lELDIANA Botany NEW SERIES, NO. 45 FLORA COSTARICENSIS William Burger, Editor Family #172 Ericaceae James L. Luteyn Robert L. Wilbur December 21, 2005 Publication 1536 PUBLISHED BY FIELD MUSEUM OF NATURAL HISTORY ! seed plants knoun or expected to occur in fosta Rica and adjacent areas, listed alpha- beticalK ,ind numbered according to the sequence of Hnglers Syllahu* ( )laeaceae < )lc.iccac Onagraceae Opiliaccae Orchiclaceae Orohaiichaeeac ( Kalidaceae Palmae Papaveraceae Passitloraceae Pedaliaceae Phytolaccaceae Pinaceae Piperaceae Pyrolaeeae Plantaginaceae Plumbaginaceac Podocarpaceae Podostemonaceae Polemoniaceae Polygalaceae Polygonaceae Pontederiaceae Portulacaceae Potamogetonaceae Primulaceae Proteaceae Punicaceae Quiinaceae Raffiesiaceae Ranunculaceae Resedaceae Rhamnaceae Rhizophoraceae Rosaceae Rubiaceae Rutaceae Sabiaceae Salicaceae Sapindaceae Sapotaceae Saxifragaceae Scrophulariaceae Simarubaceae Solanaceac Staphyleaceae Sterculiaceae Styracaceae Symplocaceae Taxaceae Theaceae Theophrastaceae ThymelaciK Tiliaceae Tovariaceae Trigoniaccae Triundaccae Tropacolaccae runicraceac Typhaceac I Iniaceac I nibelli ferae I rlicaceac V'alerianaceae X'cllo/iaccac \crbcnaceae \ lolaceae N'uaceae \och\ siaceae \\ridaceae /ingibeniccae Zygophyllaceae ^ The paper used in this publication meets the requirements of ANSI/NISO Z39.48-1992 (Permanence of Paper). FIELDIANA Botany NEW SERIES, NO. 45 FLORA COSTARICENSIS William Burger, Editor Family #172 Ericaceae •,' James L. Luteyn ' • ; •'. -••:.- . : Mary Flagler Cary Curator of Botany and Senior Curator Institute of Systematic Botany The New York Botanical Garden Bronx, New York 10458-5126 Robert L. Wilbur Department of Botany Duke University Durham, North Carolina 27706 Accepted July, 2004 Published December 21, 2005 Publication 1536 NATURAL HISTORY SURVEY DEC 2 9 2005 NATURAL HISTORY SURygy OCT ? 2006 *Y PUBLISHED BY FIELD MUSEUM OF NATURAL HISTORY Abstract A treatment of the 21 genera, 93 species, one subspecies, and four varieties of Ericaceae cur- rently known from within the political borders of Costa Rica as well as those taxa currently known from the adjacent Chiriqui highlands of western Panama is herewith presented. Seventy-one of the 93 total species, the one subspecies, and three of the four varieties are found in Costa Rica, of which 19 species, the one subspecies, and two of the four varieties are endemic there. The conser- vation status of each species is given based on 35 years of collecting experience and the numbers of herbarium specimens extant. Habitats, habits, field characters, keys, illustrations, and discussions of species relationships are given as aids to identi- fication. Resumen Se presenta un tratamiento de los 21 generos, 93 especies, una subespecie y cuatro variedades de Ericaceae conocidos dentro del limite geogrdf- ico de Costa Rica, asi como tambie"n los taxones de las tierras altas adyacentes a Chiriqui en la par- te oeste de Panama. Setenta y una de un total de 93 especies, la subespecie y tres de las cuatro var- iedades se encuentran en Costa Rica. De estas, 19 especies, la subspecie y dos de las cuatro varie- dades son ende"micas alii. Se presenta el estado de conservaci6n de cada especie basado en 35 afios de experiencia colectando en el campo y en el niimero de muestras de herbario. Se dan hdbitats, hdbitos, caracteristicas de la planta en el campo, claves, ilustraciones y discusiones sobre las rela- ciones entre las especies como guias para la iden- tificaci6n. Introduction This is the twelfth issue in the Flora Costari- censis series. The first dealt with the Piperaceae, family number 41 (Fieldiana, Bot. 35, 1971). The second included families numbered 42 through 53, Chloranthaceae through Urticaceae (Fieldiana, Bot. 40, 1977). The third issue covered the Po- aceae (Fieldiana, Bot. n.s. No. 4, 1980). The fourth issue included families numbered 54 through 70, Podostemaceae through Caryophyl- laceae (Fieldiana, Bot. n.s. No. 13, 1983). The fifth issue covered families 200 and 201, the Acanthaceae and Plantaginaceae (Fieldiana, Bot. n.s. No. 18, 1986). The sixth issue included fam- ilies 80 and 81, Lauraceae and Hernandiaceae (Fieldiana, Bot. n.s. No. 23, 1990). The seventh issue included families numbered 97 through 103, Krameriaceae through Zygophyllaceae (Fieldiana, Bot. n.s. No. 28, 1991). The eighth issue included family 202, the Rubiaceae (Fieldiana, Bot. n.s. No. 33, 1993). The ninth issue included family 113, the Euphorbiaceae (Fieldiana, Bot. n.s. No. 36, 1995). The tenth issue covered Tribe Maxil- larieae of family 39, the Orchidaceae (Fieldiana, Bot. n.s. No. 40, 1999). The eleventh issue in- cluded families numbered 193 through 197, Scro- phulariaceae through Orobanchaceae (Fieldiana, Bot. n.s. No. 41, 2000). In keeping with the original intent of Flora Costaricensis and for the biogeographical reasons discussed here, this treatment of Ericaceae in- cludes both those taxa currently known from within the political borders of Costa Rica as well as those taxa currently known from the adjacent Chiriqui highlands of western Panama. Many of the species here described and discussed are il- lustrated by means of color photos on Luteyn's Web site given in the "References" section be- low. In Figures 3-15, leaves and leafy stems are drawn to the same scale throughout. Enlarged flowers and fruits are drawn to the same scale on an individual plate unless otherwise noted. The closed scales represent centimeters, and the open scales represent millimeters. The figures are somewhat diagrammatic and repfesent the artist's (William Burger) concept of a common or char- acteristic morphology. Acknowledgments The authors wish to express our grateful ac- knowledgment to the National Science Founda- tion, whose support over the years made this study possible. J.L.L. wishes to thank The Field Museum of Natural History (Chicago) for finan- cial aid from the Robert O. Bass Visiting Scientist Fund that enabled him to work at the Field Mu- seum. Obviously a study such as the present one would be impossible without the cooperation of the curators of the many herbaria who loaned their collections for this investigation. We are very much indebted to the curatorial staff of the fol- lowing herbaria: A, AAU, BM, CAS, CR, CU, DAY, DS, DUKE, F, GB, GH, INB, K, L, LL, MICH, MIN, MO, MSC, NY, P, PH, PMA, POM, RSA, S, SCZ, TEX, US, WIS. We wish to thank the Museo Nacional de Costa Rica, the Institute Nacional de Biodiversidad (INBIO), the Field Museum of Natural History (Chicago), and the Missouri Botanical Garden for their assistance in local collecting programs and their staff and fa- cilities over the many years the writing of the manuscript has spanned. Bill Haber and Willow Zuchowski provided logistical support to J.L.L. during trips to Monteverde. For encouragement, profitable discussions, and the artistic talents of Bill Burger, which have helped greatly to embel- lish this work, we are grateful. Lucia Kawasaki helped in various aspects of the manuscript and logistically. Reviewer's comments on the manu- script are appreciated. J.L.L. wishes to thank the Missouri Botanical Garden for providing facilities to complete this manuscript. Bobbi Angell's plates (Figs. 1-2) are beautiful, as always. VI FLORA COSTARICENSIS Family #172 Ericaceae ERICACEAE By James L. Luteyn and Robert L. Wilbur REFERENCES — K. A. Kron, W. S. Judd, P. F. Ste- vens, D. M. Crayn, A. A. Anderberg, P. A. Gadek, C. J. Quinn, & J. L. Luteyn, Phylogenetic classi- fication of Ericaceae: Molecular and morpholog- ical evidence. Bot. Rev. 68: 335-423. 2002. J. L. Luteyn, Key to the subfamilies and genera of neotropical Ericaceae. Nord. J. Bot. 11: 623-627. 1991. J. L. Luteyn, Ericaceae, in G. Marling & L. Andersson (eds.), Flora of Ecuador 54: 1-404, I-VIII. 1996. J. L. Luteyn, "Neotropical blue- berries: The plant family Ericaceae." http:// www.nybg.org/bot/res/lut2. 1998. J. L. Luteyn, Diversity, adaptation, and endemism in Neotrop- ical Ericaceae: Biogeographical patterns in the Vaccinieae. Bot. Rev. 68: 55-87. 2002. J. L. Lu- teyn & R. L. Wilbur, Ericaceae, in Flora of Nic- aragua 1: 822-831. 2002. P. C. Standley & L. O. Williams, Ericales, in Flora of Guatemala. Fiel- diana, Bot. 24 (pt. 8): 73-127. 1966. P. F. Stevens, J. L. Luteyn, et al., Ericaceae, in K. Kubitski (ed.), Families of Flowering Plants, vol. VI: 145-194. 2004. R. L. Wilbur & J. L. Luteyn, Ericaceae, in R. Woodson et al., Flora of Panama. Ann. Mis- souri Bot. Card. 65: 27-144. 1978. Terrestrial or epiphytic shrubs, subshrubs, perennial herbs, or fleshy achlorophyllous mycotrophs (Monotro- pa), sometimes lianoid or rarely trees, often rhizoma- tous, sometimes arising from lignotubers (spherical to oblong, swollen hypocotyls, 0.4-1 m diam., little vas- cular tissue, mostly parenchyma); indumentum mostly of simple, unicellular hairs, also with stout, multicellular, multiseriate, usually glandular, brownish-red, clavate to capitate hairs (these sometimes fusing as fimbriae, then occurring on mature stems, leaves, or pedicels and the organ referred to as fimbriate); apical bud usually abort- ing, especially in Vaccinioideae; axillary buds scaly (outer pair elongated and rarely appearing stipular). Leaves alternate (usually spiral, rarely distichous), rarely (our species) opposite or pseudo-opposite, verticillate, whorled (pseudo-whorled- in Chimaphila), or reduced and then represented by bract-like scales (Monotropa), simple, usually petiolate, exstipulate, the blade coria- ceous to membranous, evergreen to deciduous, the mar- gin usually entire but sometimes serrulate-crenate, the venation plinerved (with a clear midrib and several strong lateral nerves) or pinnate; leaf scars with a single vascular bundle scar, the nodes with one trace and one gap (our species). Inflorescences axillary, rarely termi- nal, racemose, paniculate, fasciculate, or flowers soli- tary; floral bract 1, subtending pedicel of each flower, small or large, showy or inconspicuous, deciduous or persistent; pedicels articulate or continuous with calyx; bracteoles usually 2, usually smaller than floral bracts, borne along length of each pedicel. Flowers bisexual (rarely functionally unisexual; plants dioecious in some species of Pernettya in temperate South America), ra- dially symmetrical or slightly bilaterally symmetrical, mostly (3) 5 (7)-merous, typically obdiplostemonous, hypogynous, or epigynous and with a typically biseriate perianth; calyx synsepalous at least basally, in superior ovaries the sepals are distinct or almost distinct, imbri- cate, sometimes grading into bract-like scales (Monotro- pa), rarely fleshy and accrescent to the fruit (Gaulthe- ria), in inferior ovaries the proximal (or inferior) part of the calyx that is fused with the ovary wall is called the "calyx tube" (previous Luteyn papers often called this part the hypanthium), which is basally rounded to trun- cate or sometimes outwardly flaring and then said to be apophysate, the distal free part (of the calyx) that is borne on top of the ovary is the "calyx limb," and the portions of the limb that are distinct from one another are the "calyx lobes"; corolla sympetalous, lobed, aes- tivation imbricate or valvate, membranous to carnose, unistratose or bistratose, commonly cylindric, campan- ulate or urceolate (petals distinct in Bejaria, Monotro- pa); stamens (see Figs. 1-2) (6) 10 (14), in 2 whorls, equaling to shorter than corolla overall, equal or alter- nately unequal within themselves, twice as many as the petals (rarely as many), arising from base of ovary and then sometimes alternating with lobes of nectaries (su- perior-ovaried genera), or borne on edge of a nectarifer- ous disc (inferior-ovaried genera); filaments equal or al- ternately unequal, distinct or connate, usually straight (rarely slightly S-shaped), ligulate (sometimes broad- ened and papillose at base); connectives short and in- conspicuous in superior-ovaried genera, or thick and raised in inferior-ovaried genera, bearing white, disin- tegration tissue powder in small, elongated pouches along length and onto awns (Comarostaphylis, Gaulthe- ria, Pernettya), sometimes spurred (Comarostaphylis, Psammisia, Vaccinium); anthers inverting during devel- opment, the thecae oblong, bilocular, basally rounded or apiculate, sometimes bearing terminal awns (Gaultheria, Pernettya), in the superior-ovaried genera thecae dehisce FIELDIANA: BOTANY, N.S., NO. 45, December 21, 2005, PP. 1-107 terminally or by obliquely introrse pores (Bejaria, Gaul- theria, Pernetiya), or longitudinally (Monotropa), in all the inferior-ovaried genera each theca narrows apically into a hollow tubule through which pollen escapes, the tubules distinct or connate and dehiscing introrsely or rarely latrorsely by oblique pores or short to elongate clefts; pollen grains in tetrahedral tetrads (rarely single in Monotropa), sometimes with viscin threads (Bejaria); pistil single, the ovary superior or inferior, 4-5 (10)- carpellate, usually with as many locules as carpels or with twice as many locules as carpels (rarely loculate in lower portion and 1-locular above); placentation axile (rarely intruded parietal); ovules numerous (rarely soli- tary), anatropous to campylotroppus with a single integ- umentary layer; style usually arising from a slight de- pression, single, fluted, hollow; stigma simple (occa- sionally weakly lobed in Monotropa); nectary present at base of ovary in superior-ovaried genera and obscure (Gaultheria, Pernettya), or as 5 (Chimaphila) or 8-10 lobes (Monotropa) projecting from between filament ba- ses, or as a continuous annular or cup-shaped disc cov- ering entire top of ovary in inferior-ovaried genera. Fruit loculicidal or septicidal capsules, berries, or drupes, with a usually persistent, rarely accrescent and fleshy calyx (Gaultheria); seeds small, usually numer- ous, sometimes winged or tailed (Bejaria, Chimaphila), sometimes covered by clear, mucilaginous sheath; testa thin with elongated or isodiametric cells; endosperm fleshy; embryo straight, white or sometimes green (Sphyrospermum, Themistoclesia). Ericaceae s.l. include about 125 genera and 4500 species worldwide. Except for Antarctica, the family is cosmopolitan, inhabiting temperate regions of the world and montane areas in tropical latitudes (Luteyn, 2002). Concentrations of high species diversity occur in the tropics of both hemispheres, the Himalayas, Australia, and the Cape Region of South Africa (Kron & Luteyn, 2005). In the Neotropics, overall species richness increases near the Equator, with the highest num- bers in Colombia (ca. 300 spp.) and Ecuador (ca. 230 spp.). Forty-seven genera (ca. 70% endemic) and approximately 900 species (ca. 95% endemic) are native to the Neotropics. In this floristic treat- ment of Ericaceae for Costa Rica, we include a total of 2 1 genera with a total of 93 species, one subspecies, and four varieties. Seventy-one of the 93 total species, the one subspecies, and three of the four varieties are found in Costa Rica. Nine- teen of those 93 species, along with the one sub- species and two of the four varieties, are currently endemic to Costa Rica. Another 22 species and one variety (including Bejaria aestuans and Or- thaea panamensis) have yet to be found within the political boundaries of Costa Rica, although they do occur in the adjacent Chiriquf highlands of western Panama. We do not apologize for in- cluding this seemingly large number of "yet to be discovered" taxa in this treatment of the Erica- ceae of Costa Rica because similar life zones and habitats are present there, and, perhaps more im- portantly, the Caribbean slopes of the Cordillera de Talamanca are so poorly known botanical ly. In recent years, at least 23 species of Ericaceae, pre- viously thought to be endemic to either Costa Rica or the western Chiriquf highlands of Panama, have now been collected in western Panama or eastern Costa Rica, respectively (13 species from the Costa Rican Talamancas are now found in the adjacent western Chiriquf highlands; 10 species from the western Chiriquf highlands are now found in the eastern Costa Rica Talamancas). Fur- thermore, geologically, the western highlands of Panama, including the uplands of Bocas del Toro and Chiriquf Provinces, and parts of the newly established Comarca Ngobe-Bugle (formerly the Caribbean slopes of eastern Chiriquf Prov.), rep- resent the easternmost extension of the Cordillera de Talamanca, which originates in central Costa Rica. At the same time, the western highlands of Panama represent the easternmost distribution point for many plant species in Mesoamerica. Therefore, we anticipate that most, if not all, of these additional 23 taxa actually occur in eastern Costa Rica and that more intensive botanical ex- ploration on the Caribbean slopes of the Talaman- cas will reveal them. Two species of Satyria in- cluded within this treatment (Satyria sp. "A" and "B") are considered as "morpho" species and have yet to be formally described. Rhododendron simsii, a native to China and sometimes encoun- tered as a cultivated plant at higher elevations, is keyed but not described. The family Ericaceae is economically important in temperate regions of the world, primarily for those genera of horticultural (Erica, Rhododen- dron) and commercial food crop (Vaccinium) us- age. In the Neotropics, fruits of Vaccinium flori- bundum and Macleania rupestris are rarely found in local and commercial markets, although they are used in pies, jams, and drinks. Ecologically, the Ericaceae are usually found in the moist habitats of cool montane forests be- tween 1000 m and 3000 m elevation; in well- drained, acid soils; and where there is ample light and abundant, regularly distributed precipitation throughout the year — habitats that are also some of the most threatened ecosystems in the world (Mast et al., 1999; Knapp, 2002). They are often associated with mycorrhizal fungi and are fre- quently found as pioneers following volcanic, landslide, or road-building activities or as part of FIELDIANA: BOTANY the edge community around mature forests (Lu- teyn, 2002). The evergreen, coriaceous leaves of many Ericaceae provide an excellent substrate for mosses, hepatics, lichens, and other fungi. Also, glands and glandular areas on the mature stems, leaf surfaces, and inflorescences are often covered by fungal mycelia that probably derive nutrients from the exudates. Pollination is typically by bees in the superior- ovaried genera and by hummingbirds in the in- ferior-ovaried genera. The floral morphology of the inferior-ovaried genera, the "blueberries," in- dicates that many have a close evolutionary re- lationship with hummingbirds (Luteyn, 1998; Lu- teyn & Sylva S., 1999). Corollas of the blueber- ries house tiny, translucent mites (<1 mm long) of the genus Rhinoseius, which breed and feed in the flowers and are dispersed in the nasal cavities of hummingbirds (Col well, 1973, 1985; Nas- krecki & Colwell, 1998). In the Neotropics, the berry fruits and abundant sugar-rich nectar pro- vide sources of food for many animals. The two most important dispersal agents of Ericaceae at Monteverde (Costa Rica) include the Emerald Toucanets and Black-faced Solitaires (G. Murray, pers. comm.). Based on a combination of morphological and molecular data, the families Monotropaceae and Pyrolaceae (both in Costa Rica) and Empetraceae and Epacridaceae recognized as distinct by Takh- tajan (1980), Cronquist (1981), and others are now included within the Ericaceae sensu lato (see Anderberg, 1992, 1993; Judd & Kron, 1993; Kron & Chase, 1993; Kron, 1996; Kron et al., 2002a; Powell & Kron, 2003; and Stevens et al., 2004 for data and detailed discussions). However, generic limits in the tribe Vaccinieae are still poorly un- derstood (Luteyn & Wilbur, 1997; Luteyn, 2001; Kron et al., 2002a,b). The family is still revealing new species and new records for Costa Rica with many collecting efforts yielding novelties, emphasizing large gaps yet to be filled in our knowledge of neotropical families in general and of the floristic composition of this region. Contrary to some recent views that fieldwork in comparatively well-collected coun- tries such as Costa Rica and Panama is no longer necessary, this treatment points out once again the absolute need for field collecting and basic her- barium research. Furthermore, in view of the well- publicized destruction of the rain forest, it also demonstrates the need for continued field work in montane as well as lowland regions. Inventory work at the primary level followed by revisionary and monographic studies by specialists is still des- perately needed, and actually its support must in- crease if we are to claim a true knowledge of trop- ical ecosystems. In an effort to alert scientists and the general public as to the conservation status of the Ericaceae in Costa Rica and the adjacent high- lands of western Chiriquf Province, Panama, we have given our own personal assessment of each species herein treated based on each of our ap- proximate 35 years of collecting experience in Costa Rica and Panama, combined with the num- bers of herbarium specimens extant. The follow- ing five status categories are recognized: Endan- gered = species known from six or fewer collec- tions or only a few localities that are threatened by human intervention; Rare/Infrequent = spe- cies known from 7-12 collections but in only a few, scattered localities; Locally Common = spe- cies known from more than 12 collections but in only a few, scattered localities; Widespread = species known from many geographic localities but few collections; Common = species known from many collections and many geographic lo- calities. In Costa Rica and adjacent western Panama, Ericaceae are relatively easily recognized by the following field characters: generally woody, sun- loving shrubs, whether terrestrial or epiphytic; leaves are spiral, lacking stipules, with blades that are usually thin and red when young but coria- ceous and dark green when mature, the leaf ve- nation is often strongly plinerved, the leaf mar- gins are usually entire; nodes are unilacunar with one gap and one trace (use 10X lens); indumen- tum is rarely conspicuous; inflorescences are ax- illary but often appear terminal, the flowers are found in racemes, fascicles or are solitary, the pedicels are bibracteolate and articulate or contin- uous with the calyx; flowers are pendulous, per- fect, actinomorphic, usually 4- to 5-merous, odor- less; calyx has the sepals usually connate and per- sistent; corolla is often fleshy, usually of united petals, typically urceolate, cylindrical or some- times campanulate, often waxy and nitid, often with the refreshing taste of oxalic acid (at least in the inferior-ovaried genera), the colors of the co- rolla proper often contrast with those of the lobes or floral bracts; stamens are usually twice as many as the petals and free from them, the anthers are sometimes abaxially appendaged with awns or spurs and dehisce by apical pores or clefts often on the ends of long narrow tubules; nectar is usu- ally abundant, sweet and viscid in the base of the corolla; ovary is superior or inferior with axile LUTEYN AND WILBUR: FAMILY #172 ERICACEAE placentation, the ovules are usually many per car- pel, the style is hollow and usually impressed into the ovary, the stigma is truncate; fruits are brown and woody capsules, blue-black or rarely trans- lucent white berries, or rarely blue-black drupes, the seeds are small, numerous and golden in color while the embryo is white or less commonly green. In general, it is easy to find Ericaceae in Costa Rica. In cool, moist, montane regions they are good pioneers and opportunists, so the best places to find them are along roadside slopes, forest edg- es, and in remnant trees in pasturelands. Newly exposed land, whether due to man-made opera- tions such as road building or forest clearing, or natural events such as volcanic activity, land- slides, or tree falls, all provide Ericaceae with open to partially shaded, sparsely inhabited sites, where pollinating vectors (bees, hummingbirds) also have easy access. In terrestrial situations such as these, the shrubby plants often dominate and flourish for many years until they become over- grown, at which time they disappear or persist in lower numbers without flowering, having been lit- erally shaded out or outcompeted by more ag- gressive vegetation. Their vitality is periodically renewed after road crews cut back the dense veg- etation along road banks, thus again providing necessary sunlight for growth and flowering. A beautiful example of this situation may be seen along the interamerican highway above Cartago, where starting at about 1 200 m the showy red and white flowers and bracts of Cavendishia bracteata are found and then, at a little higher elevation, populations of Macleania rupestris present a spectacular floral display among roadside bank vegetation. Also, a more natural and historic ex- ample of this terrestrial situation may be seen at the summit of Volcan Irazu, where three species of Ericaceae (Comarostaphylis arbutoides, Per- nettya prostrata, Vaccinium consanguineum) were among the first pioneers to reestablish veg- etation there after an eruption in 1963; by 1970 they were dominant. In epiphytic situations, new- ly exposed sites are often provided by forest clear- ing or tree falls, thus providing the required amount of sunlight to lower branches or remnant trees for ericads to grow and flower. It should be emphasized here that epiphytic Ericaceae are not common within a low-light forest or on young branches of the canopy, but instead they prefer exposed to partially shaded situations on older and larger branches with usually rough bark. Also, in low-light situations within the forest, they usually become lianoid and do not flower. Both natural and disturbed open sites may be dry be- cause of increased exposure and strong winds, and here Ericaceae may supplement their moisture needs by rooting in waterlogged rotting tree stumps like remnant trees in pasturelands or in dense and water-retaining bryophyte layers; wa- ter-storing lignotubers are also quite common (for additional explanation and examples, see Luteyn, 2002). It is interesting that recent collecting activities, especially those made under the auspices of the INBIO biodiversity of Costa Rica program, have uncovered many new species or new records of Ericaceae in "lowland" (not montane) habitats below 700 m elevation. These include Cavendi- shia herrerae, C. limonensis, C. linearifolia, C. osaensis, Didonica pendula, Satyria species "A," Sphyrospermum ellipticum, and Vaccinium furfur- aceum. This strongly suggests that there may be more species to be found in the wet, lowland rain forest and that our traditional characterization of this family as montane may need to be revised. Although many of the commonly encountered Costa Rican Ericaceae have large and showy flowers to 2-3 cm long (e.g., Cavendishia brac- teata, Macleania rupestris, Psammisia ramiflora, Satyria warszewiczii) or large floral bracts to 4 cm long with contrasting colors (e.g., Cavendishia atroviolacea, C. bracteata, C. callista, etc.), quite a few also have small and inconspicuous flowers with minute, green, floral bracts and are therefore poorly collected (e.g., Themistoclesia spp.). Oth- ers occur as epiphytes high in old trees (e.g., Themistoclesia pentandra and Utleya costaricen- sis). One plant, Disterigma utleyorum, may be rarely collected because of its association with ag- gressive, biting ants. The mycotrophic Monotropa uniflora and herbaceous subshrub Chimaphila maculata are easily overlooked in the forest un- derstory under the best of conditions but are now rare and almost certainly endangered because their oak forest habitat is nearly completely de- stroyed in areas accessible to humans. A few spe- cies are rarely collected not only because they are truly very local and therefore rare in their occur- rence but also because quite a few have small, green flowers (e.g., Anthopterus costaricensis and A. revolutus, Didonica pendula, Lateropora ova- la, Themistoclesia costaricensis and T. horqueten- sis, Vaccinium costaricense, V. floccosum, V. oro- siense, and V. poasanum). Sphyrospermum dissi- mile, on the other hand, is infrequently collected because it has small, dark brick-red corollas that FIELDIANA: BOTANY FIG. 1. Stamens from the genera of Costa Rican Ericaceae. (A) Monotropa uniflora (Luteyn ft al. 1 1685, NY). (B) Chimaphila maculata (Davidse 25713, NY). (C) Bejaria aestuans (Luteyn & Lebrfin-Luteyn 11415, NY). (D) Comarostaphylis (drawn from C. discolor subsp. discolor, Luteyn et al. 1 1616, NY). (E) Gaultheria erecta (Luteyn et al. 11607, NY). (F) Pernettya prostrata (Luteyn et al. 7389, NY). (G) Satyria panurensis (Luteyn et al. 15247, NY). (H) Orthaea panamensis (Luteyn 14863, NY). (I) Cavendishia fortunensis (Luteyn 14824, NY). (Illustrations by Bobbi Angell) LUTEYN AND WILBUR: FAMILY #172 ERICACEAE FIG. 2. Stamens from the genera of Costa Rican Ericaceae. (J) Lateropora ovata (Luteyn 14810, NY). (K) Didonica pendula (redrawn from Brittonia 29: 255, fig. 1. 1977). (L) Macleania insignis (Luteyn et al. 15240, NY). (M) Macleania rupestris (spirit material ex Longwood Gardens). (N) Psammisia ramiflora (Luteyn et al. 15215, NY). (O) Gonocalyx pterocarpus (Luteyn et al. 15229, NY). (P) Thibaudia costaricensis (Luteyn 4531, NY). (Q) Anthop- terus revolutus (Luteyn 14806, NY). (R) Utleya costaricensis (Luteyn et al. 4412, NY). (S) Themistoclesia pentandra (Luteyn 14862, NY). (T) Themistoclesia smithiana (Luteyn et al. 15249, NY). (U) Disterigma humboldtii (Luteyn & Pedraza 15573, NY). (V) Sphyrospermum cordifolium (Luteyn et al. 15152, NY). (W) Vaccinium consanguineum (Luteyn et al. 15222, NY). (X) Vaccinium poasanum (Luteyn et al. 15219, NY). (Illustrations by Bobbi Angell) FIELDIANA: BOTANY Sphyrospermum buxi f ol i urn Vaccinium floribundum Disterigma humboldtii cordi folium Sphyrospermum Sphyrospermum dissimile Gonocalyx almedae Gonocalyx pterocarpus Themistoclesia VW^ 10 smithiana FIG. 3. Ericaceae shrubs with very small leaves: species of Gonocalyx, Sphyrospermum, Themistoclesia, and Vaccinium. LUTEYN AND WILBUR: FAMILY #172 ERICACEAE Vaccinium consanguineum Disterigma > pilosum Jb rA Vaccinium floribundum Gonocalyx costaricensis FIG. 4. Ericaceae shrubs with small, mostly narrow leaves: species of Bejaria, Disterigma, Gonocalyx, Pernettya, and Vaccinium. FIELDIANA: BOTANY Themistoclesia pentandra Sphyrospermum ellipticum FIG. 5. Ericaceae shrubs with smaller, mostly ovate leaves: species of Gonocalyx, Sphyrospermum, and Themis- toclesia. LUTEYN AND WILBUR: FAMILY #172 ERICACEAE Vacci ni umprosi ense Cavendishia talamancensis FIG. 6. Ericaceae shrubs with small to medium-sized leaves that are often rounded at the base: species of Cav- endishia, Macleania, and Vaccinium. 10 FIELDIANA: BOTANY Cavendishia capitulata Gonocalyx pterocarpus Disterigma utleyorum SphyrospermunTel 1 i pt i cum Macleania insignis FIG. 7. Ericaceae shrubs with small to medium-sized leaves and with rounded or obtuse apicds: species of Cav- endishia, Disterigma, Gonocalyx, Macleania, and Sphyrospermum. LUTEYN AND WILBUR: FAMILY #172 ERICACEAE 11 Vaccinium furfuraceum Vac. poasanum Gaultheria gFacilis . ^%FT > mrii Gaultheria erecta FIG. 8. Ericaceae shrubs with smaller to medium-sized leaves and acute or acuminate apices: species of Gaultheria and Vaccinium. 12 FIELDIANA: BOTANY Cavendishia chiriquiensis Cavendishia lactiviscida :.limonensis FIG. 9. Species of Cavendishia with medium-sized leaves with long-acuminate apices. LUTEYN AND WILBUR: FAMILY #172 ERICACEAE 13 Utleya costaricensis Comarostaphylisi I arbutoides Cavendishia bracteata Lateropora ovata Macleania rupestris FIG. 10. Ericaceae shrubs with medium-sized leaves and with prominent petioles: species of Cavendishia, Coma- rostaphylis, Lateropora, Macleania, and Utleya. 14 FIELDIANA: BOTANY Cavendishia compl ectens' callista FIG. 1 1 . Species of Cavendishia with medium or larger leaves with rounded or obtuse bases. LUTEYN AND WILBUR: FAMILY #172 ERICACEAE 15 Cavendishia osaensis Thibaudia- costaricensis FIG. 12. Ericaceae shrubs with larger lanceolate leaves: species of Cavendishia and Thibaudia. 16 FIELDIANA: BOTANY Satyria 's.ulbrichiana, FIG. 13. Ericaceae shrubs with larger narrowly ovate or elliptic leaves: species of Psammisia and Satyria. LUTEYN AND WILBUR: FAMILY #172 ERICACEAE 17 Cavendishia conferti flora Cavendishia callista FlG. 14. Species of Cavendishia with larger leaves with acuminate apices. 18 FIELDIANA: BOTANY Anthopterus revolutus Cavendishl lineari folia Chimaphila maculata FIG. 15. Two species of Satyria and some unusual Ericaceae of Anthopterus, Cavendishia, , Chimaphila, and Monotropa. LUTEYN AND WILBUR: FAMILY #172 ERICACEAE are usually hidden among the mosses within which it grows, whether terrestrially or on mature tree branches. Hybridization is not, in general, common in Costa Rican ericads but does occur sporadically in highly disturbed habitats where sympatry oc- curs. It has been recognized mostly in the genus Cavendishia (Luteyn, 1976c, 1983), based on field observations of plants with intermediate morphologies and then later confirmed by mea- surements of intermediate characters and of pollen viability in the laboratory. Hybrids are discussed with their parental species. [Note: Descriptions are in general comparable; however, when one or two species in a genus have striking features not found in the other species, these are mentioned only where they occur, and it can be assumed that other species in the genus do not have this feature. All measurements were tak- en from dried herbarium material unless other- wise stated by "when fresh," which means ma- terial preserved in alcohol. In this family, there is only a small degree of shrinkage once material dries. Overall calyx length is measured from the base of the calyx tube to the tip of the calyx lobes (i.e., the measurement includes both the calyx tube and limb). In similar fashion, overall corolla length is measured from the base of the corolla to the tip of the corolla lobes; corolla width is mea- sured at the widest point. For corollas, the terms unistratose and bistratose are used when one or two layers or strata (respectively) are observable in the tissue; a longitudinal section of the corolla will show this to the naked eye, but it is easily seen without sectioning in the area between the corolla lobes where a bistratose corolla has tissue between the lobes like webbing between the toes of a duck. Overall stamen length is measured from the base of the filament to the tip of the anther. Overall anther length includes the theca plus tubule lengths.] Key to Genera of Ericaceae in Costa Rica la. Plants lacking chlorophyll (whitish, pinkish or reddish, or at least not green), succulent herbs, mycotrophic Monotropa uniflora Ib. Plants with chlorophyll (green), woody subshrubs, shrubs to trees, autotrophic 2 2a. Plants subshrubs, with ± herbaceous habit Chimaphila maculata 2b. Plants shrubs or rarely trees, with woody habit 3 3a. Ovary superior; fruit usually with perianth scars at base 4 3b. Ovary inferior; fruit usually with perianth scars at apex 8 4a. Petals free, usually 7-merous, usually wide-spreading; fruit a depressed obovoid or depressed globose, septicidally 5-7-valvate capsule Bejaria aestuans 4b. Petals united, usually 4-5-merous; fruit a cylindrical septicidal or loculicidal capsule, drupe, or berry 5 5a. Corolla salverform, slightly bilaterally symmetrical; fruit a cylindrical, septicidal capsule; viscin threads mixed with pollen tetrads [infrequently cultivated in Costa Rica and not included in the text] Rhododendron simsii 5b. Corolla urceolate, cylindric-urceolate, or tubular, radially symmetrical; fruit a loculicidal capsule, drupe, or berry; viscin threads absent 6 6a. Inflorescences terminal panicles; surface of ovary and fruit papillate; fruit a drupe; leaf blade usually >3 times longer than wide, usually >10 times longer than the petiole Comarostaphylis arbutoides 6b. Inflorescences axillary racemes or flowers solitary; surface of ovary and fruit smooth; fruit a berry or capsule; leaf blade usually <3 times longer than wide, usually < 10 times longer than the petiole 7 7a. Fruit a berry, rarely with calyx becoming fleshy at base, but never surrounding the berry Pernettya prostrata 7b. Fruit a capsule, surrounded by the fleshy calyx Gaultheria 8a. Stamens strongly unequal, with filaments and/or anthers conspicuously alternately unequal ... 9 8b. Stamens equal, with filaments and anthers of equal lengths (rarely anthers inconspicuously alter- nately unequal) 11 20 FIELDIANA: BOTANY 9a. Filaments and anthers unequal; stamens as long as corolla or rarely '/r-'/i the corolla length; floral bracts usually large, showy Cavendishia 9b. Filaments equal or unequal; anthers unequal or essentially equal; stamens usually about '/, as long as corolla; floral bracts small, inconspicuous 10 lOa. Calyx limb 5-parted; staminal filaments equal and connate over entire length; anthers unequal, with tubules widening apically but otherwise hardly distinguishable from thecae; floral bracts usually persistent; inflorescences often ramiflorous Satyria lOb. Calyx limb 3-parted (in ours); staminal filaments unequal and distinct; anthers essentially equal, with tubules with parallel sides and easily distinguishable from thecae; floral bracts usually decid- uous; inflorescences from the axils of leaves Orthaea lla. Anthers dehiscing by longitudinal slits that extend the entire length of the thecae; tubules rudi- mentary, apparently nonfunctional, <0.8 mm long Laieropora 1 Ib. Anthers dehiscing by pores or slits on the tubules and not extending onto thecae; tubules conspic- uous, well-developed, > 1 mm long 12 12a. Anthers dehiscing by latrorse clefts that do not extend to tip of tubule Didonica 12b. Anthers dehiscing by terminal pores or introrse clefts (or if clefts latrorse, then extending to tip of tubule) 13 13a. Bracteoles located at apex of pedicel, clasping and surrounding calyx (and sometimes lower co- rolla) Disterigma 13b. Bracteoles located well below apex of pedicel, or if apical then not clasping calyx 14 14a. Tubules elongate, thin, about half as wide (or less) as thecae Gonocalyx 14b. Tubules about as wide as thecae, or if narrower then much shorter than thecae 15 15a. Thecae conspicuously papillate; tubules rigid, elongate- to short-conical, distinct or sometimes laterally connate or fused into one tubule; stamens often '/,-'/2 as long as corolla 16 15b. Thecae smooth to minutely papillate; tubules flexible, cylindric, elongate to short, distinct to base or fused into one tubule in Themistoclesia pentandra; stamens usually as long as corolla ... 17 16a. Connectives never spurred; tubules usually laterally connate (the septum visible) or sometimes totally fused into one; corolla cylindric or angled, gradually narrowed to apex Macleania 16b. Connectives alternately spurred, rarely all spurred, if spurs absent then connectives usually apically prolonged or thickened; tubules distinct to base; corolla cylindric, never angled and often abruptly constricted to apex, or corolla ± globose to conical (and then < 1 cm long) Psamrnisia 17a. Calyx continuous with pedicel (pedicel not jointed at apex) 18 17b. Calyx articulate with pedicel (pedicel jointed at apex) 20 18a. Calyx terete; pedicels proportionally long, thin, filiform and pendent, rarely short and 1-4 (8) mm long, but then corolla dark red and campanulate Sphyrospermum 18b. Calyx angled or winged; pedicels short or long, but thick, not filiform and pendent (rarely 9-22 mm long) 19 19a. Leaves (0.7) 1-4.5 cm long, usually distichous (spiral in T. pentandra) along stem, base of lamina without glands; calyx usually angled (winged in T. pentandra), 3.2-7 mm long; corolla terete to pentagonal, the lobes ca. '/10-'/3 overall corolla length, not strongly reflexed as to expose stamens at anthesis Themistoclesia 19b. Leaves 5-13 cm long, spirally arranged along stem, base of lamina beneath with 1 or 2 pairs of circular, concave glands; calyx 5-winged, 5-7 mm long; corolla broadly to narrowly 5-winged, the lobes ca. !/3 overall corolla length, strongly reflexed thus exposing stamens at anthesis Anthopterus 20a. Corolla ca. 4 mm long, the tube conspicuously spurred apically opposite the lobes; calyx conspic- uously winged Utleya costaricensis 20b. Corolla >7 mm long, never spurred; calyx usually terete 21 2 la. Corolla urceolate to cylindric-campanulate, the lobes imbricate or valvate; staminal filaments usu- ally distinct Vaccinium 21b. Corolla elongate-tubular, the lobes valvate; staminal filaments connate Thibaudia costaricensis LUTEYN AND WILBUR: FAMILY #172 ERICACEAE 21 Anthopterus Hook REFERENCES — J. L. Luteyn, New species and notes on neotropical Ericaceae. Opera Bot. 92: 109-113. 1987 [Anthopterus, pp. 109-113]. J. L. Luteyn, New species, new records, and neotypi- fication of some Mesoamerican Ericaceae. Brit- tonia 48: 241-249. 1996 [Anthopterus, p. 248]. J. L. Luteyn, Redefinition of the neotropical genus Anthopterus (Ericaceae: Vaccinieae), including one new species. Brittonia 48: 605-610. 1996 [1997]. Terrestrial or epiphytic shrubs. Leaves alternate (in ours), subopposite, or verticillate, petiolate, the blade with entire margin, the venation plinerved or pinnate, the base beneath with 1 or 2 pairs of circular, concave glands. Inflorescences axillary, racemose, many-flow- ered; floral bract small and inconspicuous; pedicels con- tinuous with calyx; bracteoles 2, medial to submedial (in ours). Flowers 5-merous, without odor; calyx tube broadening apically, 5-winged opposite the sinuses, limb suberect, lobes ovate to deltate; corolla sympetalous, aestivation valvate, subcylindric, cylindric-urceolate or subglobose, broadly 5-winged opposite the lobes, lobes ca. '/, overall corolla length, strongly reflexed thus ex- posing stamens at anthesis; stamens 10, essentially equal, nearly as long as the corolla, sometimes genicu- late; filaments equal, distinct or slightly connate at base, shorter than anthers; connectives lacking disintegration tissue or spurs; anthers dorsifixed, membranous, lacking awns, thecae smooth, tubules 2, distinct, about as wide as and usually longer than thecae, dehiscing by introrse, elongate clefts; pollen grains in tetrahedral tetrads, lack- ing viscin threads; ovary inferior; style filiform, about as long as the corolla; stigma truncate; nectariferous disc annular or cupuliform. Fruit a berry, usually purplish; seeds numerous, small. Anthopterus is a genus of 12 species ranging from northeastern Costa Rica to north-central eastern Peru. It is characterized by having the pedicel continuous with the calyx, and a winged calyx tube and corolla. Anthopterus costaricensis and A. revolutus are the first records of this genus in Costa Rica. Both species belong to subgen. An- thopterus, which is characterized by having alter- nate leaves, 1 or 2 pairs of circular, concave glands at the base of the lamina, a corolla that is pale yellowish-green throughout and with wings broadest basally and narrowing apically, the lobes strongly reflexed thus exposing the stamens at an- thesis, and straight stamens. Subgenus Gonandra Luteyn occurs from extreme southeastern Panama (Darien Prov.) to north-central Ecuador (Luteyn, 1996c). Because of the paucity of collections of any of the species, the range of variation within the genus is unknown. Anthopterus is morpholog- ically similar to Thibaudia and Themistoclesia, but it is separated from them by its combination of nonarticulate (i.e., continuous) calyx, conspic- uously 5-winged calyx tube, and 5-winged co- rolla. Key to the Species of Anthopterus la. Leaf blades narrowly elliptic to oblanceolate, strongly revolute, to 1.5 cm wide, not amplexicaul; petioles 3-4 mm long A. revolutus Ib. Leaf blades elliptic to oblong-elliptic, flat, 3-4.5 cm wide, clasping and amplexicaul; petioles to 2 mm long A. costaricensis Anthopterus costaricensis Luteyn, Brittonia 48: 609. 1996[1997]. Epiphytic shrubs; mature stems subterete, bluntly and broadly ribbed, glabrous; twigs subterete, complanate, bluntly angled, striate, glabrous, with scattered ovate- lanceolate, long-acuminate perulae to 10 mm long prox- imal to first leaves. Leaves with petioles subterete, strongly flattened adaxially. rugose, to 2 mm long, gla- brous; leaf blades coriaceous, subsessile, clasping or am- plexicaul, flat, elliptic to oblong-elliptic, (7) 9-12 cm long, 3-4.5 cm wide, base rounded, cordate, auriculate, apex obtuse or bluntly acute, glabrous but densely and deciduously glandular-fimbriate on both surfaces, weak- ly 5 (7)-plinerved (or pinnately nerved with 2-3 sec- ondary nerves per side), midrib impressed above and conspicuously raised beneath, lateral nerves and reticu- late veinlets slightly raised on both surfaces. Inflores- cence solitary, 12-21 -flowered; rachis subterete, angled, striate, bracteate, overall 4.5-1 1 cm long but the basal 3.5 cm usually naked and lacking fertile nodes, glan- dular-fimbriate throughout; floral bract apparently per- sistent, narrowly ovate, 3-5 mm long, apex long-acu- minate, glabrous, margin weakly glandular-fimbriate; pedicels subterete, angled, striate, 16-22 mm long (post anthesis), glandular-fimbriate; bracteoles near middle, similar to floral bract but 2.5-3 mm long. Flowers (all post anthesis) with calyx ca. 7 mm long, glabrous, glan- dular-fimbriate, tube obconical, strongly 5-winged, ca. 5 mm long, limb erect, ca. 2 mm long, lobes ovate, apic- ulate, ca. 1 .5 mm long, sinuses acute; corolla, stamens, and berry not seen. Anthopterus costaricensis is found in premon- 22 FIELDIANA: BOTANY tane forest, 1200 m elevation. It is endemic to Costa Rica and is known from only the type col- lection: Limon Province: Canton Talamanca, be- tween Cerro Chimu and Cerro Matama, L. D. Go- mez & Herrera 23557 (NY). The type was col- lected in late flower in late April. Endangered. Anthopterus costaricensis is characterized by its clasping to amplexicaul, elliptic to oblong-el- liptic, flat (not revolute) leaves, and twigs with scattered, long-acuminate perulae (to 10 mm long) proximal to the first leaves. It is morpho- logically most similar to A. pterotus (A. C. Smith) Luteyn from Colombia, a species also with sub- amplexicaul leaves but with a peculiar paniculate type of inflorescence. Although neither corollas nor stamens are known from A. costaricensis, the general morphology of the plant and especially the distinctive calyx tube place it in Anthopterus subgen. Anthopterus following Luteyn (1996c). Anthopterus revolutus (Wilbur & Luteyn) Lu- teyn, Fl. Ecuador 54: 387. 1996. Themistoclesia revoluta Wilbur & Luteyn, Ann. Missouri Bot. Card. 68: 164. 1981. Figures 2Q and 15. Epiphytic shrubs; mature stems subterete, coarsely ridged and grooved but becoming terete, glabrous, mi- nutely puberulent, or short-pubescent; twigs subterete, bluntly and coarsely angled, puberulent; axillary buds supraxillary, the scales lanceolate, acuminate, ca. 3 mm long. Leaves with petioles flattened adaxially and can- aliculate, 3-4 mm long, glabrous to inconspicuously pu- berulous to short-pubescent; leaf blades coriaceous, nar- rowly elliptic to oblanceolate, strongly revolute so as to conceal actual margin, 5-13 cm long, 0.6-1.5 cm wide, base cuneate or rounded, apex obtuse to rounded, gla- brous above or moderately to densely short-pubescent on both surfaces, glabrate, sparingly to moderately beset with reddish-brown, appressed, glandular fimbriae, 0.1- 0.2 mm long, pinnately nerved, midrib prominently im- pressed above and raised beneath, secondary nerves and reticulate veinlets raised to impressed above and mod- erately so but obscure beneath. Inflorescences 10-16- flowered, surrounded by 6-8, appressed, lanceolate to narrowly triangular, acute, 3.5-5 mm long, finely short- pubescent bracts; rachis ridged or striate, 5-10 cm long, moderately to densely short-pubescent with white to hy- aline trichomes 0.2-0.5 mm long, also with scattered glandular fimbriae; floral bract narrowly triangular to narrowly lanceolate, 2.5-4 mm long, apex long-acumi- nate, glabrous to short-pilose above, ciliate; pedicels 9- 16 mm long, moderately to densely spreading short-pu- bescent with white to hyaline trichomes 0.2-0.3 mm long, also with scattered glandular fimbriae; bracteoles medial to submedial, appressed, narrowly triangular, 2- 3 mm long, apex acute to acuminate, spreading short- pubescent. Flowers with calyx 5-6 mm long, moderate- ly spreading short-pilose throughout, also with scattered glandular fimbriae, tube obpyramidal to narrowly turbi- nate, strongly 5-angulate or winged, 4-5 mm long, limb 1-2 mm long, lobes narrowly triangular, acute to acu- minate, sometimes apiculate, 1-2 mm long; corolla cy- lindric-urceolate, narrowly "5-winged (wings broadest at base). 7-9 mm long, ca. 4-4.5 mm diam. near base in- cluding wings, pale cream-green, glabrous, lobes 2.5-3 mm long, sparingly ciliate along midrib, strongly re- flexed at anthesis exposing stamens and style; stamens 6-8.2 mm long, alternately slightly unequal; filaments equal, 1-2 mm long, sparsely ciliate and glandular-fim- briate; anthers 7.5-9 mm long, thecae 2.5-3 mm long, tapering into a short-setose, apiculate base, closely co- herent at base due to interlocking of antheridial grooves, tubules 5-6 mm long, dehiscing by a narrow cleft ca. 1.5 mm long. Berry spherical, ca. 10 mm diam. Anthopterus revolutus is found in primary moist forest to premontane forest, 900-1400 m elevation. It is distributed in Costa Rica, Panama, and Ecuador. In Costa Rica it is known from only one collection in Lim6n Province (road from Fila Dimat toward Soki via Quebrada Sha, L. D. Go- mez et al. 23871). Flowering specimens have been collected in February, March, and August (in Pan- ama). Endangered. Anthopterus revolutus is characterized by hav- ing narrow, strongly revolute leaf blades with pin- nate venation and basally cuneate bases, long- pedicellate flowers borne in elongate racemes, and glandular-fimbriate staminal filaments. It is most closely related to A. schultzeae (Sleumer) Luteyn from eastern Ecuador and adjacent Peru. Bejaria Mutis ex Linnaeus nom. & orth. conserv. REFERENCES — R. Mansfeld & H. O. Sleumer, Revision der Gattung Befaria Mutis. Notizbl. Bot. Gait. Berlin-Dahlem 12: 235-276. 1935. S. E. Clemants, (1103) Proposal to conserve 6182 Be- jaria (Ericaceae) with a conserved spelling and type. Taxon 43: 473-476. 1994. S. E. Clemants, Bejaria. Fl. Neotrop. Monogr. 66: 54-106. 1995. Terrestrial shrubs or small, slender trees, glabrous to densely glandular-setose. Leaves alternate, petiolate, the blade flat to tightly revolute, margin entire or rarely ob- scurely denticulate, the venation pinnate. Inflorescences terminal or axillary (in ours), racemose to corymbose (in ours) or rarely paniculate, few- to many-flowered, often viscid; floral bract small and inconspicuous; ped- icels continuous with calyx; bracteoles*2. Flowers (5) 7 (8)-merous, mildly fragrant only in some populations of B. aestuans; calyx synsepalous, persistent in fruit, deep- ly lobed, the lobes much longer than tube, separate or imbricate at anthesis; corolla polypetalous, aestivation imbricate, the petals showy, these suberect and imbricate (hummingbird pollinated) or (in ours) spreading (bee LUTEYN AND WILBUR: FAMILY #172 ERICACEAE 23 pollinated) at anthesis, soon deciduous; stamens twice as many as petals or more (10-20), equal, about as long as petals or longer; filaments equal, distinct, elongate, basally densely pilose or tomentose; connectives lacking disintegration tissue or spurs; anthers dorsifixed, equal, oblong-obconic, dehiscing somewhat introrsely by slightly oblique, somewhat teardrop-shaped, terminal pores, interior rim of pores whitish; pollen grains in tet- rahedral tetrads, with viscin threads; ovary superior, 6- 7-locular. with axile placentation, glabrate; style elon- gate and often bent to one side, glabrous; stigma capi- tate, 6-7-grooved; nectariferous disc surrounding ovary base, unlobed. Fruit a capsule, depressed obovoid or depressed globose, woody, septicidally 5-7-valvate from apex; seeds numerous, winged or tailed. Bejaria (formerly called Befaria) is a neotrop- ical to subtropical genus of 15 species, ranging from Georgia and Florida (U.S.A.) to Cuba, from Mexico south into Bolivia, and east to Guyana. The genus has not yet been found in Costa Rica, although one species does occur in western Chi- riqui Province, Panama. Bejaria is characterized by having a usually 7-merous corolla with free petals, nonappendaged anthers, viscin threads mixed with pollen tetrads, superior ovary, and septicidally dehiscent, capsular fruits. Within the Ericaceae, Bejaria is placed in subfamily Ericoi- deae, where it resides as an isolated genus in the tribe Bejarieae (Kron et al., 2002a). Bejaria aestuans Linnaeus, Mont. PI. 242. 1771. Bejaria glauca Bonpland in Humboldt & Bon- pland, PI. Aequinoct. 2: 118, pi. 117. 1813 (as Befaria}. Illustrated: R. L. Wilbur & J. L. Lu- teyn, Ann. Missouri Bot. Gard. 65: 39, fig. 4. 1978. S. E. Clemants, Fl. Neotrop. Monogr. 66: 75, fig. 7. 1995. Figures 1C and 4. Shrub to small tree 5-6 m tall; mature stems gla- brous, tomentose, or hispid to glandular-hispid; twigs te- rete, coarsely ribbed, densely ferrugineous-tomentose, glabrate, sometimes glandular-hispid. Leaves with peti- oles subterete, slightly winged, 5-10 mm long, ferrugin- eous-tomentose when immature, glabrate; leaf blades co- riaceous, flat or sometimes slightly revolute, elliptic to lanceolate or oblanceolate, lance-elliptic or rarely ovate, 2-5 cm long, 0.8-1.5 cm wide, base long-cuneate and decurrent, apex acute to short-acuminate, often bluntly short-mucronate, both surfaces glabrous to tomentose, hispid, or glandular-hispid when young, then often gla- brate, often glaucous beneath, midrib impressed above and conspicuously raised beneath, reticulate veinlets ob- scure on both surfaces. Inflorescences short-racemose or subcorymbose, 10-20-flowered; rachis subterete, coarsely angled, 1 .5-2.5 cm long, densely ferrugineous- tomentose to glabrate; floral bract elliptic to lanceolate or oblanceolate, 2-5 mm long, ca. 1 mm wide, some- times leaf-like and up to 30 mm long and 8.4 mm wide, apex acute to obtuse, sometimes acuminate, cililate to glandular-ciliate, ferrugineous-tomentose on both surfac- es; pedicels subterete, striate, 15-24 mm long, ca. 0.5 mm diam., glabrous, densely ferrugineous-tomentose, hispid, or glandular-hispid, glabrate; bracteoles at or be- low middle, elliptic to oblanceolate, 2-5 mm long, ca. 1 mm wide, apex acute to obtuse, sometimes acuminate, cililate to glandular-ciliate, ferrugineous-tomentose on both surfaces. Flowers erect, 7-merous; calyx 2-3 mm long, glabrous, densely ferrugineous-tomentose, hispid, glandular-hispid, glabrescent, the lobes ovate, 1 .5-2 mm long, apex obtuse, margin ciliate; corolla spreading- campanulate, the petals obovate, 15-20 mm long, 5-6 mm wide, densely ciliate at apex, white to dark rose- pink; stamens subequal to corolla or slightly exserted, 15-21 mm long; filaments 13-20 mm long, densely pi- lose in the basal third; anthers ca. 2 mm long; style subequal to corolla or exserted, 21-22 mm long, gla- brous. Capsule depressed-obloid, 7-8 mm diam. Bejaria aestuans is found in pine woods, cloud forest, dry montane forest, shrub paramo, grass- lands, and disturbed roadsides, (300) 1000-3000 (3500) m elevation (Continental Divide cloud for- est at 1200-1700 m in Panama). It is geographi- cally the most widespread and morphologically the most variable species in the genus, ranging from west-central Mexico almost continuously to Bolivia. It is very rare in El Salvador, Nicaragua, and Panama and has yet to be collected in Costa Rica. In western Panama, flowering specimens have been collected in January, May, July, and August; fruiting in January, July, and August. En- dangered (in Panama). Mansfeld and Sleumer (1935) recognized five varieties within the species that were distin- guished primarily by characters of indumentum. This character is unusable as its facies change with the age of the plant. Therefore, the varieties of B. aestuans were not recognized by Clemants (1995) in his revision of the genus. The descrip- tion above is based on Panamanian collections. In Panama, B. aestuans is characterized by having obscure floral bracts, petals that are fully separate and light to dark pink in color, stamens about equaling the corolla in length, and eglandular, fer- rugineous-tomentose pubescence that varies from dense on young parts to glabrate on older parts. The flowers have a sweet odor and are visited by bees (pers. observ.). Cavendishia Lindley nom. conserv. REFERENCES — J. L. Luteyn, The genus Caven- dishia (Vacciniaceae) in Costa Rica. Brenesia 6: 24 FIELDIANA: BOTANY 9-18. 1975. J. L. Luteyn, Ericaceae— Part I. Cav- endishia. Fl. Neotrop. Monogr. 35: 1-290. 1983. J. L. Luteyn, New species, new records, and neo- typification of some Mesoamerican Ericaceae. Brittonia 48: 241-249. 1996. J. L. Luteyn & J. F. Morales, Four new species of Cavendishia (Eri- caceae: Vaccinieae) from Costa Rica. Brittonia 48: 514-519. 1996[1997]. Terrestrial or epiphytic shrubs. Leaves alternate, pet- iolate, the blade sometimes revolute, margin entire, the venation plinerved or pinnate. Inflorescences axillary, usually solitary, racemose or subfasciculate; floral bracts usually large and showy; pedicels articulate with calyx; bracteoles 2, basal or rarely medial. Flowers 5-merous, without odor; calyx with tube apophysate or not, lobes erect or connivent after anthesis, often glandular; corolla sympetalous, aestivation valvate, cylindric, usually car- nose when fresh; stamens 10, subequal or strongly un- equal to each other, about equaling corolla in overall length or rarely 'A-'/, corolla length; filaments usually distinct (rarely slightly coherent near base), alternately unequal; connectives lacking disintegration tissue or spurs; anthers alternately unequal, lacking awns, thecae smooth or minutely papillate, tubules 2, distinct, about same width as thecae and about twice as long, dehiscing introrsely by elongate clefts; pollen grains in tetrahedral tetrads, lacking viscin threads; ovary inferior; style fili- form, usually as long as corolla, glabrous. Fruit a berry; seeds numerous, ca. 0.5-1:2 mm long. Cavendishia includes about 130 species that range from Mexico to Bolivia, east to Guyana and Brazil. It is characterized by having alternately unequal staminal filaments and anthers and usu- ally conspicuously large floral bracts. The genus has been divided into several sections and series based mostly on characters of the calyx tube base (i.e., apophysate or nonapophysate) and the types of glands found especially on the calyx lobes (see Luteyn, 1983, for details and photos). In order better to understand these infrageneric relation- ships within Cavendishia, we present below a key to the sections and series found in Costa Rica and adjacent western Panama, as well as a second key to the species. Cavendishia is morphologically most similar to Orthaea Klotzsch and Thibaudia Ruiz & Pav. ex J. St.-Hil. The genus is the most commonly collected member of the Ericaceae in Costa Rica because of its abundance and showy nature. Twenty-two species are currently known in Costa Rica. Key to the Sections and Series of Cavendishia in Costa Rica la. Calyx lobe margins eglandular or irregularly lacerate-glandular (then usually darker in color and thinner than the rest of the lobe), neither scarious, glandular-fimbriate, nor glandular-callose sect. Quereme Ib. Calyx lobe margins scarious, glandular-fimbriate, or glandular-callose 2 2a. Calyx lobe margins scarious or glandular-fimbriate, the fimbriae distinct (rarely caducous) or be- coming laterally fused and then sometimes forming a thin, supramarginal gland (sect. Cavendishia) 3 2b. Calyx lobe margins glandular-callose or entire calyx lobe glandular-callose 4 3a. Calyx lobes broadly overlapping at anthesis ser. Imbricatae 3b. Calyx lobes not overlapping at anthesis ser. Cavendishiae 4a. Calyx lobes completely glandular callose and this sometimes extending onto the calyx limb proper; plants without globular glands sect. Callisla 4b. Calyx lobes with callose glands along margins only, or the callose tissue sometimes extending onto the lobe lamina proper; if entire lobe glandular, then plants also bearing globular glands ca. 0.1- 0.2 mm diam. (sect. Engleriana) 5 5a. Sessile (or minutely peltate), subglobular glands 0.1-0.2 mm diam. along margins of floral bracts and bracteoles, rachises, pedicels, often also on the calyces, mature stems, and leaves (sometimes caducous from upper leaf surface but then leaving tiny, pustular scars, the exudate often milky- white) ser. Lactiviscidae 5b. Globular glands not present as above ser. Englerianae LUTEYN AND WILBUR: FAMILY #172 ERICACEAE 25 Key to the Species of Cavendishia la. Leaves amplexicaul, the bases strongly cordate C. complectens Ib. Leaves subsessile to long petiolate, sometimes clasping to nearly amplexicaul, but never strongly amplexicaul and with bases strongly cordate 2 2a. Margins of calyx lobes eglandular, glandular-fimbriate (fimbriae sometimes caducous), or lacerate- glandular 3 2b. Margins, apex, or all of calyx lobes glandular callose-thickened, or with intermittent stout, oblong glandular thickenings 14 3a. Calyx limb and floral bracts usually conspicuously striate; bracteoles striate, >5 mm long C. axillaris 3b. Calyx limb and floral bracts smooth; bracteoles smooth, rarely >5 mm long 4 4a. Leaf blade up to 6 (6.8) cm long, the apex obtuse, rounded, or broadly acute 5 4b. Leaf blade usually longer than 6 cm, the apex sharply acute to long-acuminate 9 5a. Base of calyx tube strongly apophysate; floral bracts 12-25 mm long 6 5b. Base of calyx tube weakly or not apophysate; floral bracts 8-14 (19) mm long 8 6a. Inflorescence pilose; rachis 3-9 cm long; pedicels 9-10 mm long; corolla 11-13 mm long C. davidsei 6b. Inflorescence glabrous (calyx tube short-pilose in C. gomezii)', rachis 0.3-2.5 cm long; pedicels 3- 8 mm long; corolla 7.5-10 mm long 7 7a. Leaves obviously petiolate, the blades neither clasping nor amplexicaul, flat; rachis 1-2.5 cm long; pedicels 6-8 mm long; calyx tube short-pilose; corolla 9-10 mm long C. gomezii 7b. Leaves with petioles inconspicuous, the blades clasping to nearly amplexicaul, strongly bullate; rachis 3-6 mm long; pedicels 3-4.5 mm long; calyx tube glabrous; corolla 7.5-8 mm long .... C. luteynii 8a. Rachis subfasciculate, < 1 cm long, 3-6 (7)-flowered; floral bracts 1 1 mm or more long; leaves flat; calyx and corolla glabrous; calyx lobes densely glandular-fimbriate C. capitulata 8b. Rachis elongate, 1-9 cm long, 8-32-flowered; floral bracts <11 mm long; leaves bullate; calyx and corolla densely pubescent with short curly hairs; calyx lobes lacerate-glandular C. talamancensis 9a. Calyx lobes broadly overlapping over entire length at anthesis; floral bracts appressed to flowers, persistent in fruit C. confertiflora 9b. Calyx lobes not overlapping at anthesis, rarely contiguous at base; floral bracts spreading, decid- uous in fruit 10 lOa. Base of calyx tube distinctly apophysate C. limonensis lOb. Base of calyx tube rounded or truncate, not apophysate 11 11 a. Calyx lobes oblong, apically rounded, marginally wavy, imbricate in bud, overlapping after an- thesis (rarely contiguous at base during anthesis) C. calycina 1 Ib. Calyx lobes triangular, apically acute (rarely obtuse), valvate in bud, sometimes overlapping after anthesis 12 12a. Calyx densely woolly, the matted hairs persistent; leaf blades persistently soft-pilose beneath . . . C. pubescens 12b. Calyx glabrous to pilose but never woolly, the hairs never matted; leaf blades glabrous to pubescent beneath 13 13a. Corolla (10) 14-23 (28) mm long, terete or nearly so; floral bracts flat or slightly concave, (11) 17-30 (40) mm long, ascending or slightly spreading; leaf blades usually flat, glabrous to pubes- cent; corolla dark pink to red with white tip; plants without odor of wintergreen . . C. bracteata 13b. Corolla 6.5-1 1(13) mm long, conspicuously bluntly angled; floral bracts conduplicate-keeled, 10- 25 mm long and ascending; leaf blades usually conspicuously bullate, glabrous; corolla orange with white tip; plants with odor of wintergreen C. quereme 14a. (From 2b) Leaf blade apex rounded, obtuse, or rarely subacute to acute; corolla <15 mm long; calyx tube and pedicel usually long-pilose, the sinuses usually ciliate; calyx lobes with intermittent, glandular thickenings which may coalesce at apex C. quercina 14b. Leaf blade apex acuminate to caudate-acuminate, or rarely acute; corolla usually >15 mm long; 26 FIELDIANA: BOTANY calyx tube, pedicel, and sinuses usually glabrous (hirsute in C. herrerae); calyx lobes either com- pletely callose- thickened, or with very distinct marginal thickenings 15 15a. Floral bract margins, bracteole margins, pedicels, rachises, and usually calyx tube with red, glob- ular or peltate glands; leaf blades with pustular or globular glands along upper surface and margin (these sometimes caducous or obscure, but leaving scars) 16 15b. Plants lacking red, globular, peltate, or pustular glands (C. endresii may have several flesh-colored, angular glands at top of pedicel) 23 16a. Leaf blades linear-lanceolate to linear-elliptic, ca. 9 times longer than broad; bracteoles 27 mm long, just beneath calyx C. pseudostenophylla 16b. Leaf blades elliptic, lance-elliptic or ovate-lanceolate, less than 4 times longer than broad; brac- teoles < 1 5 mm long, at base or middle 17 17a. Calyx tube conspicuously apophysate at base 18 17b. Calyx tube at most inconspicuously apophysate at base 20 18a. Rachis 1-1.5 mm diam. at base, glabrous, 4-7 (lO)-flowered; floral bracts lance-elliptic to oblong- lanceolate, apically acuminate, 7-11 (16) mm long; bracteoles oval to semi-orbicular, 1-4 mm long; corolla glabrous C. lactiviscida 18b. Rachis 1.2-3 mm diam. at base, moderately to densely hirsute, 14-29-flowered; floral bracts oblong to obovate, apically rounded to acute, (11) 17-26 mm long; bracteoles ovate to oblong-ovate; corolla pilose toward apex (corolla not seen in C. herrerae) 19 19a. Rachis 6-8 cm long, densely hirsute; bracteoles ovate, 1-1.5 mm long; calyx tube glabrous; calyx lobes completely callose-glandular C. ciliata 19b. Rachis 10-11 cm long, moderately hirsute; bracteoles oblong-ovate, 4-6 mm long; calyx tube hirsute; calyx lobes only marginally glandular C. herrerae 20a. Rachis 1.5-10.5 cm long (usually <5 cm long) and ca. 1-1.5 mm diam., 6-12-flowered; bracteoles linear 21 20b. Rachis usually >8 cm long and 3-5 mm diam., 1 1-26-flowered; bracteoles ovate to lanceolate or oblong-oblanceolate 22 2 la. Leaf blades flat; rachis 1.5-4.5 cm long; bracteoles linear to linear-lanceolate; corolla pilose in apical 1/3 C. chiriquiensis var. chiriquiensis 21b. Leaf blades bullate; rachis (2.5) 6-10.5 cm long; bracteoles ovate to oblong-ovate; corolla glabrous or only weakly puberulous in apical !/, C. chiriquiensis var. bullata 22a. Bracteoles 12-14 mm long, at middle of pedicel, concealing all calyx and lower part of corolla at anthesis C. fortunensis 22b. Bracteoles 1.5-2 (4) mm long, basal, never concealing calyx or corolla C. panamensis 23a. Calyx lobes glandular callose-thickened only along margin 24 23b. Calyx lobes usually completely glandular callose-thickened (sometimes glandular only in apical % in C. subfasciculata, rarely just marginally in C. megabracteata) 25 24a. Leaf blades linear C. linearifolia 24b. Leaf blades ovate or ovate-elliptic C. osaensis 25a. Floral bracts apically rounded, usually deeply emarginate 26 25b. Floral bracts apically acute to rounded, never emarginate (apiculate and rarely emarginate in C. melastomoides) 27 26a. Corolla 31-40 (43) mm long, the lobes never glandular-callose; floral bracts (20) 30-60 (70) mm long, dark red in bud but turning pink or pale purplish-red at anthesis; calyx 7-11 (13) mm long, the limb cylindric to slightly spreading C. megabracteata var. megabracteata 26b. Corolla 20-24 (30) mm long, the lobes sometimes glandular-callose; floral bracts (15) 20-30 (40) mm long, greenish to pinkish-green at anthesis; calyx 5-7 (10) mm long, the limb campanulate to cylindric G. subfasciculata 27a. Rachis (0.7) 1-2 (3) cm long, 3-8-flowered; pedicel <10 mm long, apically with several large, flesh-colored, angular glands; bracteoles essentially completely glandular callose-thickened C. endresii 27b. Rachis (2) 3-12 cm long, (4) 6-40-flowered; pedicel usually 10 mm long or more, lacking large angular glands; bracteoles glandular callose-thickened only in apical %-'/, 28 LUTEYN AND WILBUR: FAMILY #172 ERICACEAE 27 28a. Rachis 1-1.5 mm diam. at base; floral bracts 4-22 mm long, 3-12 mm wide 29 28b. Rachis (1.7) 2-5 (10) mm diam. at base; floral bracts (15) 20-40 (60) mm long, (7) 10-25 (35) mm wide 31 29a. Corolla 16-20 mm long, white; Cordillera de Tilaran, 950-1500 m C. melastomoides var. albiflora 29b. Corolla (19) 26-41 mm long, reddish-maroon or blue; Cordillera Central-Cordillera de Talamanca, (1050) 1400-2800 m 30 30a. Rachis and pedicels usually densely glandular-fimbriate; floral bracts apically rounded or acute; bracteoles lanceolate or narrowly ovate-lanceolate, glandular-callose in apical Vj,-V2', corolla reddish- maroon; Costa Rica C. melastomoides var. melastomoides 30b. Rachis and pedicels with at most few, scattered, glandular fimbriae; floral bracts apically slightly notched and callose-apiculate; bracteoles broadly ovate to semi-orbicular, callose-apiculate; corolla blue; Panama C. melastomoides var. coloradensis 3 la. Rachis and pedicels with minute, scattered, reddish, globular or clavate glands along entire length; calyx tube apophysis merely undulate at base; floral bracts usually caducous; corolla whitish to purplish-rose; bracteoles usually conspicuously 3-5-nerved C. wercklei 31b. Rachis and pedicels essentially eglandular; calyx tube apophysis usually deeply lobed at base; floral bracts persistent through anthesis; corolla white with purple-margined lobes or purple to lavender; bracteoles not conspicuously nerved 32 32a. Corolla densely sericeous (rarely glabrous), white (only lobe margins purple); floral bracts pink; rachis 4-12 cm long, 15-40-flowered; petioles densely pilose to glabrate C. callista 32b. Corolla short-pilose over entire length or only in apical half, purple; floral bracts deep violet or lavender- to maroonish-purple; rachis 3-8 (9) cm long, (7) 1 1-20 (30)-flowered; petiole glabrous even when young C. atroviolacea var. atroviolacea Cavendishia atroviolacea Luteyn var. atrovio- lacea, Brittonia 28: 49. 1976. Illustration: J. L. Luteyn, Mem. New York Bot. Card. 28(3): 121, fig. 68. 1976. Figures 11 and 12. Epiphytic, sometimes scrambling shrubs, 1-3 m tall. Leaves with petioles (4) 6-15 mm long, glabrous; leaf blades lanceolate to lance-elliptic, rarely oblanceolate or ovate, (3) 8-15 (18) cm long, 1-5 (6) cm wide, base rounded, obtuse, cuneate, or rarely subtruncate, apex acuminate, rarely ± bullate. (3) 5 (7)-plinerved, midrib inconspicuously raised in basal 1-2 cm, essentially gla- brous. Inflorescences fusiform to cylindric, (7) 11-20 (30)-flowered; rachis 3-8 (9) cm long, 2-5 mm diam., glabrous; floral bract oblong to oblanceolate, 20-40 (50) mm long, glabrous, deep violet to maroonish-purple, of- ten appearing black, rarely translucent; pedicels (6) 9- 13 (16) mm long, glabrous; bracteoles at or near base, lanceolate to linear-lanceolate rarely ovate, 1-4 (5.5) mm long, apex glandular-callose. Flowers with calyx 5.5-1 1 (13.5) mm long, glabrous, tube cylindric, coarse- ly 10-ribbed, (2) 2.5-4 (5.5) mm long, apophysate at base with rim deeply 5-lobed, limb cylindric or some- what spreading, 3-8 mm long, weakly ribbed or mi- nutely papillate, lobes 1-2 (2.5) mm long, triangular, erect after anthesis, margins glandular-callose, sinuses obtuse to concave; corolla cylindric, thinly carnose when fresh, sometimes translucent, 20-38 (45) mm long, 5-8 mm diam., short-pilose over entire length or only in apical half, sometimes viscid puberulent, whit- ish-purple at base, purple to dark purple or lavender elsewhere, lobes oblong, obtuse, 1.5-2 mm long; sta- mens 19-39 mm long; filaments alternately 3.5-6 mm and 7-14.5 mm long, subglabrous or moderately pilose; anthers alternately 18-35.5 mm and 13-27.5 mm long, thecae 6-13.5 mm long. Berry not seen. Cavendishia atroviolacea is endemic to Costa Rica and western Panama, where it is found in cloud forest, in Chusquea—Quercus forest, in wet forest, along stream banks, and rooted in remnant trees in pastured areas, (150) 400-2600 m ele- vation. It is common on the western side of Vol- can Bani, Chiriqui Province, Panama, at eleva- tions of 1 300-2500 m but has also been collected about a dozen times from scattered localities in Costa Rica from extreme eastern Puntarenas Prov- ince (Cordillera de Talamanca, Cerro Pando, 1 860 m) and Limon Province (Cordillera de Talamanca, Atlantic slopes, near border with Panama, 2200- 2600 m), to Cartago Province (Canton Turrialba, Atlantic watershed, 1200 m), to Alajuela Province (Cordillera Central, Laguna Hule road, 400-900 m), and as far west as Guanacaste Province (Cor- dillera Tilaran, Monteverde region, 700-900 m). Flowering specimens have been collected in May-September; fruiting in January and October. Endangered. Cavendishia atroviolacea is characterized by- its large, deep violet to maroonish-purple floral bracts; many-flowered elongate inflorescences; conspicuously apophysate and deeply lobed calyx 28 FIELDIANA: BOTANY tube base; and glandular-callose calyx lobes. The lower elevation collections from Alajuela Prov- ince differ from other collections by slightly shorter corollas and more bullate leaf blades. Va- riety folsomii Luteyn is a rare endemic to the low, Continental Divide forests of central Panama (Co- de and Colon Prov.). It differs from the nominal variety primarily by its shorter corollas and much more prominent leaf blade midrib. Cavendishia atroviolacea belongs to sect. Callista, which in Costa Rica/Panama is a closely knit group of spe- cies of Cavendishi characterized, in general, by plinerved leaves, large floral bracts and corollas, and strongly apophysate calyces that are usually conspicuously lobed or ribbed (Luteyn, 1983). It is difficult to say which species are more closely related because each has its own derived features. Cavendishia atroviolacea is most closely related to the widespread C. callista, with which it may also hybridize where their ranges overlap; one collection especially, Luteyn et al. 15241 from Guanacaste Province (Monteverde region), ap- pears intermediate between these two species. More collections of this rare species are needed, especially from the Alajuela Province area, before the exact limits of its variation in Costa Rica are known. Cavendishia axillaris A. C. Smith, Contrib. U.S. Natl. Herb. 28: 493. 1932. Cavendishia gaulth- erioides A. C. Smith, Ann. Missouri Bot. Gard. 28: 444. 1941. Illustrated: J. L. Luteyn, Mem. New York Bot. Gard. 28(3): 98, fig. 45. 1976. J. L. Luteyn, Fl. Neotrop. Monogr. 35: 168, fig. 57E-H. 1983. Figure 11. Terrestrial or epiphytic shrubs 0.5-1.5 (3) m tall, sometimes lianoid, glabrous when mature. Leaves with petioles 1.5-3 (5) mm long; leaf blades hard, brittle, bullate, ovate, obovate, elliptic or suborbicular, 4.5-10.5 (17) cm long, (2.5) 4-1 1 cm wide, base cuneate, obtuse or rounded and subcordate then often slightly amplexi- caul and clasping, apex obtuse or rounded, pinnately veined. Inflorescences solitary or rarely 2-5 per axil, capitate to cylindric, tightly congested, (5) 9-15 (47)- flowered; rachis 1-2 (8) cm long; floral bract concave, often persistent, striate, obovate to semi-orbicular, ap- pressed to and concealing calyx and lower portion of corolla at anthesis, 7-10 (13) mm long, pink to dark rose-red, margin densely glandular-fimbriate; pedicels to 1 mm long; bracteoles similar to floral bract, but (5) 7- 9 mm long. Flowers with calyx (4.5) 6-7.5 mm long, tube cylindric, smooth to rugose, (1.5) 2.5-3 mm long, limb spreading to campanulate, (2.5) 3.5-4 (5) mm long, conspicuously striate, lobes 1-2 mm long, ovate, trian- gular or rarely oblong, erect after anthesis, margin glan- dular-fimbriate, sinuses acute or narrowly obtuse; corol- la cylindric to bottle-shaped, (6.5) 9-10 mm long, 3.5- 4 (6) mm diam., often glandular-fimbriate apically, white to pale green in basal '/,, green to yellowish-green in middle ^,, the narrowed throat and lobes pale green, lobes oblong to narrowly triangular, obtuse, 1-2 mm long; stamens 6-8.5 mm long; filaments alternately 1.5- 2.5 mm and 2.5-3.5 mm long, short-puberulent apically; anthers alternately 5-6.5 mm and 4-6 mm long, thecae 1.5-2.5 mm long. Berry spherical, 9-10 mm diam., pur- ple to blue-black. Cavendishia axillaris 'occurs in cloud forest, elf- in forest, tropical wet forest, remnant trees in pas- tureland, and fence-row trees and along disturbed roadside slopes, 550-2700 m elevation. It is com- mon from Alajuela Province (Cordillera Tilar^n), Costa Rica, through Cocle Province, Panama, and then infrequently along the Panama/Colombia border and into northern Colombia (NW Antio- quia Dept.); one collection has been made in southern Nicaragua. Flowering occurs throughout the year; fruiting specimens have been collected in March, April, June, July, November, and De- cember. Widespread. Cavendishia axillaris is morphologically very uniform throughout its range, being characterized by pinnately veined, often strongly bullate leaf blades that are subsessile and often very hard/brit- tle-coriaceous, densely compact inflorescences, floral bracts and bracteoles that are conspicuously nerved (sclerified), pinkish, and tightly appressed to the calyx, and small and pale green to yellow- ish-green corollas. Cavendishia axillaris belongs to sen Cavendishiae, although it is morphologi- cally similar to C. complectens (ser. Imbricatae), with which herbarium specimens have sometimes been confused. Both species have pinnately veined leaf blades and rather compact inflores- cences, although the rachis of C. complectens may lengthen considerably after anthesis. They are most easily distinguished by the amplexicaul leaves and reniform and broadly imbricate calyx lobes of C. complectens (the leaves of C. axillaris are merely subsessile or rarely clasping and the calyx lobes usually triangular to ovate and only very rarely imbricate at the base). Cavendishia axillaris may hybridize with C. complectens and C. quereme since there are plants with interme- diate morphologies (Luteyn, 1976c). A common name in Alajuela Province for C. axillaris is el macho. The species is visited by hummingbirds (pers. observ.). Cavendishia bracteata (Ruiz & Pav. ex J. St. Hil.) Hoerold, Bot. Jahrb. Syst. 42: 280. 1909. Thibaudia bracteata Ruiz & Pav. ex J. St. -Hil., LUTEYN AND WILBUR: FAMILY #172 ERICACEAE 29 Expos. Fam. Nat. I: 363. 1805. T. crassifolia Benth., PI. Hartw. 65. 1840. Polyboea crassi- folia (Benth.) Klotzsch, Linnaea 24: 31. 1851. Proclesia warszewiczii Klotzsch, Linnaea 24: 35. 1851. C. crassifolia (Benth.) Hemsl., Biol. Centr.-amer., Bot. 2: 273. 1881. C. latifolia Hemsl., Biol. Centr.-amer., Bot. 2: 273. 1881. C. warszewiczii (Klotzsch) Hemsl., Biol. Centr.- amer., Bot. 2: 273. 1881. Chupalon crassifolia (Benth.) Kuntze, Rev. Gen. PI. 2: 383. 1891. Chupalon latifolium (Hemsl.) Kuntze, Rev. Gen. PI. 2: 383. 1891. Chupalon warszewiczii (Klotzsch) Kuntze, Rev. Gen. PI. 2: 383. 1891. Cavendishia costaricensis Hoerold, Bot. Jahrb. Syst. 42: 326. 1909. Cavendishia hoffmannii Hoerold, Bot. Jahrb. Syst. 42: 328. 1909. Cav- endishia smithii Hoerold, Bot. Jahrb. Syst. 42: 328. 1909. Cavendishia skutchii A. C. Smith, J. Washington Acad. Sci. 27: 308. 1937. Illustra- tion: J. L. Luteyn, Mem. New York Bot. Card. 28(3): 108, fig. 55. 1976 (as Cavendishia cras- sifolia). J. L. Luteyn, Fl. Neotr. Monogr. 35: 144, fig. 53. 1983. Figures 10, 11, and 14. Terrestrial or epiphytic shrubs 1-3 m tall; mature stems glabrous to pilose. Leaves with petioles (3) 5-1 1 (17) mm long, glabrous to pilose; leaf blades oblong, elliptic or ovate, (2.5) 4-15 (22) cm long, (1) 1.5-5 (11) cm wide, base rounded, cordate, cuneate or subtruncate, apex acute to acuminate, often abruptly, glabrous or pi- lose along veins above and below, (3) 5-7 (9)-plinerved. Inflorescences capitate and congested or elongate-cylin- dric and loosely flowered, (4) 6-20 (40)-flowered; rachis (0.5) 1-5 (8) cm long, glabrous to pilose, sometimes glandular-fimbriate; floral bract oblong, ovate, elliptic, or oblanceolate, flat or slightly concave, (11) 17-30 (40) mm long, ascending or slightly spreading, glabrous to pilose, pink to dark red, sometimes glandular-fimbriate; pedicels (1.5) 6-15 (20) mm long, glabrous or pilose, often glandular-fimbriate; bracteoles usually basal, ovate to linear, rarely uri state. 1-4 mm long, glabrous to pi- lose, sometimes glandular-fimbriate. Flowers with calyx (3.5) 4-6 (9) mm long, glabrous to pilose, often glan- dular-fimbriate, calyx tube cylindric, smooth or some- what rugose, often pentagonal, 1 .5-3 (4) mm long, limb cylindric to campanulate, usually somewhat spreading, (1.5) 2-3 (6) mm long, lobes (0.5) 1-2 (3) mm long, triangular, connivent after anthesis, margin glandular- fimbriate, fimbriae sometimes fused, sinuses concave or acute; corolla cylindric to bottle-shaped, terete or broad- ly angled, (10) 14-23 (28) mm long, 4-5 mm diam., glabrous to pilose, sometimes glandular-fimbriate, dark pink to red, lobes deltate, 1-2 mm long, white; stamens 11.5-19 mm long; filaments alternately 2-4 mm and 3.4-6.5 mm long, glabrous or pilose; anthers alternately 10-16.5 mm and 7.5-14.5 mm long, thecae 2.5-6 mm long. Berry 8-14 mm diam., spherical, blue-black, gla- brous or pilose. Cavendishia bracteata occupies montane for- est, Chusquea-Quercus-Comarostaphylis forest in Costa Rica or secondary growth, bogs, thickets, and rocky roadside slopes, (300) 1000-3200 m elevation. It is distributed from Mexico to Bolivia. Flowering and fruiting occur throughout the year. Common. Cavendishia bracteata is the most frequently encountered species of Cavendishia (and probably Ericaceae as a whole) in Costa Rica. It is espe- cially showy in the Cordillera de Talamanca, south along the PanAmerican Hwy. above Carta- go starting at about 1500 m elevation. It is char- acterized by having a nonapophysate calyx tube that is longer than the limb, glandular-fimbriate calyx lobes, and bright red corollas with white lobes. Because of its extensive geographical range, C. bracteata has many local populations that are highly variable morphologically, as is also indicated by its long list of synonyms just for Costa Rican material. Cavendishia bracteata belongs to ser. Caven- dishiae, and it is most closely related to C. pu- bescens, a species not yet found in Costa Rica but common in western Panama's Chiriqui Province along the western slopes of Volcan Bani to within a few kilometers of the border with Costa Rica. With more intensive collecting near the border, C. pubescens will almost certainly be found within Costa Rica. Cavendishia pubescens may be easily distinguished from C. bracteata by its persistently and softly, short-puberulent leaf undersurfaces (not glabrous or pilose to glabrate); calyces with woolly, matted pubescence (not glabrous or pi- lose); calyx limbs that are longer than the tubes (not shorter); corollas that are densely short-pilose and thin and soft in texture (not glabrous to dense- ly pilose and coriaceous and waxy-nitid); and densely puberulent (not glabrous to pilose) ber- ries. Cavendishia bracteata may hybridize with C. endresii, C. capitulata, and C. pubescens based on morphologically intermediate collections (Lu- teyn, 1976c). Common names for Cavendishia bracteata in Costa Rica include arraydn, colmillo, and San Miguel. In Costa Rica, the hummingbirds Eugenes ful- gens spectabilis, Lampornis calolaema, and Pan- terpe insignis forage populations of C. bracteata for nectar and probably pollinate it in the process (pers. observ.; Col well, 1973; Luteyn, 1998). The flower-piercer Diglossa plumbea also visits C. bracteata to rob nectar (Colwell, 1973, as Cav- endishia smithii), which is the reason for the small holes often seen at the base of the corolla. 30 FIELDIANA: BOTANY Cavendishia callista J. D. Smith, Bot. Gaz. 20: 5, pi. II. 1895. C. longiflora J. D. Smith, Bot. Gaz. 37: 420. 1904. C. bullata A. C. Smith & Standl., Contrib. U.S. Natl. Herb. 28: 453. 1932. Illustration: J. L. Luteyn, Mem. New York Bot. Card. 28(3): 120, fig. 67. 1976. J. L. Luteyn, Fl. Neotrop. Monogr. 35: 262, fig. 77. 1983. Figures 11 and 14. Terrestrial and epiphytic shrubs 1.5-3 m tall; mature stems glabrate or persistently pilose; twigs pinkish to flesh-colored and glaucous, glabrous to pilose-hispid. Leaves with petioles (4) 6-13 (22) mm long, glabrous to densely pilose; leaf blades ovate, elliptic, oblong or rarely obovate, (4.5) 7-23 (35) cm long, (1) 2.5-10 (13) cm wide, base obtuse, rounded or truncate, often cor- date, apex acute to acuminate, sometimes abruptly short- acuminate, glabrous to pubescent on lamina above, gla- brous or densely hispid beneath, often scabrous, (3) 5 (9)-plinerved, often strikingly bull. no. Inflorescences elongate-cylindric, 1 5—40-flowered, viscid; rachis stout, 4-12 cm long, ca. 4-5 mm diam., glabrous or sericeous; floral bract oblong, obovate, or oblanceolate, (15) 20- 47 (60) mm long, 10-20 (30) mm wide, glabrous, pink to deep rose; pedicels 6-15 (20) mm long, glabrous to sericeous; bracteoles basal to submedial, 1-5 (8) mm long, apex glandular-callose, usually glabrous. Flowers with calyx 5-9 (11) mm long, glabrous or infrequently pilose-hispid, tube cylindric, (1.5) 2-4 (5.5) mm long, coarsely ribbed, rugose, often muricate, strongly apoph- ysate at base, limb cylindric or spreading to campanu- late, 3.5-6 (8) mm long, smooth or striate, often muri- cate, lobes 1-2.5 (4) mm long, triangular, erect after an- thesis, green, glandular-callose, sinuses obtuse to broad- ly acute; corolla cylindric, (15) 18-30 (42) mm long, 4-8 mm diam., densely sericeous (rarely glabrous), white to pearl-white or with a pinkish hue, lobes white with purple margins; stamens (13)1 8-34 mm long; fil- aments alternately (1.5) 3-4.5 (6) mm and (4.5) 6-12 mm long, distinct or weakly coherent at base; anthers alternately (11) 16-32 mm and (8.5) 12-26 mm long, thecae 5-20 mm long. Berry spherical, 8-13 mm diam., purple to blue-black. Cavendisia callista is found primarily in cloud forest, woods, and disturbed roadsides, 200-2000 m elevation. It is distributed in Guatemala, Nic- aragua, Costa Rica, central Panama, Colombia, Ecuador, the Guianas, and Brazil. Flowering spec- imens have been collected from December through August but vary locally; fruiting in April- August. Widespread. Cavendishia callista is characterized by having twigs that are pinkish to flesh-colored and glau- cous; often strikingly bullate leaf blades, rose- pink colored floral bracts, rachis, pedicels, and ca- lyx; elongate rachis; conspicuously ribbed and apophysate calyx tube; calyx lobes that are green and glandular-callose thickened; and corollas that are densely sericeous (rarely glabrous) and pearl- white with purple-margined lobes. In leaf mor- phology and leaf and stem pubescence, C. callista is highly variable, but the above combination of features characterizes the species. It is one of the most beautiful cavendishias in Costa Rica. Within Mesoamerica and Cavendishia sect. Callista, C. callista is most closely related to C. atroviolacea and C. wercklei based on inflorescences many- flowered, floral bracts large and apically acute, obtuse, or rounded (never emarginate); rachises elongate, thick and stout; pedicels usually >10 mm long and lacking apical angular glands; and bracteoles glandular apically (not completely). It may be distinguished from C. wercklei by its eglandular rachis and pedicels and from C. atro- violacea by its usually sericeous, white corollas (not short-pilose and purple). A common name in Costa Rica is colmillo de perro. The species is visited by hummingbirds (pers. observ.). Cavendishia calycina A. C. Smith, Ann. Missou- ri Bot. Card. 28: 447. 1941. Terrestrial or epiphytic shrubs to 5 m tall; mature stems subterete, smooth to slightly striate, glabrous; twigs subterete or bluntly angled, striate or ridged, gla- brous. Leaves with petioles subterete, flattened adaxi- ally, rugose, 4-7 (13) mm long, 1-3 mm diam., gla- brous, blade sometimes narrowly decurrent along upper '/3 causing it to appear slightly winged; leaf blades thin- to thick-coriaceous, lanceolate, elliptic, ovate- or ob- long-elliptic, 6-17.5 cm long, 2-6.5 cm wide, base broadly cuneate or rounded, apex long-acuminate, gla- brous, 5-7-plinerved from near base, midrib raised and thickened in basal 1 .5 cm, then impressed above, raised and prominent beneath, lateral nerves impressed or be- coming plane apically above and prominent beneath, re- ticulate veinlets impressed or slightly raised above and inconspicuous beneath or conspicuously elevated be- neath and then leaves appearing slightly bullate. Inflo- rescences capitate-globose and (6) 8-10-flowered or elongate-cylindric and 10-40-flowered, with numerous, imbricate, coriaceous, smooth, glabrous to densely ap- pressed pilose, broadly ovate to oblong-ovate bracts to 20 mm long at base; rachis subterete, bluntly angled, rugose, 1.5-20 cm long (but still in bud), 3-4.5 mm diam., glabrous, nodes congested or widespread; floral bract chartaceous to membranaceous, oblong-elliptic, 20-36 mm long, 16-25 mm wide, apex rounded, gla- brous, pink; pedicels slightly rugose, 3-10 (13) mm long, 1-2 mm diam., glabrous; bracteoles basal, char- taceous, oblong or ovate-oblong, keeled, slightly auric- ulate, 4-8.5 mm long, ( 1 .5) 3-5 mm wide, apex round- ed, margin glandular-fimbriate and broadly scarious. Flowers with calyx 7-15 mm long, glabrous, tube cy- lindric or somewhat campanulate, slightly rugose, 1.5- 3 mm long, basally rounded but shallowly undulate, bearing glandular fimbriae, limb cylindric somewhat spreading, 5.5-1 1 mm long, bearing scattered glandular fimbriae, lobes oblong, 3-8 mm long, 1 .5-4 mm wide, LUTEYN AND WILBUR: FAMILY #172 ERICACEAE 31 apex rounded and apiculate, margin undulate, broadly scarious and glandular-fimbriate, separate at anthesis but imbricate in bud and after anthesis when then erect and curling around base of style; corolla cylindric or bottle- shaped, slightly carnose, 18-31 mm long, ca. 5-7 mm diam., glabrous, dark pink to red, lobes deltoid, ca. 1- 1.5 mm long, widely flaring, white; stamens 14.5-22 mm long; filaments alternately 2.5-5 mm and 5.5-10 mm long, glabrous or densely short-pilose adaxially in apical half; anthers alternately 13-18 mm or 1 1-14 mm long; thecae 4-6 mm long. Mature berry not seen, but immature to at least 1 1 mm diam. Cavendishia calycina is endemic to Chiriqui Province, Panama, where it is found in montane cloud forest, 1100-2200 m elevation. It has not yet been collected in Costa Rica. Flowering oc- curs throughout the year; immature fruits are rath- er more sporadic. Locally Common. Based on the protologue (Smith, 1941), Cav- endishia calycina is characterized by having a ca- lyx limb longer than the tube, calyx lobes that are long, oblong, glandular-fimbriate, and somewhat undulate-margined and that remain erect after an- thesis while curled around the style base, and by elongate, oblong bracteoles. The calyx lobes are noticeably imbricate before and after anthesis but are separate (or at best contiguous in the basal half) during anthesis. The collections from west- ern Panama (region of Boquete) show two forms: one, like the type, has capitate-globose inflores- cences with ca. 6-10 flowers; the other has elon- gate-cylindric inflorescences with 10-40 flowers that are characterized by an elongate rachis (4-10 cm vs. ca. 1.5 cm long) and calyx limbs (7-11 mm vs. 5-6 mm long). Until more collections are available, we have chosen not to recognize those populations with elongate-cylindric inflorescences as distinct. Because of the imbricate nature of the calyx lobes before and after anthesis, C. calycina may be keyed under both ser. Cavendishiae and Imbricatae in Luteyn (1983). Cavendishia capitulata J. D. Smith, Bot. Gaz. 25: 147. 1898. Illustration: J. L. Luteyn, Mem. New York Bot. Card. 28(3): 101, fig. 48. 1976. J. L. Luteyn, Fl. Neotrop. Monogr. 35: 168, fig. 57A-D. 1983. Figure 7. Erect, much-branched, spreading, usually epiphytic shrubs 0.5-1.5 (2) m tall, with twigs glabrous or dense- ly hirsute with spreading white or yellowish trichomes 0.5 mm long, frequently also glandular-fimbriate. Leaves with petioles 1-3 (5) mm long, glabrate; leaf blades obovate, oblanceolate, elliptic, oblong-elliptic or ovate, 1.5-3 (6) cm long, 0.5-1.5 (3) cm wide, base cuneate or rounded, apex rounded, obtuse or rarely ta- pering to a blunt tip, 3-5 (7)-plinerved. Inflorescences capitate, congested, 3-6 (7)-flowered; rachis 1-3 (7) mm long, glabrous; floral bract obovate, shallowly emargin- ate, 11-14 (19) mm long, glabrous, margin sparsely glandular-fimbriate, purple to reddish-purple, frequently glandular-fimbriate; pedicels (0.5) 1.5-2 (5) mm long, glabrous or pilose, rarely glandular-fimbriate; bracteoles basal, (1) 3-6 (8.5) mm long, margin densely glandular- fimbriate with fimbriae to 0.8 mm long. Flowers with calyx 4.5-7.5 mm long, glabrous, tube cylindric or bar- rei-shaped, often pentagonal, smooth or minutely rugose, 1-3 mm long and slightly apophysate at base with a narrow rim or collar, limb erect, somewhat spreading or campanulate, 1.5-4 mm long, lobes 0.5-1.5 mm long, triangular, erect or somewhat connivent after anthesis, margin glandular-fimbriate with fimbriae to 0.2 mm long, sinuses concave or obtuse, rarely acute; corolla cylindric, narrowed at base, constricted at apex, 10-16 mm long, 4-5 mm diam., glabrous, basal '/4 white or pale red, middle half purple or reddish-purple, apical '/4 plus lobes white, lobes triangular, obtuse, ca. 1 mm long; stamens 9-14.5 mm long; filaments alternately 1.5-3 mm and 3-6 mm long, glabrous or pubescent; anthers alternately 7.5-12.5 mm and 5.5-10 mm long, thecae 2- 3 mm long. Berry spherical, 5-9 mm diam., purple to blue-black. Cavendishia capitulata is found in tropical wet forest, premontane wet forest, montane cloud for- est, thickets along roadside slopes, and remnant trees in pastureland, 350-2700 m elevation. It oc- curs in Costa Rica, Panama, and northern Colom- bia (Antioquia Dept.). Flowering occurs through- out the year; fruiting from December to July. Widespread. Cavendishia capitulata is characterized by its small leaves that are apically obtuse or rounded; capitate, 3-4-flowered inflorescences; and corol- las that are purple or reddish-purple with white lobes. When living, its leaves are often densely congested, imbricate, and point backward along the mature stems, although in size and shape there is considerable variation. In floral characters the species is very uniform. Cavendishia capitulata belongs to ser. Cavendishiae and therein is likely most closely related to C. pilosa Luteyn, a north- ern Colombian endemic (Antioquia Dept.). Al- though the two species ranges overlap in Antio- quia, they are not sympatric. They both have sim- ilar small leaves with apex rounded or obtuse, in- florescences few-flowered, rachises short and congested, pedicels very short and apically swol- len, oblong bracteoles that may extend beyond the calyces, calyx tubes basally truncate or short-ri- mose, and relatively short corollas. The two spe- cies are very distinct, however, and may be easily separated by C. capitulata being glabrous (not pi- lose) and having fewer flowered inflorescences 32 FIELDIANA: BOTANY (3-7 vs. 10-12 flowers), a shorter rachis (1-7 mm vs. 1-2 cm long), and longer floral bracts (11-19 mm vs. ca. 9 mm long) that are spreading not appressed and concealing flowers. Cavendishia capitulata may hybridize with C. bracteata, based on morphologically intermediate collections (Lu- teyn, 1976c). The species is visited by the hum- mingbird Lampornis castaneoventris calolaemus in Costa Rica (pers. observ.; Luteyn, 1998). Cavendishia chiriquiensis A. C. Smith, Ann. Missouri Bot. Card. 28: 449. 1941. Figure 9. Weakly erect, epiphytic shrubs to 3 m tall. Leaves with petioles 4-8 mm long, glabrate; leaf blades elliptic or ovate, sometimes bullate, 4.5-9 cm long, 1.5-3.5 cm wide, base cuneate, sometimes attenuate, apex long-acu- minate or caudate-acuminate, glabrous or puberulous at base of midrib, 3-5 (7)-plinerved, newly unfolding leaves often with dense caducous, globular glands ca. 0.1-0.2 mm diam. on adaxial surface, these leaving mi- nute reddish or blackish papillate scars on mature leaves. Inflorescences loosely 6-12-flowered, the buds with vis- cid, white exudate; rachis 1.5-10.5 cm long, thin, deli- cate, ca. 1-1.5 mm diam., sometimes flexuous, glabrous, sometimes with tiny, globular glands 0.1-0.3 (0.5) mm diam. along entire length or concentrated basally; floral bract oblong to oblanceolate, 15-16 mm long, glabrous, pink to red, margin with 11-12 sessile globular glands, to 0.5 mm diam., scattered along edge, these often ca- ducous; pedicels 4-7 mm long, glabrous but with few globular glands and cilia apically; bracteoles basal, lin- ear to linear-lanceolate or ovate to oblong-ovate, 1-2 (4) mm long, apex callose glandular and this deciduous, marginally with globular glands. Flowers with calyx 3.5-4.5 mm long, glabrous, tube cylindric or somewhat pentagonal, smooth or rugose, 1 .5-2 (2.5) mm long, in- conspicuously apophysate at base and with globular glands along margin of apophysis (not obscuring tube surface), limb campanulate to spreading-erect, rugose, ribbed, 1.5-2.5 mm long, with scattered glands similar to those on tube, lobes 0.5-1.5 mm long, triangular, sometimes apiculate, erect after anthesis, completely to only marginally glandular-callose, sinuses concave or flat; corolla tubular, 16-24 mm long, 4-4.5 mm diam., bluish- to reddish-purple at base, purple apically, gla- brous to pilose in apical '/,. lobes triangular, obtuse, 1- 2 mm long; stamens 14.5-21 mm long; filaments alter- nately ca. 3 mm and 5-7.5 "mm long, glabrous or pilose; anthers alternately 13-20 mm and 9.5-16.5 mm long, thecae 3-6.5 mm long. Berry not seen. Cavendishia chiriquiensis is found in the Cor- dillera de Tilar£n and Cordillera de Talamanca of Costa Rica and Panama, 700-1900 m elevation. Endangered. Cavendishia chiriquiensis is characterized by having an often lianoid habit, lanceolate leaf blades with long-acuminate to caudate-acuminate apices, pink to red floral bracts, apophysate calyx tube, and bluish- to reddish-purple corollas. It be- longs to ser. Lactiviscidae and is probably most closely related to C. panamensis, with which it shares densely glandular-margined floral bracts and bracteoles. Cavendishia chiriquiensis also shows morphological similarities with C. endresii, both species having sometimes flexuous rachises, angular to subspherical glands at the articulation of the calyx and pedicel, and calyx lobes some- times completely glandular thickened. The differences between the two varieties of Cavendishia chiriquiensis are given in the follow- ing key: Key to the Varieties of Cavendishia chiriquiensis la. Leaf blades flat; rachis 1.5-4.5 cm long; bracteoles linear to linear-lanceolate; corolla pilose in apical '/, C. chiriquiensis var. chiriquiensis Ib. Leaf blades bullate; rachis (2.5) 6-10.5 cm long; bracteoles ovate to oblong-ovate; corolla glabrous or only weakly puberulous in apical '/, C. chiriquiensis var. bullata Var. chiriquiensis. Illustrated: J. L. Luteyn, Mem. New York Bot. Card. 28(3): 113, fig. 60. 1976. Leaves with petioles densely white puberulous, gla- brate; leaf blades flat, 3-5-plinerved, midrib, lateral nerves and veinlets impressed to slightly raised and con- spicuous above, all veins raised beneath but usually only midrib conspicuous. Inflorescence rachis 1.5-4.5 cm long; bracteoles linear to linear-lanceolate. Calyx tube slightly apophysate; corolla pilose in apical '/3. Variety chiriquiensis is found in montane for- est, elfin forest, and mossy forest and along forest trails, 1800-1900 m elevation. It is distributed in Panama, on the eastern slopes of Volcdn Baru, and in Costa Rica, where it is known only from Lim6n Province (Parq. Nac. Cord. Talamanca, Herrera & Gamboa 5993, F). FJowering speci- mens have been collected in December, January, and July; fruits are unknown. This variety is vis- ited by the hummingbird Lampornis castaneov- entris castaneoventris in Panama (pers. observ.; Luteyn, 1998). LUTEYN AND WILBUR: FAMILY #172 ERICACEAE 33 Var. bullata Luteyn, Ann. Missouri Bot. Card. 68: 157. 1981. Leaves with petioles inconspicuously puberulent; leaf blades usually strongly bullate, 5-7-plinerved with mid- rib and lateral nerves moderately to strongly impressed above and conspicuously raised beneath, veinlets slight- ly raised above and usually obscure beneath. Inflores- cence rachis (2.5) 6-10.5 cm long; bracteoles ovate to oblong-ovate. Calyx tube moderately to conspicuously apophysate; corolla glabrous or only weakly puberulous in apical '/,. Variety bullata is found in cloud forest, along the Continental Divide in Chiriqui and Code Provinces, Panama, 700-1750 m elevation. In Costa Rica, it is known only from the Cordillera Tilar6n in the Monteverde Cloud Forest Reserve (Haber 4705 and Haber & Bello 7376). Flowering specimens have been collected in July, August, and September; fruits are unknown. Cavendishia ciliata Luteyn, Brittonia 28: 45. 1976. Illustration: J. L. Luteyn, Mem. New York Bot. Card. 28(3): 110, fig. 57. 1976. Fig- ure 12. Terrestrial and epiphytic shrubs to 1 m tall, with twigs glabrous to weakly pilose, with tiny, spherical glands. Leaves with petioles 3-6 mm long, pilose; leaf blades elliptic to oblong-elliptic, (5) 1 1-15 (19) cm long, (2) 4-7.5 cm wide, base obtuse to rounded, apex acu- minate (sometimes abruptly), glabrous but with scat- tered, reddish, pustular glands, 0.1-0.2 mm diam., along upper surface, hirsute beneath, 3 (5)-plinerved. Inflo- rescences cylindric, congested, 1 8-29-flowered, viscid; rachis 6-8 cm long, 1 .5-3 mm diam., densely spreading- hirsute, also with globular glands; floral bract oblong to obovate, (11) 18-26 mm long, base auriculate, apex ob- tuse to rounded, margin ciliate and glandular-fimbriate, dark pink; pedicels spreading, (7) 12-16 mm long, hir- sute, also with globular glands; bracteoles basal, ovate, 1-1.5 mm long, slightly pubescent, apex and margin with globular glands. Flowers with calyx 4.5-5.5 (7.5) mm long, tube 1.5-2.5 mm long, conspicuously apoph- ysate at base with margin deeply 5-lobed and also bear- ing globular glands, limb cylindric-campanulate, 2.5-3.5 (5) mm long, basally pilose, lobes ca. 1 mm long, tri- angular, glabrous, erect after anthesis, completely glan- dular-callose, sinuses flat; corolla cylindric, (18) 20- 21.5 mm long, 6-7.5 mm diam., swollen at base, sharply constricted and ca. 3.5 mm at throat, pilose apically, pinkish-purple but paler and almost whitened at base and apically, lobes ca. 1.5 mm long, strongly reflexed; sta- mens 14-15.5 mm long; filaments alternately 2.5-3 mm and 4-5.5 mm long, apically strigose; anthers alternately 12.5-13.5 mm and 10.5-11.5 mm long, thecae 5-5.5 mm long. Berry not seen. Cavendishia ciliata is found in disturbed cloud forest, 1400-1700 m elevation. It is endemic to Costa Rica and is known from only four collec- tions in Cartago Province from the vicinity of El Muneco and also along CR Hwy. 230 between Trinidad and Coliblanco. Flowering specimens have been collected in May-July; fruits are un- known. Endangered. Cavendishia ciliata is characterized by having a stout, densely spreading-hirsute rachis; basally auriculate floral bracts with ciliate and glandular- fimbriate margins; a conspicuously apophysate and deeply lobed calyx tube; and calyx lobes that are glandular thickened over the entire surface. All parts of its inflorescence have the dark pink to red, globular to disc-shaped sessile glands char- acteristic of C. sen Lactiviscidae, but it apparently has no close relatives there and cannot be con- fused with other members. Cavendishia complectens Hemsl. van complec- tens, Biol. Cent.-amer., Bot. 2: 272. 1881. Chu- palon complectens (Hemsl.) Kuntze, Rev. Gen. PI. 2: 383. 1891. Illustration: J. L. Luteyn, Mem. New York Bot. Card. 28(3): 96, fig. 43. 1976. J. L. Luteyn, Fl. Neotrop. Monogr. 35: 118, fig. 48G. 1983. Figure 11. Epiphytic shrubs to 2 m tall, sometimes subscandent (or rarely terrestrial), glabrous throughout. Leaves with petioles to 5 mm long; leaf blades subsessile, amplexi- caul, oval to suborbicular, (5) 8-16 (18) cm long, (4) 5.5-10 (13) cm wide, base deeply cordate with lobes rounded and usually overlapping, apex obtuse to round- ed, pinnately veined. Inflorescences loosely cylindric, usually solitary but with as many as 3 per axil, 8-80- flowered; rachis (1.2) 2-9 (20) cm long; floral bract el- liptic to ovate-lanceolate, (10) 11-16 (20) mm long, smooth when fresh, often becoming ribbed when dry, ascending and often appressed at anthesis but usually spreading after anthesis, dark pink to red, margin glan- dular-fimbriate; pedicels (2) 4-9 (11) mm long; bracte- oles similar to floral bract, ( 1 ) 3-4 (7) mm long. Flowers with calyx 4-10 mm long, tube hemispheric, smooth or minutely muricate, 1-3.5 (6.5) mm long, at most with a very narrow basal rim, limb cylindric or campanulate, 2.5-6.3 mm long, lobes (1.5) 2-3 (4) mm long, reni- form, broadly ovate, broadly imbricate, the tips spread- ing at anthesis, connivent and tightly curling around base of style after anthesis, marginally glandular-fimbri- ate; corolla cylindric or bottle-shaped, 6-12 mm long, 4-6.5 mm diam., multicolored (white at extreme base then distally purplish-black, then green in midsection, then white lobes), lobes oblong, obtuse, 2-2.5 mm long; stamens 4.5-8.5 mm long; filaments alternately 2-3 mm and 2.7-4.3 mm long, scattered pubescent; anthers al- ternately 2.5-7 mm and 2.2-6.5 mm long, thecae 2-4 mm long. Berry spherical, 9-13 mm diam., purple to blue-black. Cavendishia complectens van complectens is 34 FIELDIANA: BOTANY found in cloud or mossy forest, disturbed forest edges, remnant trees in pastureland, fence-row trees, and thickets and on weedy roadside slopes, 800-1800 m elevation. It is distributed from Nic- aragua to northwestern Colombia (Choc6 Dept.). Flowering and fruiting occur sporadically throughout the year. Widespread. Cavendishia complectens is one of the easiest species of Cavendishia in Costa Rica to recognize because of its semi-orbicular, deeply cordate, strongly amplexicaul leaves. Variety complectens is widespread, whereas var. striata (A. C. Smith) Luteyn and var. cylindrica Luteyn are restricted to South America (Luteyn, 1983). Cavendishia complectens belongs to ser. Imbricatae, character- ized by calyx lobes that are broadly imbricate at anthesis. Therein it is somewhat isolated, showing no apparent close relationships with any of the other species in the series. Although the inflores- cences of C. complectens var. cylindrica (Colom- bia) closely resemble those of C. confertiflora in having elongate rachises, congested flowers, and persistent floral bracts (and bracteoles) that are strongly concave and tightly clasp the flowers at anthesis, this seems to be only a matter of con- vergence. Cavendishia complectens may hybrid- ize with C. axillaris, some collections being mor- phologically intermediate (Luteyn, 1976c). Cav- endishia complectens is visited by the humming- bird Lampornis calolaema in Costa Rica (pers. observ.; Luteyn, 1998). Cavendishia confertiflora A. C. Smith, Phyto- logia 1: 210. 1937. Illustration: J. L. Luteyn, Mem. New York Bot. Card. 28(3): 106, fig. 53. 1976. Figure 14. Terrestrial shrubs to 3 m tall, sometimes epiphytic, glabrous throughout. Leaves with petioles 6-10 mm long; leaf blades oblong, ovate-elliptic or oblong-ellip- tic, (8) 10-15 (20) cm long, (3) 3.5-5 (7) cm wide, base rounded or obtuse, apex acuminate, 5 (7)-plinerved. In- florescences cylindric, tightly congested, 13-32-flow- ered; rachis 3-6.5 cm long; floral bract semi-orbicular to broadly oval, tightly appressed to and covering calyx and lower half of corolla at anthesis, persisting to fruit, concave, 10-17 mm long, margin glandular-fimbriate, pink to dark red; pedicels ca. 2 mm long; bracteoles basal, clasping lower half of calyx tube, 4.5-5 mm long, margin glandular-fimbriate. Flowers with calyx 7-9.5 mm long, glandular-fimbriate, tube obconic to hemi- spheric, bluntly 5-angled, slightly rugose, 2.5-3 mm long, limb erect or spreading, 4-6.5 mm long, lobes 3- 5.5 mm long, imbricate at base, oblong, connivent and tightly enclosing style after anthesis, margin glandular- fimbriate; corolla bottle-shaped, 13-17 mm long, ca. 8 mm diam. at broadest point, dark pink at base and white at apex, lobes oblong or triangular, ca. 1.5 mm long, white; stamens 9.5-11 mm long; filaments alternately 2-3 mm and 3.5-5 mm long; anthers alternately 8-9 mm and 7-8 mm long, thecae 2.5-3.5 mm long. Berry spherical, not seen mature, but at least 10 mm diam. Cavendishia confertiflora may be found in ev- ergreen forest, primary forest, roadside thickets, and disturbed areas, 1 200-3200 m elevation. It is endemic to a small area of the Cordillera de Tala- manca, north of the General Valley in San Jose Province, Costa Rica. Flowering specimens have been collected in January, November, and Decem- ber; immature fruiting in December and January. Locally Common. Cavendishia confertiflora is characterized by having cylindric inflorescences that are composed of many congested imbricate flowers, tightly ap- pressed and persistent floral bracts (and usually bracteoles), and imbricate calyx lobes. It belongs to ser. Imbricatae, where it is morphologically most similar to C. zamorensis A. C. Smith, an Ecuadorean endemic. In Costa Rica, sterile ma- terial of C. confertiflora may be confused with C. bracteata, and in leaf alone it is virtually impos- sible to distinguish these taxa. They also share similar glands on the calyx lobes and bracteoles. Cavendishia confertiflora also resembles C. ca- lycina (a western Panamanian endemic), both spe- cies having proportionally long calyx limbs, elon- gate-imbricate calyx lobes, and clasping bracte- oles; none of these features, however, are well de- veloped in C. calycina. The species are easily distinguished by the appressed, persistent floral bracts and tightly congested flowers of C. confer- tiflora. Plants are visited by hummingbirds (pers. observ.). Cavendishia davidsei Luteyn, Brittonia 48: 241, fig. 1. 1996. Figures 6 and 7. Epiphytic shrubs; mature stems terete, striate to ribbed, glabrate; twigs complanate, striate to ribbed, densely short-pilose with white hairs, glabrate. Leaves with petioles broadly flattened and canaliculate adaxi- ally, rugose, 3-5 mm long, densely short-hirsute, gla- brate; leaf blades coriaceous, somewhat bullate, elliptic- ovate, elliptic to oblong-elliptic, rarely ovate, (2) 4-6 cm long, 1 .5-2.5 cm wide, base rounded, cordate to sub- cordate, sometimes somewhat clasping, apex acute, ob- tuse to narrowly rounded, glabrous to short-hirsute at base of nerves above, moderately and persistently short- hirsute beneath, also with scattered, minute, glandular fimbriae on both surfaces, 5-plinerved from above base, midrib raised and thickened in basal 2-10 mm, then im- pressed apically above, conspicuously raised beneath, lateral nerves impressed above (the inner pair joined with midrib for 2-10 mm) and conspicuously raised be- LUTEYN AND WILBUR: FAMILY #172 ERICACEAE 35 neath, reticulate veinlets slightly impressed above but conspicuously raised beneath. Inflorescences loosely cylindric, racemose, 8-32-flowered, surrounded at base by a series of ovate, acute bracts to 2 mm long; rachis angled, slender, striate, 3-9 cm long, densely short-pi- lose with white hairs; floral bract flat, elliptic, sometimes somewhat oblanceolate, 1 8-25 mm long, base tapering, apex acute, short-pilose on upper surface, margin glan- dular-fimbriate apically, pink to red; pedicels striate, 9- 10 mm long, densely short-pilose, eglandular; bracteoles basal, elliptic-oblong, 2-3 mm long, apex obtuse, short- pilose, margin glandular-fimbriate. Flowers with calyx 4-5.5 mm long, moderately short-hirsute, tube cylindric, ribbed, 1.5-2.5 mm long, strongly apophysate basally with margin undulate to lobed, limb cylindric-campan- ulate, 2.5-3 mm long, lobes deltate, ca. 1.3 mm long, apex acute, eglandular, erect after anthesis, sinuses rounded; corolla cylindric, 11-13 mm long, ca. 4 mm diam., dark violet to purple, short-pilose, lobes erect, ovate-deltate, to 1 mm long, acute, white; stamens over- all equal, 11-12 mm long; filaments distinct, alternately ca. 3 mm and 5.2 mm long, glabrous to weakly short- pilose; anthers alternately ca. 9 mm and 8 mm long, thecae alternately ca. 3 mm and 4.5 mm long. Berry not Cavendishia davidsei is found in montane (ap- parently Quercus) cloud forest, 2000-2600 m el- evation. It is distributed in Costa Rica, where it is found on the Continental Divide of Cerro Bekom (Puntarenas-Limon Prov. border: Davidse et al. 25736, Davidse & Herrera 26204) and on Cerro Hoffman (Limon Prov., close to the border with Panama: Davidse et al. 28601), and Cerro Frantz- ius (Puntarenas Prov.: Davidse et al. 28552). In Panama it has been collected once in Bocas del Toro Province. It is known from a total of five collections. Flowering specimens have been col- lected in March and September; fruits are un- known. Endangered. Cavendishia davidsei is distinguished by its short-pilose petioles, elongate rachis, multiflow- ered inflorescences, elongated floral bracts that are flat and tapering at base, eglandular pedicels, con- spicuously apophysate calyx tube, calyx lobes that are erect after anthesis, short-pilose corolla, and limited geographical range. Cavendishia davidsei belongs to sect. Quereme and is morphologically similar to C. talamancensis and C. gomezii. A comparison of its important characters along with other Costa Rican members of sect. Quereme is given in Table 1. Cavendishia endresii Hemsl., Biol. Cent.-amer, Bot. 2: 273. 1881. Chupalon endresii (Hemsl.) Kuntze, Rev. Gen. PI. 2: 383. 1891. C. gluti- nosa Hoerold, Bot. Jahrb. Syst. 42: 321. 1909. Illustration: J. L. Luteyn, Mem. New York Bot. Card. 28(3): 116, fig. 63. 1976. J. L. Luteyn, Fl. Neotrop. Monogr. 35: 247, fig. 74A-C. 1983. Figure 9. Terrestrial and epiphytic shrubs to 1-2.5 m tall, with twigs puberulent and usually also glandular-fimbriate. Leaves with petioles 2.5-5 (7) mm long, puberulent adaxially, rarely glabrate; leaf blades ovate, elliptic- or oblong-lanceolate, (3.5) 5-8.5 (11) cm long, 1-2 cm wide, base cuneate, apex long-acuminate, pinnately veined or conspicuously (3) 5 (7)-plinerved, midrib pu- berulent, glabrate above. Inflorescences fusiform, 3-8- flowered, viscid; rachis (0.7) 1-2 (3) cm long, glabrous; floral bract oblong to oblanceolate, 15-20 (25) mm long, apex rounded to acute, margin ciliate, purple to reddish- purple; pedicels (3.5) 5-9 mm long, glabrous, but api- cally surrounded by flesh-colored, angular disc-shaped glands; bracteoles basal, ovate-lanceolate, 0.5-1.5 mm long, glabrous, completely glandular-callose except for small, central basal portion. Flowers with calyx 3-4 mm long, glabrous, tube bluntly 5-angled or -ribbed, 1-1.5 mm long, conspicuously apophysate, limb campanulate or rarely cylindric, smooth, (1.5) 2-2.5 (3) mm long, lobes 0.5-1 mm long, triangular, erect after anthesis, completely glandular-callose, sinuses concave to almost flat; corolla cylindric, slightly constricted at base, slight- ly narrowed at throat, (10) 15-20 mm long, 2.5-5 mm diam., glabrous, white at base and upper V4-V3, purple or reddish-purple or rarely rose in middle, lobes white with purple or rose tips and margins; stamens 12.5-16 mm long; filaments alternately 1.5-3.5 mm and 3.5-5.5 mm long, glabrate; anthers alternately 10.5-15 mm and 8.5- 12.5 mm long, thecae 2-4 mm long; style exserted at anthesis. Berry spherical, 10-12 mm diam., blue-black. Cavendishia endresii is occasionally found in cloud forest, 1000-1700 m elevation. It is endem- ic to the Central Valley of Costa Rica and extreme western Panama (one collection, Chiriqui Prov.: Fortuna Dam area). Flowering specimens have been collected in February-September; fruiting specimens in May-July. Locally Common. Cavendishia endresii is characterized by leaves that are usually pinnately veined, short rachis bearing ca. 3-8 flowers, short-ciliate floral bracts that are reddish-purple to purple in color, angular and flesh-colored pedicellar glands, almost com- pletely glandular-callose thickened bracteoles, and purple or reddish-purple corollas with white bases and lobes. The very large, angular disc-shaped, flesh-colored glands at the top of the pedicel are diagnostic for this species. Cavendishia endresii belongs to sect. Callista, where it has no close relatives in Mesoamerica. Instead, it is related to the Colombian C. violacea A. C. Smith and C. aurantiaca Luteyn, with which it has in common angular glands at the ped- icel tips and (usually) pinnately veined leaves. Cavendishia endresii may hybridize with C. 36 FIELDIANA: BOTANY i 1 2 a > ^ j S 8 2? .D o _u Q 9 15 „ O. c « n -s oo" c "5b _0 ^ c T3 CJ- o " •Q >« 5 o "3 ^ ^ •^ O oo 2 Tf Q "5 ° ^ « ;2 . Q a. *> C/l ^^ WJ ^« ^^ B • ^ «3 f^ c S i i/^ J^ y ^3 ••^ >^ i— o 00 ^ U 1 ^ ** "y "* o ig — C "0 §•2 •« ^. .h ^•N QJ C — ,*••»«, Jv — '— Q ^— s &* Tj O ^ 3 1 — to -C 53- r-* _o | .E es c3 n •c K, £ 5" 0 0 V 1 U Cavendis querem u H | ^§1 -3 •^ K "2 7 8 § ^~ v> •=^0 3 •C — o 00 1 2 — S ~ c U U i ^ _w _w 00 S c u £ .S l! <^> W C 1 C C M ll 5 3 J9 ^^^ i* p2 — i. 4i Q "^*" — •O — ^™ W) O O O j-^ 00 02 O SfsO — (E o; !2U < o ~ Q. W5 C t« "v eO W5 ! 3 ^ 1 sal 78'! 1" 2 § 00 o nts ll e ca C 1 2^0 2<£ °JG x> < £ O sll 11 "g •2 M) 3 -~ <* EL I a | z. 8 °O 00 § u 1 1 i s u I O 0 8 J3 1 O e S _~ 3 >^ 00 ^, i >» °u "3 1 c -r > y >-, u U -C 3 O J g "3 r^ Q. o 'ob o o «1 1 S u 'S. "u •7 ^ '5.^ S- nj C "s- "1 I 5; a .5 "3 « _« PQ Puberul if! 111 1! Present W s! ^ 5-s. c u GO 3^ !5 o 3 CO M > Petiole pubescence Flowers/inflorescence Rachis length (cm) Rachis pubescence •s ^ III - - ~- O 0 0 EEE Pedicel length (mm) Pedicel pubescence M •o "H '•5 Bracteole shape 5 'S £ 1 1 ! > w o 0 S3 5-° M OO V V X X X ji'j^.^ « « cs UUU Calyx sinuses (at anthesi Corolla pubescence LUTEYN AND WILBUR: FAMILY #172 ERICACEAE 37 wercklei, C. bracteata, and C. quereme (Luteyn, 1976c). A common name around San Jose Prov- ince is colmillo. This species is visited by hum- mingbirds (pers. observ.). Cavendishia fortunensis Luteyn in Wilbur & Lu- teyn, Ann. Missouri Bot. Card. 68: 157. 1981. Figure 1. Epiphytic or terrestrial shrubs; mature stems subterete or bluntly angled, slightly striate, glabrous, with scat- tered pustular glands to 0.3 mm diam., reddish-brown. Leaves with petioles subterete, rugose, 3-5 mm long, 2.5-3.5 mm diam., glabrous or hispid only on adaxial surface, bearing globular glands 0.2 mm diam. apically; leaf blades coriaceous, bullate, lanceolate or elliptic, (5) 7-14.5 cm long, 2-5.5 cm wide, base obtuse or narrowly rounded, apex acuminate (sometimes abruptly short-cau- date-acuminate), glabrous but with scattered black ses- sile, pustular glands 0.1 mm diam. along upper surface, these often caducous leaving a reddish punctate scar, 5 (7)-plinerved with innermost pair of lateral nerves aris- ing 1 .5 cm above base, midrib and lateral nerves deeply impressed above and conspicuously raised beneath, re- ticulate veinlets slightly impressed above and raised be- neath. Inflorescences loosely cylindric, to ca. 35-flow- ered, lowest few nodes sterile, flowers with white, viscid exudate; rachis subterete, bluntly angled, striate, 1 1-25 cm long, 3-4 mm diam., glabrous, bearing scattered, globular or angular glands ca. 0.2 mm long; floral bract ovate, oblong or oval-elliptic, 21-25 mm long, 10-12 mm wide, base narrowed, truncate and clasping, apex rounded, margin crisped and bearing 12-20, dark red, globular glands, 0.1-0.2 mm diam. on each side, gla- brous or weakly ciliate, pink; pedicels swollen apically, striate, 5-7 mm long, ca. 0.75 mm diam., glabrous, bear- ing globular glands concentrated apically; bracteoles medial, clasping or nearly concealing calyx and lower '/, of corolla at anthesis, oblong-oblanceolate, 10-14 mm long, 6-8 mm wide, margin crisped and bearing dark red, globular glands scattered along edge. Flowers with calyx ca. 3.5-5 mm long, glabrous, tube cylindric, ob- scurely ribbed, 1.5-2.5 mm long, truncate basally, cov- ered by globular or angular glands 0.2 mm diam., pink, limb spreading-campanulate, 2-3 mm long and covered by globular glands, lobes broadly triangular, ca. 1 mm long, erect after anthesis, completely glandular-callose in apical 7, and covered by globular glands, sinuses flat to concave; corolla thin-carnose, cylindric, slightly nar- rowed apically, 18-20 mm long, 4-6 mm diam., gla- brous without but sparsely pilose within, pearl white to pinkish when fresh, lobes triangular, ca. 1 mm long, apex acute or obtuse; stamens 15-19 mm long; fila- ments alternately 2.5-4.3 mm and 5-7 mm long, the short ones sparsely pilose adaxially at apical tips, the long ones densely pilose adaxially in apical half; anthers alternately 14.5-17.5 mm and 1 1.5-15 mm long, thecae 5-8 mm long. Berry not seen. Cavendishia fortunensis is known only from premontane cloud forest at the Fortuna Dam site in Chiriqui Province, Panama, 1100-1800 m el- evation. Flowering specimens have been collected in April and May; fruits are unknown. Locally Common. Cavendishia fortunensis is characterized by its bullate leaves; elongate, viscid inflorescences; nonapophysate calyx tube; and abundance of globular to spherical glands. It is a typical mem- ber of sen Lactiviscidae and is superficially sim- ilar to C. albopicata Luteyn (NW Colombia) and C. pseudostenophylla, the three having bracteoles medial along the pedicel. However, we feel it is more closely related to C. panamensis and C. ar- izonensis Luteyn (Veraguas Prov., Panama) be- cause of its overall coarse habit, nonapophysate calyx tube, and globular to irregularly spherical glands of the calyx tube that are so abundant as to nearly obscure the surface from view. It also has in common with C. arizonensis bullate leaves and with C. panamensis bracteoles with globular glands. From these two species, C. fortunensis differs by the medial (not basal) insertion of the larger bracteoles (10-14 mm vs. 1.5-4 mm long) and glandular calyx limb. Cavendishia gomezii Luteyn, Brittonia 48: 244. 1996. Epiphytic shrubs; mature stems terete to subterete, sometimes bluntly angled, striate, short-pilose, glabrate; twigs subterete, complanate, bluntly to sharply angled, striate, moderately short-pilose with white hairs. Leaves with petioles subterete, flattened adaxially, rugose, 1.7- 3.3 mm long, densely short-pilose, glabrate; leaf blades coriaceous, ovate, ovate-elliptic, rarely oblong-ovate, 2- 3.5 cm long, 1-1.7 cm wide, base rounded, subcordate, apex acute, obtuse, or narrowly rounded, margin slightly thickened and slightly revolute, sparsely but conspicu- ously short-pilose along margins and at base of midrib beneath, glabrate, also bearing deciduous, minute, glan- dular fimbriae on both surfaces, 3 (5)-plinerved from base, midrib plane or usually raised above and conspic- uously raised beneath, lateral nerves and reticulate vein- lets conspicuously raised above and slightly raised but inconspicuous beneath, sometimes only midrib notice- able beneath. Inflorescences cylindric, racemose, 7-20- flowered; rachis sharply angled, 1-2.5 cm long, gla- brous; floral bract flat, smooth, oblong to oblanceolate, 12-15 mm long, 3-7 mm wide, base tapering to obtuse, apex rounded, eglandular, ciliate apically, red (fide la- bel); pedicels ribbed, 6-8 mm long, eglandular, gla- brous; bracteoles basal, oblong-elliptic, 2-3 (5) mm long, apex rounded to acute, margin densely and decid- uously glandular-fimbriate. Flowers with calyx 4-5 mm long, tube cylindric, ribbed, 1.5-1.6 mm long, base broadly apophysate with the margin flaring, undulate to lobed and short-pilose, limb campanulate-spreading, 2.4-2.5 mm long, glabrous, lobes oblong-ovate, apicu- late, ca. 1 mm long, margin lacerate-glandular, erect af- ter anthesis, sinuses acute; corolla cylindric, 9-10 mm long, ca. 3.5 mm diam., red to vermillion, glabrous, lobes ovate, ca. 1 mm long, apex acute, white; stamens 38 FIELDIANA: BOTANY equal overall, 8-8.5 mm long; filaments distinct, alter- nately ca. 3.5 mm and 1.5 mm long, glabrous; anthers alternately ca. 6.3 mm and 7.5 mm long, thecae alter- nately ca. 3 mm and 2.5 mm long; style slightly exsert- ed, glabrous. Berry not seen. Cavendishia gomezii is endemic to the montane cloud forest of the Cordillera de Talamanca in Bo- cas del Toro Province, Panama, 2400-2900 m el- evation. Although having been collected four times (on one trip) close to the Costa Rican bor- der, it has not yet been collected within the polit- ical boundaries of Costa Rica. Flowering speci- mens have been collected in March; fruits are un- known. Endangered. Cavendishia gomezii is characterized by small, flat, subcordate leaf blades; essentially glabrous inflorescence parts; broadly apophysate calyx tube with the margin flaring; and lacerate-glandular ca- lyx lobes that are erect after anthesis. It belongs to subgen. Cavendishia sect. Quereme following Luteyn (1983). In leaf size and shape and general appearance of the inflorescence, C. gomezii re- sembles C. talamancensis; it differs from that spe- cies by the characters mentioned in the key and Table 1. In the available collections, there is no mention of the plant having the odor of winter- green, a general feature of sect. Quereme. This new species was named to honor Luis Diego G6- mez, whose contributions to Costa Rican natural history have been numerous. Cavendishia herrerae Luteyn & J. F. Morales, Brittonia48: 517. 1996[1997]. Figure 12. Pendent, epiphytic shrubs; mature stems bluntly an- gled, striate, moderately short-hirsute, glabrate, also with scattered, globular glands ca. 0.2-0.3 mm diam.; twigs complanate, sparsely hirsute with white trichomes 0.7- 0.9 mm long. Leaves with petioles rugose, 4-6 (7) mm long, sparsely short-hirsute, glabrate; leaf blades coria- ceous, stiff, lanceolate or elliptic-lanceolate, 8-18 cm long, 1.2-3.5 cm wide, base cuneate or obtuse, apex gradually acuminate, glabrous above, sparsely hirsute along midnerve beneath, glabrate, also with few globular glands 0.1-0.2 mm diam. along margin and on blade above, 3 (5)-plinerved, midrib and inner pair of lateral nerves strongly impressed and conspicuous above, raised and conspicuous beneath, reticulate veinlets slightly impressed on both surfaces. Inflorescences elongate-cylindric, racemose, ca. 14-flowered; rachis striate, ribbed, 10-11 cm long, moderately hirsute and with a few, globular, sessile glands; floral bract oblong or oblong-obovate, 17-21 mm long, apex obtuse or acute, margin ciliolate and bearing few, sessile, globular glands on margin and abaxial surface, pinkish-lilac; ped- icels striate, rugose, 11-14 mm long, densely hirsute, also with few, scattered, globular glands; bracteoles bas- al, striate, oblong-ovate, 4-6 mm long, marginally cili- olate and also bearing few, sessile, globular glands, the tip also glandular-thickened. Flowers with calyx ca. 9 mm long, tube ca. 2 mm long, pink, moderately hirsute, the base apophysate, moderately undulate and bearing few, globular or disc-shaped glands, limb spreading- campanulate, ca. 7 mm long, hirsute basally, lobes tri- angular, ca. 2 mm long, 1 .8-2 mm wide, erect after an- thesis, each margin with an oblong to crescent-shaped callose gland, these contiguous at apex, sinuses acute to obtuse; corolla (lilac fide label) not seen; stamens not seen. Berry not seen. Cavendishia herrerae is endemic to the Carib- bean slope of Costa Rica (Lim6n Prov.). It is known only from the type specimen (Cant6n de Limon, El Progreso, Valle de la Estrella, Matama ridge, trail between Cerro Muchilla and Cerro Avioneta, vie. Rio Suruy, Herrera & Chacon 2623), which was collected in rain forest at 700 m elevation and lacked corollas. The type collec- tion was flowering in April; fruits are unknown. Endangered. Cavendishia herrerae is characterized by hav- ing globular glands in its vegetative and floral parts, a basally apophysate calyx tube, and glan- dular-margined calyx lobes. It belongs to Caven- dishia sect. Engleriana ser. Lactiviscidae, where it is morphologically similar to C. ciliata. It dif- fers from C. ciliata by its narrower leaf blades, leaf apex gradually (not abruptly) acuminate, lon- ger and only moderately (not densely) hirsute ra- chis, shorter floral bracts that are acute (not obtuse or rounded) at the apex, larger bracteoles, hirsute (not glabrous) calyx tube and limb, calyx lobes that are callose glandular only along each margin (not the entire lobe), and its different geography and altitude. Cavendishia herrerae was named af- ter Gerardo Herrera, who collected many new and interesting species of Ericaceae in Costa Rica. Cavendishia lactiviscida Luteyn, Brittonia 28: 46. 1976. Illustration: J. L. Luteyn, Mem. New York Bot. Card. 28(3): 1 1 1, fig. 58. 1976. Fig- ure 9. Terrestrial or epiphytic shrubs to 1 m tall; mature stems and twigs with scattered, subsessile, globular glands to 0.4 mm diam., densest near nodes and on youngest branches. Leaves with petioles (4) 6-7 (10) mm long, hirsute, glabrate; leaf blades bullate, elliptic to lance-elliptic, 4-9 cm long, 1-3 cm wide, base cu- neate, apex long caudate-acuminate *(often abruptly), base of midrib above hirsute but glabrate, adaxial sur- face and margins of mature leaves with caducous, scat- tered, reddish, pustular glands 0.1-0.2 mm diam., 5-pli- nerved. Inflorescences elongate-cylindric, loosely 4-7 ( 1 0)-flowered, the basal several nodes sterile; rachis 4- 9 (15) cm long, thin, 1-1.5 mm diam. at base, glabrous, LUTEYN AND WILBUR: FAMILY #172 ERICACEAE 39 with globular glands; floral bract foliaceous, lance-ellip- tic to oblong-lanceolate, 7-11 (16) mm long, conspicu- ously nerved, pale green, apex acuminate, margin ciliate and with globular glands to 0.2 mm diam.; pedicels (14) 19-25 mm long, glabrous, with globular glands; brac- teoles basal, 1-4 mm long, oval to semi-orbicular, mar- gin glandular as floral bracts. Flowers with calyx 4-6 mm long, glabrous, tube short-cylindric, 10-ribbed, 1.5- 2 mm long, basally apophysate with the margins undu- late and bearing globular glands which secrete a milky exudate, limb erect or wide-spreading to campanulate, smooth to slightly rugose, 2.5-4 mm long, eglandular, lobes (0.5) 1 (1.5) mm long, triangular, erect after an- thesis, each side with an oblong callose gland, sinuses concave to flat; corolla carnose with walls 1.5-2 mm thick when fresh but drying membranous, cylindric, slightly constricted at base, constricted at throat, when immature and prior to anthesis apex broadly rounded and nipple-shaped, 19-29 mm long, ca. 9.5 mm diam., glabrous, greenish-red, lobes ca. 0.5 mm long; stamens 15-20 mm long; filaments alternately 3.2-4.5 mm and 6.3-7.5 mm long, glabrous to weakly short-pilose; an- thers alternately 12.5-17 mm and 10-14 mm long, the- cae 2.5-5 mm long. Berry not seen. Cavendishia lactiviscida is found in cloud for- est, 1200-1850 m elevation. It is endemic to the Cordillera Tilaran, Costa Rica, where it has been collected fewer than 10 times. Flowering speci- mens have been collected in February, May, June, and July; fruits are unknown. Endangered. Cavendishia lactiviscida is characterized by having bullate and caudate-acuminate leaf blades; loosely flowered inflorescences; foliaceous, con- spicuously nerved and pale green floral bracts; long pedicels; basally undulate and apophysate calyx tube; greenish-red corollas; and globular to pustular glands scattered along mature stems, leaves, and inflorescences. Although it superfi- cially resembles C. chiriquiensis, its relationships within Cavendishia ser. Lactiviscidae are obscure. The plants are visited by the hummingbird Do- ryfera ludovicae in Costa Rica (pers. observ.; Lu- teyn, 1998). a series of broadly ovate bracts to 2 mm long; rachis sharply angled, 1.5-2.5 cm long, glabrous, eglandular, the basal 3-4 nodes often sterile; floral bract condupli- cate or concave, oblong-obovate, 7-16 mm long, apex rounded, glabrous, eglandular, at first surrounding floral buds but becoming strongly reflexed through and after anthesis; pedicels striate, 7-10 mm long, glabrous; brac- teoles basal, oblong, 1 .5-2 mm long, apex rounded, gla- brous, margin conspicuously glandular-fimbriate in api- cal !/3. Flowers with calyx 4-5 mm long, glabrous, tube cylindric, broadly and bluntly ribbed, 1.5-2 mm long, basally shallowly but distinctly apophysate, limb erect- spreading to cylindric-campanulate, 2.5-3 mm long, lobes ovate, ca. 1 mm long, erect after anthesis, margin lacerate-glandular, darker in color and apparently thinner in texture than lamina proper, sinuses acute; corolla cy- lindric, slightly narrowing at throat, 14-16.5 mm long, ca. 5 mm diam., lobes ovate-apiculate, ca. 1 mm long; stamens overall equal, ca. 13.5 mm long; filaments dis- tinct, alternately ca. 4.5 mm and 2 mm long, glabrous; anthers alternately ca. 10.7 mm and 12.3 mm long, the- cae alternately ca. 4.7 mm and 3 mm long. Berry not seen. Cavendishia limonensis is found in lowland rain forest, 500-700 m elevation. It is endemic to Costa Rica (Limon Prov.), where it is known only from two collections from Canton de Talamanca, the Alto Uren area, along the road between Que- brada Chaho and Cororina (Herrera 3321) and at Cororina, between Rio Uren and Quebrada Chaho (Fco. Solis et al. 37). Flowering specimens have been collected in July; fruits are unknown. En- dangered. Cavendishia limonensis is characterized by its leaves that lack glandular fimbriae, petioles that are adaxially strongly canaliculate, eglandular flo- ral bractes that are conduplicate or concave and strongly reflexed at anthesis, apophysate calyx tube, and calyx lobes with lacerate-glandular mar- gins. It belongs to Cavendishia sect. Quereme and is most similar to C. laurifolia (Mexico-Guate- mala, Panama-Colombia), differing by the fea- tures mentioned in Table 2. Cavendishia limonensis Luteyn, Brittonia 48: 244, fig. 2. 1996. Figure 9. Epiphytic shrubs; mature stems terete, striate, gla- brous; twigs bluntly angled, striate, densely white pu- berulent, glabrate. Leaves with petioles strongly cana- liculate adaxially, 4-6 mm long, densely puberulent, glabrate; leaf blades coriaceous, ovate to ovate-elliptic, 3.5-5.6 cm long, 1.2-1.8 cm wide, base rounded to broadly obtuse, apex long-acuminate, glabrous, 3 (5)- plinerved, midrib weakly impressed in basal '/2-'/3' be- coming plane to slightly raised in apical '/j-1/,, above, lateral nerves and reticulate veinlets slightly raised above, all nerves slightly raised beneath. Inflorescences loosely racemose, 6-9-flowered. surrounded at base by Cavendishia linearifolia Luteyn & J. F. Morales, Brittonia 48: 516, fig. 1 E-F. 1996[1997]. Fig- ure 15. Pendent, epiphytic shrubs; mature stems terete or subterete, striate, glabrous; twigs terete, glabrous. Leaves with petioles rugose, 1-3 mm long, glabrous; leaf blades coriaceous, linear, strongly revolute, (6) 7.5- 15 cm long, 0.3-0.7 cm wide, base cuneate, apex acute, glabrous, but punctate on both surfaces from deciduous fimbriae borne in slight depressions, apparently pinnate- ly nerved, midrib impressed above, raised and conspic- uous beneath, secondary nerves obscure (if present), re- ticulate veinlets obscure to inconspicuous but slightly raised when present. Inflorescences subspherical, race- 40 FIELDIANA: BOTANY TABLE 2. Salient morphological differences between Cavendishia laurifolia, C. limonensis, and C. quereme. Cavendishia laurifolia Cavendishia limonensis Cavendishia quereme Leaf size (cm) (3.5)6-13 X (1)2-4 3.5-5.6 X 1.2-1.8 (3.5)6-11(14) X (1) 1.5-4(5) Petiole shape Subterete Canaliculate adaxially Subterete or terete Flowers/inflorescence (8) 17-32(45) 6-9 (8) 12-20(25) Rachis length (cm) 4-15(20) 1.5-2.5 2-5 (6) Pedicel length (mm) (9) 14-22 (26) 7-10 (5)8-16(19) Pedicel glands Densely glandular-fim- Absent Absent briate Bracteole glands Densely glandular-fim- Glandular-fimbriate in Glandular-fimbriate briate apical ft Calyx glands Densely glandular-fim- Absent Marginally lacerate- briate glandular Calyx sinuses (at anthesis) Flat to concave Acute Acute Calyx lobes (post-anthesis) Connivent Erect Connivent Corolla length (mm) (6.5)8-10(12) 14-16.5 7-11(13) Corolla shape (cross section) Pentagonal Cylindrical Pentagonal mose, 4-1 1 -flowered; rachis smooth, 1-2 cm long, gla- brous; floral bract obovate or oblong-obovate, 14-24 mm long, base cuneate or attenuate, apex obtuse, gla- brous, dark pink when fresh; pedicels striate, sometimes rugose, 4-6 (7) mm long, glabrous; bracteoles basal, ob- long, ca. 0.5 mm long, margin with 3-4 oblong, callose- fimbriae at tip. Flowers with calyx 6-8 mm long, gla- brous, dark rose, tube smooth, 1 .5-2.5 mm long, basally apophysate with margin somewhat undulate, limb tu- bular-campanulate, smooth to striate, 4-5.5 mm long, lobes triangular, 1.5-3 mm long, erect after anthesis, marginally glandular-thickened with 1-2 thickened fim- briae basally and these coalescing into a supramarginal gland on either side at apex, sinuses obtuse; corolla cy- lindric, narrowed at throat, 1.6-1.8 cm long, 3.5-5 mm diam., glabrous, pink, lobes erect, triangular, 1-1.5 mm long, apex acute, white; stamens overall equal, 13-18 mm long; filaments alternately ca. 2.5 mm and ca. 4 mm long, glabrous; anthers alternately ca. 1 .7 cm and 1 .5 cm long, thecae ca. 4-5 mm long. Berry not seen. Cavendishia linearifolia is found in lowland rain forest, 100-200 m elevation. It is endemic to the Osa Peninsula of Puntarenas Province, Costa Rica, where it is known from only two collections around Golfito, along the road to Rio Bonito (Herrera 5037) and along the road between Gol- fito and Villa Briceno (Hammel et al. 18398). Flowering collections have been made in January; fruits are unknown. Endangered. Cavendishia linearifolia is characterized by its linear leaves. Only two other species of Caven- dishia, C. stenophylla (endemic central Panama) and C. pseudostenophylla (endemic western Chi- riqui Prov., Panama), have linear leaves with which it could be confused, but in both those spe- cies the leaf blade width is double that of C. li- nearifolia (1-2.5 cm vs. 0.3-0.7 cm). Cavendishia linearifolia belongs to Cavendishia sect. Engler- iana sen Englerianae and is similar to C. osaen- sis, differing by its long, linear leaves; smaller bracteoles with strictly terminal glands (not scat- tered along entire margin); and calyx lobe glands that are more highly coalescent at the apex. More collections of both species are needed before their interspecific relationships can be more fully un- derstood. Cavendishia luteynii J. F. Morales in Luteyn & Morales, Brittonia 48: 518. 1996[1997]. Figure 11. Epiphytic shrubs; mature stems subterete to bluntly angled, smooth to somewhat striate, glabrous to glabrate, with scattered, globular glands ca. 0.1 mm diam.; twigs subterete, glabrous. Leaves with petioles inconspicuous, rugose, less than 1 .5 mm long, glabrous; leaf blades co- riaceous, strongly bullate, lanceolate to oblong-lanceo- late, 2.4-6.8 cm long, 1.3-4.2 cm wide, base clasping to nearly amplexicaul, apex acute to scarcely mucronate, glabrous above and glabrate beneath, also with few glan- dular fimbriae, 5-7-plinerved, midrib and lateral nerves strongly and conspicuously impressed above and raised and conspicuous beneath, reticulate veinlets conspicu- ously impressed on both surfaces. Inflorescences elon- gate-cylindric, racemose, 9-18-flowered; rachis some- what angled, striate, ribbed, 3-6 cm long, glabrous, eglandular; floral bract oblong to oblong-elliptic, 15-18 mm long, base obtuse, apex gradually acuminate, margin ciliolate, the tip callose glandular, pinkish-red to red; pedicels striate, rugose, 3-4.5 mm long, glabrous to gla- brate; bracteoles basal, linear-oblong, 3.5-4.5 mm long, margin ciliolate. Flowers with calyx ca. 5.2-6.2 mm long, reddish, glabrous, tube ca. 2.2-2.5 mm long, ba- sally strongly apophysate with a conspicuous collar or rim, eglandular, limb narrowly campanulate, ca. 2.5-3.5 LUTEYN AND WILBUR: FAMILY #172 ERICACEAE 41 mm long, lobes triangular, ca. 0.8 mm long, ca. 1.7 mm wide, eglandular, erect after anthesis, sinuses acute to obtuse; corolla cylindric, 7.5-8 mm long, ca. 2 mm ilium., greenish-white to greenish-yellow, lobes erect, triangular to ovate-triangular, ca. 0.8 mm long, apex acute; stamens 6-7 mm long; filaments alternately ca. 1 mm and 2-2.3 mm long, glabrous to weakly short- pilose; anthers alternately ca. 5.8-6 mm and 4.5-5 mm long, thecae alternately ca. 1.8 mm and 2.7 mm long. Berry not seen. Cavendishia luteynii is endemic to the premon- tane forest, 2200 m elevation, in the Cordillera de Talamanca (Limon Prov.), Costa Rica. It is known only from the following two collections: Canton de Talamanca, Cordillera de Talamanca, Cerro Carguicacha to Cerro Biricuacua, between Ujarras and San Jose Cabecar (Herrera 6140) and Canton de Talamanca, Quebrada Kuisa (J. Bittner 1820). Flowering specimens have been collected in April; fruits are unknown. Endangered. Cavendishia luteynii is characterized by its strongly bullate leaf blades that are basally clasp- ing to nearly amplexicaul, short rachis, floral bracts with acuminate and callose-glandular tips, basally strongly apophysate calyx tube, and eglan- dular calyx lobes. It belongs to Cavendishia sect. Quereme and is related to C. talamancensis, dif- fering from it by clasping to nearly amplexicaul, strongly bullate leaves; conspicuously reticulate leaf veinlets; glabrous calyx and corolla; and strongly apophysate calyx tube. Cavendishia megabracteata Luteyn var. mega- bracteata, Brittonia 28: 49. 1976. Terrestrial or epiphytic shrubs to 2.5 m tall, often viney in habit; mature stems terete, minutely striate, gla- brous, the base to 5 cm in diam.; twigs terete to bluntly angled, striate, glabrous, reddish-brown. Leaves with petioles often flattened adaxially, rugose, rarely striate, 4-10 (14) mm long, 1-2.5 (3) mm diam., glabrous to densely puberulent adaxially when young, often glab- rate; leaf blades ovate, lanceolate, lance-elliptic or ob- long, (4) 8-12 (15) cm long, (1) 2.5-7 cm wide, base obtuse, rounded or truncate, apex acuminate and often abruptly so. often puberulous at base of midrib and lat- eral nerves on upper surface, 5-7-plinerved, midrib in- conspicuously raised and thickened through basal 1 cm otherwise weakly impressed above, inner pair of lateral nerves moderately or weakly impressed above but some- times plane or raised apically, outer lateral nerves and veinlets raised and usually very conspicuous above rare- ly impressed, all nerves raised beneath but veinlets often obscure. Inflorescences obconic to spherical in bud, (2) 5-9 (12)-flowered, the basal nodes often sterile; rachis coarsely and bluntly angled, striate, (0.6) 0.8-2 (3.5) cm long, ca. 2 mm diam., glabrous, rarely with clavate glands or cartilaginous teeth at base; floral bract trans- lucent, oblong, obovate, or rarely subrotund, (2) 3-6 (7) cm long, (1) 1.5-2.5 (4.5) cm wide, base narrowed and somewhat clasping, apex rounded, usually deeply emar- ginate, with minute, red, clavate, glandular fimbriae abaxially, dark red in bud but pale purplish, bright rose, or dark pink when mature; pedicels swollen apically, striate, (8) 10-16 (19) mm long, (0.5) 1 (1.5) mm diam., glabrous, rarely weakly glandular apically; bracteoles with or without prominent midrib, ovate, lanceolate, lance-elliptic, or nearly linear, 1.5-2 (5) mm long, 0.5- 1.5 (4) mm wide, apex glandular-callose. Flowers with calyx glabrous, 7-11 (13) mm long, tube cylindric, ribbed, 2-5 mm long, 3-5 mm diam., apophysate and deeply lobed at base with lobes straight to flaring out- ward and extending well below articulation, limb cylin- dric to slightly spreading, minutely papillate, (4) 5-7.5 (9) mm long, lobes triangular, (1) 1.5-2 (2.5) mm long, erect after anthesis, tips completely glandular-callose or callose apically, centrally, or only marginally, sinuses subacute when fresh, but drying obtuse to concave, rare- ly flat; corolla cylindric, slightly constricted at base and throat, often translucent, 31-40 (43) mm long, 6-9 mm diam., glabrous or rarely pilose apically, tube dull or glossy white, often pink to pinkish at base grading through violet to pale purple apically, lobes oblong, ob- tuse, 1 .5-2 mm long, reflexed, white sometimes margin- ally purple; stamens 24.5-36 mm long; filaments dense- ly puberulous apically, alternately 2.5-7.5 mm and 7.5- 13.5 mm long; anthers alternately 21.5-32.5 mm and 16-26.5 mm long, thecae 6-13 mm long. Immature ber- ry subspherical. Cavendishia megabracteata var. megabracteata is found in montane cloud forest, on the slopes of Volcan Baru and Cerro Horqueta, western Chiri- qui Province, Panama, 1 700-2500 m elevation. It has not yet been found in Costa Rica. Flowering specimens have been collected from December to August; immature fruiting specimens in January. Endangered. Cavendishia megabracteata is characterized by its large, apically emarginate, reddish floral bracts; obconic or spherical inflorescences (when in bud); basically whitish corollas; and glandular- callose calyx lobes. Both the floral bracts and the corollas are extremely thin and translucent when dry. The size, shape, and texture of its floral bracts alone serve to distinguish this species from most other Mesoamerican species of Cavendishia. Cav- endishia megabracteata is distinguished from oth- er members of Cavendishia sect. Callista by its emarginate floral bracts (see discussion under C. atroviolacea). The related C. megabracteata var. attenuata Luteyn is endemic to Cocle Province, Panama. Cavendishia melastomoides (Klotzsch) Hemsl., Biol. Cent.-amen, Bot. 2: 273. 1881. Socratesia melastomoides Klotzsch, Linnaea 24: 23. 1851. C. klotzschiana Niedenzu, Bot. Jahrb. Syst. 1 1 : 42 FIELDIANA: BOTANY 206. 1890 (nom. super.). Chupalon melasto- moides (Klotzsch) Kuntze, Rev. Gen. PI. 2: 383. 1891. C. graebneriana Hoerold, Bot. Jahrb. Syst. 42:324. 1909. Figure 9. Epiphytic shrubs, 0.5-3 m tall; mature stems pendent, terete, smooth, glabrous, often appearing glaucous; twigs glabrous or pilose, with reddish, caducous glands. Leaves with petioles flattened adaxially, rugose, (3) 4.5- 9 mm long, glabrous or densely short-pilose, glabrate; leaf blades ovate or elliptic, 3-8.5 ( 1 0) cm long. ( 1 ) 1.5- 4 cm wide, base rounded, truncate or cuneate, apex long- acuminate or caudate-acuminate, glabrous, 3 (5)-pliner- ved, midrib impressed and prominent above becoming plane apically, raised and conspicuous beneath, lateral nerves slightly raised on both surfaces or rarely im- pressed above, obscure or conspicuous beneath, reticu- late veinlets slightly raised on both surfaces but ± ob- scure. Inflorescences loosely (4) 6-10 (IS)-flowered, the basal nodes often sterile, all parts viscid; rachis some- times ribbed, (2) 3-8 (15) cm long, 1-1.5 mm diam. at base, glabrous, often glandular-fimbriate; floral bract of- ten caducous, smooth or ribbed, oblong, oblanceolate or elliptic, rarely semi-orbicular, 4-22 mm long, 3-12 mm wide, apex often apiculate and glandular-callose. rarely emarginate, glandular-fimbriate, glabrous, rose to pale green, often translucent when fresh; pedicels ribbed, (7) 10-21 (27) mm long, glabrous, often glandular-fimbri- ate; bracteoles basal to medial, smooth or conspicuously nerved, ovate, ovate-lanceolate and often cochleariform or semi-orbicular, 1-5 mm long, 0.5-6 mm wide, apex glandular-thickened. Flowers with calyx (3) 4.5-7.5 mm long, glabrous, tube smooth or rugose, pentagonal or ribbed, 1-2 (3) mm long, with basal apophysis deeply 5-lobed and perpendicular to calyx axis, limb cylindric or somewhat spreading, often conspicuously striate, (2) 2.5-5 (6.5) mm long, lobes triangular, (0.5) 1-1.5 mm long, erect after anthesis, completely glandular-callose, sinuses obtuse or concave and almost flat; corolla thin- carnose, cylindric, slightly narrowed at throat, some- times curving upward apically (var. melastomoides), 16- 41 mm long, 3-7 mm diam., viscid or not, glabrous or weakly pilose, dull white, blue or dark reddish-maroon, lobes triangular, 1-2.5 mm long, apex obtuse; stamens 16-34.5 mm long; filaments alternately 2.5-6 mm and 5.5-12.5 mm long, glabrous or pilose; anthers alternate- ly 14-32 mm and 11.5-25 mm long, thecae 3-8 mm long. Berry spherical. 11-14 mm diam.. maroon to blue- black. Cavendishia melastomoides is found in Costa Rica and western Panama, 950-2800 m elevation. Cavendishia melastomoides is characterized by having relatively small and narrow, acuminate leaf blades with long petioles, loosely flowered and viscid inflorescences, thin rachis, caducous floral bracts, broadly apophysate calyx tube, and maroon or white corollas. Within sect. Callista, C. melastomoides is probably most closely related to C. wercklei based on similarly (often) glandu- lar-fimbriate rachises and pedicels, bracteoles that are glandular-callose in the apical '/3 to "A, and often caducous floral bracts. Cavendishia melas- tomoides differs by its thinner rachis (1-1.5 vs. 2-4 mm diam.), smaller floral bracts (4-22 X 3- 12 vs. 20-30 X 10-20 mm), apically curved co- rollas, and calyx tube apophysis that is deeply 5- lobed (not merely undulate) at the base. Caven- dishia melastomoides is separable into three mor- phologically distinct and geographically isolated varieties, vars. melastomoides and alhiflora from Costa Rica and var. coloradensis from western Panama. Although the two Costa Rican varieties are distinguished by numerous morphological characters, perhaps indicating specific status to some, we have chosen to recognize them as va- rieties, emphasizing the features that unite them as a species rather than their differences. Al- though the differences between vars. melastomo- ides and albiflora are relatively stable, the two varieties do show some minor convergence in leaf size and shape and in corolla length and color in the area between Zarcero and Quesada (Alajuela Prov.), where their ranges meet. Nevertheless, there should be no difficulties in separating them. Both varieties are visited by hummingbirds (pers. observ.). The varieties are distinguished by the following key: Key to the Varieties of Cavendishia melastomoides la. Corolla 16-20 mm long, white; Cordillera de Tilarsin, 950-1500 m C. melastomoides var. albiflora Ib. Corolla (19) 26-41 mm long, reddish-maroon or blue; Cordillera Central-Cordillera de Talamanca, 1400-2800 m 2 2a. Rachis and pedicels usually densely glandular-fimbriate; floral bracts apically rdunded or acute, eglandular; bracteoles lanceolate or narrowly ovate-lanceolate, glandular-callose in apical V^-V^ co- rolla reddish-maroon; Costa Rica C. melastomoides var. melastomoides 2b. Rachis and pedicels usually with only a few, scattered, glandular fimbriae or none; floral bracts apically rounded, slightly notched, callose-apiculate; bracteoles broadly ovate to semi-orbicular, callose-apiculate; corolla blue; Panama C. melastomoides var. coloradensis LUTEYN AND WILBUR: FAMILY #172 ERICACEAE 43 Van albiflora Luteyn, Brittonia 28: 51. 1976. Il- lustration: J. L. Luteyn, Mem. New York Bot. Card. 28(3): 118, fig. 65. 1976. Inflorescence rachis (2) 3-4 (6) cm long, eglandular; floral bract caducous, oblong or elliptic, 4-7.5 (10.5) mm long, apex rounded and usually slightly emarginate; pedicels 9-13(15) mm long, eglandular; bracteoles nar- rowly ovate-lanceolate, glandular-thickened in apical '/j-'A. Corolla 16-20 mm long, glabrous, apically straight, white; stamens 16-18 mm long. Variety albiflora is found in disturbed areas of montane cloud forest and on remnant trees in pas- tureland, 950-1500 m elevation. It is endemic to the Cordillera de Tilaran of central Costa Rica. Flowering collections have been made in July- September; fruiting in September-November. En- dangered. Var. coloradensis Luteyn, Brittonia 28: 401. 1976. Inflorescence rachis (2) 2.5-7 cm long, with or with- out scattered clavate glandular fimbriae; floral bract ca- ducous or persistent, oblong to somewhat oblanceolate, 6-22 mm long, apex rounded or slightly notched, cal- lose-apiculate; pedicels 10-19 mm long, with or without clavate glandular fimbriae; bracteoles broadly ovate to semi-orbicular, callose-apiculate. Corolla 29-32 mm long, glabrous or weakly pilose in apical '/>, rarely api- cally arcuate, blue; stamens 27-29 mm long. Variety coloradensis is found in premontane to Continental Divide cloud forest, 1050-1750 m el- evation. It is endemic to the Chiriqui highlands of Panama, where is has been collected at Cerro Col- orado and in the vicinity of the Fortuna Dam site. Flowering collections have been made in all months from January through October; fruits un- known. Locally Common. Var. melastomoides Inflorescence rachis (3) 5-8 (15) cm long, usually scattered to densely glandular; floral bract caducous, ob- long, oblanceolate, 6-14 (20.5) mm long, apex rounded or acute, never emarginate, pale green often with pinkish margins; pedicels (7) 10-21 (27) mm long, usually glan- dular as rachis; bracteoles narrowly ovate-lanceolate, glandular-callose in apical '/,-'/,. Corolla (19) 26-37 (41) mm long, glabrous, apical portion often distinctly curved upward, dark reddish-maroon; stamens 19.5- 34.5 mm long. Variety melastomoides occurs in primary cloud forest, disturbed forest, remnant trees in pasture- land, or fence-row trees, 1400-2800 m elevation. It is endemic to the Cordillera Central and Cor- dillera de Talamanca, Costa Rica. Flowering col- lections have been made in April-July; fruiting in June-September. This variety is visited by the hummingbird Lampornis calolaema in Costa Rica (Luteyn, 1998). Widespread. Cavendishia osaensis Luteyn & J. F. Morales, Brittonia 48: 514, fig. 1 A-D. 1996[1997]. Fig- ure 12. Epiphytic shrubs; mature stems subterete, bluntly an- gled, smooth or minutely striate, nitid, glabrous; twigs subterete, slightly angled, smooth, glabrous. Leaves with petioles rugose, 2-4 mm long, weakly short-pilose, glabrate; leaf blades coriaceous, ovate, lanceolate to ovate-elliptic, 7-11.5 cm long, 1.3-3.2 cm wide, base obtuse or rounded, apex gradually tapering and acumi- nate, glabrous, also with deciduous, minute, glandular fimbriae beneath, 3 (5)-plinerved, midrib impressed above, raised and conspicuous beneath, lateral nerves slightly raised but relatively inconspicuous above, slightly raised beneath, reticulate veinlets slightly raised and conspicuous above, slightly raised but obscure be- neath. Inflorescences subspherical, congested, race- mose, 4-1 1 -flowered, surrounded at base by a series of ovate to oblong, glabrous bracts; rachis angled, 5-10 mm long, glabrous; floral bract smooth but venation conspicuous on both surfaces, obovate or oblong-ellip- tic, 16-24 mm long, base cuneate, apex obtuse or round- ed, glabrous, pinkish-lilac; pedicels sometimes ribbed, 5.5-7 mm long, glabrous; bracteoles basal, ovate, ca. 1 mm long, marginally with thick cartilaginous teeth. Flowers with calyx 5-7 mm long, glabrous, tube cylin- dric, slightly rugose, 1.5-2 mm long, base apophysate with a relatively smooth margin, limb spreading-cam- panulate, 3.5-5 mm long, lobes slightly striate, long- triangular, 2.5-3 mm long, erect after anthesis, margin scarious and bearing ca. 3-4 oblong to fimbriate glands basally, these becoming thickened cartilaginous teeth apically, sinuses obtuse; corolla cylindric, narrowed at throat, 1.8-2.2 cm long, 3-4.5 mm diam., basal % red and apical !/3 white, lobes erect, somewhat ovate-trian- gular, 0.7-1 mm long, white; stamens overall equal, ca. 18 mm long; filaments alternately 2-3 mm and ca. 4 mm long, glabrous or weakly puberulent apically; an- thers alternately ca. 17 mm and ca. 14.5 mm long, thecae ca. 4-5 mm long. Berry not seen. Cavendishia osaensis is found in lowland rain forest, 200-750 m elevation. It is endemic to the Osa Peninsula of Costa Rica, where it is known from only the following five collections: Canton de Golfito, Jimenez, Dos Brazos de Rfo Tigre, along trail toward the union of Quebrada Pate- mazo and Quebrada Porsillego (G. Herrera 4636) and at Cerro Rincon (G. Herrera 4157); Canton de Osa, Reserva Forestal Golfo Dulce, Quebrada Vaquedano (R. Aguilar & G. Herrera 806), along the trail to Cerro Brujo, from Quebrada Vaque- 44 FIELDIANA: BOTANY dano to the top (A. Fernandez 161); and Rancho Quemado, Sierpe, around the lakes (J. Marin & F. Quesada 2612). Flowering specimens have been collected in January and November; fruits are unknown. Endangered. Cavendishia osaensis is characterized by hav- ing a glabrous habit, gradually long-tapering and acuminate leaves, calyx lobes that are most of the length of the limb with glands that are broken up into ca. 3 or 4 oblong to spherical portions. Its relationships are not clear, although it belongs to Cavendishia ser. Englerianae and keys closest to C. engleriana var. engleriana. Cavendishia panamensis Luteyn, Brittonia 28: 47. 1976. Erect, often arching, terrestrial shrubs 2.5-3 m tall; mature stems terete, glabrous, ca. 1 cm diam., the base ca. 2.5 cm diam.; twigs bluntly angled or ribbed, mi- nutely striate, glabrous, often bearing tiny, sessile, black- ish, spherical glands. Leaves with petioles slightly flat- tened adaxially, rugose, (2.5) 6-8 (10) mm long, 2-2.5 (3.5) mm diam., glabrous or infrequently puberulent; leaf blades elliptic, lance-elliptic to ovate-lanceolate, 9- 16 cm long, 3-6 cm wide, base acute to narrowly obtuse, rarely rounded, apex acuminate to somewhat caudate acuminate, glabrous but often with caducous, scattered, reddish, pustular glands 0.1-0.2 mm diam. along upper surface that leave a reddish or blackish papillate scar, 5 (7)-plinerved, innermost lateral nerves often arising 2- 3 cm above base, midrib impressed above, raised and conspicuous beneath, lateral nerves impressed above be- coming plane to slightly elevated apically with outer lat- eral nerves often obscure beyond middle, raised and conspicuous beneath, veinlets raised and usually con- spicuous above, obscure to slightly raised beneath. In- florescences loosely cylindric, 1 1-21 (37)-flowered, vis- cid, often basal nodes sterile; rachis flattened, often flex- uous apically, conspicuously ribbed and minutely striate, glabrous or rarely puberulent, (3) 8-17 cm long, to 5 mm diam. at base, pink to dark pink when fresh with minute, peltate, globular, reddish glands along length and especially at base; floral bract oblong to oblanceo- late, rarely obovate, (2) 3-5 cm long, (0.6) 1-2 cm wide, narrowed to a slightly clasping base, apex rounded or narrowly obtuse to acute, pink, glabrous or sparsely cil- iate along apical '/, of margin, marginally with 7-23 dark red, globular, peltate or sessile glands ca. 0.7 mm diam.; pedicels swollen apically, striate, (11) 15-19 mm long, 1-1.5 mm diam., pink, glabrous, with scattered, globular, reddish glands; bracteoles basal, ovate to lanceolate, 1.5-2 (4) mm long, 1-2 mm wide, pink with scattered, dark red, globular glands at apex and margins. Flowers with calyx glabrous, 4-6 mm long, tube cylindric or spreading, rugose to strongly ribbed, 1 .5-2 mm long, ca. 2.5 mm diam., usually with dark red, globular to disc- shaped, peltate glands obscuring surface, limb cylindric- campanulate, (2.5) 3-3.5 mm long, with scattered, glob- ular to disc-shaped, peltate glands, lobes triangular, ob- tuse, 1-1.5 mm long, ca. 1.5 mm wide, erect after an- thesis, margin glandular callose thickened with glands contiguous at lobe apices, sinuses concave; corolla cy- lindric, constricted at base, narrowed at throat, 2.5-3.5 cm long, 8-10 mm diam., glabrous or pilose but gla- brous at constricted base, white, lobes triangular, obtuse, 1-2 mm long; stamens 27-30 mm long; filaments slightly coherent at base, strigose apically, alternately 4- 6 mm and 10-13.5 mm long; anthers alternately 17-20.5 mm and 25-27 mm long, thecae 7.5-1 1 mm long. Berry not seen. Cavendishia panamensis is endemic to cloud forest in western Chiriquf Province, Panama (Vol- c£ Baru), 1830-1980 m elevation. It has not yet been found in Costa Rica. Flowering specimens have been collected in February, April, and May; fruits are unknown. Endangered. Cavendishia panamensis is characterized by its glandular-margined floral bract and bracteoles and nonapophysate calyx tubes the surfaces of which are nearly covered by the globular glands. It be- longs to Cavendishia ser. Lactiviscidae, where it differs from C. chiriquiensis by a basic difference in calyx lobe glands — in C. panamensis the cal- lose-thickened portion of the calyx lobe is strictly marginal, while in C. chiriquiensis the entire ca- lyx lobe is callose thickened and appears occa- sionally to break off. Cavendishia pseudostenophylla Luteyn, Britton- ia 28: 46. 1976. Terrestrial or epiphytic shrubs, sometimes lianoid; mature stems terete, minutely striate, puberulent; twigs subterete to bluntly angled, minutely striate, puberulent. Leaves with petioles subterete, rugose, 2-3.5 mm long, 1-2 mm diam., densely puberulent; leaf blades linear- lanceolate to linear-elliptic, strongly revolute, 9-14 cm long, 1-1.5 cm wide, base broadly obtuse or rounded, slightly cordate, apex long-attenuate, acute, glabrous. 3 (5)-plinerved, midrib deeply impressed above and raised and conspicuous beneath, lateral nerves and retic- ulate veinlets raised and conspicuous above and slightly raised but obscure beneath. Inflorescences loosely cy- lindric, viscid, 1 2-20-flowered, lowest nodes sterile; ra- chis long-tapering, bluntly angled, minutely striate, 5- 15 cm long, ca. 1-1.5 mm diam., glabrous, bearing scat- tered globular glands; floral bract prominently veined, often somewhat ribbed, oblong or oval elliptic, (13) 19- 27 mm long, (5) 7-11 mm wide, base narrowed and rounded, apex rounded or acute, apex short-ciliate, pink, margins bearing (3) 7-12 (per side) sessile, globular glands to 0.2 mm diam.; pedicels to 8 mm long, ca. 1 mm diam., glabrous, bearing few globular glands; brac- teoles apical, completely enveloping flowers in bud, clasping calyx and corolla at anthesis, oblong, oblong- elliptic or oblanceolate, ca. 25 mm long, ca. 10 mm wide, base rounded, apex broadly rounded, glabrous but margin short-ciliate and bearing up to 15 globular glands. Flowers with calyx 6.5-7.5 mm long, glabrous, tube cylindric, slightly rugose, 1.5-2 mm long, ca. 3.5 mm diam., base slightly expanded and bearing globular LUTEYN AND WILBUR: FAMILY #172 ERICACEAE 45 glands around margins and onto surface, limb campan- ulate or erect-spreading, smooth or slightly ribbed, 5-6 mm long, lobes triangular, ca. 2 mm long, margins con- spicuously glandular callose-thickened, the thickenings nearly contiguous at apex of lobe, sinuses obtuse or rounded; corolla cylindric, ca. 32 mm long, ca. 6 mm ilia in., pink, densely short-pilose in apical '/,, lobes tri- angular, ca. 2 mm long; stamens 27-28 mm long; fila- ments alternately 4-4.5 mm and 7.5-8 mm long, short- pilose in apical 'A; anthers alternately ca. 24-24.5 mm and 20 mm long, thecae 6-7 mm long. Berry not seen. Cavendishia pseudostenophylla is known only from cloud forest in the Bocas del Toro/Chiriquf Province border area and tropical wet forest north of Santa Fe (Veraguas Prov.), Panama, 300-1 100 m elevation. It has only been collected eight times in Panama and is not yet known from Costa Rica. Flowering specimens have been collected in April and May; fruits are unknown. Endangered. Cavendishia pseudostenophylla is characterized by its leaves that are about nine times longer than wide, inflorescences and flowers with globular glands, prominently veined floral bracts, long and apical bracteoles that clasp the calyx and corolla at anthesis, a slightly apophysate calyx tube, and calyx lobes that are glandular callose-thickened. It is very reminiscent of C. stenophylla A. C. Smith (Cocle and Panama Prov., Panama). Closer examination of the leaves of C. pseudostenophyl- la, however, shows them to be plinerved and api- cally acute, in contrast to the obscurely pinnately nerved and apically obtuse leaves of C. steno- phylla. Other characters, especially differences in bracteole size and position, also distinguish the two species. Cavendishia pubescens (Kunth in H.B.K.) Hem- sley, Biol. Cent.-amer., Bot. 2: 273. 1881. Thi- baudia pubescens Kunth in H.B.K., Nov. Gen. Sp. 3: 273. 1819. Proclesia pubescens (Kunth in H.B.K.) Klotzsch, Linnaea 24: 33. 1851. Chupalon pubescens (Kunth in H.B.K.) Kuntze, Rev. Gen. PI. 2: 383. 1891. Epiphytic and terrestrial shrubs 1-3 m tall; mature stems subterete or bluntly angled, ridged, moderately short-pilose to glabrate; twigs flattened and coarsely an- gled, ridged, densely short-pilose with whitish or grayish trichomes ca. 0.5 mm long. Leaves with petioles terete to often adaxially flattened, rugose, sometimes coarsely ridged, (6) 9-15 mm long, 2-3.5 mm in diam., densely short-pilose or puberulent; leaf blades elliptic, oblong- elliptic, lanceolate or ovate, (5.5) 14-20 (30) cm long, (1.5) 3.5-9 (12.5) cm wide, base cuneate, obtuse, round- ed, cordate or truncate, apex long-acuminate with a blunt or sharply acute tip, usually densely puberulent to short- pilose on both surfaces with soft, white trichomes, per- sistent beneath but often glabrate above, 5-7-plinerved, midrib impressed above and prominently raised beneath, lateral nerves impressed above but plane apically, raised and prominent beneath, veinlets plane to slightly raised or conspicuously impressed above but obscure beneath. Inflorescences capitate to obovate, 7-1 1 -flowered, en- circled at base by densely matted or appressed short- pilose bracts; rachis bluntly angled, striate, rugose, per- sistently densely short, white pilose, sometimes with glandular fimbriae, 1-2.5 (4.5) cm long and 2-3 mm in diam., rarely with cartilaginous teeth at base; floral bract muricate, elliptic to oblanceolate, 17-30 mm long, 7-13 (17) mm wide, base narrowed, apex rounded or rarely emarginate, pale pinkish-green or whitish-green through dark pink to red, often with small, red, clavate glands abaxially, glabrous or only ciliate, to sparsely or densely short, white and usually appressed pilose both abaxially and marginally, sometimes fimbriate along margin; ped- icels bluntly angled, densely short-pilose, often fimbri- ate, 6-14 mm long and 1-1.5 mm in diam.; bracteoles oblong, linear, or subulate, 2-4.5 (5) mm long, ca. 1 mm wide, densely pubescent or only marginally ciliate, mar- gin glandular-fimbriate, the fimbriae rarely fusing api- cally. Flowers with calyx (4.5) 5.5-8 mm long, pubes- cent with densely matted, white or tan trichomes, glan- dular-fimbriate, tube cylindric, usually shorter than limb, (2) 2.5-4 mm long, limb campanulate or spreading, (2.5) 3-5 mm long, lobes oblong, oblong-triangular, broadly triangular or deltoid, (1) 1.5-3 mm long, 1.5-1.7 mm wide, apex obtuse or broadly rounded, sometimes then apiculate, connivent or curling after anthesis, rarely re- flexed, marginally glandular-fimbriate, the fimbriae rare- ly fusing apically, sinuses obtuse; corolla cylindric or bottle-shaped, slightly constricted at base, narrowed at throat, (12) 15-20 (22) mm long, 7-8 mm in diam., white or tinted pink, with age becoming red, moderately to densely short-pilose with white or tawny trichomes, lobes triangular, ca. 1-2 mm long, wide-spreading at an- thesis; stamens 10-15.5 mm long; filaments alternately (1.8) 2.5-4 mm and 3.5-6 mm long; anthers alternately 9.5-13 mm and 8-11 mm long, thecae 2.5-4 mm long. Berry spherical, densely puberulent, 8-14 mm in diam., dark maroon to blue-black. Cavendishia pubescens is found in relatively dry exposed slopes and thickets, although a few labels indicate its presence in humid or rain forest, (700) 1200-2100 (3000) m elevation, in western Panama (within 20 km of the border with Costa Rica), and from Venezuela to Bolivia. It has not yet been found in Costa Rica. Flowering and fruit- ing occur throughout the year. Rare/Infrequent (in Panama). Cavendishia pubescens is readily distinguished by its pale pinkish to rose or whitish-green floral bracts, densely short-pilose, white to pinkish- white corollas, densely pilose to matted or woolly calyces of which the limb is longer than the tube and the lobes are oblong and obtuse, and persis- tently soft-puberulent or pilose leaf undersurfaces and fruits. The densely matted pubescence of the 46 FIELDIANA: BOTANY calyx is usually enough to characterize the spe- cies. Cavendishia pubescens belongs to ser. Cav- endishiae, where it is most closely related to C. bracteata (see that species). Cavendishia pubes- cens may hybridize with C. bracteata (Luteyn, 1976c). The species is visited by hummingbirds (pers. observ.). Cavendishia quercina A. C. Smith, Contrib. U.S. Natl. Herb. 28: 450. 1932. Illustration: J. L. Lu- teyn, Mem. New York Bot. Card. 28(3): 109, fig. 56. 1976. Figure 6. Terrestrial or epiphytic shrubs 0.5-2 m tall; mature stems densely puberulent, glabrate. Leaves with petioles (1) 2—4 (6) mm long, densely short-puberulent, often glabrate; leaf blades oblong, oblong-ovate or ovate, 2.5- 7.5 cm long, (1) 1.5-3.5 cm wide, base rounded and subcordate, clasping, apex rounded, obtuse or often sub- acute, usually densely short-pilose on both surfaces but persistent only on the upper surface, 3-5 (7)-plinerved. Inflorescences loosely cylindric, 12-20 (53)-flowered; rachis 2-12 cm long, glabrous or densely hirsute; floral bract oblong or oblong elliptic, (9.5) 11-18 (23) mm long, base rounded to obtuse, not clasping, apex obtuse or rounded, glabrous or margin short-ciliate, margin glandular fimbriate, red; pedicels (6.5) 10-14 (18) mm long, sparsely to densely hirsute or rarely glabrate, dark pink but turning greenish; bracteoles oblong or oblong- ovate, 1-3 (5.5) mm long, glabrous, ciliate, or short- pilose at apex, margin densely glandular-fimbriate. Flowers with calyx (5.5) 6-8 (9) mm long, sparsely to densely hirsute or rarely glabrous, red, tube short-cylin- dric, often 5-ribbed, rugose, 1.5-2.5 (3) mm long, ba- sally apophysate, limb cylindric, spreading-erect or rare- ly campanulate, smooth, rugose or sometimes ribbed, (3.5) 4-6 (6.5) mm long, lobes 1-2.5 mm long, narrowly or sometimes broadly triangular, erect after anthesis, marginally short-ciliate and provided along each side with 3-5, intermittent, stout, glandular-thickenings that may coalesce, sinuses concave, usually ciliate; corolla cylindric, slightly constricted at base, (8.5) 11.5-15 (17.5) mm long, 2.5-4.5 mm diam., densely pilose api- cally, rarely glabrous, pale pinkish at base becoming pale green above, lobes whitish to pinkish-green, lobes ca. 2 mm long, oblong; stamens 8-15 mm long; fila- ments alternately 2-5 mm and 3-8 mm long, pilose api- cally; anthers alternately 5-9.5 mm and 6.5-12 mm long, thecae 1-4 mm long. Berry spherical, at least 8- 9 mm diam., weakly pilose, dark blue-black. Cavendishia quercina is infrequently found in cloud forest, 1050-2300 m elevation. It is distrib- uted from the eastern Cordillera de Tilardn of Costa Rica east through the Central Valley and Cordillera de Talamanca to western Chiriquf Prov- ince (Volcan Bani), Panama. Flowering occurs throughout the year but is most abundant from October to March; fruiting specimens have been collected in January, February, and April through June. Widespread. Cavendishia quercina is characterized by its short, oblong-ovate leaf blades with subcordate bases and rounded or subacute apices, hirsute ra- chises, pedicels and calyces, short-pilose to al- most woolly corollas, stout calyx lobe glands, and densely glandular-fimbriate bracteoles. It is easily recognized in the field by the leaves, the deep red floral bracts and calyces, and the pale green co- rollas with whitish tips. It belongs to C. ser. Cav- endishiae, where it is most similar to C. ruiz-ter- anii Luteyn from Venezuela. It is also similar to C. capitulata, in which occasionally the glandular fimbriae of the calyx lobes seem to fuse to form the stout, more or less oblong glands and by sim- ilar extremes in leaf variation. A comparison of the important characters of C. quercina, along with other Costa Rican members of sect. Quere- me, is given in Table 1 . Cavendishia quereme (Kunth in H.B.K.) Benth. & Hook.f., Gen. PI. 2: 570. 1876. Thibaudia quereme Kunth in H.B.K., Nov. Gen. Sp. 3: 274, table 256. 1819. Polyboea quereme (Kunth m H.B.K.) Klotzsch, Linnaea 24: 31. 1851. Chupalon quereme (Kunth in H.B.K.) Kuntze, Rev. Gen. PI. 2: 384. 1891. Illustration: J. L. Luteyn, Mem. New York Bot. Card. 28(3): 104, fig. 51. 1976. J. L. Luteyn, Fl. Neotrop. Mon- ogr. 35: 247, fig. 74D-F. 1983. Figures 9 and 14. Terrestrial or epiphytic shrubs 1-2.5 m tall; mature stems glabrous and with odor of wintergreen. Leaves with petioles terete or subterete, (3) 5-7 (9) mm long, glabrous at maturity; leaf blades bullate, elliptic, lance- elliptic or lanceolate, rarely ovate, (3.5) 6-11 (14) cm long, (1) 1.5-4 (5) cm wide, base acute or obtuse or rounded and short-attenuate, usually somewhat short-de- current on petiole, apex acuminate or caudate-acumi- nate, glabrous, 5 (7)-plinerved. Inflorescences loosely cylindric, (8) 12-20 (25)-flowered; rachis 2-5 (6) cm long, glabrous; floral bract conduplicate and slightly keeled, ascending, oblong, elliptic or lanceolate, 10-25 mm long, base rounded to obtuse, slightly clasping, apex acute, margin glandular-fimbriate, glabrous, pink or red; pedicels (5) 8-16 (19) mm long, glabrous, pinkish- green; bracteoles basal, lanceolate, oblong or linear-lan- ceolate, rarely ovate, 1-2.5 (5) mm long, 0.5-1 (2) mm wide, apex acute or obtuse, margin glandular-fimbriate. Flowers with calyx 2.5-5 mm long, 'glabrous, tube ba- sically cylindric but pentagonal with broad, blunt angles, smooth or rugose, 1 .5-2 mm long, limb cylindric, slight- ly spreading or campanulate, 2-3.5 mm long, lobes 1- 1 .5 mm long, ovate or oblong, acute to bluntly apiculate, connivent after anthesis, marginally thin lacerate-glan- ilul.it. sinuses acute; corolla bottle-shaped, pentagonal, LUTEYN AND WILBUR: FAMILY #172 ERICACEAE 47 7-1 1 (13) mm long, 3-6 (7) mm diam., glabrous, whit- ish at constricted base, then becoming orange or reddish- orange through the middle half and then white at throat, lobes ca. 1 mm long, white; stamens alternately 5.5-7.5 mm and 8-9.5 mm long; filaments alternately 1.5-2 (3.5) mm and (2.5) 3.5-5 mm long; anthers alternately (4.5) 5.5-7.5 mm and (3.5) 5-6.5 mm long, thecae 1.5- 3 mm long. Berry spherical, 7-1 1 mm diam., blue- black. Cavendishia quereme is found as an epiphyte in rain forest and montane cloud forest and as a terrestrial shrub in disturbed areas along trails or roadsides, 600-1700 m elevation. It has a disjunct distribution from the Central Valley (Alajuela Prov.) of Costa Rica to western Panama (Chiriqui Prov.) and then again in the Cordillera Occidental (Valle and Narino Depts.) of Colombia. Flowering and fruiting occur sporadically throughout the year. Locally Common. Cavendishia quereme is a very pretty species characterized by nitid, coriaceous, bullate leaf blades; dark pink floral bracts; a bottle-shaped co- rolla that is bluntly pentagonal and bright orange with white lobes; and stamens that are conspicu- ously alternately unequal to each other in length. One of its most distinctive characters is the very strong odor of wintergreen (methyl-salicylate), which is emitted by the vegetative parts of the plant (Luteyn, 1976c; Luteyn et al., 1980). The odor is so distinctive that it can often be smelled in the field before it is seen, and it persists in the herbarium even after the plants have been dried by heat whether pretreated with formaldehyde or ethanol or preserved in FAA. Cavendishia quer- eme is morphologically remarkably uniform de- spite its disjunct distribution. There is so little in- terpopulational variation that specimens from Co- lombia, Costa Rica, and Panama are indistinguish- able. Cavendishia quereme belongs to C. sect. Quereme, where it is presumably most closely re- lated to C. laurifolia (Klotzsch) Benth. & Hook.f. from Guatemala, Panama, and adjacent Colombia, both taxa having similar habits, pronounced bot- tle-shaped and bluntly pentagonal corollas, lacer- ate-glandular calyx lobes, and stamens that are conspicuously alternately unequal in overall length. Cavendishia quereme differs from C. laur- ifolia by its bullate and glabrous leaf blades (not flat and puberulous); the absence of glandular fim- briae from corollas, calyces, and pedicels; con- duplicate-keeled and ascending floral bracts (not concave and perpendicular to recurved from ra- chis); shorter pedicels (8-16 mm vs. 14-22 mm long); and corolla color (orange with white vs. red with white). A comparison of the important char- acters of C. quereme, along with other Costa Ri- can members of sect. Quereme, is given in Table 1. Cavendishia quereme putatively hybridizes with C. endresii and C. axillaris in Costa Rica and western Panama (Luteyn, 1976c). A common name in Costa Rica (San Jose Prov.) is colmillo. This species is visited by hummingbirds (pers. ob- serv.). Cavendishia subfasciculata Luteyn, Brittonia 28: 50. 1976. Terrestrial or epiphytic shrubs 0.7-3 m tall, some- times lianoid; mature stems terete, smooth or minutely striate, glabrous, usually with a thin, whitish waxy layer; twigs subterete to bluntly angled, striate, often coarsely ridged, glabrous. Leaves with petioles usually flattened adaxially, rugose and often coarsely ridged, 7-13 mm long, 1.5-2 mm diam., glabrous to weakly pilose adax- ially when young, often glabrate; leaf blades coriaceous, lanceolate to lance-elliptic, (3) 5-9 (1 1) cm long, (1)2- 4 cm wide, base obtuse or rounded, often short-decurrent onto petiole, apex often abruptly short-acuminate, gla- brous or often with numerous, short trichomes at base of midrib adaxially, usually glabrate, 5 (7)-plinerved, midrib plane or more commonly weakly impressed above, lateral nerves usually raised and conspicuous or rarely weakly impressed above and raised beneath, re- ticulate veinlets raised and conspicuous to obscure above and obscure beneath. Inflorescences obconic to spherical in bud, (2) 3-6 (9)-flowered; rachis flattened, bluntly angled, striate, viscid, (0.3) 0.6-1.2 (3.2) cm long, glabrous, usually bearing minute, cartilaginous teeth basally; floral bract smooth or rarely slightly ribbed, oblong to oblanceolate or oblong-lanceolate, (1.5) 2-3 (4) cm long, 0.5-1 (2) cm wide, base narrowed and somewhat auriculate, apex rounded or obtuse and usually emarginate or when young with the notch almost completely filled with callose tissue, glabrous, usually bearing minute, red, glandular fimbriae abaxially, pale green but suffused with pink along margins and basally, usually translucent; pedicels swollen apically, ridged, (6) 8-13 (-15) mm long, (0.5) 1-1.5 mm diam., glabrous, occasionally with cartilaginous teeth at apical tip; brac- teoles linear-lanceolate, 1-2.5 (3) mm long, 0.5-1 mm wide, apex acuminate and glandular callose-thickened in apical '/2-%. Flowers with calyx 5-7 (10) mm long, gla- brous, tube cylindric, weakly rugose, strongly ribbed, (1.5) 2-3 (4) mm long, basally apophysate and deeply lobed with these lobes straight and extending to or just beyond articulation, limb campanulate to cylindric, smooth or weakly ribbed and minutely papillate, (2.5) 3.5-4.5 (6) mm long, lobes triangular, (0.7) 1-1.5 (2) mm long, 2-3 mm wide, sometimes flaring but later erect, glandular callose thickened throughout or only in apical %, sinuses obtuse, concave or flat; corolla thinly carnose, cylindric, slightly constricted at base and nar- rowed to throat, often translucent, 20-24 (30) mm long, 5-7 mm diam., glabrous or rarely pilose, whitish at base and apex but otherwise pinkish-violet to rose-red, lobes triangular or oblong, ca. 1 mm long, reflexed, apex acute and often callose thickened, white with pinkish or violet 48 FIELDIANA: BOTANY margins; stamens 19-24.5 mm long; filaments alter- nately 2.5-5 mm and 5-10 mm long, densely pilose; anthers alternately 13-20 mm and 17-22 mm long, the- cae 4.5-7 mm long. Berry ca. 9 mm diam., blue-black. Cavendishia subfasciculata is endemic to west- ern (Chiriquf Prov.) and central (Veraguas Prov.) Panama, where it is found in premontane and montane cloud forest, 1100-2600 m elevation. It has not yet been found in Costa Rica. Flowering specimens have been collected from December to August; immature fruits in March. Rare/Infre- quent. Cavendishia subfasciculata is characterized by having capitate, few-flowered inflorescences, emarginate and translucent floral bracts, apophy- sate and deeply lobed calyx tubes, glandular-cal- lose calyx lobes, and relatively short corollas with callose-thickened lobes. It has two slightly differ- ent color phases that are correlated with geogra- phy. In western Chiriquf Province (Panama), around Boquete, the floral bracts are greenish to pinkish-green and the corollas pinkish-violet to rose-red, while in eastern Chiriquf Province, around the Fortuna Dam site, and in Veraguas Province, the bracts are pink and the corollas pink to white. Within sect. Callista, C. subfasciculata is likely most closely related to C. morii Luteyn and its allies, all normally having glandular-cal- lose corolla lobes. However, it also shows mor- phological similarities to C. megabracteata, based on emarginate floral bracts; few-flowered, obcon- oid to spherical inflorescences; similarly colored floral bracts and corollas; and glandular-callose corolla lobes. Cavendishia subfasciculata may be distinguished from C. megabracteata by its over- all smaller size, its fuller and more dense habit, its usually green floral bracts (pinkish or pale pur- ple bracts of C. megabracteata), slightly decurrent leaf bases, and leaf venation. Cavendishia talamancensis Luteyn, Brittonia 28: 43. 1976. Illustration: J. L. Luteyn, Mem. New York Bot. Card. 28(3): 102, fig. 49. 1976. Fig- ure 6. Terrestrial or epiphytic shrubs 0.5-2 m tall; mature stems glabrous; twigs densely short-pilose, all pubes- cence curly. Leaves with petioles 3-4 (6) mm long, densely puberulent; leaf blades bullate. oblong or ob- long-ovate, (2) 3.5-5.5 (7) cm long, (1.5) 2-4 (4.5) cm wide, base rounded and subcordate, apex rounded or ob- tuse, densely short-pilose along nerves and margins when young, 3-5 (7)-plinerved. Inflorescences short-cy- lindric, compact, 8-32-flowered; rachis 1-9 cm long, densely crisped, short-pilose; floral bract ribbed, oblong or oblong-elliptic. 8-10 mm long, base rounded, some- what clasping rachis, apex rounded and often shallowly emarginate. glabrous or densely short-pubescent, margin ciliate and glandular-fimbriate, purplish; pedicels (1.5) 4.5-7 mm long, densely short-pilose, rarely glabrate, usually with tiny, black, clavate glands apically; brac- teoles basal, ovate or subulate. (0.5) 1-2 mm long, usu- ally densely short-pilose and also densely glandular-fim- briate marginally. Flowers with calyx 6-9 mm long, usually densely short-pilose, rarely glabrate, tube barrel- shaped, strongly rugose, 2-3 (3.5) mm long, basally short-apophysate, limb cylindric or somewhat spreading, rugose. 3.5-6 mm long, lobes 1.5-2 mm long, oblong- ovate or ovate, usually somewhat connivent after anthe- sis, margin lacerate-glandular, sinuses acute; corolla cy- lindric, 11.5-14 mm long, 4-5 mm diam., pilose api- cally, rarely glabrate, pink, pinkish-white or purplish, lobes oblong-deltate to triangular, acute, ca. 1-1.3 mm long; stamens 8.5-10.5 mm long; filaments alternately 2.5-3.5 mm and 3.5-5.5 mm long, pilose apically; an- thers alternately 6.5-8.5 mm and 5-7.5 mm long, thecae 1-3 mm long. Berry not seen. Cavendishia talamancensis is found in mon- tane, Quercus cloud forest, wet thickets, and steep rocky road cuts, 2000-3000 m elevation. It is a rare species, endemic to the Cordillera de Tala- manca (Cartago and San Jose Provs.) .in Costa Rica. Flowering specimens have been collected in December-February and April-August; immature fruiting specimens only in January. Rare/Infre- quent* Cavendishia talamancensis is characterized by leaf blades that are small, bullate. oblong, or ob- long-ovate, basally subcordate and apically round- ed or obtuse; densely crisped-pilose inflorescenc- es (rachis, floral bracts, pedicels, and calyces); and lacerate-glandular calyx lobes. Its morpholog- ical features are compared with other members of Cavendishia sect. Quereme in Table 1. Although without close relatives, it is vegetatively similar to C. quercina but differs by the characters of calyx lobe glands and tube. Occasional popula- tions of C. talamancensis are reported to emit the odor of wintergreen from crushed leaves, which may indicate a relationship with C. quereme. Cavendishia wercklei Hoerold, Bot. Jahrb. Syst. 42: 325. 1909. Illustration: J. L. Luteyn, Mem. New York Bot. Card. 28(3): 1 19, fig. 66. 1976. Terrestrial or epiphytic shrubs to 2 m tall; mature stems usually glabrous but sometimes scattered spread- ing hirsute; twigs frequently with reddish, clavate glands. Leaves with petioles (5) 6-11 (12) mm long, densely short-pilose, occasionally glabrous; leaf blades lanceolate, lance-elliptic, oblong or rarely ovate, (4) 7- 12 (15) cm long, (1) 2-4 (5) cm wide, base rounded, cuneate or obtuse, apex long-acuminate, weakly pilose LUTEYN AND WILBUR: FAMILY #172 ERICACEAE 49 along nerves and lamina base and then glabrate, or pu- bescence persisting along nerves on adaxial surface, 3- 5-plinerved. Inflorescences loosely cylindric, (6) 9-12 ( 19)-flowered; rachis (3) 5-7 (9) cm long, 2-4 mm diam. at base, glabrous, viscid, densely covered by sessile to subsessile. globular or clavate, reddish glands ca. 0.2 mm long; floral bract oblong to obovate, (15) 20-30 (35) mm long, apex obtuse to rounded and rarely apiculate, usually caducous, usually conspicuously nerved, gla- brous, rose to pinkish-purple, abaxially with minute sub- sessile clavate reddish glands; pedicels (9) 11-18 mm long, viscid, glabrous, usually with reddish, sessile glands over entire length or only at base, rarely eglan- dular; bracteoles ovate or oblong-lanceolate, 2-4.5 (5) mm long, apex acute and glandular-thickened, usually conspicuously 3-5-nerved. Flowers with calyx 6-9 (11) mm long, glabrous, viscid, tube cylindric, coarsely ribbed, 2-3.5 (5) mm long, basally apophysate with mar- gin undulate, limb cylindric to campanulate, smooth to ribbed, rarely minutely papillate, (3.5) 4-6 (6.5) mm long, lobes 1.5-2 mm long, triangular, obtuse or acute, erect after anthesis, completely glandular-thickened and this extending onto limb surface, sinuses obtuse, con- cave or rarely flat; corolla cylindric, narrowed at base, slightly constricted at throat, (25) 31-38 mm long, 6-8 mm diam., viscid, short-pilose for apical % or sometimes velutinous, whitish to purplish-rose, lobes 1.5-2 mm long, oblong, obtuse; stamens 26-34 mm long; fila- ments alternately 3.5-5.5 mm and 8-11.5 mm long, pi- lose apically; anthers alternately 25.5-31 mm and 20.5- 24 mm long, thecae 5.5-10.5 mm long. Berry spherical, when immature at least 15 mm diam. and dark maroon. Cavendishia wercklei is found in premontane forest, 1000-1500 m elevation. It is a rare species, endemic to the Central Valley of Costa Rica, hav- ing been collected mostly at La Palma de San Jose and in the vicinity of Vara Blanca (Alajuela Prov.); it is known from fewer than 10 collections. Flowering specimens have been collected in May and June. Endangered. Cavendishia wercklei is characterized by its stout rachis; large, usually caducous floral bracts; glandular-fimbiate rachis and pedicels; 3-5- nerved bracteoles; undulate apophysis base; and whitish to pinkish-rose, apically short-pilose co- rollas. Within C. sect. Callista, it is related to C. melastomoides and C. callista and is distinguished by the characters mentioned above, as well as col- or, particularly that of the corollas and floral bracts. Cavendishia wercklei apparently hybrid- izes with C. endresii (Luteyn, 1976c). The species is visited by hummingbirds (pers. observ.). Chimaphila Pursh REFERENCE — L. J. Dorr, Chimaphila. Fl. Neo- trop. Monogr. 66: 37-46. 1995. Herbs or subshrubs, erect (rarely decumbent), the mature stems simple or branched at the base. Leaves alternate to subopposite, clustered in pseudo-whorls along stem, petiolate, the blade coriaceous, margin ser- rate, the venation pinnate. Inflorescences terminal, a pe- dunculate corymb or subumbel, or rarely flowers soli- tary; peduncles naked but subtended by persistent bud scales of winter bud from which it emerges; pedicels subtended by deciduous or persistent (then adnate) bracts, slightly recurved in flower, erect in fruit; brac- teoles lacking. Flowers 5-merous, actinomorphic, fra- grant; calyx persistent, with sepals distinct or shortly connate at base, lobes imbricate at base; corolla crater- iform, aestivation imbricate, the petals distinct, spread- ing to reflexed, margin fimbriolate; stamens 10, equal, distinct, lacking appendages; filaments dilated at or be- low middle; connectives lacking disintegration tissue or spurs; anthers with thecae smooth, tubules short, ± spreading, dehiscing by obliquely and slightly introrse pores; pollen in tetrads; ovary superior, 5-locular, de- pressed-globose, subtended by a 5-lobed, nectariferous disc; style massive, subsessile, fleshy, obconical or tur- binate, embedded in summit of ovary; stigma broad, pel- tate, rounded, entire or with 5 undulate lobes. Fruit an erect, loculicidal capsule, dehiscence starting from both above and below, proceeding toward middle, the sutures smooth, lacking connecting threads; seeds numerous, small, winged; embryos undifferentiated, without devel- oped cotyledons. The genus Chimaphila consists of five species distributed in Eurasia, the West Indies, and North America south to Panama. One species occurs in Costa Rica. Chimaphila is distinct from all other Costa Rican Ericaceae, being characterized by its herb-like habit, leaves in pseudo-whorls, broadly crateriform corollas with distinct petals, staminal filaments conspicuously dilated, and superior ova- ry. Chimaphila maculata (L.) Pursh, Fl. Amer. Sept. 1: 300. 1814. Pyrola maculata L., Sp. PI. 396. 1753. C. costaricens Andres, Verh. Bot. Vereins Prov. Brandenburg 56: 66. 1914 (nom. nud.). C. guatemalensis Rydberg, N. Amer. Fl. 29(1): 32. 1914. Figures IB and 15. Herb-like, suffrutescent subshrubs 10-53 cm tall. Leaves spreading, clustered in 2-5 pseudo-whorls of 3- 5 leaves each along stem, the pseudo-whorls separated by 1-7 cm long internodes, with petioles 3-13 mm long; leaf blades thick-coriaceous, lanceolate, oblong-lanceo- late, ovate-lanceolate or ovate, 2-10 cm long, 0.8-3 cm wide, base obtuse, rounded or widely cuneate, apex acute or acuminate, margin sharply and remotely serrate, almost spine-tipped, dark green above and marked with a broad, whitish line along midrib and larger secondary veins, pale reddish-purple below. Inflorescences (1)2- 5-flowered; peduncles 7-11 cm long, finely papillate; pedicels 20-30 mm long, finely papillate; floral bract linear to linear-lanceolate, 2-4 mm long, apex acute, the margin dark brown, papillate near base, slightly dentic- 50 FIELDIANA: BOTANY ulate. Flowers nodding to divergent; calyx 3-6 mm long, lobes ovate, 2-5 mm long, apex obtuse to rounded, margin finely dentate, ciliate, reflexed in fruit; corolla broadly crateriform, petals obovoid to orbicular, narrow- ing at base, 8-9 mm long, ca. 6 mm wide, margin ciliate, waxy white to pinkish; stamens 6-8 mm long; filaments ca. 4 mm long, the dilated portion villous with broad, flat, translucent hairs; anthers 2.5-3.5 mm long, pink, thecae 1.5-2 mm long, tubules 1-1.5 mm long. Capsule erect, depressed-spherical, 5-10 mm long and in diam., green turning black. Chimaphila maculata is found on the forest floor in montane (including Quercus) forest, 2000-3000 m elevation. It is distributed from eastern North America south through Mexico to western Panama (Bocas del Toro Prov.). In Costa Rica and adjacent western Panama, it is known from only six collections from both slopes of the Cordillera de Talamanca. Flowering specimens have only been collected in March, April, July, and August; fruiting in August, September, and December-January. Rare/Infrequent. Chimaphila maculata is characterized by leaves that are lustrous or shiny dark green above with a broad, whitish line along the midrib and larger secondary nerves and pale reddish-purple beneath and coarsely serrate margins. Throughout its ex- tensive range, there is much variation in leaf mor- phology and degree of maculation, with numerous populations, especially in Mexico, being formally recognized (Rydberg, 1914; Camp 1939; Dorr, 1995). One variable species is herein recognized. Chimaphila maculata is probably most closely re- lated to C. umbellata (L.) W. P. C. Barton, another species of widespread temperate and boreal dis- tribution but with isolated populations in the mountains of Mexico, Guatemala, and on the is- land of Hispaniola. The two species differ most conspicuously in that C. maculata has maculated leaf blades that are coarsely serrate the entire length (vs. green leaf blades that are finely serrate, often only in the apical margins in C. umbellata) and villous staminal filaments (vs. glabrous or pa- pillate). Comarostaphylis Zuccarini REFERENCE — G. M. Diggs, Comarostaphylis. Fl. Neotrop. Monogr. 66: 146-193. 1995. Erect to spreading, rarely trailing shrubs to small trees; bark exfoliating. Leaves alternate, petiolate. the blade bifacial, usually coriaceous, margin entire or toothed, the venation pinnate. Inflorescences usually terminal, paniculate (in ours); pedicels continuous with calyx; bracteoles 2. basal -to apical. Flowers (4) 5-mer- ous, apparently without odor; calyx synsepalous. persis- tent in fruit, lobes much longer than tube, separate or slightly imbricate at anthesis, appressed to corolla, spreading to reflexed in fruit; corolla sympetalous, aes- tivation imbricate, cylindric to nearly globose, glabrous or pubescent, lobes much shorter than tube; stamens (Fig. ID) 10, equal; filaments dilated near base; connec- tives with disintegration tissue present, with 2, abaxial, recurved, horn-like spurs;' anthers ovoid, the surface smooth, dehiscing by 2, introrse pores/slits '/4-'/, to near- ly '/, as long as anther; pollen grains in tetrahedral tet- rads, lacking viscin threads; ovary superior, papillate; ovule 1 per locule. Fruit drupaceous, globose, juicy, the surface warty, granular or papillate, the nutlets united into a solid, spheroidal, thick-walled, stone-like endo- carp; seed I per nutlet, 1-1.5 mm long. Comarostaphylis consists of 10 species distrib- uted in the United States (especially southern Cal- ifornia), south through Mexico to western Panama (Chiriqui Prov.). It is characterized by its superior ovary and drupaceous, warty fruit. One species occurs in Costa Rica. Comarostaphylis arbutoides I null.. Edward's Bot. Reg. 29: 30. 1843. Arctostaphylos arbu- toides (Lindl.) Hemsl., Biol., Cent.-amer., Bot. 2: 278. 1881. Figure 10. Erect to spreading, much-branched, evergreen shrubs to small trees 1-20 m tall; bark gray, tan, reddish-brown or brown; twigs glabrous and sometimes glaucous or glandular hirsute to ferruginously tomentose. the tri- chomes glandular or eglandular. Leaves with petioles 4- 20 mm long, glabrous and sometimes glaucous or with indumentum as twigs; leaf blades lanceolate to elliptic, ovate or obovate, 4.5-12.5 cm long. 1-4.5 cm wide, base cuneate, apex obtuse to acute, margin sometimes undulate or with a few serrations, upper surface glabrous or scattered pilose, lower surface glabrous or with fer- ruginous eglandular tomentum to nearly wooly. midrib glabrous or with glandular or eglandular hairs margin- ally plane or revolute, rarely with a few serrations. In- florescences terminal, densely paniculate. 3.5-13.5 (18) cm long, the rachis, pedicels, bracts, bracteoles and ca- lyx glabrous or tomentose to glandular hirsute; floral bract lanceolate, nearly plane to navicular or carinate, 2.5-10 mm long, apex acuminate, ciliate or not; pedicels 5-8 mm long; bracteoles basal to nearly apical. 1.5-4 mm long, ciliate or not. Flowers with calyx lobes 1.7- 2.5 mm long, 1 .4-2 mm wide, triangular to ovate, acute to acuminate, glabrous or pubescent, trichomes glandu- lar or not. often ciliate, strongly reflexed in fruit; corolla membranous, unistratose. broadly urceolate, 5.7-8 mm long, 4-6 mm diam.. glabrous or pubescent, greenish- white to cream, pale yellow or white, sometimes suf- fused with pink, lobes 1-1.7 mm long, the margins often bluntly erose; stamens 2.5-3.2 mm long; filaments 2.4- 3 mm long, villous; anthers 1-1.5 mm long, the spurs LUTEYN AND WILBUR: FAMILY #172 ERICACEAE 51 0.4-1 mm long. Drupe ovoid to globose, 4-6 (7) mm diam., conspicuously warty, glabrous. Comarostaphylis arbutoides ranges from Mex- ico (Chiapas) to western Panama. It is found in montane Quercus forests, cloud forests, and pa- ramo, 1350-3800 m elevation, in the understory and as an important species in succession. This species "forms arbutoid ectomycorrhizal associ- ations with members of the Agaricales, Boletales, and Russulales. The roots of the plant take on a definitive morphology and anatomy. Above tim- berline, in the pdramo zone of the Talamancas, the plant can form extensive thickets, or in some sites, semi-open mono-dominant stands occur" (Hall- ing, pers. comm. and http://www.nybg.org/bsci/ res/hall/arbutoides.html). Comarostaphylis arbutoides is distinguished from its congeners by its large and entire leaf blades, densely paniculate inflorescences, and southern distribution. It is morphologically most similar to C. discolor (Hooker) Diggs, from which it is separated by minor leaf blade differences (shape, serrations) and slightly denser inflores- cences; C. discolor also grows several hundred kilometers to the north (Diggs, 1995). It has two subspecies that are distinguished by the following key: Key to the Subspecies of Comarostaphylis arbutoides la. Leaf blades ferruginous tomentose beneath; twigs, petioles, and calyx lobes pubescent; Cordillera de Talamanca to W Panama subsp. arbutoides Ib. Leaf blades glabrous beneath; twigs, petioles and calyx lobes glabrous; central volcanic highlands of Costa Rica subsp. costaricensis Subsp. arbutoides. Comarostaphylis chiriquensis Camp, Ann. Missouri Bot. Card. 26: 297. 1939. C. sleumeri Suess., Bot. Jahrb. Syst. 72: 282. 1942. Illustration: R. L. Wilbur & J. L. Luteyn, Ann. Missouri Bot. Card. 65: 36, fig. 3. 1978 (as Arctostaphylos). Shrubs or small trees to 20 m tall; twigs and petioles hirsute to ferruginously tomentose, the trichomes glan- dular or eglandular. Leaves with petioles pubescent; leaf blades usually densely ferruginous tomentose beneath, the midrib trichomes often glandular. Inflorescences with rachis, pedicels, bracts and bracteoles tomentose to glandular hirsute. Flowers with calyx tomentose, the lobes with glandular or eglandular pubescence; corollas glabrous to tomentose; ovary pubescent or more rarely glabrous. Subspecies arbutoides ranges from the central highlands of Chiapas, Mexico to western Panama. In Costa Rica and Panama, it occurs in montane cloud forests (Quercus} and paramo, in the Cor- dillera de Talamanca of Costa Rica to western Panama (Chiriqui Prov., Volcan Bani), 1350- 3800 m elevation. Flowering and fruiting occur throughout the year. The subspecies is character- ized by ferruginous pubescence. The fruit is eaten in Honduras. The plants are visited by small, bumblebee-like Hymenoptera in Guatemala (Diggs, 1995). Common. Subsp. costaricensis (Small) Diggs, Brittonia 38: 344. 1986. Comarostaphylis costaricensis Small, N. Amer. Fl. 29: 89. 1914. Arctostaph- ylos costaricensis (Small) Standl., Publ. Field Mus. Nat. Hist., Bot. Ser. 4: 320. 1929. A. ar- butoides van costaricensis (Small) Wilbur & Luteyn, Ann. Missouri Bot. Card. 65: 36. 1978. Shrubs to 5 m tall; twigs and petioles glabrous, often glaucous. Leaves glabrous beneath. Inflorescences gla- brous or with slight pubescence. Flowers with calyx lobes, corollas, and ovary glabrous. Subspecies costaricensis occurs in the montane cloud forest and in the dense ericaceous scrub vegetation at the edge of volcanic craters, 2500- 3450 m elevation. It is endemic to Costa Rica, where it is abundant on the crater rim of Volcan Irazii (Cartago Prov.), and extends down into the forest of the Cordillera Central at Palmira (Ala- juela Prov.) and Volcan Barva (Heredia Prov.). Flowering and fruiting occur throughout the year. The subspecies is characterized by its glabrous habit and often glaucous twigs and petioles. Com- mon names in Costa Rica include anonillo (He- redia), arraydn (Cartago), and madrono (Carta- go). Locally Common. Didonica Luteyn & Wilbur REFERENCES — J. L. Luteyn & R. L. Wilbur, New genera and species of Ericaceae (Vaccinieae) 52 FIELDIANA: BOTANY from Costa Rica and Panama. Brittonia 29: 255- 276. 1977. J. L. Luteyn, A synopsis of the genus Didonica (Ericaceae: Vaccinieae) with two new species. Syst. Bot. 16: 587-597. 1991. Epiphytic shrubs, with mature stems glabrous to var- iously pubescent. Leaves alternate, petiolate, the blade coriaceous, flat to involute, margin entire or remotely crenate, the venation plinerved. Inflorescences axillary, flowers solitary or racemose; floral bract and bracteoles usually small and inconspicuous, margin glandular-fim- briate; pedicels continuous with calyx. Flowers 5-6- merous; calyx synsepalous, tube terete or apparently bluntly angled, limb usually terete and campanulate, lobes broad, relatively inconspicuous, erect; corolla sympetalous, aestivation valvate, thin or succulent, bis- tratose (in ours), cylindric-campanulate, usually green- ish; stamens 10-12, equal; filaments distinct; connec- tives lacking disintegration tissue or spurs; anthers lack- ing awns, thecae finely papillate, incurved at base, tu- bules 2, distinct, rigid, slightly shorter than thecae, somewhat spreading, dehiscing by oval, latrorse clefts which do not extend to tip of tubule; pollen grains in tetrahedral tetrads, lacking viscin threads; ovary inferior; style filiform, about as long as corolla and included, or slightly exserted, glabrous. Fruit a berry; seeds numer- ous. Didonica consists of four species endemic to Panama and adjacent eastern Costa Rica. Virtually nothing is known about their biology, but the ge- nus as a whole, as well as each of the four species, must be considered in extreme danger of extinc- tion because the localities and habitats in which they are found are undergoing severe alteration. Only one species is currently known in Costa Rica. The morphological relationships of Didon- ica are presumed to be with genera such as Ma- cleania, Psammisia, and Mycerinus because of similar short, broad, rigid anther tubules or with Vaccinium sect. Oreades Sleumer because of green flowers and sometimes latrorse anther de- hiscence (Luteyn & Wilbur, 1977). As yet, there have been no DNA studies of this genus. Didon- ica, however, represents a rather distinct evolu- tionary line, and we are still unable to suggest with confidence its relationships to other Vacci- nieae. Key to the Species of Didonica la. Leaves imbricate, subsessile, clasping to amplexicaul, the blades deeply cordate at base, the margins entire; petioles to 2.5 mm long; mature stems hirsute; pedicels ca. 3 mm long .... D. crassiflora Ib. Leaves well separated, not imbricate, obviously petiolate, the blades short-attenuate or short-cuneate at base, the margins remotely crenate to essentially entire; petioles 5-15 mm long; mature stems glabrous or puberulent; pedicels (90) 150-205 mm long D. pendula Didonica crassiflora Luteyn, Syst. Bot. 16: 592, fig. 4. 1991. Epiphytic shrubs; mature stems subterete, striate, hir- sute with white hairs 0.5-0.7 mm long, glabrate; twigs subterete, striate to ribbed, densely hirsute with white hairs to 1.5 mm long; axillary buds with outer pair of scales broadly ovate, mucronate, ca. 2 mm long. Leaves subsessile with petioles subterete, broadly canaliculate adaxially, to 2.5 mm long, 2.5-3 mm diam., densely hirsute; leaf blades coriaceous, densely imbricate, flat, amplexicaul with lobes slightly overlapping, ovate, 3.4- 7 cm long, 1.7-3.7 cm wide, base rounded and deeply cordate, apex acuminate, sparsely hirsute over both sur- faces and densely hirsute basally along nerves beneath. 5 (9)-plinerved from near base, midrib and inner 1-2 pairs of lateral veins impressed above and conspicuously raised beneath, outer 2 pairs of lateral veins plane to slightly raised above and beneath but obscure. Inflores- cences with flowers solitary, surrounded at base by ca. 5 ovate, acute, marginally hirsute and glandular-fimbri- ate (inflorescence) bracts to ca. 1 mm long; floral bract ovate, ca. 1.5 mm long, apex acute to obtuse, sparsely short-pilose, margin with few, glandular fimbriae; pedi- cels subterete, broadening apically, ca. 3 mm long, ca. 1.5 mm diam., hirsute with white hairs ca. 1 mm long; bracteoles medial, ovate, ca. 1.5 mm long, densely pi- lose, apex acuminate. Flowers 6-merous; calyx 8-8.5 mm long, densely hirsute all over with white hairs to ca. 0.6 mm long, tube poorly developed, campanulate. ca. 2 mm long, ca. 4 mm diam., limb broadly campanulate, smooth, ca. 6 mm long, ca. 1 1 mm diam. at apical tip, lobes broadly deltate, 1 .2-2 mm long, 3.5-4 mm wide at base, sinuses concave; corolla succulent when fresh, coriaceous, cylindric-campanulate, 14-15 mm long, 10- 1 1 mm diam. at apical tip, pale green, nearly glabrous at base where covered by calyx limb but densely short- tomentose apically with pale, reddish, flat hairs to 0.7 mm long, lobes triangular-deltate, 3-3.5 mm long, ca. 4 mm wide at base, sparsely short-pilose within, apex acute; stamens 12, ca. 1 1 mm long; filaments 3-4 mm long, abaxially pilose; anthers 9.5-10 mm long, thecae 6.5-7.5 mm long, tubules ca. 3 mm long, dehiscing by oval clefts ca. 1 .5 mm long; style slightly exserted, 1 5- 16 mm long. Berry not seen. . Didonica crassiflora is known only from the type, which was collected along a stream on white, sandy soil with a very thin humus layer at the Fortuna Dam site, Chiriquf Province, Panama, LUTEYN AND WILBUR: FAMILY #172 ERICACEAE 53 1200 m elevation. It has not yet been collected in Costa Rica. The type was flowering in February. Endangered. Didonica crassiflora is characterized by its densely imbricate, amplexicaul leaves with hirsute blades; solitary flowers that are subsessile and 6- merous; hirsute calyces and pedicels; and tomen- tose, succulent corollas. It is morphologically sim- ilar to D. subsessilis Luteyn (endemic to Veraguas Prov., Panama), which is probably also its closest relative. Both species have short-petiolate, am- plexicaul to clasping leaves, solitary flowers, and succulent corollas. They differ by D. crassiflora having mature stems hirsute (not glabrous); leaves amplexicaul with the blades flat and entire mar- gined (not leaves clasping with blades flat to in- volute and margins remotely crenate); flowers 6- merous (not 5-merous); calyces and corollas hir- sute and tomentose, respectively (not glabrous); and bracteoles inconspicuous and ca. 1 .5 mm long (not conspicuous, concealing calyces and 7-8 mm long). Didonica pendula Luteyn & Wilbur, Brittonia 29: 255, fig. 1. 1977. Figure 2K. Epiphytic shrubs 0.7-1 m tall, arising from a ligno- tuber; mature stems subterete, smooth or minutely stri- ate, glabrous or puberulent; twigs subterete, striate, gla- brous or puberulent; axillary buds with outer pair of scales narrowly ovate to linear-acicular, 2.5-5 mm long, long-acuminate. Leaves with petioles subterete, rugose, narrowly winged laterally over entire length, 5-15 mm long, 1.5-2 mm diam., glabrous; leaf blades coriaceous, slightly concave and bullate to flat, ovate to lanceolate- elliptic, (4.5) 6-18.5 cm long, (1.5) 2-5.4 cm wide, base rounded to obtuse and short-attenuate or short-cuneate, apex short- to long-acuminate, glabrous but bearing de- ciduous glandular fimbriae beneath, margin remotely crenate to essentially entire, weakly 5-7-plinerved (in ours) but appearing pinnately nerved with inner pair of lateral nerves arising about 1 .5-4 cm above base, midrib conspicuous above in raised and thickened basal ca. 5 mm. then impressed apically, raised and prominent be- neath, lateral nerves inconspicuous above, thin and slightly raised entire length, raised and conspicuous be- neath, reticulate nerves slightly raised on both surfaces but somewhat conspicuous only beneath. Inflorescences with flowers solitary or in 1 -flowered racemes, some- times two racemes arising from same node, subtended by several, narrowly lanceolate to linear, long-acuminate bracts ca. 2 mm long; radii s (when present) subterete, thin, 1.5-10 mm long, the basal 1-2 nodes sterile; floral bract ovate, ca. 1-1.8 mm long, glabrous but margin glandular-fimbriate; pedicels long-pendent, minutely ribbed, (9) 15-20.5 cm long, 0.3-0.8 mm diam. through- out most of length, then greatly expanded apically near articulation to 5-8 mm diam. (15 mm diam. in fruit), glabrous; bracteoles 2.5-3 cm from base, subopposite, 1 .5-4 mm long, aristate, glabrous, margin glandular-fim- briate. Flowers 5-6-merous; calyx 8-12 mm long, gla- brous, tube obconic, muricate, 3.5-5 mm long, ca. 4-5 mm diam. at base, limb campanulate, conspicuously veined, 3.5-7 mm long, ca. 12-13 mm diam. at apical tip, lobes broadly ovate, apiculate, 1-2.5 mm long, ca. 5 mm wide at base, sinuses acute to concave; corolla thin-carnose when fresh, bistratose, chartaceous, cylin- dric-campanulate, 16-25 mm long, 13-25 mm diam. at apical tip, glabrous, pink to greenish-white, lobes broad- ly ovate, 2.5-5 mm long, 5-6 mm wide at base, apex acute to rounded, glabrous internally; stamens 10 or 12, shorter than corolla, ca. 12-18 mm long; filaments ca. 4.5 mm long, ca. 2 mm wide at base, ciliate; connectives short-pilose; anthers ca. 10 (17?) mm long, thecae 6.5- 7.5 mm long, tubules 3-3.5 mm long, pilose adaxially and on inner surfaces, dehiscing by oval clefts 1 .5-2 mm long [authors unable to examine stamens from 25 mm long corolla]; style 16-18 mm long. Berry spherical, to 27 mm diam., translucent white. Didonica pendula is distributed in eastern Cos- ta Rica and west-central Panama, in cloud forest, 600-2600 m elevation. Five collections are known from Costa Rica (Limon Prov., Caribbean slope: road from Fila Dimat to Soki, L. D. Gomez et al. 23865; Zona Protectora Barbilla, SE of Si- quirres, M. Grayum et al. 7946; and Sabanas de Durika, Parque Nacional La Amistad, Cerro Bir- icuacua, G. Herrera 6227. Puntarenas Prov., Pa- cific slope: Goto Brus, Zona Protectora Las Ta- blas, E. Navarro 838) and three from Panama. Flowering specimens have been collected in Jan- uary, July, and October; fruiting specimens in January and October. Endangered. Didonica pendula is characterized by leaves that are widely spaced and obviously petiolate, with blades basally broadly cuneate to short-at- tenuate and marginally remotely crenate to essen- tially entire; inflorescences with flowers solitary (or racemose but with only one flower maturing); and (usually) extremely long pedicels. The pedi- cels of some populations are the longest known in the Ericaceae, although one recent collection from Panama Province, Panama, has pedicels 9.5- 13 cm long. The six collections of this species show differences in twig pubescence, leaf size and shape, pedicel length, and number of flower parts. For example, the Panamanian populations (includ- ing the type) have thick-coriaceous, slightly con- cave, slightly bullate, ovate, short-acuminate leaf blades, 5.5-8.5 cm long, with the margins re- motely crenate, and the venation conspicuously 5 (7)-plinerved with the inner pair of lateral nerves arising from near base. The Costa Rican popula- tions have thin-coriaceous, flat, not bullate, lan- ceolate-elliptic, long-acuminate leaf blades, 10- 18 cm long, with margins essentially entire, and FIELDIANA: BOTANY the venation obscure and appearing pinnate- nerved, with the inner pair of lateral nerves aris- ing well above the base. We do not, however, con- sider these differences sufficient to warrant taxo- nomic recognition at this time. Additional collec- tions are sorely needed. Disterigma (Klotzsch) Niedenzu REFERENCES — A. C. Smith, The genera Sphy- rospermum and Disterigma. Brittonia 1 : 203-232. 1933. R. L. Wilbur, A synopsis of the genus Dis- terigma (Ericaceae: Vaccinieae) in Mexico and Central America with the description of two pre- viously undescribed species. Bull. Torrey Bot. Cl. 119: 280-288. 1992. Compact, bushy to straggly, terrestrial or epiphytic shrubs. Leaves alternate, often congested, petiolate, the blade coriaceous, usually less than 3 cm long, the margin entire or crenate, the venation usually obscurely pliner- ved. Inflorescences axillary, of solitary or 1-6-clus- tered, subsessile flowers; floral bract 1 to several, mi- nute, subcoriaceous; pedicels short or lacking, articulate with calyx; bracteoles 2, apical, immediately subtending and embracing calyx tube. Flowers (3) 4 (5)-merous, without odor, aestivation valvate (or rarely imbricate in calyx); calyx tube campanulate to short cylindric, the lobes suberect; corolla sympetalous, subcylindric or campanulate-cylindric, (3) 4-5-lobed; stamens equal, (5-6) 8, usually twice as many as or rarely equal to corolla lobes; filaments distinct, ligulate, longer or short- er than anthers; connectives lacking disintegration tissue or spurs; anthers abaxially attached near base, membra- nous, lacking awns, thecae smooth, tubules 1 or 2, de- hiscing by elongate, elliptical, introrse clefts or subter- minal pores; pollen grains in tetrahedral tetrads, lacking viscin threads; ovary inferior, (3) 4 (5)-locular; style fi- liform, about as long as corolla or slightly exserted, gla- brous; stigma truncate; nectariferous disc fleshy, annu- lar-pulvinate. Fruit a thick-walled, coriaceous berry, blue-black or translucent white; seeds numerous, some- times mucilaginous. Disterigma is a genus of about 35 species dis- tributed from southern Mexico to Bolivia and east to Guyana (Mt. Roraima). It is characterized by solitary or few-flowered clusters of sessile to sub- sessile flowers and two apical bracteoles that sub- tend the calyx tube and usually surround the entire calyx and sometimes also part of the corolla. The genus is morphologically most similar to Vaccin- ium, from which the bracteole feature distinguish- es it. Four species are currently known from Costa Rica. Disterigma is currently being revised in a Ph.D. study by L. Paola Pedraza (New York Bo- tanical Garden). Key to the Species of Disterigma la. Flowers 3-merous; corolla deep red; stamens 6 D. trimerum Ib. Flowers 4-merous; corolla white (to pinkish), pale greenish, or red; stamens 8 or 10 2 2a. Anther tubules 1; leaf blades with a terminal apiculus 1-2 mm long D. utleyorum 2b. Anther tubules 2; leaf blades not apiculate 3 3a. Leaf blades 4-20 mm wide, the apex rounded or obtuse (rarely acute); calyx tube 4-angled; calyx lobes eciliate; corolla 4.7-8 (14.5) mm long; flowers 1-3 per axil 4 4a. Leaf blades mostly 6-12 mm long, 4-7 mm wide; flowers mostly 1 (2 or 3) per axil; corolla 5-8 (14.5) mm long D. humholdtii 4b. Leaf blades mostly 15-30 mm long, 10-20 mm wide; flowers (1) 2 (3) per axil; corolla 4.7-7.5 mm long D. alaternoides 3b. Leaf blades 0.8-3 mm wide, the apex acute; calyx tube terete; calyx lobes ciliate; corolla 6-12 mm long; flowers always solitary 5 5a. Leaf blades 0.8-1.2 mm wide; calyx lobes ca. 1.5-2.2 mm long; corolla 6-8 mm long; staminal filaments 3.7-5 mm long D. fortunense 5b. Leaf blades 1.5-3 mm wide; calyx lobes 2.5-3 mm long; corolla 10-12 mm long; staminal filaments 7-9 mm long 6 6a. Corolla lobes 2.2-3 mm long; leaf blades 1 .5-2.2 mm wide, glabrous or at least glabrate; stamens 8, with filaments 7-9 mm long, twice as long as the anthers . . D. hammelii 6b. Corolla lobes ca. 1.5-1.7 mm long; leaf blades 2.5-4 mm wide, moderately short- pilose; stamens 10, with filaments ca. 7 mm long, less than 1.5 times longer than anthers D. pilosum LUTEYN AND WILBUR: FAMILY #172 ERICACEAE 55 Disterigma alaternoides (Kunth in H.B.K.) Nie- denzu, Bot. Jahrb. Syst. 11: 224. 1889. Vaccin- ium alaternoides Kunth in H.B.K., Nov. Gen. Sp. 3: 265. 1818. Metagonia alaternoides (Kunth i/i H.B.K.) Nuttall, Trans. Amer. Phil. Soc. N.S. 8: 266. 1843. Terrestrial or epiphytic shrubs 0.5-1.2 m tall (ours); mature stems terete or subterete, sometimes bluntly an- gled, nitid, glabrous; twigs subterete, angled, densely and finely spreading short-pubescent to hirsutulous with hyaline or grayish to brownish, ± straight trichomes 0.2-0.6 mm long. Leaves with petioles subterete, stout and usually broadly flattened adaxially, 1-3 mm long, 1-1.5 mm diam., densely spreading short-pubescent; leaf blades coriaceous, with internodes mostly '/J-TJ length of adjacent leaves, broadly elliptic, (15) 18-30 mm long, 10-20 mm wide, base rounded, apex obtuse to very broadly rounded or rarely acute, margin entire, glabrous or occasionally sparingly puberulous when young, also with inconspicuous appressed, amber-colored, glandular- fimbriae ca. 0.2 mm long beneath, obscurely 3-5-pli- nerved from base, midrib plane above and slightly raised beneath, lateral nerves usually plane above and slightly raised but usually obscure beneath, reticulate veinlets obscure on both surfaces. Inflorescences fasciculate, ( 1 ) 2 (3)-flowered (in ours); floral bract 1, semi-orbicular to broadly ovate, to 3 mm long, apex obtuse; pedicels sub- terete, ribbed, 2-3 mm long, puberulous to glabrate, sometimes also with scattered, minute, glandular fimbri- ae; bracteoles reaching to base of calyx lobes, imbricate at base, semi-orbicular, to 3 mm long, apex rounded, glabrous. Flowers 4-merous; calyx ca. 3-4 mm long, glabrous, tube cylindric, quadrangular, ca. 1 .5 mm long, limb erect-spreading to campanulate, ca. 1.5 mm long, lobes deltate, ca. 1.6 mm long, apex acute, eciliate, si- nuses acute; corolla thin-carnose, cylindric, ca. 4.7-7.5 mm long, 2-2.5 mm diam., glabrous or sometimes spar- ingly appressed puberulous, white, pinkish or pale red, to rarely even crimson, lobes deltate, 1.5-2 mm long, apex acute; stamens 8, ca. 6 mm long; filaments 2-4 mm long, densely hirsutulous; anthers 1.2-3 mm long, tubules 2, distinct, equaling or slightly exceeding thecae. Berry to 10 mm diam., translucent white or translucent pinkish, reddish-purple, or wine-red (unknown in our range). Disterigma alaternoides ranges from Panama, south through the Andes to Bolivia, and east to Guyana and is found in premontane to montane cloud forest and paramo, (1600) 2400-3400 (3900) m elevation. It has not yet been found in Costa Rica. The Panamanian collections were flowering in March, April, June, and August (Wil- bur, 1992). Endangered (if present). Disterigma alaternoides is characterized by its broad, apically rounded to acute (in South Amer- ica) leaves; inflorescences with usually paired, 4- merous flowers (in ours); and white to pinkish co- rollas. Morphologically it is most similar to D. humboldtii, with which it has in common broad and usually rounded leaves, 4-merous flowers, and white corollas. The differences between these species are mentioned in the key, and although few, they do seem to hold within our range. Al- though mostly distributed in South America, D. alaternoides has been reported in Chiriquf Prov- ince, Panama (Wilbur, 1992). More collections are needed from Panama, and it may be possible that only one species, D. alaternoides, should be rec- ognized. Disterigma fortunense Wilbur, Bull. Torrey Bot. Cl. 119: 286. 1992. Epiphytic or terrestrial, erect to pendent subshrubs 0.3-0.4 m tall, often scrambling and rooting where stems touch moist soil; mature stems subterete, somewhat ridged and grooved, moderately to densely short-pilose with whitish to tawny trichomes 0.5-1 mm long, these especially dense at raised nodes; internodes very short, usually 1-2 mm long, mostly less than Vg-V4 length of nearest leaf. Leaves with petioles 0.5-0.7 mm long, sub- terete, somewhat canaliculate adaxially, rugose, glabrous but appearing puberulous at base due to long nodal tri- chomes; leaf blades coriaceous, strongly spreading from stems often at about a right angle, needle-like, very nar- rowly linear, (4.5) 7-9 (11) mm long, 0.8-1.2 mm wide, 8-10 times as long as wide, base ± rounded or very shortly tapering, apex acute, margin entire to very mi- nutely serrulate, ciliolate when young but becoming gla- brous on both surfaces, obscurely 3-5-plinerved from near base although inner pair of lateral nerves arising ca. 2-3 mm above base, midrib and lateral nerves plane to weakly raised above but moderately weakly raised beneath, reticulate veinlets obscure. Inflorescences with flowers solitary; floral bracts ca. 5, to ca. 0.8 mm long, ovate, ± orbicular to oblong, apex rounded, copiously short-ciliate, closely imbricate; pedicels ca. 1 mm long or less; bracteoles ca. 0.8-2 mm long, ± orbicular or oblong, copiously short-ciliate, broadly imbricate and broadly clasping base of calyx. Flowers 4-merous; calyx ca. 3.2-4 mm long, tube ca. 1-1.8 mm long, ca. 1 mm diam., campanulate to somewhat globose, ± terete, smooth, glabrous, limb spreading, ca. 1 .6-2.2 mm long, lobes ± erect, narrowly deltate to triangular-lanceolate, ca. 1.5-2.2 mm long, apex acute, ciliate in apical V2-^3 but otherwise glabrous, sinuses acute; corolla thin-car- nose, ± fusiform in bud but cylindric to somewhat in- fundibuliform when mature with base slightly tapering and mid- to apical swelling, 6-8 mm long, 2-3.6 mm diam., pink to white, essentially glabrous but with a few, apical trichomes, lobes triangular, 1-1.5 mm long, apex acute, strongly reflexed, with few microscopic tri- chomes; stamens 8, ca. 6 mm long, exserted from throat of corolla for ca. 1 mm; filaments 3.7-5 mm long, dense- ly ciliate, obviously longer than anthers; anthers ca. 2.5- 2.7 mm long, thecae slightly papillate, ca. 1-1.2 mm long, tubules 2, distinct, ca. 1.5-1.7 mm long, dehiscing by oblique, elongate pores; style exserted, ca. 8-8.5 mm long. Berry spherical, ca. 5.2 mm diam., translucent white with erect, persistent lobes. 56 FIELDIANA: BOTANY Disterigma fortunense is a rare endemic in pre- montane cloud forest at the Fortuna Dam site of western Chiriqui Province, Panama, at ca. 1000- 1 100 m elevation. It has not yet been collected in Costa Rica and is known from only six collec- tions. Flowering collections have been made in January and February; fruiting only in January. Endangered. Disterigma fortunense is characterized by hav- ing needle-like, narrow leaves; small 4-merous flowers; and pink to white corollas. It belongs to a small group of species of Disterigma that nor- mally have a scandent, "wiry", epiphytic habit; short internodes; and narrowly lanceolate to near- ly linear leaf blades less than 5 mm wide. This group ranges from western Panama to central Peru and includes D. agathosmoides (Wedd.) Nied., D. fortunense, D. hammelii, D. luteynii Wilbur, D. panamense Standley, D. pilosum, and D. weberbaueri Hoerold. Apparently D. fortunense is most closely related to the South American (Co- lombia and Ecuador) D. agathosmoides, and more collections and field observations are needed to distinguish the species. One collection (Luteyn 14808, NY) was associated with stinging ants. Disterigma hammelii Wilbur & Luteyn, Ann. Missouri Bot. Gard. 68: 160. 1981. Straggly, pendent, epiphytic shrubs; mature stems thin, subterete, irregularly ridged and grooved, moder- ately spreading to appressed short-pilose with hyaline to tawny trichomes; twigs subterete, angled, ridged, dense- ly spreading short-pilose with tawny to reddish-brown trichomes 0.4-1 mm long; internodes short, almost all less than '/4 length of subtending leaves and usually 1- 2 mm long. Leaves with petioles subterete, flattened adaxially, 0.5-1 mm long, glabrous or nearly so; leaf blades coriaceous, elliptic to lance-elliptic, (4) 5-7 (8) mm long, 1.5-2.2 (3) mm wide, mostly about 2.5 times as long as wide, base gradually tapering into petiole, apex acute in general outline but actual apex obtuse, margin entire, glabrate on both surfaces and margin or ciliolate at apex, rarely with a few appressed, reddish, glandular fimbriae on lower surface, venation indistinct except midvein slightly elevated beneath. Inflorescences axillary, of solitary flowers; floral bracts 5-8, mostly 0.8-1.8 mm long, ovate to broadly elliptical, tawny to dull reddish-brown, scarious; pedicels ca. 0.5-0.8 mm long, glabrous, obscured from view by bracts and brac- teoles; bracteoles broadly clasping, covering calyx to base of lobes, ovate to broadly oblong, striate, appar- ently concave, 2-2.7 mm long, ca. 2-2.5 mm wide, apex broadly rounded, margin scarious. Flowers 4-merous; calyx ca. 3.8-4 mm long, glabrous to sparingly short- pilose, tube terete, campanulate to obconic, ca. 1.4-1.8 mm long, 1-1.4 mm cliam.. limb spreading, ca. 2.8-3.3 mm long, lobes strongly ascendent, striate, narrowly lan- ceolate with incurved margin, ca. 2.5-3 mm long, apex acute, margin entire, ciliate for apical '/,-'/, but otherwise glabrous, sinuses acute; corolla cylindric, sometimes somewhat quadrate, 10-12 mm long, 2.5-3.2 mm diam., pale pink, nearly glabrous but usually with a few, ap- pressed. glandular trichomes ca. 0. 1 mm long just above middle, lobes ascendent to more typically reflexed, 2.2- 3 mm long, lanceolate, acute, spreading, short-pubescent with hyaline trichomes; stamens 8. ca. 10-11 mm long, about as long as corolla; filaments ca. 7-9 mm long, conspicuously longer than anthers, sparingly hyaline short-pilose with trichomes 0.2-0.6 mm long; anthers ca. 3.7-4.3 mm long, thecae ca. 1.2-1.5 mm long, tubules 2. distinct, ca. 2.5-2.8 mm long; style exserted, ca. 13- 14 mm long, glabrous. Berry not seen. Disterigma hammelii occurs in montane and Continental Divide cloud forest, 1800-2200 m, and is known only from three collections in Chi- riquf-Bocas del Toro Provinces, Panama. It has been collected near but not yet within the borders of Costa Rica. Flowering specimens have been collected in February and March; fruits are un- known. Endangered. Disterigma hammelii also belongs with those Disterigma species that normally have a scandent, "wiry," epiphytic habit; short internodes; and narrowly lanceolate to nearly linear leaf blades less than 5 mm wide, as mentioned in the discus- sion above. From this group D. hammelii may be distinguished by its longer corolla, especially its proportionally long staminal filaments, and by its exserted style. Disterigma humboldtii (Klotzsch) Niedenzu, Bot. Jahrb. Syst. 11: 224. 1889. Vaccinium humboldtii Klotzsch, Linnaea 24: 57. 1851. V. pachyphyllum Hemsl. Biol. Centr.-amer., Bot. 2: 275. 1881. D. pachyphyllum (Hemsl.) S. F. Blake, J. Washington Acad. Sci. 16: 365. 1926. Figures 2U and 3. Bushy, compact to straggly, terrestrial or epiphytic shrubs (0.2) 0.5-1 (1.2) m tall; mature stems terete to subterete, sometimes conspicuously ribbed and bluntly angled due to raised nodes; twigs subterete, bluntly an- gled, striate, densely short-pilose to setose with ferru- ginous to brown hairs 0.5-1.2 mm long, or moderately to densely pubcrulent, glabrate. Leaves with petioles subterete, broadly flattened adaxially, 1-2 mm long, weakly to moderately short-pilose and this sometimes onto base of midrib beneath; leaf blades coriaceous, ovate, elliptic, to suborbicular, revolute near base, 6-12 (16) mm long. 4-7 (9) mm wide, bas^e rounded, apex obtuse to rounded, margin entire, sometimes with a dense but short, tuft of hairs at apex, otherwise glabrate, also bearing scattered, appressed, glandular fimbriae be- neath to ca. 0.2 mm long, obscurely 3-plinerved from base, midrib slightly impressed above and sometimes slightly raised beneath, all other venation completely ob- scured on both surfaces. Inflorescences of solitary flow- LUTEYN AND WILBUR: FAMILY #172 ERICACEAE 57 ers or 2-3 in a fascicle; floral bract 1 (3), ovate, less than 1 (2.2) mm long: pedicels to 2 mm long, glabrous; bracteoles clasping calyx to base of lobes, broadly ob- long-ovate, 1.5-4 mm long, broader than long, apex rounded to truncate. Flowers 4-merous; calyx ca. 3.3 mm long, glabrous, tube campanulate, 4-angled, ca. 1 mm long, limb spreading. 2-3.5 mm long, lobes erect to slightly spreading, ovate-deltate, 1.3-2.5 mm long, apex acute, glabrous or sparsely appressed pubescent to ciliate, sinuses acute; corolla thin-carnose, cylindric, 5- 8 (14.5) mm long, 2-4 (5) mm diam., glabrous to spar- ingly appressed pilose externally and rarely moderately short-pilose internally, white or sometimes suffused with rose or pink, lobes erect to horizontally spreading, del- tate, ca. 1 mm long, apex acute; stamens 8, 5.8-11.5 mm long, about equaling corolla but often exposed at anthesis when corolla lobes become reflexed; filaments 2.5-4.5 (7.5) mm long, moderately to densely hispidu- lous to appressed pilose (or villous) in apical half with whitish or hyaline hairs; anthers 2.4-5.5 mm long, the- cae 1.2-2 mm long, tubules 2, distinct, 1.3-3.5 mm long, dehiscing by elongated clefts about half the tubule length; style slightly exserted, glabrous. Berry spherical, crowned by persistent calyx limb, ca. 5 mm diam., trans- lucent white to translucent purple. Disterigma humboldtii is found in premontane cloud forest to subparamo, and pastures, 300- 3200 m elevation, from Mexico to Ecuador and east to Guyana. In Costa Rica it is encountered in the Cordilleras Tilaran, Central and Talamanca. Flowering occurs throughout the year; fruiting specimens have been collected in July, October, and December. Widespread. Disterigma humboldtii is characterized by its apically obtuse to rounded leaf blades, usually 1 (3)-flowered inflorescences, and 4-merous flow- ers. The relationships of D. humboldtii lie near D. alaternoides as discussed above. The berries are reportedly sweet. Flowers are visited by hum- mingbirds (J.L.L., pers. observ.), but bees are probably the primary pollinators. Disterigma pilosum Wilbur, Bull. Torrey Bot. Cl. 119: 285. 1992. Figure 4. Pendent, densely clustered, epiphytic shrubs; mature stems subterete, angled to ribbed, swollen at nodes; twigs subterete, angled, striate, densely spreading short- pilose with tawny hairs 1-1.2 mm long, the new inno- vations with persistent, striate, oblong, apically obtuse to rounded, ciliolate perulae to 4 mm long; internodes short, almost all less than '/, length of nearby leaves, mostly 2-6 mm long. Leaves with petioles subterete, 0.5-1.5 mm long, sparingly to moderately short-pilose; leaf blades coriaceous, thick when fresh but puckering and wrinkling on drying, elliptic to lance-elliptic, 8-15 mm long, 2.5-4 mm wide, mostly 3-4 times as long as wide, base gradually tapering and rounded, apex taper- ing to a blunt tip, margin entire, moderately short-pilose on both surfaces and margins, obscurely 3-plinerved from base, midrib and lateral nerves sometimes obscure above but raised beneath, reticulate nerves obscure above but slightly raised beneath. Inflorescences axillary, of sol- itary flowers; floral bracts 5-9, ovate to broadly orbicular, apex rounded, glabrous to short-pilose to minutely cilio- late, scarious, mostly 0.6-1.5 mm long, concealing pedi- cel; pedicels ca. 0.4-1.5 mm long; bracteoles broadly clasping, broadly ovate to ovate-oblong or ± orbicular, 1.8-3 mm long, apex rounded, margin scarious, ciliate. Flowers 4-merous; calyx ca. 4.2-4.5 mm long, somewhat striate, sparingly puberulous to short-pilose, tube terete, obconic, ca. 1 .5 mm long, limb ca. 2.7-3 mm long, lobes erect or strongly ascendent, narrowly lanceolate, acute with incurved margins, 2.5-3 mm long, sinuses acute; corolla cylindric to somewhat swollen medially, 10-12 mm long, 1.5-3 mm diam., sparingly short-pilose exter- nally, whitish to pale pink, lobes erect to somewhat re- flexed, lanceolate, acute, 1.5-1.7 mm long; stamens 10, ca. 12 mm long; filaments ca. 7 mm long, longer than anthers, swollen at base, scattered long-pilose in apical '/2; anthers ca. 5 mm long, thecae ca. 1 .5 mm long, tubules 2, distinct, ca. 3.5 mm long, dehiscing by clefts ca. 1.5 mm long. Berry not seen. Disterigma pilosum is found on the wet Carib- bean slope of the Continental Divide in Quercus forest, 2300-2700 (3000) m elevation. It is en- demic to the Cordillera de Talamanca near the Costa Rica/Panama border and is known only from three collections: Limon Province, Costa Rica (Davidse et al. 28937 and Morales 5801), and Bocas del Toro Province, Panama (T. Antonio 1630, sterile). Flowering specimens have been collected in September; fruits are unknown. En- dangered. Disterigma pilosum is characterized by its gen- erally dense to moderately short-pilose, narrow el- liptic to lance-elliptic leaf blades; 4-merous flow- ers with relatively long (2.5-3 mm) calyx lobes; and 10 stamens (not the normal eight). Disterigma pilosum is seemingly related to those species of Disterigma that normally have a scandent, "wiry" epiphytic habit; short internodes; and nar- rowly lanceolate to nearly linear leaves less than 5 mm wide, as mentioned in the discussion of D. fortunense above. It keys closest to D. hammelii, but until more collections of these taxa are made, their relationships are uncertain. Disterigma trimerum Wilbur & Luteyn in Lu- teyn & Wilbur, Brittonia 29: 258. 1977. Epiphytic or terrestrial, bushy shrubs 0.5-1.5 (2) m tall to small trees ca. 4 m tall (fide label); mature stems terete to subterete, conspicuously fissured, glabrous; twigs subterete, sharply ridged and grooved, indistinctly 3-angled to irregularly 4-angled, glabrous or minutely puberulous, sparingly glandular-fimbriate; internodes '/5-3/5 as long as the surrounding leaves. Leaves with 58 FIELDIANA: BOTANY petioles subterete, broadly canaliculate adaxially. 3-5 mm long, glabrous to puberulent; leaf blades coriaceous, broadly elliptic to somewhat obovate or spatulate, (0.8) 1.2-2 cm long, (4) 6-14 mm wide, base cuneate. apex obtuse to broadly rounded, short-retuse, margin entire to obscurely serrulate, glabrous above, evenly but sparingly bearing appressed, amber to reddish-brown, glandular fimbriae 0.2-0.3 mm long beneath, 3-5-plinerved from near base, midrib and lateral nerves raised (sometimes conspicuously) basally but nearly plane apically above, raised (sometimes conspicuously) beneath but often ob- scure on both surfaces, reticulate veinlets slightly raised but usually obscure on both surfaces. Inflorescences ax- illary, of solitary (rarely 2), subsessile flowers; floral bract 1, triangular-acuminate to deltate or acutely ob- long, to ca. 1 .5 mm long, ca. 1 .5 mm wide, glabrous or ciliate, margin glandular-fimbriate; pedicels 2-3 mm long, densely glandular-fimbriate at articulation; bracte- oles 1.5-1.8 mm below base of calyx tube, subopposite, imbricate, broadly clasping, semi-orbicular, ca. 1.5-2.5 mm long, 1 .8-2 mm wide, scarcely overlapping base of calyx tube, ciliate, margin glandular-fimbriate. Flowers 3-merous; calyx ca. 4-5 mm long, glabrous, tube terete, cylindric, ca. 2-3.5 mm long, 2-2.5 mm diam., limb spreading, smooth, ca. 1.5-2.5 mm long, lobes erect, broadly deltate, ca. 0.5-1.3 mm long, apex acute, ecilia- te, sinuses concave; corolla carnose, bistratose, cylindric but abruptly tapering at base, 5.6-9 mm long, 2.5-4 mm diam., deep red, glabrous but sometimes bearing few, ascendent, glandular fimbriae to 0.1-0.2 mm long in basal half, lobes erect, oblong-deltoid, 1.5-2 mm long, 1-1.2 mm wide, apex acute; stamens 6, ca. 7 mm long, shorter than corolla; filaments distinct, ca. 2.5-3.5 mm long, glabrous; connectives sometimes sparsely pilose along margin; anthers ca. 4.8-5.3 mm long, thecae 2.3- 2.8 mm long, tubules 2, distinct, 2.5-2.8 mm long, de- hiscing by introrse, elongate, oval slits ca. 1 .5 mm long; style equaling corolla or slightly exserted. Berry at first greenish-white turning dark blue (in two collections re- portedly translucent white). Disterigma trimerum is endemic to the premon- tane, wet montane, and Continental Divide forest in eastern Costa Rica and western Panama, (1 100) 1500-2240 m elevation. It has been collected only twice in Costa Rica (Limon Prov., Dtto. Bratsi, Cordillera de Talamanca, Caribbean slopes, Cuen- ca del Sixaola, area of Rfo Lari and Rio Coe"n, Bittner 1816 and Aguilar 1157). Flowering spec- imens have been collected in January-July; fruit- ing in January-February and May-July. Endan- gered. Disterigma trimerum is characterized by having usually obovate, serrulate leaf blades with round- ed and shortly retuse tips; broadly clasping, semi- orbicular bracteoles that are located below the apex of the pedicel and scarcely reach the base of the calyx tube; and 3-merous flowers with deep red corollas. The trimerous flowers are unknown anywhere else in Disterigma. Herbarium speci- mens indicate that mature fruit color varies from generally dark blue to rarely translucent white; this needs further investigation. Morphological and molecular studies show that D. trimerum is isolated from all other species in the genus. Disterigma utleyorum Wilbur & Luteyn in Lu- teyn & Wilbur, Brittonia 29: 259. 1977. Fig- ure 7. Epiphytic shruhs 20-45 cm tall; mature stems terete, glabrous; bark cracking irregularly and exfoliating in thin strips; twigs subterete to terete, moderately to densely puberulent. leaves congested, with petioles sub- terete, flattened adaxially, 1-2 (4) mm long, puberulent; leaf blades thick and fleshy when fresh but becoming thin and wrinkled when dry. oblong, obovate, somewhat spatulate, or rarely oblong-elliptic, slightly revolute, ( 1 ) 1.5-2.5 (3) cm long, (0.7) 1-1.8 cm wide, base broadly cuneate to obtuse, apex obtuse to broadly rounded, with a deciduous or persistent, weakly short-pilose apiculum 1-2 mm long, margin entire, both surfaces minutely pu- berulent but also with minute, glandular fimbriae ca. 0.1 mm long, 5 (7)-plinerved from base, midrib and lateral nerves conspicuously raised above and beneath, reticu- late veinlets slightly raised on both surfaces. Inflores- cences axillary, of solitary, subsessile flowers; floral bract apparently 1. ovate, ca. 0.5-1.5 mm long, apex rounded, puberulent, ciliate; pedicels subterete, striate, 0.5-2 mm long; braceoles subopposite, fused below into a cupule 0.5-1 mm long that loosely surrounds the basal part of the calyx tube, broadly oblong to ovate, 1 .5-2 mm long, 2-3 mm wide, minutely appressed puberulous. Flowers 4-merous; calyx ca. 4.5-5.5 mm long, tube cy- lindric, ribbed, 2-3 mm long, puberulous, limb campan- ulate, 1-3 mm long, lobes erect, lanceolate to narrowly triangular, 1.5-2.2 mm long, ca. 1 mm wide at base, apex acute to acuminate, minutely puberulous, sinuses acute; corolla thin-carnose, somewhat funnelform or cy- lindric-campanulate, 5.5-10 mm long, 3.5-9 mm apical diam., white, glabrous but sparingly glandular-fimbriate. lobes oblong-elliptic to broadly deltate, 3.5-5 mm long, ca. 3-4 mm wide, apex acute; stamens 8, ca. 5-6 mm long, about half as long as corolla; filaments 1.5-2.5 mm long, weakly ciliate; anthers 2.5-4.5 mm long, minutely papillate over entire thecae and apically onto base of tubules, thecae 1.5-2.5 mm long, basally shortly appen- diculate, tubule 1, ca. 1-2.3 mm long, dehiscing by a single, obliquely-terminal, circular pore; style 4.8-5.3 mm long. Berry spherical, ca. 7 mm diam., translucent white. Disterigma utleyorum is found in lowland rain- forest to premontane cloud forest, seemingly in the tops of large trees, (450) 700-1200 m eleva- tion. It is a rare species, and is sporadic in its distribution, having been collected twice from one locality in Costa Rica (vicinity of Cariblanco, He- redia Prov.), once in Panama (Prov. Veraguas), once in Colombia, and about six times from four localities in Ecuador. Flowering specimens have been collected in February-May, August, Octo- LUTEYN AND WILBUR: FAMILY #172 ERICACEAE 59 her, and November; fruiting in August and Octo- ber. Endangered. Disterigma utleyorum is easily identified and is unique in the genus in its combination of obovate to spatulate leaf blades that are basally cuneate and 5-plinerved, inflorescences with solitary, 4- merous flowers, and anthers with a single tubule. Its relationships within the genus are obscure. In Costa Rica and Ecuador (perhaps everywhere?), D. utlevorum always seems to be associated with colonies of biting ants; furthermore, the mature stems seem to radiate from a large, moist ball of earth and vegetable debris 30 cm or so in diameter capped by a dense, mat-forming acrocarpous moss. The details of this intriguing commensal re- lationship between ericad and insect are at present unknown. Gaultheria Linnaeus REFERENCE — J. L. Luteyn, Gaultheria. Fl. Neo- trop. Monogr. 66: 384-488. 1995. Erect shrubs (in ours), prostrate undershrubs, or rare- ly small trees; indumentum of simple, unicellular or multiseriate, multicellular eglandular or gland-tipped hairs. Leaves alternate, evergreen, petiolate, the blades usually coriaceous, the margin usually serrate or crenate, the venation pinnate. Inflorescences axillary, racemes or flowers solitary, these then multibracteate at base; ped- icels articulate with calyx and subtended by a single floral bract; bracteoles 2 to several, basal to apical. Flowers 5-merous, without odor; calyx synsepalous, lobed nearly to base, usually fleshy and accrescent after anthesis; corolla sympetalous, aestivation imbricate, ur- ceolate to campanulate, or sometimes cylindric-urceo- late, white to pinkish or reddish; stamens 10, ca. '/2-% corolla length; filaments distinct, glabrous or pubescent; connectives with disintegration tissue along connective and onto awns, lacking spurs; anthers bilocular, each loc- ule dehiscing introrsely by an apical pore and with 2, terminal, papillate awns, thecae papillate, tubules lack- ing; pollen grains in tetrahedral tetrads, lacking viscin threads; ovary superior, 5-locular; ovules 5-10; style about as long as corolla. Fruit a 5-valved, loculicidal capsule surrounded by the usually white or dark blue- black, fleshy, accrescent calyx; seeds numerous, ca. 1- 1.5 mm long, wingless. Gaultheria consists of ca. 115 species distrib- uted in a circum-Pacific ring from Japan and Chi- na; south through Malesia, southern Australia, and New Zealand; and then from the southern tip of South America north through the Andes, Me- soamerica, the West Indies, and to the U.S.A. and Canada. There are two species in Costa Rica. Gaultheria is characterized by a superior ovary and capsular fruit. At maturity the capsule is com- pletely surrounded by a fleshy, accrescent calyx; thus, the dispersal unit appears berry-like. The ge- nus is traditionally distinguished from Pernettya by its capsular fruit (vs. a true berry in Pernettya). Furthermore, the calyx of Pernettya normally re- mains dry and basal to the berry. Although the tips of the calyx lobes of Pernettya sometimes become fleshy, the calyx always remains small, distinct, and located at the base of the berry, never accrescent and surrounding it (Luteyn, 1995). Several recent authors have chosen to unite Per- nettya under Gaultheria, reasoning that in all oth- er morphological characters (save the calyx and fruit features), Pernettya is not distinct from Gaultheria (Stevens, 1971; Middleton & Wilcock, 1990; Kron et al., 2002a; Stevens et al., 2004). Molecular data are somewhat inconclusive, and too few taxa have been examined; however, the studies of Powell and Kron (2001) and Kron et al. (2002a) do support the inclusion of Pernettya within Gaultheria s.l. In this treatment of the Cos- ta Rican Ericaceae, we maintain the genera as dis- tinct. Key to the Species of Gaultheria la. Branchlets conspicuously strigose with straight, rigid, appressed (more or less spreading) hairs; leaf blades appressed-strigose beneath, elliptic to ovate-elliptic; indumentum eglandular . . G. gracilis Ib. Branchlets glabrous to densely white puberulent and also moderately to densely hirsute; leaf blades glabrous to moderately hirsute beneath, elliptic to subrotund; indumentum eglandular to glandular . G. erecta Gaultheria erecta Vent., Descr. PI. Nouv. 5, pi. 5. 1800. G. odorata Bredem. ex Willd., Ges. Naturf. Freunde Berlin Neue Schriften 3: 425. 1801. Brossea erecta (Vent.) Kuntze, Rev. Gen. PI. 2: 388. 1891. G. odorata Kunth in H.B.K., Nov. Gen. Sp. 3: 285. 1819, non Bredem. ex Willd. (1801). Brossea odorata (Kunth in H.B.K.) Kuntze, Rev. Gen. PI. 2: 388. 1891. G. 60 FIELDIANA: BOTANY odorala Kunth in H.B.K. var. costaricensis, J. D. Smith, Bot. Gaz. 20: 292. 1895. C. costari- censis (J. D. Smith) Small, N. Amer. Fl. 29: 79. 1914. G. glandulifera Small, N. Amer. Fl. 29: 78. 1914. G. donnellii Sleumer, Notizbl. Bot. Gart. Berlin-Dahlem 12: 125. 1934. G. poasana Sleumer, Notizbl. Bot. Gart. Berlin-Dahlem 12: 127. 1934. G. subrotunda Sleumer, Notizbl. Bot. Gart. Berlin-Dahlem 12: 284. 1935. Fig- ures IE and 8. Terrestrial, erect to arching or spreading, rhizomatous, shrubs to 3 m tall, often epiphytic in Costa Rica; mature stems sometimes glaucous, glabrous to sparsely white puberulent, also often persistently hispid-hirsute with eglandular or gland-tipped hairs. Leaves with petioles 1-8 mm long, puberulent (glabrate) and often hirsute with eglandular or gland-tipped hairs; leaf blades flat to sometimes bullate, sometimes clasping, ovate, elliptic, to subrotund, (2) 5-11 cm long, (1) 2.5-6.5 cm wide, base rounded to broadly cuneate, often subcordate to deeply cordate, apex acute or short-acuminate, glabrous to puberulent along veins above, also sparsely or mod- erately hirsute on lamina or veins with often gland- tipped hairs, sparsely to densely hirsute beneath with eglandular or minutely gland-tipped hairs, both surfaces often glabrate, margin remotely and shallowly crenate- serrate, pinnately veined. Inflorescences axillary, race- mose, 10-26-flowered; rachis 2-12 cm long, moderately to densely puberulent, also moderately to densely hirsute with eglandular or gland-tipped hairs; floral bract con- cave, cochleariform, ovate to ovate-elliptic or obovate, (3.7) 8-16 mm long, (1.6) 3-6 mm wide, apex acute to acuminate, sparsely puberulent, margin sparsely ciliolate and often glandular-pubescent, white to pale greenish or sometimes tinted with rose; pedicels (3) 8-13 mm long, pubescent as rachis; bracteoles basal or medial, narrowly elliptic-ovate to linear-ovate, (1.5) 5-7 mm long, (0.5) 1-1.5 mm wide, apex long-acuminate, weakly puberu- lent, margin ciliolate, often bearing gland-tipped hairs. Flowers with calyx (3) 4-6 (9) mm long, glabrous to densely puberulent, sometimes glabrate, also often mod- erately to densely hirsute with eglandular or glandular setae, lobes (1.5) 3-5.5 (7) mm long, erect to sometimes reflexed, triangular to ovate, acute to long-acuminate, densely ciliolate; corolla ovate-urceolate to cylindric-ur- ceolate, (4.5) 6-7 (8) mm long, 3-4.6 mm diam., gla- brous to sparsely or densely short-pilose, also often sparsely to densely hirsute to strigose with eglandular or gland-tipped hairs, white to red; stamens (2.7) 5-6 mm long; filaments (2.1) 3.5-5 mm long, pilose; anthers 1.2- 2.8 mm long, awns short and sometimes inconspicuous; ovary glabrous to densely short-pilose. Fruiting 7-12 (15) mm diam., glabrous to sparsely pilose, sometimes also sparsely hirsute with gland-tipped hairs, blue-black. Gaultheria erecta is found in open or shrubby areas such as road banks, landslides, or meadows; rocky places; montane and Quercus forest; edges of cloud and elfin forest; paramo and subparamo thickets; and rarely in boggy areas, (1200) 2000- 3200 (3700) m elevation (in Costa Rica). It has the broadest geographical range of any species in the genus, being distributed from north-central Mexico through Central America into the Andes of South America to northern Argentina and also rarely in southeastern Brazil. In Costa Rica it is commonly found as an epiphyte (less frequently terrestrial), whereas in the rest of its range it is always found as a terrestrial shrub. Flowering and fruiting occur throughout the year. Common. In Costa Rica, Gaultherici erecta is character- ized by its usually epiphytic habit, large leaves with blades that are oblong-elliptic and nitid, su- perior ovary, and capsular fruit that is surrounded by fleshy, accrescent calyx lobes. The relationships of Gaultheria erecta are un- certain, but it is morphologically similar to several South American species, including G. rigida Kunth, G. santanderensis A. C. Smith, G. hapal- otricha A. C. Smith, and G. setulosa N. E. Brown. Within Central America, it is morphologically most similar to the Panamanian endemic G. chi- riquensis Camp, a species that is not expected in Costa Rica. From the only other species of Gaul- theria in Costa Rica and adjacent western Pana- ma, G. gracilis, with which it may be remotely related, it is easily distinguished by the characters mentioned in the key. In South America, Gaul- theria erecta apparently hybridizes with Pernettya prostrata and several other species of Gaultheria (see Luteyn, 1995), but thus far there is no evi- dence for hybridization in Costa Rica. Fruits are eaten by birds. Gaultheria gracilis Small, N. Amer. Fl. 29: 77. 1914. C. barbata Small, N. Amer. Fl. 29: 77. 1914. G. setosa Small, N. Amer. Fl. 29: 77. 1914. G. gracilis Small var. intermedia Sues- senguth & Goeppinger, Bot. Jahrb. Syst. 72: 282. 1942, nom. nud. Illustration: R. L. Wilbur & J. L. Luteyn, Ann. Missouri Bot. Gard. 65: 86, fig. 8. 1978. Figure 8. Terrestrial or sometimes epiphytic shrubs 0.4-2 m tall; mature stems glabrous, but twigs moderately to densely appressed-strigose with eglandular hairs, often glabrate. Leaves with petioles 4-7 mm long, strongly canaliculate adaxially, short-puberulent and glabrate, of- ten also strigose; leaf blades flat or somewhat bullate, elliptic to ovate-elliptic, (3) 4-6 (8.5] cm long, ( 1 ) 1 .5- 2.5 (3.5) cm wide, base acute or cuneate to obtuse, apex long-acuminate to acute, sparsely to densely appressed strigose above with deciduous hairs, conspicuously and usually densely appressed strigose beneath, margin mi- nutely serrulate with each tooth terminating in an ap- pressed bristle, pinnately nerved. Inflorescences race- mose, 6-9 (14)-flowered; rachis (2) 3-7 (8) cm long. LUTEYN AND WILBUR: FAMILY #172 ERICACEAE 61 moderately to densely white-short-pilose or densely stri- gose with usually eglandular hairs; floral bract appar- ently continuous with pedicel, conspicuously striate, cu- cullate, strongly divergent, elliptic to spatulate, 3-10 (14) mm long, ciliate, puberulous or glabrous; pedicels 5-10 (12) mm long, moderately to densely white-short- pilose; bracteoles apparently continuous with pedicel, medial, linear to narrowly oblong, 2-4.5 mm long, cil- iolate and sometimes densely short-pilose. Flowers with calyx 3.5-5 mm long, glabrous or rarely densely stri- gose, lobes 2-3 mm long, ovate to deltoid, acute to acu- minate, ciliate; corolla urceolate to cylindric-urceolate, 4-8 mm long, 3-4.3 mm diam., glabrous or moderately to densely strigose, rarely strigose with eglandular or minutely gland-tipped hairs, greenish or white to rose; stamens 2.3-3 mm long; filaments 1.5-2.2 mm long, glabrous or weakly pilose; anthers 1-1.3 mm long, awns prominent; ovary glabrous or weakly short-pilose. Fruiting to 9 mm diam., glabrous, blue-black. Gaultheria gracilis is found in cloud and elfin forests, high-elevation bogs, disturbed forest, and remnant trees in pasturelands, (1100) 1400-2700 (3150) m elevation. It is endemic to Costa Rica and adjacent western Panama. Flowering speci- mens have been collected in January, February, June, July, and October-December; fruiting in February, June-August, and December. Wide- spread. Gaultheria gracilis is characterized by having strigose branches, long-acuminate leaf blades, strongly canaliculate petioles, densely white pi- lose rachises and pedicels, and linear and medially located bracteoles. The plants are said to be small trees in boggy areas, but no heights have been given on herbarium labels. There is some varia- tion in its indumentum. The branches, which are normally appressed strigose, may have spreading hairs (= G. barbata), or the corolla may be gla- brous (= G. setosa) to densely strigose, but these variations are matters of degree and may often occur within populations. Gaultheria gracilis is morphologically most similar to G. strigosa Ben- tham and G. insipida Bentham, both from South America. There is a somewhat remote relationship with G. erecta, also found in Costa Rica, as in- dicated by a similarity in the shape and texture of the floral bracts. Gonocalyx Planch. & Lindl. REFERENCES — J. L. Luteyn, Notes on Neotrop- ical Vaccinieae (Ericaceae). I. Gonocalyx — a ge- nus new to Central America. Brittonia 28: 37-41. 1976; J. L. Luteyn, Gonocalyx amplexicaulis (Er- icaceae): A new Panamanian blueberry. Syst. Bot. 15: 745-747. 1990. Epiphytic shrubs, rarely small terrestrial trees or herbs. Leaves alternate, sessile to petiolate, the blade coriaceous, flat to revolute, with margin entire or cre- nate, the venation obscurely plinerved. Inflorescences axillary, flowers solitary, in pairs, or in loose, few-flow- ered racemes; floral bract small; pedicels articulate with calyx; bracteoles 2, usually basal. Flowers (4) 5-merous; calyx synsepalous, tube terete, bluntly angled, or nar- rowly winged opposite the sinuses, lobes apiculate; co- rolla sympetalous, aestivation valvate, membranous to thinly carnose, usually bistratose, tubular, subcylindric, cylindric-campanulate, or globose-urceolate; stamens 10, equal (rarely apparently alternately unequal), nearly as long as the corolla; filaments distinct or connate; con- nectives lacking disintegration tissue or spurs; anthers lacking awns, thecae minutely papillate, tubules 2, dis- tinct, longer and thinner than thecae, dehiscing by in- trorse, minute, subterminal, oblique pores or short slits; pollen grains in tetrahedral tetrads, lacking viscin threads; ovary inferior; style filiform, included or ex- serted, glabrous; stigma truncate or punctiform. Fruit a berry; seeds numerous. Gonocalyx is a small genus of 1 1 species dis- tributed in Costa Rica, Panama, Colombia, and the West Indies. It is characterized by having ped- icels articulate with the calyx; tubular or cylindric corollas; isomorphic stamens with long, thin tu- bules; and dehiscing by nearly terminal, oblique pores or short slits. Its generic relationships are still uncertain, although Luteyn (1976b) suggested a relationship to Siphonandra Klotzsch. Five spe- cies are currently known to occur in Costa Rica. Key to the Species of Gonocalyx la. Flowers solitary or in pairs; anthers 1 1-19 mm long 2 Ib. Flowers in racemes; anthers 7-1 1 mm long 3 2a. Staminal filaments connate; leaf blades ovate to pyriform; pedicels 1-3 (10) mm long; calyx 5-8 (10) mm long G. almedae 2b. Staminal filaments distinct; leaf blades elliptic to obovate; pedicels 5-9 (13) mm long; calyx 3-4.5 mm long G. pterocarpus 62 FIELDIANA: BOTANY 3a. Rachis 3-7 cm long; bracteoles large, 13-16 mm long, apical; calyx tube conspicuously ribbed at base or narrowly winged; calyx lobes 3-5 mm long G. megabracteolatus 3b. Rachis 0.9-1.8 cm long; bracteoles small, 1-2 mm long, basal; calyx tube terete or bluntly pentag- onal, or base slightly expanded as a rim; calyx lobes 0.3-2 mm long 4 4a. Leaves sessile, amplexicaul G. amplexicaulis 4b. Leaves petiolate, not amplexicaul 5 5a. Mature stems, rachis, pedicels, calyx, corollas and fruits glabrous; floral bract ca. 1 mm long; staminal filaments distinct; plants epiphytes G. costaricensis 5b. Mature stems, rachis, pedicels, calyx, corolla and fruits short-pilose; floral bract 5-20 mm long; staminal filaments connate; plants terrestrial G. liliae Gonocalyx almedae Luteyn, Brittonia 28: 403, fig. 1. 1976. Figure 3. Epiphytic shrubs 1-3 m tall; mature stems coarsely and bluntly angled, glabrous, reddish-brown; bark thin, cracking longitudinally; twigs terete to subterete. puber- ulent, grayish- to reddish-brown; axillary buds awl- shaped, 1 .5-3 mm long, appearing stipular. Leaves with petioles subterete, 1.5-2.5 mm long, puberulent adaxi- ally; leaf blades coriaceous, ovate, lanceolate to pyri- form, 1-3 cm long, 0.5-1 cm wide, base rounded or obtuse, apex obtusely acuminate, margin crenate with each tooth gland tipped, glabrous, obscurely 3-5-pliner- ved, only midrib prominent. Inflorescences axillary, flowers solitary; floral bract ovate, 2-2.5 mm long, apex acuminate, margin glandular-fimbriate; pedicels 1-3 mm long (in one case 10 mm), glabrous or puberulent; brac- teoles basal, concealing pedicel, ovate, 1-2.5 mm long, apex acuminate, margin glandular-fimbriate. Flowers 5- merous with calyx puberulent or glabrous, 5-8 (10) mm long, 5-winged, tube 2.5-5 mm long, limb erect-patent, sometimes striate, 2.5-4 mm long, lobes long-triangular, acute, 1.5-2 mm long, sinuses rounded to broadly ob- tuse; corolla thin-carnose when fresh, bistratose, cylin- dric, 21-26 mm long, 2.5-4.5 mm diam., glabrous or puberulent externally but pilose at throat and lobes in- ternally, also bearing scattered glandular fimbriae, bright red, lobes triangular 1-2 mm long; stamens 22-24 mm long, sometimes apparently alternately unequal; fila- ments connate into a tube 5-6.5 mm long, glabrous; an- thers 18-19 mm long, thecae 3.5-4 mm long, tubules ca. 13-15 mm long, dehiscing by oblique pores ca. 0.5 mm long; style exserted, 24-27 mm long. Berry not seen. Gonocalyx almedae is found in premontane cloud forest, 1000-1600 m elevation. It is distrib- uted in Costa Rica (Cordillera de Talamanca) and western Panama (Chiriquf Prov.) and is infre- quently collected (ca. six collections). Flowering specimens have been collected in March, October, and December; fruits are unknown. Endangered. Gonocalyx almedae is characterized by having ovate, lanceolate, to pear-shaped leaf blades; sol- itary, pentamerous flowers; tiny bracteoles; and connate staminal filaments. It is reminiscent of and probably most closely related to G. pterocar- pus but is distinguished from the latter by its leaf shape; shorter pedicels; longer, puberulent calyx and corolla; connate filaments; and poroid anther dehiscence. Gonocalyx amplexicaulis Luteyn, Syst. Bot. 15: 745, fig. 1. 1990. Epiphytic shrubs, somewhat scandent; mature stems terete, weakly striate, glabrous, nitid; twigs terete, gla- brous. Leaves sessile, strongly amplexicaul; leaf blades subcoriaceous, broadly ovate to subrotund, 7-8.5 cm long, 4.6-6.5 cm wide, base rounded, strongly and deep- ly cordate with lobes overlapping, apex sharply acumi- nate, margin thin, scarious and minutely toothed in api- cal half, glabrous but both surfaces bearing minute, de- ciduous, glandular fimbriae ca. 0.3 mm long, plinerved with 11-13 nerves arising from near base, arcuate and anastomosing near margin, midrib and lateral nerves plane to slightly raised on both surfaces, reticulate vein- lets obscure and ± plane. Inflorescences racemose, 3- 4-flowered; rachis subterete, weakly striate, 0.9-1.7 cm long, glabrous; floral bract persistent, ovate, 1-1.8 mm long, apex acuminate, margin glandular-fimbriate; ped- icels subterete, striate becoming ribbed apical ly, 16-20 mm long, glabrous but bearing scattered, glandular fim- briae ca. 0.3 mm long apically; bracteoles persistent, nearly basal, ovate, 1-1.2 mm long, acuminate, margin glandular-fimbriate. Flowers (not fully mature) 5-mer- ous; calyx 4.5-5 mm long, glabrous, tube cylindric-cam- panulate. strongly hut bluntly angled opposite sinuses, 2.5-3 mm long, limb campanulate. 2-2.3 mm long, lobes bluntly apiculate, 0.3-0.5 mm long, sinuses con- cave; corolla carnose, bistratose, apparently globose-ur- ceolate, ca. 1 1 mm long, glabrous but bearing scattered, glandular fimbriae throughout, orange (fide label), lobes bluntly triangular or deltate, ca. 1 .5 mm long; stamens equal, ca. 8-9.5 mm long; filaments ca. 1.4 mm long, glabrous; anthers 8-9 mm long, thecae ca. 3 mm long, incurved at base, tubules ca. 6 mm long, dehiscing by tiny, oblique, terminal pores; style included, ca. 5 mm long. Berry not seen. Gonocalyx amplexicaulis is krtown only from the type collection, from Continental Divide pre- montane cloud forest at the Fortuna Dam site, