIGURE 1. a, Hillia triflora var. pittieri, habit with flower; from Nee 9733 (F). b, H. triflora var. triflora, habit with flowers; from Gomez 2230 (F). c, stipule; from Burger & Gentry 8569 (F). d, leaf of H. triflora var. triflora, showing narrow form; from Taylor & Skotak 4708 (DUKE). e, two capsules of H. triflora var. triflora; from Croat 46855 (MO). f, seed of H. triflora var. triflora; from Taylor & Skotak 4708 (DUKE). g, flower of H. longifilamentosa; from Allen 4744 (F). h, flower of H. allenii; from Allen 3558 (F). a-e, g to same scale.

26 1989] TAYLOR: HILLIA REVISION 27

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d 28 SELBYANA [Volume 11

that have been described since 1852 with R. tri­ capitate in Hillia, in contrast to bilobed in Rav­ flora. nia, but some species described in Hillia have This study is a revision of Ravnia throughout bilobed styles (Steyermark, 1974: 165-178). All its range. Newly presented here are the extension authors have agreed that the calyx of Ravnia is of the geographic range, northward to southern persistent. Wernham (1916) contrasted this with Mexico and south to Colombia; expanded de­ the deciduous calyx of Hillia. However, Standley scriptions of all taxa; the revival ofSteyermark's (1938) used the persistent calyx of Hillia to sep­ two Panamanian species; the recognition of R. arate this genus from Cosmibuena. Calyx per­ pittieri Standley as a variety (but not a consis­ sistence therefore does not appear to be a con­ tently narrow-leaved one) of Hillia triflora; and sistent characteristic of Hillia. the transfer of all members of the genus to a The corollas of Hillia are white or green, fun­ newly-erected subgenus of Hillia, with the ap­ nelform to salverform, and often sweetly scented. propriate new combinations. A list of the spec­ These flowers are apparently adapted to moths imens examined, with determinations, is ap­ or other night-flying pollinators. In contrast, pended. Ravnia has bright red to dull pink, apparently This study is based on examination of her­ odorless corollas with a funne1form or tubular barium specimens from A, DUKE, F, GH, MO, NY, shape, which are apparently adapted for polli­ and SEL. Plants of Hillia are fleshy and shrinkage nation by hummingbirds. The corollas of both of all parts except woody stems accompanies the Hillia and Ravnia have a similar funnelform preparation of dried specimens. Calculations shape, and can be separated only by color. (The based on the dimensions ofthe fresh plants noted unique, swollen corollas of R. triflora are here on the labels of some specimens place this regarded as a derived character within these three shrinkage at about 8-10%. Measurements of soft red-flowered species.) Corolla color, then, is the parts given here are taken from dried specimens, only characteristic that remains to separate Rav­ and should be adjusted accordingly for fresh ma­ nia from Hillia. terial. This distinction between two closely related groups does not seem to merit recognition at the THE GENERIC PLACEMENT OF RA VNIA generic level. Other genera of Rubiaceae contain species that are apparently adapted to pollination Steyermark's (1952) suggestion that Ravnia is by different agents, with the corresponding vari­ allied to Coutarea is supported only by the fun­ ation in corolla color (for example Crusea Cham. nelform shape of the corolla in R. panamensis & Schldl., Hamelia Jacq., Isertia Schreber, and and R. longifilamentosa, and the six-merous Faramea Aublet). This variation does not weak­ flowers of some members of both genera. Cou­ en their identity. Therefore, Ravnia is here com­ tarea is a genus of terrestrial shrubs with small bined with Hillia. However, the Ravnia group stipules and zygomorphic flowers, and further of species shares corolla color and a relatively differs from Ravnia and Hillia in its glabrous restricted geographic range; these data may be seeds. As Schumann suggested (1891), Cosmi­ interpreted to indicate that these species are most buena is similar vegetatively and florally to species closely related to each other. Thus Ravnia is re­ of Ravnia and Hillia, but likewise has glabrous tained at the subgeneric level. The appropriate seeds. Hillia is the only genus in the Rubiaceae new combinations are made below. other than Ravnia with plumose seeds. In ad­ dition, Hillia shares floral and vegetative fea­ MORPHOWGY AND GEOGRAPHIC tures, except flower color, with Ravnia. DISTRIBUTION OF HILLIA SUBG. RA VNIA Without mature flowers, Hillia is difficult to separate from Ravnia. Previous authors have at­ The three species of Hillia subg. Ravnia C. M. tempted to distinguish them by the aestivation Taylor are succulent, usually epiphytic shrubs of the corolla lobes (Standley, 1921, 1938); the with relatively few, straggling, limber branches form of the stipules (Schumann, 1891); the form that may extend to I m or more in length. The of the style (Wernham, 1916); the degree of calyx stems are terete with smooth brown bark. Al­ persistence (Wernham, 1916); and the corolla though the plants are usually epiphytic, they are color and form (Dwyer, 1980). Corolla aestiva­ occasionally encountered in terrestrial sites, usu­ tion is generally described as convolute in Hillia ally in deep leaf mold or rotting wood. These in contrast to imbricate in Ravnia. However, all may be originally epiphytic plants that persist specimens of Ravnia examined have convolute after falling to the ground (F. Putz, pers. comm.). corolla lobes. The stipules have been described These species are found in wet forest from about as bilobed in Ravnia, in contrast to entire in 100 to 2,400 m elevation, from southern Mexico Hillia, but only entire stipules were seen in the to northern Colombia (FIGURE 2). Hillia triflora specimens examined. The style is supposedly (Oersted) c. M. Taylor is common both in ma- 1989] TAYLOR: HILLIA REVISION 29

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15

FiGURE 2. a, distribution of Hillia triflora var. triflora (circles) and H. allenii (square). b, distribution of H. triflora var. pittieri (circles) and H. longifilamentosa (triangles).

ture forest and in remnant woodlots and pasture the stamens alternate and adnate to the lobes. edges throughout this elevational range; the other The free portion of the calyx is divided to the two species, H. longifilamentosa (Steyerm.) C. base into triangular lobes, sometimes with a set M. Taylor and H. allenii C. M. Taylor, are known ofreduced lobes alternating with the five to seven only from remnant montane forests at about larger lobes. It was this set of reduced lobes that 1,000 to 1,600 m elevation. Standley used to distinguish "Ravnia pittieri" The leaves ofthese plants are simple, opposite, from Hillia trijlora, but such supernumerary lobes subsessile to shortly petiolate, and succulent to are found frequently in all species and are not coriaceous. The lingulate, large, interpetiolar correlated with any other feature. The corolla stipules are membranaceous, usually markedly tube may be slightly to strongly funnelform, or paler than the foliage, and very quickly decidu­ tubular and usually inflated in the middle por­ ous. The leaf blades are elliptic to lanceolate or tions while relatively constricted at the base and ovate, entire, and usually acuminate. The leaf apex (FIGURE la, b). The corolla lobes are equal blades of Hillia trif/ora vary widely in shape, in number to the calyx lobes, and are convolute from extremely narrow to quite broad, but the in aestivation, not imbricate as previously re­ leaves of any individual