NEW GENERA AND SPECIES OF ERICACEAE (VACCINIEAE) FROM COSTA RICA AND PANAMA 1 JAMES

L. LUTEYN AND ROBERT L. WILBUR

Luteyn, James L. (The New York Botanical Garden, Bronx, NY 10458) and Robert L. Wilbur (Department of Botany, Duke University, Durham, NC 27706). New genera and species of Erieaceae (Vaccinieae) from Costa Rica and Panama. Brittonia 29" 255-276. 1977.--Two new monotypic genera, Didoniea and Utleya, are described, with D. p e n d u l a from Panama and U. costarieensis from Costa Rica. Disterigma trimera (Panama), D. u t l e y o r u i n (Costa Rica, Panama, Colombia, and Ecuador), Lateropora santafeensis (Panama), Lysielesia p a n a i n e n s i s (Panama), Maeleania talamancensis (Costa Rica), T h e m i s t o e l e s i a costarieensis (Costa Rica) and T. h o r q u e t e n s i s (Panama), V a c c i n i u m costarieense and V. orosiense (both from Costa Rica) and V. jefense (Panama) are all described as new. New combinations are provided for the Panamanian Vaccinium f l o e e o s u m (~Symphysia Jloccosa) and the West Indian V a e c i n i u m r a e e m o s u m (~-~Symphysia racemosa). Keys are provided for the Central American species of Disterigma and Themistoclesia, the species of Lateropora and Lysiclesia, and the Costa Riean and Panamanian species of Vaccinium. Six new species are illustrated.

Recent collections and the preparation of treatments for the Ericaceae for both the Flora of Panama and the Flora Costaricensis have brought to light several new genera and species which we wish to describe now. These are treated below in alphabetical sequence. DIDONICA LUTEYN & WILBUR,gen. nov. Frutex epiphytieus. Folia alterna coriacea petiolata plinervia. Flores axillares solitarii penduli. Pedicellus cum calyce continuus. Calyx 5-1obus. Corolla campanulato-cylindrica, 5 vel 6-1oba. Stamina 10 vel 12; filamenta distincta; connectivum ecalcaratum; loculi antherarum leviter granulates; tubuli antherarum 2 ad basim distincti aliquantum rigidi per rimas ovales latrorsas apicem tubulorum non attingentes dehiscentes.

Didoniea pendula Luteyn & Wilbur, sp. nov. (Fig. 1). Folia laneeolato-elliptica vel ovata, (4.5) 6-9 • (1.5) 2-5 cm, basi rotundata vel obtusa, breviter attenuata, apice acuminata. Pedicelli 15-18 cm longi. Bracteae florales ovatae, 1 mm longae. Calyx 10-12 mm longus, glaber; tubus obconicus, 3.5-5 mm longus; limbus 6.5-7 mm longus lobis inclusis; lobi ovati 2.5 mm longi. Corolla 22 mm longa, glabra. Stamina 12 mm longa; filamenta 4.5 mm longa; antherae 10 mm longae tubulis inclusis; tubuli 3-3.2 mm longi. Stylus 16-18 mm longus, glaber.

Epiphytic shrub, 0.75-1 m tall. Mature stems subterete, smooth or minutely striate, glabrous or puberulent, dark blackish-red when dry. Immature stems and twigs of new growth subterete, striate, glabrous or puberulent, brown or gray when dry. Leaves lance-elliptic or ovate, (4.5) 6-9 • (1.5) 2-5 cm, glabrous, basally rounded or obtuse, short attenuate, apically short or long acuminate, marginally entire, green to olive-green when dry, glandular-fimbriate beneath but fimbriae soon deciduous; 5 (7)-plinerved, midrib impressed above (although thickened and raised proximal 1 We should like to acknowledge our appreciation of the financial support to R.L.W. from NSF Grants 13815 and DEB-76-1085, an NIH Biomedical Sciences Research Grant and the Duke University Research Council which made possible the necessary field work represented here. BRITTONIA 29: 255-276.

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1-1.5 cm) and raised beneath, lateral nerves and veinlets slightly raised on both surfaces; pseudostipules 2 mm long. Petiole subterete, rugose, glabrous, laterally slightly winged, 5-10 mm long and 1.5-2 mm diam, dark reddish-brown when dry. Rachis 1.5 mm long. Pedicel terete, minutely ribbed, glabrous, 15-18 cm long and 0.3-0.5 mm diam. Floral bract ovate, 1 mm long, marginally glandular-fimbriate. Bracteoles located near the base of the pedicel, aristate, 1.5 mm long, marginally glandular-fimbriate. Calyx glabrous, 10-12 mm long; tube obconical, muricate, 3.55 mm long; limb campanulate, conspicuously nerved, 6.5-7 mm long including the lobes; lobes broadly ovate, apiculate, 2.5 X 5 ram; sinus acute. Corolla campanulate-cylindric, glabrous, --- 22 mm long, pink when living (fide Dressler 5170), thin and membranaceous when dry; lobes broadly ovate, 2.5 mm )< 6 ram. Stamens 12 mm long; filaments ciliate, 4.5 mm long, 2 mm broad at the base and narrowing to the connective which itself extends to the base of the tubules; anther including tubules 10 mm long; thecae 6.8-7.5 mm long, minutely granular; tubules slightly spreading, 3-3.2 mm long, pilose on inner surfaces, dehiscing by oval, latrorse clefts 1.5-2 mm long but not extending to the apex. Style glabrous, 16-18 mm long; stigma peltate. Fruits not seen. TYPE: PANAMA. VERAGUAS: vicinity of third branch of Rio Santa Maria, 10-14 km NW of Santa F4, 650-750 m, 10-11 Oct 1975, Robert L. Dressler 5170 (HOLOTYPE: DUKE). Didonica is characterized by solitary pendent flowers on an elongate pedicel continuous with the calyx and anthers with short somewhat rigid tubules dehiscing by oval, latrorse clefts not extending to the apex of the tubule. In 1949, A. C. Smith attached a note to the Allen 3760 collection in which he recognized the plant as representing a new genus "suggestive of Semiramisia in habit and in characters of calyx, corolla, etc., differing in its very short anther-tubules which dehisce by oval lateral clefts." In general habit, in leaf size and shape, in the calyx continuous with pedicel, and in the elongate campanulate-cylindric corolla, the plant does resemble Semiramisia speciosa. However the resemblance goes no further for in the essential features of stamen type the new genus stands alone in the Vaccinieae. The most distinctive feature of the stamens is the manner of dehiscence. Pollen is released latrorsely through short, oval clefts which do not extend to the apex of the tubules (see Fig. 1). Also, the tissue in this region of the tubule is extremely thin although the tubule itself is rather rigid. Other staminal characters which distinguish Didonica from Semiramisia are the short, broad somewhat rigid tubules and the proportionately long and slightly granular thecae. This is in contrast to Semiramisia which has extremely long, narrow flexible tubules which dehisce by short introrse oblique pores, and proportionately short and strongly granular thecae. The new genus resembles Macleania, Psammisia and Mycerinus in the short (or rarely elongate), rigid tubules, but these genera have strongly granular thecae as well as a very different dehiscence type which does not suggest a close relationship. Didonica would seem to represent a rather distinct evolutionary line and we are unable to suggest with confidence its relationships to any other genera. Traditionally, the genera of the Ericaceae have been named for historical or mythological figures. In keeping with this, the new genus is named for Dido the Tyrian princess, founder and queen of Carthage, and said by Virgil to be the lover of Aeneas. The specific epithet refers to the extremely long and thread-like, pendent pedicels which are nowhere else repeated in the Vaccinieae. There is some morphological variation between the two known collections: Allen 3760 from the Province of Cocl~ has glabrous twigs, ovate short-acuminate leaves 3-5 cm broad, and pentamerous floral parts; in contrast, Dressler 5170 from the

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FIG. 1. Didonica pendula. A. Habit, X 0.5. B. Corolla opened to show six stamens, X 2. C. Stamen, ventral view, X 6.5. D. Stamen, lateral view, X 6.5. Drawn from Dressler 5170.

Province of Veraguas has puberulent twigs, narrowly lanceolate long-acuminate leaves to 2 cm broad, and flowers with 5-lobed calyces, 6-lobed corollas and 12 stamens. We feel these differences are probably insignificant and are not unexpected in view of the geographic separation of the plants. Certainly, much more material must be collected and examined before a more thorough knowledge of the morphological variation and understanding of the phylogenetic position of the genus can be attained.

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Additional specimen seen. PANAMA. CocL~: E1 Valle de Ant6n; crest of Cerro Pajita, 1100 m, Allen 3760 (MO).

DISTERIGMA

(KLoTZSCIt) NIEDENZU

R e c e n t collections h a v e r e n d e r e d obsolete a brief a c c o u n t of this small genus in C e n t r a l A m e r i c a published b y one of us ( W i l b u r , 1974). T w o new species are described below along w i t h a k e y which we hope will aid in the i d e n t i f i c a t i o n of the C e n t r a l A m e r i c a n r e p r e s e n t a t i v e s of the genus. T h e r e c e n t l y described G u a t e m a l a n D i s t e r i g m a m a y a n u m L u n d e U ( W r i g h t i a 5 : 85. 1975) is considered b y us to be no m o r e t h a n a s y n o n y m of D i s t e r i g m a h u m b o l d t i i ( K l o t z s c h ) N i e d e n z u . C o n t r a r y to L u n d e l l ' s opinion, all C e n t r a l A m e r i c a n species of D i s t e r i g m a are o f t e n e p i p h y t i c a n d there seems to be no consistent difference b e t w e e n D . h u m b o l d t i i a n d D . m a y a n u m in the size of the corollas. KEY TO THE CENTRAL AMERICAN SPECIES OF

Disterigma

1. Lobes of calyx and corolla 3; corolla deep red; stamens 6 .......... D. trimerum Wilbur & Luteyn 1. Lobes of calyx and corolla 4; corolla white or pale greenish. 2. Thecae of anthers both emptying into a single tubule +__ 1.5 mm long; leaves mostly more than 1 cm long and wide and with an apical apiculus 1-2 mm long .................. ..................................................................................................... D. utleyorum Wilbur & Luteyn 2. Thecae of anthers each emptying into its own tubule 2 mm long or longer; leaves mostly 1 cm or less long and wide and non-apiculate. 3. Leaf blades 2.5 times as long as wide or less and 4 mm wide or wider; calyx lobes eciliate; calyx tube +__ 4-angled in cross-section; corolla tube 6-8 mm long; Guatemala, Costa Rica, Panama (Chiriqui) and northern South America ......................... ........................................................................................ D. humboldtii (Klotzsch) Niedenzu 3. Leaf blades 3 or more times as long as wide and 2.5-4.5 mm wide; calyx lobes ciliate; calyx tube terete or nearly so in cross-section; corolla tube 4-6 mm long; endemic to Panama (but unknown from Chiriqui). 4. Calyx lobes 2-2.5 mm long; leaves typically widest at or near the middle, usually + linear, mostly + 4-times as long as wide; Prov. de Dari6n .. D. panamense Standley 4. Calyx lobes 1.5-1.7 mm long; leaves typically widest decidedly above the middle, _ oblanceolate, mostly + 3-times as long as wide; Prov. de Panam~_ D. luteynii Wilbur Disterigma

trimerum

W i l b u r & L u t e y n , sp. nov.

Frutex epiphyticus vel terrestris 0.5-1.5 (2) m altus; ramulis + triangularibus, glabris; ramis teretibus. Petioli 3-5 mm longi, glabri. Lamina coriacea, ~ integra, obovata vel spatulata vel elliptica, (0.8) 1.2-1.8 cm X (4) 6-12 mm, apice obtusa vel rotundata, basi cuneata, supra glabra et subta sparse strigosa. Calycis tubus • teres, glaber, + 3.5 X 2.0--2.2 mm; calycis lobi 3, ~_ erecti, + 0.5 mm longi, integri, glabri. Corolla rubra, sparse strigillosa; tubus cylindraceus, 7-7.5 X -----2.5 ram; lobi 3, erecti, acuti, ~ 1.5-2 mm X 1-1.2 mm. Stamina 6, corollae paullo breviora, 5.0-5.3 mm longa. E p i p h y t i c or terrestrial, b u s h y shrubs 0.5-1.5 (2) m tall. B r a n c h l e t s sharply ridged and grooved and r a t h e r indistinctly 3 - a n g l e d to v e r y i r r e g u l a r l y 4-angled, g l a b r o u s or v e r y s p a r i n g l y strigillose or m i n u t e l y p u b e r u l e n t ; m a t u r e branches bec o m i n g terete, grayish to blackish, conspicuously fissured. I n t e r n o d e s f r o m a b o u t 1/~_90 as long as the s u r r o u n d i n g leaves. Petioles 3 - 5 m m long, glabrous, f l a t t e n e d a b o v e . L e a v e s coriaceous, b r o a d l y elliptic to s o m e w h a t o b o v a t e or e v e n spatulate, (0.8) 1.2-1.8 cm • (4) 6-12 m m , basally c u n e a t e , apically obtuse to b r o a d l y r o u n d e d , entire to v e r y o b s c u r e l y and m i c r o s c o p i c a l l y serrulate, d a r k green a b o v e a n d p a l e green b e n e a t h w h e n fresh, glabrous above, b e n e a t h e v e n l y b u t s p a r i n g l y b e s e t with appressed, a m b e r - c o l o r e d (when d r y ) t r i c h o m e s 0.2-0.3 m m long; 3 - 5 p l i n e r v e d , midrib and l a t e r a l veins slightly e l e v a t e d to n e a r l y plane on b o t h surfaces.

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Flowers solitary, axillary, subsessile. Pedicel disarticulating just beneath base of calyx, 2-3 mm long, with 1-3 deltoid to acutely oblong bracts near the base. Bracteoles subopposite, semiorbicular, + 1.5 • 1.8-2 mm, broadly clasping, ciliate, located + 1.5-1.8 mm below and scarcely overlapping the base of the calyx. Hypanthium terete, shortly cylindric, glabrous, + 3.5 • 2-2.2 mm; calyx lobes 3, not very pronounced, erect, broadly deltoid, acute, + 0.5 mm long, marginally entire, non-ciliate, glabrous. Corolla deep red, thinly fleshy; corolla tube +- cylindric, abruptly tapering into a basal portion 2-2.3 • about 1-1.4 mm, the longer portion of tube 5-5.5 • about 2.5 mm with a few ascendent trichomes 0.1-0.2 mm long; corolla lobes 3, erect, oblong-deltoid, acute, 1.5-2 x 1-1.2 mm. Stamens 6, + 1 mm shorter than the corolla-tube; filaments 2.5 mm long, flattened, glabrous; anthers granular, 2.52.8 mm long, each theca tapering into a slender tubule + 2.5-2.8 mm long with an introrse elongate-oval slit 1.5 mm long. Style glabrous, slender, extending + 1.O-1.2 mm beyond the closely enveloping stamens or about to the top of the corolla tube. Berry reportedly at first greenish-white turning dark blue. TYPE: PANAMA. CI-IIRIQUi: slopes approaching Cerro Horqueta between 1500 and 1800 m starting about 6.6 km NNE of Boquete, Wilbur & Luteyn 19321 (I-IOLOTYPE: DUKE; ISOTYI~ES: CAS, DUKE, F, GH, MICH, MO, NY, PMA, US). Except for this species, trimerous flowers are not known in Disterigma. The deep red corollas of D. trimerum are unlike those of any other Central American representative of the genus. Hence there seems to be no question but that this plant is an undescribed Central American ericad; the only question is its proper generic placement, a question which arises repeatedly among the tropical Ericaceae. Additional specimens examined. PANAMA. CItlRIQUI: cloud forest on Caldera-Chiriquicito Trail between Q. Hondo and divide, Kirkbride & Duke 939 (DUKE) ; heavily forested slopes along a ridge of the western side of Cerro Horqueta, alt. • 1700-1800 m, Luteyn 3750 (DUKE, NY); premontane cloud forest along ridge E of Cerro Horqueta, elev. 1700-1900 m, Luteyn & Wilbur 4611 (DUKE, NY, US) ; Cerro Hornitos, 40 km NW of Gualaca, cloud forest dominated by Quercus, Podocarpus and Drirnys at 2238 malt., Mori & Bolton 7470 (DUKE, MO).

Disterigma u t l e y o r u m

Wilbur & Luteyn, sp. nov.

Frutex epiphyticus 3-4.5 dm altus; ramuli teretes puberuli. Petioli 1-3 (4) mm longi, puberuli. Lamina succulenta, obovata vel spatulata vel elliptica, apice apiculata, rotundata vel obtusa, puberula, (1.0) 1.5-2.5 (3.0) X (0.1) 1-1.8 cm. Flores axillares solitarii erecti. Pedicelli 1-2 m m longi, puberuli. Bracteae connatae, 1.5-2.0 mm longae. Hypanthium 2-3 X 1.5-1.8 ram, puberulum; calycis lobi 4, erecti, acuti, puberuli, 1.6-2.2 mm longi. Corolla alba; tubus cylindraceus, 4-5.5 mm longus; lobi 4, oblongo-elliptici, acuti, 3.5-5 mm longi. Stamina 8; filamenta distincta alba, ciliata, 1.5-2.5 mm longa; antherae glabrae, non-granulares 2.5-3.5 mm longae; tubulus antherarum I, ___ 1.5 mm longus.

Epiphytic shrub 3-4.5 dm tall. Branchlets subterete to terete, moderately to densely scurfy-puberulent. Internodes short and mostly about 3/~o-% as long as the nearest leaf. Petioles 1-2 (4) mm long and about 1-1.2 mm wide, very short-puberulent. Venation with 5 prominently elevated veins radiating from the petiole. Leaves thick and fleshy when fresh becoming wrinkled when dried, oblong-obovate to somewhat spatulate or rarely oblong-elliptic, (1.0) 1.5-2.5 (3.0) x (0.7) 1-1.8 cm, basally broadly cuneate to rarely obtuse, apically obtuse to broadly rounded often with a pointed apiculus 1-2 mm long, marginally entire and slightly revolute, both surfaces very minutely or even microscopically puberulent and also with amber-colored strigillose trichomes 0.1 mm long; 5-plinerved with the prominently elevated veins arising from the base. Flowers solitary in the axils of the upper leaves, subsessile. Pedicels 1-2 mm long. Bracteoles subopposite, broadly oblong to ovate, --- 1.5-2 •

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2-3 mm, fused the lower 0.5-1 mm into a loosely clasping cupule, minutely appressed-puberulous. H y p a n t h i u m cylindric, 2-3 mm long, pale green in flower, finely and minutely puberulous; calyx rim 1-1.2 m m long; calyx lobes 4, erect, lanceolate to narrowly triangular, acute, 1.6-2.2 • 1 ram, minutely puberulous. Corolla white, cylindrical or barrel-shaped to somewhat funnelform at maturity, glabrous to very sparingly spiculate; corolla tube 4-5.5 x 3.5-4 ram; corolla lobes 4, oblong-elliptic, acute, 3.5-5 • 3 ram. Stamens 8, + 5 mm long or just a little shorter than the corolla-tube at anthesis; filaments white, sparingly ciliate, 1.5-2.5 mm long; anthers smooth, 2.5-3.5 mm long, the 2 thecae emptying into a single orange tubule +- 1.52.0 mm long which open by a single, subterminal but obliquely introrse circular pore. Style glabrous, 4.8-5.3 • 0.6-0.8 ram; stigma slightly bilobed. Berry reportedly "transluscent white." TYPE: COSTA RICA. HEREDIA: pastured and cut-over slopes at Colonia Virgen del Socorro at + 900 m, Wilbur 21706 (HOLOTYPE" D U K E ; ISOTOPES: CAS, CR, F, GH, LL, M I C H , MO, NY, PMA, WIS, etc.). This very interesting but little known species is known from recently collected specimens made at low montane elevations in Costa Rica, Panama, Colombia and Ecuador. Botanical collectors among whom we are no exception have expended a disproportionate amount of effort on the higher elevation of tropical and subtropical countries while neglecting the lower slopes which only now are yielding a surprising number of novelties. This species is biologically interesting in that it seemingly is always intimately associated with a large colony of ants which defend their home environment with great tenacity and vigor. Their persistent defense and painful bites have perhaps dissuaded some collectors from making specimens. The specimens seen by us form a picket fence surrounding a large, moist ball of earth and vegetable debris a foot or so in diameter capped by a dense mat-forming acrocarpus moss. The details of the intriguing commensal relationships between ericad and insect are at present unknown. The species is named in honor of John and K a t h y Utley who collected the specimens first seen by us and months later guided one of us back to the plant. Disterigma utleyorum is easily overlooked and although we were standing under the small tree in which the ericad was growing epiphytically, it was undetected until Kathy's sharp eye spotted it. Additional specimens examined. COSTA RICA. HEREDIA: vicinity of Colonia Virgen del Socorro, 2-4 km E of Cariblanco on the finca of St. Carlos Molina, John & Kathy Utley 2830 (DUKE, F, NY, MO, US). PANAMA. VERACUAS: 5 mi W of Santa F6 on road past Escuela Agricola Alto Piedra on Pacific side of divide, alt. 800-1200 m, Liesner 942 (MO). COLOMBIA. NARINO: Altaquer, 1200 m, M. B. & R. Foster 2114 (COL). ECUADOR. PICHINCHA: downed tree along the Rio Chiguilpe, 7 km S of Sto. Domingo and 6 km E on side road from highway Quevedo-Sto. Domingo, 450 m elev., Dodson, Hilton & Morgan 6061 (SEL). L A T E R O P O R A A. C. SMITH The dehiscence of the anthers in this genus is unique in the Vaccinieae, the pollen sifting out through a longitudinal slit in each theca instead of through terminal tubules. The two known species are little-collected Panamanian endemics. KEY TO Lateropora 1. Calyx lobes 2-3 mm long and 4-5 mm wide basally; corolla lobes +__ 3 mm long and 4.5-5 mm wide basally; petioles 0.5-1.5 cm long; Chiriqui ................ L. ovata A. C. Smith 1. Calyx lobes 1.5-2 mm long and 2.0-2.2 mm wide basally; corolla lobes 1.5-1.8 mm long and • 2 mm wide basally; petioles 3-5 mm long; Veraguas _ L. santafeensis Wilbur & Luteyn

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Lateropora santafeensis Wilbur & Luteyn, sp. nov. Frutex epiphyticus ca. 0.5 ra altus. Folia coriacea, elliptica, 3-8.5 X 2-4 cm; petioli glabri, 3-5 mra longi et 2.0-3.5 mm diam. Inflorescentia axiUaris, umbellato-racemosa, 2-3 cm longa; bracteae ovatae vel suborbiculares, 2.0-2.2 ram longae; pedicelli 5-13 ram longi, glabri; bracteoli ciliati, ovati vel suborbiculares, 2-2.5 • 1.8-2.2 ram. Hypanthia cum pedicello articulata, campanulata, 3-4 mm X 4-5 ram, glabra; lobi calycis 5, erecti, triangulares, acufi, ciliati, 1.5-2 tara longi. Corona urceolata, 4-5 • 5-6 mm, extus strigillosa; lobi corollae 5, triangulates, acuti, 1.5-1.8 ram longi, intus floccosi. Stamina 10; filamenta 2.5-3 mm longa, supra pilosa; thecae granulosae, • 3 ram longae; tubuli rudimentarii, 0.3 ram longi, de latere dehiscentes. Epiphytic shrub about 0.5 m tall with rather stout, glabrous, purplish branchlets and grayish branches. Lea/-blades coriaceous, elliptic, 3-8.5 • 2.0-4.5 cm, - acutely tapering to both apex and base, entire with a thickened, slightly revolute margin, glabrate or with appressed, reddish-brown, apparently glandular, strigillose trichomes 0.2-0.3 m m long especially on the lower surfaces; pinnately veined (or very indistinctly plinerved). Petioles glabrous, 3-5 • 2.0-3.5 mm. Inflorescence axillary, of compactly umbelliform racemes; rachis 8-12 mm long. Floral bracts ovate to suborbicular, 2.0-2.2 m m long and wide, ciliate, somewhat keeled. Pedicels somewhat clavate, 5-13 m m long and • 1 m m in diam at the middle, glabrous, often with an inconspicuous fringe of glandular trichomes 0.1-0.2 m m long distally at the point of articulation with the calyx. Bracteoles subopposite, broadly ovate to suborbicular, 2-2.5 • 1.8-2.2 ram, ciliate, keeled, appressed, located in the lower proximal quarter, ttypanthium campanulate, 3-4 • 4-5 m m at anthesis, glabrous, clearly disarticulating from the pedicel by a marked groove; calyx limb glabrous, • 0.5-0.7 mm long; calyx lobes 5, erect, deltoid, acute, 1.5-2 • 2-2.2 m m , shortly ciliate. Corolla broadly cylindric to urceolate, • 4-5 m m long and medially 5-6 ram, pale greenish, sparingly and inconspicuously strigillose with trichomes 0.1-0.2 m m long; corolla lobes 5, spreading to weakly reflexed, triangular, acute, 1.5-1.8 x 2 ram, densely tangled-villous within with white trichomes 0.6-1.0 m m long. Stamens 10; filaments flattened, - 2.5-3 m m long, broadened basally to • 1 ram, attached submedially just above the strongly bent base of the thecae and densely pilose distally; anthers conspicuously granular, • 3 m m long, the strongly incurved base • 1 mm long or % the length of the thecae; dehiscence by a slit extending from apex to the pronounced basal curve. TYPE: P A N A M A . V~RACTJAS: road beyond Escuela Agricola Alto Piedra, above Santa F6, 800-1000 m, Luteyn & Wilbur 4575 (HOLOTYPE: D U K E ) . L Y S I C L E S I A A. C. S~ITH

Lysiclesia panamensis Luteyn & Wilbur, sp. nov. (Fig. 2). A L. caudata foliorum forma, raraulis, rhachidi calycisque lobis puberulis, rhachidi pedicellisque glanduliferis, et rhachidi breviori paucifloro differt. "Lax shrub, almost vine-like" (fide label data). Branches terete to bluntly angled, smooth, short-pilose becoming glabrate when mature, glaucous, reddish-brown when dry. Petioles subterete, flattened adaxially, rugose, short pilose, 2.5-3 mm long and 1 m m diam. Leaves ovate-elliptic, (2.5) 3-3.5 • (1.2) 1.5-2 cm, basally obtuse or rounded, apically abruptly short acuminate, apiculate, glabrous, obscurely 3 (5)-plinerved, midrib nearly plane above but raised beneath, lateral nerves and veinlets slightly raised and prominent above, obscure beneath. Inflorescence axillary, racemose, 2-3 flowered; rachis short-pilose, bluntly angled, 1.5-2.2 cm long and 1 mm diam, with scattered capitate, glandular hairs 1 m m long. Floral bracts lanceolate, 1.5 m m long, ciliate. Pedicels subterete, short-pilose, 15-17 mm long and

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FIG. 2. Lysiclesia panamensis. A. Habit, X 0.7. B. Calyx and pedicel showing bracteoles, elongate calyx lobes, style and glandular hairs, X 1. C. Corolla opened to show four stamens, X 1. I). Stamens in dorsal view, X 5. Drawn from Mori 6" Kallunki 3602.

about 1 m m diam, with scattered capitate, glandular hairs. Bracteoles ovate-lanceolate, acuminate, 3-4 X 1.5 ram, strongly nerved, ciliate and provided with capitate, glandular hairs. Calyx short-pilose, 36-37 m m long, reddish-brown; tube obscured by wings, --- 4.5 m m long; limb short-campanulate, 2 mm long, 3-parted, the lobes triangular, acuminate, extended slightly below the base of the tube, 10 m m broad at base, their margins and lamina with scattered capitate, glandular hairs. Corolla

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pink, cylindric b u t narrowing at throat and with lobes flaring, membranaceous, shortpilose (except a t very base) without and at throat within, 35 m m long; lobes triangular, obtuse, 1 m m long. Stamens a l t e r n a t e l y 10 and 12 m m long; filaments pilose distally, alternately 5 and 8.5 m m long; anthers including tubules slightly unequal, alternately 5 and 5.5 m m long; thecae 2.5-3 mm long; tubules with terminal or (in shorter stamens) slightly oblique pores. Style pilose distally, • 36 m m long. Berry not seen. TYPE: P A N A M A . PANAMa: Cerro Jefe, cloud forest dominated b y Clusia spp. and Colpothrinax cookii. Along trail on ridge running N E of summit, 1000 m elev., Mori & Kallunki 3602 (HOLOTYPE: M O ; ISOTYPE: D U K E ) . T h e new species m a r k s a significant extension of the range of its v e r y distinct but extremely rare genus. Three species of Lysiclesia are now recognized, each known only from its type collection. KEY TO THE SPECIES OF Lysiclesia 1. Branchlets, petioles and rachis glabrous; leaves long-acuminate, 6-7 cm long; stamens 8, 6-8 mm long; plants from 2400-2800 m elev.; Colombia .................. L. caudata A. C. Smith 1. Branchlets, petioles and rachis puberulent; leaves acute, abruptly or shortly acuminate, 2-3.5 cm long; stamens 10, 8-12 mm long; plants from 1000-2000 m elev.; Colombia or Panama. 2. Raehis short-hirsute; calyx lobes glabrous on external surfaces and ciliate with eglandular hairs; corolla basally glabrous and distally sparsely pilose; Colombia ................ .......................................................................................................... L. minor A. C. Smith 2. Rachis puberulent and glandular; calyx lobes puberulent on external surfaces and glandular-ciliate; corolla uniformly puberulent; Panama .... L. panamensis Luteyn & Wilbur M A C L E A N I A HOOKER Macleania talamancensis

Wilbur & L u t e y n , sp. nov. (Fig. 3).

Frutex terrestris 0.5-1 m altus. Petioli 2-3 mm longi, puberuli. Lamina foliorum coriacea, obovata vel oblongo-elliptica, minutissime serrulata, 2.5-4.7 X 1.2-1.8 (2.3) cm. Racemi axillares, compacti; pedunculi, rachides et pedicelli pilosuli vel puberuli; pedicelli articulati 6-10 mm longi; bracteae lineares vel lanceolatae puberulae 3-4 mm longae; bracteolae 2-3 mm longae. Hypanthia campanulata vel cylindrica puberula 2.5-3 X 2.5-3 mm; limbus calycium 0.8-1 mm longus; lobi calycium 5, puberuli acuti 1.2-1.8 mm longi. Corolla puberula, subrosea vel rubella; tubus cylindricus, 8-10 X 4-5 mm; lobi 5, 1.5-2 mm longi, reflexi, pilosuli. Stamina 10; filamenta distincta 2.0-2.5 mm longa; thecae granulosae + 2.5 mm longae; tubuli 2, 3.5-4 mm longi; pori obliqui introrsi + 0.6 mm longi. Shrub 0.5-1 m tall. Leaves coriaceous, imbricated, obovate to more typically oblong-elliptic, mostly 2.5-4.7 x 1.2-1.8 (2.3) cm, acute to apically rounded with a 1-2 m m long apiculus, seemingly entire b u t actually indistinctly and minutely serrulate, glabrous on the upper surface except finely whitish-appressed-pilosulous near the base and inconspicuously ciliate especially in lower quarter, lower surface indistinctly and sparingly beset with both appressed reddish-brown glandular trichomes • 0.2 mm long and also appressed whitish-pilosulous basally. Petioles 2-3 mm long, moderately puberulous. Inflorescence a dense, axillary raceme with softly pilosulous or puberulous rachises and pedicels. Floral bracts appressed, linear to lanceolate, 3 - 4 mm long, puberulent. Pedicels distinctly articulate just beneath the hypanthinm, 6-10 m m long. Bracteoles appressed, linear to narrowly lanceolate, 2-3 mm long, puberulent. Hypanthium campanulate to cylindric, • 2.5-3.0 long and in diam, m o d e r a t e l y puberulous with whitish trichomes • 0.1-0.2 m m long; calyx limb • 0.8-1 m m long, slightly flaring; calyx lobes triangular, acute, 1.2-1.8 mm long, puberulous externally and internally; sinus broadly rounded. Corolla tube

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Fro. 3. Macleania talamancensis. A. Habit, X 0.5. B. Flower, • 5. C. Stamen, ventral view, X 9. D. Stamen, lateral view, • 8. Drawn from Burger & Liesner 7401.

cylindric, 8-10 • 4-5 mm, pink to red, externally appressed- to spreading-puberulous with moderately abundant hyaline trichomes - 0.4 mm long and internally moderately to densely pilose with hyaline trichomes; corolla lobes reflexed, triangular, 1.5-2 mm long, white, densely pilosulous both externally and internally. Stamens 10; Jilaments distinct, attached to the receptable, +-- 2.0-3.0 mm long, glabrous proximally but marginally with appressed-pilosulous hyaline trichomes dis-

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tally; connective 1.0-1.2 mm long; anther sacs 2.5-3.5 mm long, coarsely granular, basally strongly incurved; tubules 2, separate to the base, 3.5-4.5 m m long opening by oblique, introrse pores - 0.6 mm long. Berry not seen. TYPE: COSTA RICA. SAN Jos~: elfin forest of dead oak trees 4-10 m tall along the trail from Canahn to Chirrip6 via Los Angeles north of Rio Talari at 3200-3400 m, Burger 6" Liesner 740I (HOLOTYPE: D U K E ; ISOTYeES: F, NY, MO). This only recently collected species is strikingly different from all other Central American species of Macleania. I n M. talamancensis and M. rupestris (H.B.K.) A. C. Smith [including M. glabra (Klotzsch) Hoerold] each anther sac terminates in a tubule, while all other Central American species have a single tubule serving the paired anther sacs. Macleania talamancensis, known only from the upper slopes of Costa Rica's highest mountain, can be distinguished from the widespread M. rupestris by the following key. 1. Hypanthium and corolla tube glabrous or very nearly so; bracteoles broadly deltoid to oblong, _ 1-2 times as long as wide; anther sacs 4-7 mm and tubules 3.5-6 mm long ............................................................................................................................................... M. rupestris 1. Hypanthium and corolla tube densely pilosulous or puberulous; bracteoles linear to narrowly lanceolate, +_ 4-6 times as long as wide; anther sacs 2.5-3.5 and tubules 3.54.5 mm long ............................................................................................................. M. taIamancensis Additional specimens examined. COSTA RICA. SAp Josg: Cordillera de Talamanca, Cerro Chirrip6 massif, forest just below the p~tramo with many dead trees, elev. 3200 m, Davidse & Pohl 1594 (DUKE). T H E M I S T O C L E S I A KLOTZSCH T h e m i s t o e l e s i a e o s t a r i e e n s i s Luteyn & Wilbur, sp. nov. Frutex epiphyticus. Folia ovata, (1.6) 2-2.5 )< (0.8) 1-2 cm. Flores axiliares solitarii. Pedicellus cum calyce eontinuus glaber 1-2 mm longus. Calyx 4-4.5 mm longus hispidulus; tubus obconicus 5-angulatus 3 mm longus; limbus 1-1.5 mm longus lobis inclusis. Corolla cylindrica 5-6 mm longa hispidula. Stamina 10, 5 mm longa; filamenta alternatim 2.8-3 mm et 3-3.2 mm longa hispidula; antherae 2.5 mm longae basi appendiculatae. Stylus 5-6 mm longus. Pendent, epiphytic shrub with branches to 0.5 m long. Stems terete, minutely striate, tan to reddish-brown when dry, hispidulous with reddish-brown hairs 0.6-0.8 mm long becoming glabrous with age. Leaves distichous, flat or concave, ovate, (1.6) 2-2.5 • (0.8) 1-2 cm, basally rounded or subcordate, apically acute or short acuminate, marginally entire, hispidulous on upper surface especially near margins, glabrous beneath, brown above and olive-green or light brown beneath when dry; obscurely 5-plinerved, the midrib slightly impressed proximally and raised distally above and slightly impressed but inconspicuous beneath, lateral nerves and veinlets slightly raised but inconspicuous above and totally obscure beneath. Petiole subterete, densely hispidulous, 1-1.5 mm long. Flowers axillary, solitary with 4 tiny inflorescence bracts at base. Floral bract ovate, marginally ciliate, 1.5 mm long. Pedicel terete, glabrous, 1-2 mm long, completely concealed by the 2, ovate to hemispheric distally hispidulous bracteoles. Calyx somewhat glaucous, copiously hispidulous with translucent or reddish-tinged hairs 1-1.2 mm long; tube obconic, bluntly 5-angled, 3 mm long; limb 1-1.5 mm long including apiculate lobes. Corolla cylindric, narrowed at the throat, 5-6 • 2.2 mm, hispidulous throughout, white; lobes 1 mm long. Stamens 10, 5 mm long; filaments alternately slightly unequal, 2.8-3 mm and 3-3.2 mm long, hispidulous over entire length; anther 2.5 m m long, the tubules slightly longer than the thecae; thecae with a minute (0.1 mm long) finger-like appendage protruding from the base. Style 5-6 mm long, glabrous. Berry not seen. TYPE: COSTA RICA. CARTAGO: steep slopes along roadside on property of ICE

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hydroelectric plant • 15-20 km beyond suspension bridge over the Rio Grande de Orosi, 1300-1800 m elev., Luteyn, Wilbur & K. Utley 4407 (HOLOTYPE: D U K E ; ISOTYPES: F, MO, N Y ) . Themistoclesia horquetensis and T. costaricensis are only the third and fourth species in the genus to be described from Central America, T. pentandra and T. smithiana, both from Costa Rica and Panama, being previously known. These four Central American species, together with the Colombian T. an]racta Sleumer, T. compta Sleumer and T. cuatrecasasii A. C. Smith, form a distinct group characterized by axillary, solitary (or rarely paired), subsessile flowers. The newly described species are most closely related to T. cuatrecasasii, however, differing most significantly in the characters mentioned in Table 1. To facilitate the identification of the Central American species, we provide the following key. K E Y TO T H E C E N T R A L A M E R I C A N S P E C I E S OF

Themistoclesia

1. Stamens 5; calyx distinctly 5-winged; leaves apically obtuse ................ T. pentandra Sleumer 1. Stamens 10; calyx bluntly 5-angled, not winged; leaves apically acute or acuminate. 2. Corolla 2.5-3.5 mm long; leaves less than 1 em long, basally cuneate ......................... ........................................................................................................ T. smithiana (Standl.) Sleumer 2. Corolla 5-6 mm long; leaves more than 2 cm long, basally rounded, subtruncate or subcordate. 3. Bracteoles ovate to hemispheric, completely concealing the glabrous pedicel; corolla cylindric, hispidulous throughout; stamens 5 mm long, filaments hispidulous, thecae basally appendiculate; Costa Rica ........................... T. costaricensis Luteyn & Wilbur 3. Bracteoles awl-shaped, never concealing the hispidulous pedicel; corolla urceolate, hispidulous distally; stamens 2.5 mm long, filaments glabrous, thecae without basal appendage; Panama ......................................................... T. horquetensis Luteyn & Wilbur T h e m i s t o c l e s i a h o r q u e t e n s l s Luteyn & Wilbur, sp. nov. Frutex epiphyticus. Folia ovata (2.5) 3-4.5 X 1.5-2.5 cm. Flores axillares solitarii. Pedicellus cum calyce continuus hispidulus 1.5-2.5 mm longus. Calyx 4.5-5 mm longus hispidulus; tubus obconicus 5-angulatus 2-3 mm longus; limbus 1.5-1.75 mm longus lobis inclusis. Corolla urceolata 5-5.5 mm longa basi glabra apice bispidula. Stamina 10, 2.5 mm longa; filamenta 1.5-2 mm longa glabra; antherae 1-1.2 mm longae. Stylus 3.5 mm longus. Epiphytic shrub about 5 dm tall. Stems subterete or bluntly angled, minutely striate becoming smooth, glaucous, pale reddish-brown when dry, hispidulous with translucent hairs 0.2-0.3 mm long becoming glabrous with age. Leaves distichous, ovate, (2.5) 3-4.5 • 1.5-2.5 cm, basally broadly rounded or subtruncate, apically short acuminate, apiculate, marginally entire, obscurely short-pilose above especially near margins and glabrous beneath, pale green above and brown beneath when dry but with a pale greenish-brown area 1 mm broad along the entire margin beneath; 3 (5)-plinerved with the nerves obscure but slightly impressed above and only the midrib slightly raised beneath. Petiole subterete, flattened above, hispidulous as branchlets, 2-3 mm long and 1 mm diam. Flowers axillary, solitary with 2-4 minute inflorescence bracts at base. Floral bract ovate, marginally ciliate, 0.3-0.5 mm long. Pedicel angled, striate, glaucous, hispidulus, 1.5-2.5 mm long. Bracteoles awlshaped, ciliate, 0.5 mm long, located above the middle. Calyx glaucous, sparsely hispidulous with translucent hairs 0.2 mm long; tube obconic, bluntly 5-angled, 2-3 m m long; limb 1.5-1.75 m m long including the apiculate lobes. Corolla urceolate, strongly constricted at throat, 5-5.5 • 2.2 ram, glabrous beneath but upper part of throat and lobes hispidulous; lobes • 0.5 mm long. Stamens 10, 2.5 m m long; Jilaments slightly but distinctly alternating 1.5 and 2 mm long, glabrous, attached near base of anther; anther 1-1.2 mm long with tubules equaling thecae. Style 3.5 mm long, glabrous. Berry - 5 mm diam.

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TABLE I THE MAJOR DI~'FERENCES BETWEEN Themistoclesia cuatrecasasii, Themistoclesia horquetensis, AND

ThemistocIesia costaricensis T. cuatrecasasii

Characters

T. horquetensis

T. costaricensis

Branchlet pubescence

reddish brown hairs 0.5 mm long

translucent hairs 0.20.3 mm long

reddish brown hairs 0.6-0.8 mm long

Leaves

2-3.5 X 1-2 cm long; basally subcordate; glabrous

2.5-4.5 X 1.5-2.5 cm long; basally rounded or subtruncate; marginally copiously hispidulous

2-2.5 X 1-2 cm long; basally rounded or subcordite; marginally copiously hispidulous

Pedicel

1 mm long; glabrous

1.5-2.5 mm long; hispidulous

1-2 mm long; glabrous

Bracteoles

ovate to hemispheric; glabrous ; concealing pedicel

awl-shaped; ciliate throughout ; never coneealing pedicel

ovate to hemispheric; distally hispidulous ; concealing pedicel

Corolla

cylindric, 11-12 mm long; hasally glabrous and hispid distal half; carmine

urceolate, 5-5.5 mm long; upper throat and calyx lobes hispidulous; white (?)

cylindric, 5-6 mmlong; hisp'_dulous throughout; white

9

2.5

5

Filaments

6 mm long; equal; hispidulous distal half

1.5 or 2 mm long; slightly alternately unequal; glabrous

3 mm long; slightly alternately unequal; hispidulous entire length

Anthers

4 mm long; tubules longer than thecae; thecae without basal appendage

1.2 mm long; tubules equaling thecae; thecae without basal appendage

2.5 mm long; tubules longer than thecae; thecae with basal appendage

3.5

5-6

Stamen length (mm)

Style length (mm)

10-12

TYPE: P A N A M A . CHIRIQUi: S slopes o f C e r r o H o r q u e t a , N of B c q u e t e , 2200 m elev., Wilbur, Teeri & Foster 13491 (HOLOTYPE: D U K E ) . A d i s c u s s i o n of t h e r e l a t i o n s h i p s of T. horquetensis is i n c l u d e d w i t h t h a t of T.

costaricensis. UTLEYA

WILBUR & LUTEYN, gen. nov.

Frutex epiphyticus. Folia alterna coriacea petiolata pinnatinervia. Inflorescentia axillaris breviter racemosa pauciflora. Calyx cum pedicello articulatus alis 5 conspicuis e hasi ad apicem adscendentibus cum lohis alternantihus praedito. Corolla carnosa urceolata calcaribus 5 conspicuis lohis oppositis praedita. Stamina 10 aequilonga; filamenta distincta; connectivum ecalcaratum; loculi antherarum leviter granulates basi introrsum curvi; tubuli antherarum 2 ad basim distincti flexibiles per rimas ovales aliquantum introrsas elongatas dehiscentes. Ovarium 5-1oculare; ovula numerosa. Utleya costarieensis

W i l b u r & L u t e y n , sp. n o v . ( F i g . 4 ) .

Folia elliptica vel ovato-elliptica, apice breviter acuminata et basi cuneata et decurrentes, 2.5-5 (6))< (1) 1.5-3.5 cm; petioli 4-12 mm longi, glabri. Inflorescentia 3-6 mm longa; bracteae ovatae, 1 mm longae; pedicelli 12-14 mm longi, glabri; bracteoli 1-1.5 mm longi. Calyx 5.5-8 mm longus, glaher; tubus obconieus 2-2.5 mm Iongus; limbus 2 mm longus lobis inclusis; lobi apiculati 0.5 mm longi; alae 2-3 mm latae. Corolla 4 mm longa, glabra; calcaria 1.5 mm lata. Stamina 3 mm longa; filamenta 2 mm longa, pilosa; antherae 1.5-1.75 mm longae; tubuli 0.5-0.75 mm longi.

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Epiphytic shrub 1 m tall. Mature stems glabrous, subterete and usually with narrow, corky wings; bark gray. Immature stems and twigs of new growth glabrous, sharply angled, 3-5-winged with wings to 1 mm broad, reddish-brown when dry. Leaves alternate, exstipulate but with pseudostipules in the form of subulate bud scales 2-2.5 mm long; lamina broadly elliptic to ovate-elliptic, 2.5-5 (6) x (1) 1.53.5 cm, basally cuneate and decurrent onto the petiole, apically short-acuminate with the apex ultimately either acute or obtuse, glabrous on both sides but with scattered, reddish, glandular fimbriae, margins slightly revolute, entire or distally crenate; pinnately nerved, midrib very prominent above and raised proximal %-1/2 but plane to slightly impressed distally, carinate beneath, lateral nerves arcuate and anastomosing near the margins, slightly impressed above but raised beneath, veinlets reticulate and slightly raised on both sides. Petiole 4-12 mm long, sharply angled, canaliculate or carinate above, carinate beneath, glabrous but with scattered glandular fimbriae. Inflorescence racemose, 3-4 flowered; rachis glabrous, 3-6 mm long, sharply angled, with 2-4 tiny (less than 1 mm long), ovate bracts at base. Floral bracts ovate, 1 mm long. Pedicels articulate with the calyx, subterete, striate, angled, 12-14 mm long and 0.5 mm in diameter, glabrous but with scattered glandular fimbriae. Bracteoles basal, ovate, acute, 1-1.5 mm long, glandular-fimbriate marginally. Calyx 5.5-8 mm long and 7-9 mm across the top including the wings, broadly 5-winged, the wings alternate with and produced beyond the lobes, glabrous but glandularfimbriate basally; tube obconic, 2-2.5 mm long; limb spreading, 2 mm long including the lobes; lobes apiculate, 0.5 mm long; wings 2-3 mm broad. Corolla light pink with white lobes, fleshy, urceolate, strongly constricted at the throat, 4 mm long and 6 mm diameter including the spurs, pentagonal in cross-section, glabrous without but densely pilose within at throat, corolla tube bearing behind each lobe vertically oriented, ovate-obtuse laterally compressed spurs about twice as tall as the lobes proper, spurs 1.5 mm broad at widest point; lobes triangular, 1 mm long, slightly reflexed at anthesis. Stamens 10, equal, 3 mm long; filaments distinct, 2 mm long, densely long-pilose in distal 89 anthers 1.5-1.75 mm long; thecae strongly curved inwards at the base, 1 mm long, surface slightly granular; tubules 0.5-0.75 mm long, dehiscing introrsely although somewhat latrorsely their entire length and somewhat onto the theca itself. Style 2 mm long, glabrous, included. Fruit not seen. TYPE: COSTA RICA. CARTAGO: steep roadside slopes on property of ICE hydroelectric plant - 15-20 km beyond the suspension bridge over the Rio Grande de Orosi, elev. 1300-1800 m, 20 Dec 1974, Luteyn, Wilbur, Utley & Utley 4412 (HOLOTYPE: DUKE; ISOTYPES: DUKE, F, GH, MICH, MO, NY, US). Utleya costaricensis is characterized by winged stems, fleshy flowers with articulate, winged calyces whose wings alternate with the calyx lobes, corollas with spurs opposite the corolla lobes, and stamens dehiscing by introrse or slightly latrorse oval slits extending the entire length of the tubules and slightly onto the sac itself (see Fig. 4). The combination of characters possessed by this little-collected remarkable genus are so unlike those of any described one that we are uncertain as to its affinities. At first glance the winged calyx recalls that of Themistoclesia pentandra Sleumer, also from Costa Rica. In Themistoclesia, however, the calyx is continuous with the pedicel, staminal dehiscence is by short terminal oval pores, and the seeds are covered by a translucent, mucilaginous sheath (this last character being known to occur in the Neotropics only in Themistoclesia and Sphyrospermum). (Note: Mature fruits were not available for the new genus but the immature seeds, at least, lacked the mucilaginous sheath.) Nevertheless, Themistoclesia may be its closest relative. The stout, rather coarse anthers and long-pilose filaments and connectives are reminiscent of

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FIG. 4. Utleya costaricensis. A. Habit, X 0.7. B. Flower, X 4. C. Calyx viewed from above, X 4. D. Corolla viewed from above, X 4. E. Calyx in longitudinal section, style removed, X 5.5. F. Stamen, ventral view, X 13. G. Stamen, lateral view, X 13. Drawn from Luteyn et al. 4412.

certain species of Vaccinium [e.g.V. floccosum (L. O. Williams) Wilbur & Luteyn], although nowhere in that genus are the calyces winged, and anther dehiscence is generally either by terminal pores or relatively short oval clefts. The manner in which the somewhat latrorsely dehiscing clefts extend proximally onto the sac begins to approximate that seen in Lateropora; however, in that genus the anthers seemingly lack tubules and dehiscence extends the entire length of the theca. To our knowledge, latrorse dehiscence occurs in only two other vacciniaceous species--Vaccinium jeJense and Didonica pendula (both newly described herein), where however, the

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t u b u l e s a r e m o r e e l o n g a t e a n d t h e c l e f t s are r e s t r i c t e d to t h e d i s t a l %. T h e n a t u r e of t h e corolla s p u r s in U t l e y a is of s o m e i n t e r e s t . A s d e s c r i b e d a b o v e , t h e s e s p u r s a r e v e r y c o n s p i c u o u s , f l e s h y a n d solid ( n o t h o l l o w ) , a n d a r e s i t u a t e d o n t h e c o r o l l a t u b e o p p o s i t e t h e corolla lobes. S p e c i m e n s of M a c l e a n i a p e n t a p t e r a H o e r o l d m a y somet i m e s b e a r a p i c u l a t e s p u r s d i r e c t l y o n t h e tip of t h e corolla lobes, b u t t h e s e a r e of a v e r y d i f f e r e n t t y p e a n d a r e i n n o w a y c o m p a r a b l e to t h e p r o n o u n c e d s p u r s of U t l e y a . Additional specimens examined. COSTA RICA. CARTAGO: disturbed primary forest high over Rio Grande de Orosi, 8 km S of Tapanti, 1500 m alt., Lent 923 (F, NY); 10 km S of Tapanti along new road on E slope above the Rio Grande de Orosi, 1600 m elev., Burger & Burger 7548 (F, NY) ; Tapanti hydroelectric project S of Orosi, 1400-1600 m, Utley & Utley 2105 (DUKE, F, MO, NY) ; Tapanti hydroelectric project, 4.7 km beyond gate, Utley & Utley 4360 (DUKE) ; road from Tapanti to Taus and Tausito, 3 km NE of Rio Grande de Orosi, 1300-1500 m, Utley & Utley 5025 (DUKE).

VACCINIUM LINNAEUS KEY TO THE COSTA RICAN AND PANAMANIAN SPECIES OF V a c c i n i u m 1. Leaves entire; corolla reddish or pale greenish-yellow; flowers solitary in the axils of the leaves or borne in an umbelliform cluster; anthers spurless. 2. Corolla 4-lobed, red or reddish-purple; stamens 8, shrublets creeping, vine-like, closely clasping the bark; hypanthium and pedicel continuous (i.e. non-articulate) ................ ....................................................................................................................... 1. V. dissimile Blake 2. Corolla 5-lobed, mostly pale greenish-yellow but often tinged with pink; erect shrubs or trees or, if epiphytic vines, neither creeping nor closely clasping the bark; hypanthium and pedicels jointed (i.e. articulate). 3. Calyx lobes 5-6 mm long; principal veins of leaf blades moderately spreading-hirsute beneath; hypanthium and calyx lobes both hirsute and with glandular, capitate protuberances ....................................................................... 2. V. je]ense Luteyn & Wilbur 3. Calyx lobes 3 mm or less long; principal veins of leaf blades at most puberulent or glandular-strigillose beneath; hypanthium and calyx lobes glabrous to inconspicuously puberulous. 4. Calyx lobes 2-2.5 mm long; staminal tubules • 1 mm long; endemic to Panama ................................................................... 3. V. floccosum (L. Williams) Wilbur & Luteyn 4. Calyx lobes 0.2-1.2 mm long; staminal tubules 2-5.5 mm long. 5. Staminal tubules 3.5 mm long or longer; corolla internally glabrous to sparsely villous ......................................................................................... 4. V. poasanum Donn. Sin. 5. Staminal tubules 1.8-2.2 mm long; corolla moderately to densely pilose within. 6. Leaves basally acutely tapering to rounded; calyx lobes 0.5-0.8 mm long; Prov. de Heredia and San Jos~ in the Cordillera Central ........................... ................................................................................. 5. V. costaricense Wilbur & Luteyn 6. Leaves basally rounded to more typically cordate and amplexicaule; calyx lobes 0.8-1.2 mm long; Prov. de Cartago along the Rio Grande de Orosi upstream from Tapanti ........................................... 6. V. orosiense Wilbur & Luteyn 1. Leaves crenate-serrulate; corolla white or tinged with pink or red; flowers borne in a secund raceme; anthers spurred .................................................... 7. V. consanguineum Klotzsch

Vaccinium costaricense W i l b u r & L u t e y n , sp. nov. ( F i g . 5). Arbuscula epiphytica vel terrestris, 1-3 m a l t a . Folia obscure 3-5-plinervia, coriacea, ovata vel lanceolata vel elliptica, apice acuta vel acuminata et basi cuneata vel rotundata, (2.5) 4-8 (11) X (1.5) 2.5-4.5 (6.2) cm; petioli 1-2 mm longi, glabri. Inflorescentia axillaris, umbellato-racemosa, 2-4 (5) cm longa; bracteae ovato-lanceolatae vel lanceolatae, 1-3 mm longae; pedicelli 1-3 mm longi, glabri vel puberuli; bracteolae 0.8-1.5 (2) mm longae. Hypanthia cum pedicello articulata, campanulata, 2.5-3 mm diam, glabra vel puberulenta; lobi calycis 5, triangulares, 0.5-0.8 mm longi, acuti. Corolla cylindrica vel urceolata, 6-8 mm longa, intus pilosa; lobi corollae 5, triangulares, 1-2 mm longi, acuti. Stamina 10, thecae granulosae, ___ 1.8 mm longae; tubuli • 2 mm longi. T e r r e s t r i a l or m o r e t y p i c a l l y e p i p h y f i c s h r u b s 1 - 3 m tall w i t h g r a y i s h to r e d d i s h b r o w n , g l a b r o u s b r a n c h l e t s . L e a v e s coriaceous, o v a t e or l a n c e o l a t e to elliptic, ( 2 , 5 ) 4 - 8 ( 1 1 ) • ( 1 . 5 ) 2.5-4.5 ( 6 . 2 ) cm, b a s a l l y a c u t e l y t a p e r i n g to r o u n d e d , a p i c a l l y

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FIc. 5. Vaccinium costaricense. A. Habit, X 0,5. B. Flower, X 3.5. C. Stamen, ventral view, X 13. D. Stamen, lateral view, X 13. Drawn from Utley & Utley 1277.

acute to acuminate, entire, marginally slightly thickened and perhaps appearing slightly revolute, glabrous to moderately but inconspicuously appressed glandularstrigillose both above and more especially beneath; indistinctly 3-5-plinerved and also somewhat weakly arcuately veined. Petioles 1-2 mm long, glabrous. InJlorescences axillary (but often superficially appearing terminal), umbelliform racemes; rachis 1-2.5 cm long, glabrous to very sparingly and minutely puberulent. Floral bracts persistent, scale-like, ovate-lanceolate to lanceolate or triangular, 1-3 mm long, glabrous to minutely ciliate. Pedicels mostly 1-3 cm long, glabrous to inconspicuously puberulous. Bracteoles subopposite, appressed, persistent, lance-ovate to lanceolate, 0.8-1.5 (2) mm long, glabrous to indistinctly ciliate, located in the lower

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third of pedicel. H y p a n t h i u m clearly disarticulating from the pedicel by means of a sharply delineated groove often indistinctly fimbriate with glandular trichomes - 0.1 m m long, campanulate, - 2-2.5 • 2.5-3 mm, glabrous to inconspicuously although often densely, minutely puberulent; calyx limb • 0.5-0.8 m m long; calyx lobes 5, triangular, acute, 0.5-0.8 m m long, glabrous to inconspicuously puberulent. Corolla broadly cylindric to somewhat urceolate, 6-8 • 4-7 mm, pale greenish-yellow, externally glabrous and internally moderately to densely pilose especially in the throat and on the lobes with sordid, white trichomes; corolla lobes 5, erect to spreading or somewhat reflexed, triangular, acute, 1-2 mm long. Stamens 10; filaments flattened, 3 m m long, slightly adherent to the very base of the corolla tube and attached to the anther-sacs in their lower quarter, slightly ciliate; anther sacs (thecae) granular, • 1.8 mm long, golden, basally markedly inwardly curved with an apiculate tip; tubules stout, - 2 mm long each with slightly flaring terminal pore. TYPE: COSTA RICA. HEREDIA: vicinity of Cerro Chompipe between Rio Las Vueltas and the Rio Nuevo at an elev. of 1800-2000 m • 12 km N N E of San Rafael, Wilbur & L u t e y n 18579 (HOLOTYPE: D U K E ; ISOTYPES: CR, F, MO, N Y ) .

Distribution: Endemic to the cordillera between the northeastern slopes of Volc~n Barba, Prov. de Heredia, and the western slopes of Volc~n Irazfi, Prov. de Cartago. Additional specimens examined. COSTA RICA. HEREBY: primary forest near Rio Las Vueltas, NE of Volc~in Barba, 1950 m, Lent 2645 (DUKE, NY, PMA); heavily cut forests between Rio Las Vueltas and Quebrada Cabrfl, vicinity of Cerro Chompipe, 1700-2000 m, Luteyn & Wilbur 4438 (DUKE, F, MO); along Calle Zurqui near the border with San Jos6 Prov., 1580 m in pastures with remnant trees, Luteyn & Wilbur 4454 (DUKE, F, NY); along the west fork of the upper Rio Para Blanco beyond the road terminus of Calle Zurqul ~- 6-8 km NE of San Isidro de Heredia, Utley 690 (DUKE, F) ; SW fork of the upper Rio Par~ Blanco on roadside banks and an open pasture, lower slopes of Cerro Zurqui at 1500-1650 m, Utley & Utley 1277 (DUKE, MO, NY); Cerro Zurqui, lower pastured slope 1500-1700 m, Utley 3667 (DUKE); dense cloud forest and grown over abandoned pastures 12 km NE of San Rafael at • 2000 m mostly between Rio Las Vueltas and Quebrada Grande, Wilbur, Almeda & Luteyn 15955 (DUKE); vicinity of Cerro Chompipe between Rio Las Vueltas and Rio Nuevo at -4- 1800-2000 m, Wilbur & Luteyn 18576 (DUKE, F, MO, NY, US) ; pastured slopes above the Rio Pardi Blanco on the lower slopes of Cerro Zurqui • 7 km NE of San Josecito at • 1600-1800 m, Wilbur & Luteyn 18629 (DUKE), 18644 (DUKE, F), 18654 (DUKE). SAN Jos~: Open pastures with scattered trees on NW flanks of Volc~in Irazfl • 1850 m • 3-6 km beyond Las Nubes in the vicinity of Cascajal, Almeda 2626 (DUKE) ; wooded slopes • 7 km NE of Cascajal at -4- 1650 m, Wilbur 19815 (DUKE) ; pastured slopes and thickets at Alto La Palma ~ 15 km NE of San Jos~ at -~ 1480 m, Wilbur 20339 (CR, DUKE, F, MICH, MO, NY, PMA, US, WIS).

V a c c i n i u m f l o e e o s u m (L. O. Williams) Wilbur & Luteyn, comb. nov. Hornemannia floccosa L. O. Williams, Brittonia 18: 248. 1966. TYPE: PANAMA. CHmlQUI: Boquete District, rain forests, Bajo Chorro, 6000 ft, Davidson 104 (HOLOTY~: F; ISOTYPE:

MO). Symphysia ]loccosa (L. 0. Williams) L. O. Williams, Phytologia 24: 158. 1972. This species is known only from the Boquete District of Chiriqui, Panama. Although known for little more than a decade, it has been assigned to three different genera. Its relationships are clearly with Vaccinium poasanum, V. costaricense and V. orosiense to which we would also refer the West I n d i a n plant known most recently as Symphysia racemosa (Vahl) Stearn. V a c c i n i u m j e f e n s e L u t e y n & Wilbur, sp. nov. (Fig. 6). Frutex epiphyticus vel terrestris 2 m altus. Petioli 5-9 mm longi. Lamina coriacea, integra, elliptica vel ovato-elliptica, (4.5) 8-12 (13.5) X (2.5) 3-7 cm, apice acuminata, basi rotundata et

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FIc. 6. Vaccinlum je#nse. A. Habit, X 0.5. B. Flower with pedicel partially drawn, X 3.5. C. Stamen, ventral view, X 5.5. D. Stamen, lateral view, X 5.5. A drawn from Mori et al. 6910, B-I) drawn from Luteyn & Kennedy 3955.

cordata, pinnafinervia. Infiorescentia racemosa vel subcorymbosa; rhachis 1-1.5 cm longa, glabra vel sparse hirsuta, dense glanduloso-fimbriata; bracteae florales lineari-lanceolatae, 3-9 mm Iongae; pedicelli dense glanduloso-fimbriati, 15 mm longi. Calyx 7-9 m m longus, sparse hirsutus; tubus 2-2.5 mm longus; limbus campanulatus 5.5-7 mm longus lobis inclusis; lobi 5-6 toni longi. Corolla cylindrica, sparse hirsuta, 11-12 mm longa. Stamina 8 mm longa; filamenta 3 mm longa, dense sericea; antherae 7 m m longae; tubuli 4 m m longi. Stylus 9-10 ram longus, glaber.

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Viney epiphyte or arching terrestrial shrub to 2 m. M a t u r e stems green becoming reddish-brown when living but drying brown, subterete, glabrous; bark thin, cracking into longitudinal strips. I m m a t u r e branches and twigs of new growth subterete, glabrous to densely hirsute distally. Leaves coriaceous, entire, elliptic or ovateelliptic, (4.5) 8-12 (13.5) • (2.5) 3-7 cm, basally rounded and cordate, apically acuminate, dark green above and paler beneath when fresh but drying olive- to grayish-brown, glabrous above and sparsely to moderately hirsute beneath along the principal veins, lower surface also with numerous minute glandular fimbriae; pinnately veined with 3 pairs of lateral nerves, the midrib with lateral and secondary veins impressed above and raised beneath causing the blades to be bullate, proximal 1-2 cm of midrib conspicuously thickened above; pseudostipules (bud scales) lanceolate, subulate, 3 mm long, hirsute. Petiole terete, rugose, hirsute becoming glabrous, 5-9 mm long and 2-2.5 mm in diameter. Inflorescence axillary but often appearing terminal, racemose or subcorymbose, 8-17 flowered, wholly green when living (including rachis, pedicels, bracteoles, calyx, and corolla); rachis 1-1.5 cm long, subterete, reddish-brown, glabrous or sparsely hirsute, densely glandular-timbriate. Floral bracts linear-lanceolate, 3-9 • 1 mm, hirsute, glandular-fimbriate, conspicuously nerved. Pedicels terete, striate, sparsely hirsute, densely glandularfimbriate, 15 x 1 mm but extending to 26 mm in length after anthesis, articulate with the calyx. Bracteoles linear-lanceolate, 5-8 • 0.5 mm, hirsute, glandular-timbriate. Calyx 7-9 mm long, sparsely to moderately hirsute and glandular-fimbriate; tube cylindric, 10-ribbed, slightly angled opposite the sinuses, 2-2.5 • 2-3 mm; limb campanulate, with lobes conspicuously nerved, 5.5-7 mm long including the lobes; lobes oblong or ovate, acute, often basally slightly imbricate, 5-6 • 1.75-3 mm. Corolla cylindric, sparsely hirsute and glandular-fimbriate (especially on surface of lobes), 11-12 • 5.5-7.5 mm; lobes triangular, 3 • 3 mm. Stamens 10, 8 mm long; filaments distinct, 3 mm long, basally widened, marginally densely sericeous especially onto the connective, translucent when fresh; anthers • 7 mm long, strongly curved inwards at the base; thecae granular, 3 mm long, orangish-brown when fresh; tubules 4 mm long, sparsely pilose, dehiscing by laterally oriented apical slits - 1.5 mm long, yellowish-brown when fresh. Style included, glabrous, 9-10 mm long. TYPE: PANAMA. PANAMa: at the top of Cerro Jefe, 1000 m elev., L u t e y n & K e n n e d y 3955 (HOLOTYPE: DUKE; ISOTYPES: MO, NY). Vaccinium je/ense is an unusual species which cannot be closely associated with any other Neotropical Vaccinium. It is similar to V. fioccosum (Panama), V. poasanum (Guatemala-Panama), V. costaricense and V. orosiense (both from Costa Rica), and V. racemosum (West Indies) in the short racemose, umbellate to subcorymbose inflorescences with green or yellowish corollas; however, it differs most significantly in its pinnately veined leaves, glandular inflorescence, elongate bracteoles, and anther tubules which dehisce through latrorse clefts. This lateral dehiscence seems to be unique in the genus and may eventually provide the basis for recognition of a new section within Vacdnium. From V. /loccosum, V. costaricense and V. orosiense, the new species further differs in its longer calyx lobes, internally glabrous corolla, and anther tubules nearly twice as long. Additional specimens examined. PANAMA. PANAMa: una milla depu~s de La Eneida cerca de Cerro Jefe, • 750 m, Correa & Dressler 955 (DUKE, MO, PMA); La Eneida near Cerro Jefe, Dressler 3541 (PMA) ; path beyond the village of La Eneida at ___ 1000 m, Luteyn & Foster 1124 (DUKE, F, MO, US); cloud forest along trail on ridge running NE from summit of Cerro Jefe • 1000 m, Mori & Kallunki 6080 (DUKE); premontane rain forest in vicinity of Altos de Pacora, 15-20 km WNW of Cerro Azul, • 800 m, Mori, Dumont & Carpenter 6910 (MO) ; cloud forest on Cerro Jefe at • 100o m, Mori 7110 (MO); top of Cerro Jefe, Tyson, Dwyer & Blum

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4344 (DUKE, MO, SCZ); new road leading N from summit of Cerro Jefe, Witherspoon 8558 (DUKE, MO). Vaceinium

o r o s i e n s e Wilbur & Luteyn, sp. nov.

Arbuscula epiphytica vel terrestris, 1-2 malta. Folia obscure 3-5-plinervia, coriacea, ovata vel ovato-lanceolata, apice acuta et basl cordata vel rotundata, (2) 4-9 (14) • (1.5) 3-5 (7) cm; petioli ___ 1 mm longi, glabri. Inflorescentia axillaris, umbellato-racemosa, 2-4 em longa; bracteae triangulares, 1-2.2 mm longae; pedicelli 1.5-2.5 cm longi, glabri; bracteolae 0.8-1.5 mm longae. Hypanthium cum pedicello articulatum, campanulatum, 2-3 )< 2.5-4 ram, glabrum; lobi calycis 5, triangulares, 0.8-1.2 mm longi, acuti, glabri. Corolla cylindrica vel urceolata, 6-8 mm longa, intus pilosa; lobi corollae 5, triangulates, 1-1.2 mm longi. Stamina 10; filamenta 3 mm longa, villosa; thecae granulosae, ~ 1.5-1.8 mm longae; tubuli • 2 mm longi. Epiphytic or terrestrial shrub 1-2 m tall with olivaceous to purplish, glabrous branchlets. Leaves coriaceous, ovate to ovate-lanceolate, basally rounded to more typically cordate and amplexicaule, apically acute, (2) 4-9 (14) • (1.5) 3-5 (7) cm, entire, marginally slightly thickened, glabrous to sparsely and inconspicuously appressed, glandular-strigillose. Petioles - 1 mm long, glabrous. Inflorescence axillary (but often superficially appearing terminal), umbelliform raceme; rachis glabrous, 1-1.5 cm long. Floral bracts persistent, scale-like, triangular to ovate-lanceolate, 1-2.2 mm long, glabrous to minutely ciliate. Pedicels glabrous, 1.5-2.5 cm long. Bracteotes subopposite, appressed, persistent, deltoid to lance-ovate, 0.8-1.5 mm long, glabrous to inconspicuously ciliate, located in the lower quarter of pedicel. Hypanthium clearly disarticulating from the pedicel by means of a sharply delineated groove, campanulate to urceolate, • 2-3 • 2.5-4 ram, glabrous; calyx limb --x- 0.5-0.8 mm long; calyx lobes 5, triangular, acute, 0.8-1.2 mm long. Corolla cylindric to urceolate, 5-8 • 5-7 ram, yellowish-green often suffused with maroon, externally glabrous and internally moderately to densely pilose especially on the lobes and the upper half of the tube with white trichomes; corolla lobes 5, spreading or reflexed, triangular, acute, 1-1.2 mm long. Stamens 10; filaments flattened, • 3 mm long, weakly adherent to the very base of the corolla tube and attached medially to the anther sac, villous; anther sacs (thecae) granular, • 1.5-1.8 m m long, golden, basally inwardly curved and with an apiculate tip; tubules stout, • 2 • 0.5 m m each with a slightly introrsely oblique terminal pore. T~PE: COSTA RICA. CARTACO: steep roadside slopes on property of I C E hydroelectric plant • 15-20 km beyond suspension bridge over Rio Grande de Orosi, elev. 1300-1800 m, Luteyn, Wilbur & K. Utley 4411 (HOLOTYP~: D U K E ; ISOTYP~S: CR, F, MO, NY, US). Additional specimens examined. COSTA RICA. CARTACO: wet roadbank at Tapanti hydroelectric project, Utley 3263 (CR, DUKE, F, MO, NY) ; road from Tapanti to Taus and Tausito "+ 3 km NE of Rio Grande de Orosi at Tapanti, 1300-1500 m, Utley 5041 (DUKE, F, NY). Vaecinium

racemosum

(Vahl) Wilbur & Luteyn, comb. nov.

tfornemannia racemosa Vahl, Skr. Nat.-Selsk. Kj6b. 6: 120. 1810. Symphysia racemosa (Vahl) Stearn, Taxon 21: 111. 1972. A more detailed s y n o n y m y of this West Indian species can be found in the papers by Smith (1935) and Stearn (1972) and will not be repeated here. I t might seem premature to submerge Symphysia in Vaccinium since the range of diversity within that genus at present suggests that the much needed revisionary studies probably will result in its wholesale dismemberment. Until the world-wide revision is completed, however, it would seem prudent not to dismember the genus on a regional basis as has frequently been its past fate. Clearly there seems to be no way in which such species

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as the Central American Vaccinium poasanum Donnell Smith can be generically sepa r a t e d from either the recently described P a n a m a n i a n Symphysia floccosa (L. O. Williams) L. O. Williams, or the W e s t I n d i a n Symphysia racemosa (Vahl) Stearn, or a t least two of the three Costa Rican species described in this paper. Therefore for t e m p o r a r y convenience we are placing all of them in Vaccinium. Sleumer (1941) t r e a t e d Vaccinium poasanum Donnell Smith as the t y p e of a monotypic Vaccinium sect. Oreades Sleumer. Klotzsch (1851, p. 60) established a monotypic Vaccinium subg. Scytanthemum Klotzsch for V. imrayi Hooker, a later synonym of Symphysia racemosa (Vahl) S t e a m or V. racemosum. U n f o r t u n a t e l y in the same paper Klotzsch maintained two taxonomic synonyms of this same species as the only members of the genus Symphysia. ACKNOWLEDGMENTS W e would like to express our thanks to Page R. Allen and Gabrielle R a u m b e r g e r for preparing the illustrations, and to R u p e r t B a r n e b y and A r t h u r Cronquist for their critical and helpful review of the manuscript. LITERATURE CITED

Sleumer, H. 1941. Vaccinioideen-studien. Bot. Jahrb. Syst. 71: 375-510. Smith, A. C. 1935. Hornemannia, a much-described West Indian genus. Amer. J. Bot. 22: 9-11. Steam, W. T. 1972. The generic name Hornemannia and its diverse applications. Taxon 21: 105-I 1I. Wilbur, R . L . 1974. The Central American species of the genus Disterigma (Ericaceae: Vaceinieae). Bull. Torrey Bot, Club I 0 1 : 245-249.