plant are generally sim­ ported (Standley, 1921, 1938; Schumann, 1891; ilar. Many individuals ofthis species show a pro­ Dwyer, 1980). At anthesis the corolla lobes are nounced anisophylly on flowering branches, al­ strongly reflexed. The stamens are exserted in though adjacent sterile shoots may be virtually one species, H. longifilamentosa, but partially isophyllous. The unequal leaves are usually sim­ included in the other two species. The stigma is ilar in width, so that the shorter leaf is propor­ bilobed and borne on a style approximately equal tionately broader than the larger leaf at the same in length to the corolla tube or, in H. longifila­ node. Isophyllous plants without flowers are dif­ mentosa, to the filaments. ficult to assign to species. The mature "cigar-shaped " (Dwyer, The flowers are borne terminally, either singly 1980) is chartaceous, brown, and septicidal, re­ or, in some individuals of Hillia trijlora, in di­ leasing numerous small, flattened, narrowly chasia. The terminal (central) flower of a group rhomboidal seeds each surrounded by a marginal of three is always slightly larger than the sub­ wing and with a tuft of long brown trichomes at tending flowers in its pedicel, hypanthium, calyx, one end. The old reflexed valves of the capsule and corolla, while the two subtending flowers are persist on the pedicels. The capsules and seeds similar to each other in size. The flowers are five­ of the different species are difficult to distinguish. to seven-merous but usually six-parted in all Because of the large number of specimens ofHi/­ species. fia trif/ora seen relative to the other two species, As in most Rubiaceae, the ovary is bicarpellate isophyllous fruiting specimens are here placed and inferior, and the corolla forms a tube with provisionally in that species. 30 SELBYANA [Volume 11

TAXONOMY Ravnia triflora Oersted, Vidensk. Meddel. Dansk Naturhist. Foren. Kjobenhavn 1852: 50. 1852. Key to the Subgenera of Hillia TYPE: Costa Rica, Cartago: near Cartago and Candelaria, 1,800-2,400 m, Oersted 11696 [c, 1. Corolla green or white, funnelforrn or salverform. lectotype, selected by Standley (1938); F!, MO!, ...... subg. Hillia. NY! photographs]. 1. Corolla red to orange, funnelform or tubular and then usually inflated in the middle...... Hillia triflora is easily separated from other ...... subg. Ravnia. Hillia species by its bright red, tubular, usually inflated corollas. The corolla lobes and some­ Hillia Jacq. subg. Ravnia (Oersted) C. M. Taylor, times also a short apical portion of the tube are comb. nov. strongly reflexed at maturity to form a ring around the mouth of the corolla, and the tube is some­ Ravnia Oersted, Vidensk. Meddel. Dansk N aturhist. times curved. Foren. Kjobenhavn 1852: 49.1852. TYPE: Rav­ This species is generally characterized as a plant nia triflora Oersted. of higher elevations (Standley, 1938; Dwyer, Succulent epiphytic shrubs. Leaves opposite, 1980) but it is not uncommon at the La Selva simple, entire, sometimes anisophyllous; stipules Biological Station in Costa Rica, at 100 m ele­ membranaceous, lingulate, deciduous. Flowers vation, and should be expected in similar areas. terminal, solitary or in three-flowered dichasia A wide range of leaf form is found in both bracts reduced or lacking. Calyx lobes 5-7, re~ varieties of H. triflora. In particular, the leaves mote, triangular, sometimes with smaller lobes may vary in proportionate width from very nar­ interposed. Corolla funnelform, or tubular and row (5-8 times as long as wide) to relatively wide then usually inflated in the middle, red to orange, (to 1.7 times as long as wide). This variation in with lobes 5-7, convolute. Stamens 5-7, the an­ leaf form shows no correlation with any other thers oblong to linear, longitudinally dehiscent. characteristic, nor any discernible pattern of geo­ graphic distribution. Thus, in spite of its visual S~yle equalling or exceeding corolla tube, stigma bllobed; ovules numerous, ascending. Fruit a impact, this character seems to have no taxo­ septicidal capsule; seeds flattened, with a mem­ nomic significance. Standley originally separated branaceous, continuous marginal wing and a sin­ ~is "Ravnia pittieri" from H. triflora by, in part, gle tuft of trichomes at the micropylar end. Its extremely narrow leaves (Standley, 1921). Three species; in wet forests at 100-2,400 m These narrow leaves are well within the range of elevation; southern Mexico to northern Colom­ variation observed in typical H. triflora, how­ bia. ever, and for that reason Standley'S species was combined with H. triflora by later workers (Stan­ dley, 1938; Dwyer, 1980). Here, "R. pittieri" is recognized as a variety based on other charac­ Key to Species and Varieties of teristics, anisophylly and strongly inflated co­ Hillia subg. Ravnia rollas, which are fortuitously present on Stand­ ley's type specimen. 1. Corolla tubular, usually inflated in the middle and constricted at the base and apex. 2. Leaves at each node approximately equal in size on flowering shoots; mature corolla tube mod­ lao H. triflora var. triflora. erately inflated at most, up to 1.6 times as broad at the broadest portion as at the apex...... Succulent, epiphytic or rarely terrestrial shrubs, · ...... 1a. H. triflora var. triflora. glabrous throughout, with slender branches to 1 2. Leaves at each node strongly anisophyllous on m long; bark smooth, brown to grey. Leaves with flowering shoots, the larger leaf at least 1.5 times blades coriaceous, 4.5-16.5 cm long, 1-5.8 cm as long as the shorter leaf; mature corolla tube strongly inflated, 1.8 or more times as broad at wide, nearly isophyllous, narrowly lanceolate to the broadest portion as at the apex...... elliptic or slightly ovate, 1.7-5.1(-8.1) times as · ...... lb. H. triflora var. pittieri. long as wide, obtusely angled to cuneate at the 1. Corolla funnelforrn, broadest at the apex. base, acuminate at the apex with the tips 0.5-2 3. Corolla lobes ca. 0.6 cm long; free portion of cm long, willi secondary veins 3-5 on each side the filaments to 1 mm long. . ... 2. H. allenii. of t~e the costa, acutely angled with it, nearly 3. Corolla lobes 1.3-1.7 em long; free portion of straIght? obscure; petioles stout, 0.1-0.7(-1.2) cm the filaments 9-10 mm long ...... long; stlpules 2.8-4.3 cm long, lanceolate to lin­ · ...... 3. H. longifilamentosa. gulate or oblanceolate, obtusely or acutely angled an~ sometimes also acuminate. Flowers solitary 1. Hillia triflora (Oersted) C. M. Taylor, comb. or III three-flowered dichasia borne on stout pe­ nov. duncles to 0.2cm long, with the terminal flower 1989] TAYLOR: HILLIA REVISION 31 slightly larger than the lateral flowers; bracts usu­ The typical variety of Hillia trijlora is known ally lacking, when present ca. 3 mm long, ovate, from Chiapas in southern Mexico, and from Nic­ acute; pedicels stout, 0.1-0.6 em long. Hypan­ aragua to western Panama. This species might thium 0.2-0.8 em long, obconic. Calyx lobes 5- be expected in Guatemala, El Salvador, and 7 but usually 6, 0.3-1.3 cm long, linear to very Honduras as well. narrowly triangular or oblaneeolate, acute or sometimes rounded, sometimes interposed with 1b. HiHia triftora var. pittieri (Standley) C. M. linear or triangular lobes to 1 mm long. Corolla Taylor, comb. nov. red, membranaceous, tubular or usually inflated in the middle and constricted at the base and Ravnia pittieri Standley, N. Amer. Flora 32: 114. apex, 1.1-1.6 times as broad in the middle as at 1921. TYPE: Costa Rica, Alajuela. Santa Clara, the apex, the tube 4-6.5 cm long, the lobes 5-7 Pittier 13461 (NY!, holotype; F! photograph). but usually 6, 0.2-0.8 cm long, triangular to very Lagenanthus parviflorus Ewan, [Gentianaceae] Mu­ broadly so, acute to rounded, strongly reflexed tisia 4: 5. 1952. TYPE: Panama, Coele. Region at anthesis. Stamens with the filaments inserted N of EI Valle of Anton, 1,000 m, Allen 3601 (MO!, holotype). [Maas, 1980] in the upper portion of the corolla tube, the an­ thers partially exserted, 3-5 mm long, yellow, Similar to the typical variety, but the paired oblong to linear. Style equalling the corolla tube leaves of flowering shoots anisophyllous, the or usually exceeding it by 1-3 mm, stigma lobes blades of the larger leaves 1.5-4(-5.2) times as 1-2 mm long. Capsule 4.5-9.5 cm long, cylin­ long as that of the shorter leaves and usually also drical, brown. Seeds brown, narrowly rhomboi­ proportionately narrower; stipules 2.3-7.1 em dal, 0.8-2.5 mm long, with a membranaceous, long; calyx lobes 0.2-1.1 cm long; corollas with complete marginal wing and a single tuft of dark tubes strongly inflated, 1.8-3.3(-4.3) times as brown trichomes 1.5-3 cm long at one end broad in middle portion as at apex, 3-7 cm long (FIGURE 1b-f). (FIGURE 1a). REPRESENTATIVE SPECIMENS EXAMINED. Mexico. ADDITIONAL SPECIMENS EXAMINED. Costa Rica. CHIAPAS: settlement ofTolimfm 38 km W [sic] ofHuix­ ALAJUELA: La Paz de San Ramon, Brenes 2 (NY), 1,000 t1a on the road to Motozintla, Breedlove & Thorne m, Brenes 4523 (F). CARTAGO: Hacienda La Marina, 31213 (MO); municipio Angel Albino Cozo, slope along Rio San Rafael, Canton de Aguas Zarcas, 450-500 m, Rio Cuztepeques near Finca Cuztepeques, 2,400 ft [77 4 Williams et al. 29105 (F). HEREDIA: Estacion Carrillo, m], Shilom Ton 3853 (MO). Nicaragua. MATAGALPA: de la fila al canon del Rio Sucio, 700-450 m, Chacon W slope of Cerro El Picacho, 13°00'N, 85°55'W, Ste­ & Herrera 1602 (MO). Panama. CocLE: 7 km N of EI vens 22725 (MO). RIVAS: Isla de Ometepe, NW slope Cope, Alto de Calvario, New Works at Aserradero Riv­ of Volciin Maderas, ca. 11°27'N, 85°31 'W, ca. 800- er, 1,700 m, Folsom 2340 (MO), 750-800 m, Folsom 1,000 m, Stevens 6555 (MO). Costa Rica. ALAJUELA: 4934 (MO). COLON: ridge between Rio Piedras and Rio La Palma de San Ramon, 1,250 m, Brenes 4097 (F, Gatun waterbeds, trail from end of Santa Rita Ridge NY), 1,075-1,100 m, Brenes 4974 (F), 1,050-1,100 m, road 5-8 km SW of Cerro Bruja, 9°27'N, 79°36'W, Brenes 5434 (F), 1,050 m, Brenes 5598a (F), without 700-800 m, Sytsma et al. 4291 (MO). DARIEN: N slopes elevation, Brenes 6270 (NY), 21162 (NY). ALAJUE­ of Cerro Pirre, 700-950 m, Mori & Kallunki 5466 (MO, LA-PuNTARENAS BORDER: on and near Continental Di­ NY). PANAMA: 5-10 km NE of Altos de Pacora, Yellow vide ca. 2-5 km E-SE of Monteverde, 100IS'N, 84°46'W, Fever Research Camp, 600 m, Mori & Kallunki 3402 1,580-1,700 m, Burger & Gentry 8661 (F). CARTAGO: (MO). VERAGUAS: vicinity of Santa Fe, slopes of Cerro 8 km S of Tap anti, above Rio Grande de Orosi, 1,550 Tute, 2,500 ft [806 m], Allen 4382 (F). Panama-Colom­ m, Lent 1225 (F, MO, NY). HEREDIA: Finca La Selva, bia border. Alto de Nique, southernmost peak of Cerro the OTS field station at Puerto Viejo de Sarapiqui, 100 Pirremassif, J,300-1,520m, Gentryetal. 28640 (MO). m, Folsom 10044 (DUKE). PUNTARENAS: along Rio Bel­ Colombia. CHoco: N ridge of Alto de Buey E-SW [sic] la Vista NW of Las Alturas, 8°57'N, 82°51'W, 1,450- ofEI Valle, 500-100 m, Gentry & Fallen 17395 CMO). 1,600 m, Davidse 23405 (MO). SAN JOSE: NE of San Jeronimo, below La Palma, Rio La Hondura drainage, PHENOLOGY. Collected in flower and fruit Rio Claro Valley, 1,000-1,200 m, Burger & Burger throughout the year, although more commonly 8694 (F). Panama. CHIRIQui: Boquete region, Cerro in February and August. Horqueta, von Hagen & von Hagen 2168 (F, NY). HABITAT AND DISTRIBUTION. Wet forests, PHENOLOGY. Collected in flower and fruit woodlots, pasture edges and remnant , 100- throughout the year, although more commonly 1,700 m. Costa Rica to northern Colombia in March through May and again in August, No­ (FIGURE 2b). vember, and December. This variety is separated from var. trijlora by HABITAT AND DISTRIBUTION. Wet forests, its more strongly inflated corollas and its mark­ woodlots, pasture edges, and remnant trees, edly anisophyllous leaves on flowering shoots. (100-) 800-2,400 m. Southern Mexico, Nicara­ The leaves may be isophyllous on sterile branch­ gua to western Panama (FIGURE 2a). es of the same plant. A few specimens seen have 32 SELBYANA [Volume 11

strongly inflated corollas but nearly isophyllous Although mature seeds have not been seen, leaves; these are placed in var. pittieri here. this plant appears to be best placed in Hillia subg. Hillia triflora var. pittieri is found almost Ravnia. Vegetatively it is indistinguishable from throughout Costa Rica and Panama, and in the other species of Hillia. The red corollas and the area of Colombia adjacent to Panama. To a large occurrence of this species only in montane Costa extent it is sympatric with the typical variety, but Rica and Panama suggest that it is related to H. in any given area one variety may greatly pre­ triflora. dominate in number and distribution of collec­ Hillia allenii is similar to H. psammophila tions. Busby (pers. comm.) notes that at least in Steyerm. of Venezuela, which can be separated the Monteverde area in Costa Rica, the two va­ from H. allenii by its corollas with tubes 6 cm rieties are not strictly sympatric; the typical va­ long. Hillia allenii is also similar to an as yet riety is found at slightly higher elevations, while undescribed species of Hillia from Costa Rica var. pittieri is restricted to lower elevations. (Taylor & Hammel, in prep.). This latter can be A wide range of leaf form is found in var. recognized by its lack of calyx lobes. Both H. pittieri. Similarly, the broadness of the swollen psammophila and the undescribed new species corolla varies greatly. The variation in these fea­ have pale green corollas. tures shows no correlation with geographic dis­ tribution or any other characteristic. 3. Hillia longifilamentosa (Steyerm.) c. M. Tay­ lor, comb. nov. 2. Hillia allenii C. M. Taylor, nom. nov. Non Ravnia longifilamentosa Steyerm., Ceiba 3(1): 21. Hillia panamensis Standley. 1952. TYPE: Panama, Chiriqui. Upper southern slopes of Quebrada Velo, vicinity of Finca Ler­ Ravnia panamensis Steyerm., Ceiba 3(1): 22. 1952. ida, 1,631 m, Allen 4744 (F!, holotype; NY!, is­ TYPE: Panama, Cocle. North Hills, El Valle de otype). [A further specimen was reported to have Anton, P. H. Allen 3558 (F!, holotype; NY!, iso­ been deposited at MO, but has not been located.] type). [The name of the collector and the num­ ber of the type specimen were omitted from the Succulent, epiphytic, straggling or sometimes original publication, although the locality, date, climbing shrubs to 6 m tall, glabrous throughout; and location of the specimens were presented bark smooth, brown to grey. Leaves with blades in detail, and the specimens were clearly marked coriaceous, 6.5-15.5 em long, 1.8-7.3 em wide, as types. A further specimen was reported to elliptic, 2.6-3.4 times as long as wide, obtusely have been deposited at MO, but has not been angled to cuneate at base, acute to slightly acu­ seen.] minate at the apex, with secondary veins 5-6 on Succulent, epiphytic shrubs ca. 1.5 m tall, gla­ each side of costa, acutely angled with it, nearly brous throughout; bark smooth, brown. Leaves straight, obscure; petioles stout, 0.2-0.8 cm long; with blades coriaceous, 3.8-10.5 cm long, 1.5- stipules 2.2-3.7 cm long, elliptic to oblanceolate, 3.7 em wide, elliptic, 2.5-2.8 times as long as obtusely to acutely angled. Flowers solitary; bracts wide, rounded at base, acute to slightlyacumi­ 2(-3) mm long, triangular, acute; pedicels stout, nate at the apex, with secondary veins 7-9 on 1 mm long. Hypanthium 0.4-1.0 cm long, ob­ each side of costa, acutely angled with it, nearly conic to ellipsoid. Calyx lobes 6, 0.6-1.4 cm long, straight, obscure; petioles stout, 0.1-0.2 cm long; triangular to narrowly so or lingulate to oblan­ stipules 1-1.2 em long, oblong to lingulate (pos­ ceolate, acute, sometimes interposed with linear sibly broken on specimen examined). Flowers or triangular lobes to 1 mm long. Corolla orange­ solitary; peduncles and bracts not seen. Hypan­ red to salmon pink or white flushed with pink, thium ca. 4 mm long, obconic. Calyx lobes 6, ca. membranaceous, funnelform and widely flaring, 6 mm long, lingulate to triangular, acute. Corolla the tube 3.2-4.2 em long, the lobes 6, 1.3-1.7 salmon pink, membranaceous, funnelform and em long, triangular to lingulate, rounded to widely flaring, the tube ca. 2.5 cm long, the lobes somewhat acutely angled, strongly reflexed at an­ 6, ca. 0.6 em long, triangular, acute. Stamens thesis. Stamens with the filaments inserted near with the filaments inserted near the apex of the the apex of the corolla tube, the free portion 9- corolla tube, the free portion to 1 mm long, the 10 mm long, the anthers ca. 5 mm long, well anthers ca. 5 mm long, partially included. Style, exserted. Style approximately equalling the fil­ stigma, and fruit not seen (FIGURE 1h). aments. Capsules to ca. 10 cm long. Seeds not seen (FIGURE Ig). PHENOLOGY. Collected in flower in late June. ADDITIONAL SPECIMENS EXAMINED. Costa Rica. HABITAT AND DISTRIBUTION. Wet forest, at ALAJUELA: region of Zarcero, Austin Smith A24 (F). about 1,000 m elevation. Western Panama HEREDIA: south slope ofVo1can Barba, above San Jose (FIGURE 2a). de la Montaiia, 1,600 m, Hatheway 1499 (NY). Panama. 1989] TAYLOR: HILLIA REVISION 33

CHIRIQui: along road to Fortuna Dam site north of STEYERMARK, J. A. 1952. New Rubiaceae from Pan­ Gualaca 22.7 miles beyond bridge south of Rio Esti, ama. Ceiba 3(1): 18-22. 11.8 miles north of Los Planes de Homito, 10.7 miles --. 1974. Rubiaceae. In T. LASSER, eds., Flora N of junction to tunnel, 1,400 m, Croat 48688 (MO); de Venezuela 9: 1-2070. Instituto Botanico, Ca­ vicinity of Fortuna Dam, along trail from highway down racas, Venezuela. to reservoir, 8°45'N, 82°15'W, 1,100 m, McPherson VEGETER, 1. H. 1983. Collectors. N-R. In F. A. 9134 (MO). STAFLEU, ed., Index herbariorum, Part 2, Reg. Veg. 109: 577-803. PHENOLOGY. Collected in flower in April, July­ VERDCOURT, B. 1958. Remarks on the classification August, and November. of the Rubiaceae. Bull. Jard. Bot. de l'Etat, Bru­ HABITAT AND DISTRIBUTION. Wet forest at xelles 28: 209-290. WERNHAM, H. F. 1916. Tropical American Rubi­ 1,100-1,631 m. Costa Rica (Cordillera Central) aceae VII. J. Bot. (London) 54: 322-334. and western Panama (FIGURE 2b). As in Hillia allen ii, the fruits of H. longifila­ mentosa have not been seen. This species is placed ApPENDIX in Hillia for similar reasons. Specimens of Hillia subg. Ravnia Examined Specimens are listed alphabetically by prin­ ACKNOWLEDGMENTS cipal collector. Determinations are indicated by the numbers in parentheses, which correspond I thank the curators of the following institu­ to the numbers of the species in the text. Aster­ tions, who very kindly and promptly made spec­ isks indicate type collections. All collections cit­ imens available: A, DUKE, F, GR, MO, NY, and SEL; ed in the text are included here, with additional and library staffs of the Institute of Tropical For­ collections of var. not cited estry of the V.S.D.A. Forest Service Southern Hillia triflora triflora in the text. Forest Experiment Station, particularly Mrs. Joanne Feheley, and OfMO for help in obtaining Allen, P. H. 3601* (Ib); 3558* (2); 4382 (lb); 4744* literature. I also thank Denisse Rijo, of the Uni­ (3). versity of Puerto Rico, who assembled the rec­ Almeda, F., Jr. 669 (la); 2037 (Ia). ords of exsiccatae. I am indebted to those who Antonio, T. 2094 (lb); 2809 (la); 4026 (Ib); 4105 (la); contributed through discussions and comments 4139 (la). on the manuscript, notably John Dwyer, Francis Breedlove, D. 31213 (la). Putz, D. Jean Lodge, Bill Busby, Roy Gereau, Brenes, A. M. 2 (lb); 3576 (Ia); 3988b (la); 4097 (la); 4523 (lb); 4974 (la); 5434 (Ia); 5598a (la); 6270 two anonymous reviewers, and in particular Mi­ (la); 13473 (la); 14877a (la); 21162 (Ia). chael Madison, who had begun this project him­ Burger, W. C. 6727 (Ia); 8596 (la); 8661 (Ia); 8694 self at one time and generously contributed his (la); 10794 (Ia). preliminary conclusions and comments. Busby, W. H. 626A (la); 627A (la). Carvajal U., A. 167 (la); 455 (la). Chacon, I. A. 1602 (lb). LITERATURE CITED Churchill, H. W. 5974 (lb). Croat, T. B. 14341 (lb); 25944 (Ib); 26673 (la); 33474 BREMEKAMP, C. E. B. 1966. Remarks on the position, (la); 33496 (la); 33907 (l b); 34048 (1 b); 46855 the delimitation, and the subdivision of the Ru­ (la); 47103 (la); 48688 (3); 48767 (I a); 48852 (la); biaceae. Acta Bot. Neerl. 15: 1-33. 49012 (lb). DWYER, J. D. 1980. 179. Rubiaceae. In R. Davidse, G. 23405 (la); 23754 (la); 28279 (Ia). E. WOODSON, JR. AND R. W. SCHERY, eds., Flora Dressler, R. L. 5019 (I b). of Panama, Part IX. Ann. Missouri Bot. Gard. 67: Dryer, V. J. 780 (la). 1-522. Duke, J. A. 5341 (lb); 14997 (Ib). MAAS, P. J. M. 1980. On the true identity of Lage­ Folsom, J. P. 2340 (Ib); 4361 (lb); 4934 (lb); 6117 nanthus parviflorus Ewan (Gentianaceae). Ann. (la); 10044 (la). Missouri Bot. Gard. 68: 685-686. Foster, R. 1892 (Ib). OERSTED, A. S. 1852. Centralamerikas Rubiaceer. Vi­ Gentry, A. 6971 (lb); 7032A (lb); 17395 (lb); 28640 densk. Meddel. Dansk Naturhist. Foren. Kjo­ (I b). benhavn 1852: 23-61. Gomez P., L. D. 2230 (la); 20197 (la); 20937 (Ia). SCHUMANN, K. 1891. Rubiaceae. In A. ENGLER AND Grayum, M. H. 3227 (Ia). K. PRANTL, eds., Die naturlichen Pflanzenfamilien Haber, W. 419 (Ia); 1158 (la). IV(4): 1-194. von Hagen, C. 2168 (Ia). STANDLEY, P. C. 1921. Rubiaceae, Part 1. N. Amer. Hammel, B. 7475 (la). Flora 32(1): 1-86. Hatheway, W. H. 1499 (3). --. 1938. Rubiaceae. In Flora of Costa Rica. Knapp, S. 3491A (lb). Pub!. Field Mus. Nat. Hist., Bot. Ser. 18: 1-1379. Koptur, S. SK314 (la). 34 SELBYANA [Volume 11

Lankester, C. H. 1293 (Ib). Skutch, A. F. 3203 (la). Lawton, R. O. 1I51 (la). Smith, A. A24 (3); H305 (la); A559 (la); A662 (la); Lent, R. W. 1225 (la); 1740 (la); 1912 (la); 2858 (la). NY810 (la); H897 (la). Liesner, R. L. 876 (lb); 14875 (la); 15000 (la). SoIls R., F. 375 (la); 521 (la). Luteyn, J. L. 3030 (la); 4464 (la). Stevens, W. D. 6555 (la); 22725 (la). McPherson, G. 9134 (3). Stolze, R. G. 1571 (la). Meerow, A. 1055 (la). Stork, H. E. 1367 (la). Molina R., A. 17237 (lb). Sytsma, K. 4291 (lb); 4348 (lb); 4656 (lb). Moreno, P. 7417 (la). Taylor, C. M. 2980 (la); 4708 (lb). Mori, S. 2600 (Ib); 2636 (Ib); 3402 (Ib); 5466 (lb); Taylor, J. 11502 (I b); 17639 (la). 6913 (Ib). Tonduz 7466 (Ia). Murphy, H. 1343 (Ia). Utley, J. 2408 (la); 3659 (la); 4087 (lb). Nee, M. 9733 (lb). Valerio, M. 1217 (la). Oersted, A. S. 11696* (la). White, G. 33 (Ia); 39 (la). Palmer, C. W. 157 (Ia). Wilbur, R. L. 12169 (la); 15802 (la); 18628 (la); 18763 Pittier, H. 13461 * (lb). (la); 18868 (la). Raven, P. H. 21800 (la); 21907 (la). Williams, L. O. 16379 (la); 29105 (lb). Shilom Ton, A. 3853 (la). Woodson, R. E., Jr. 559 (la).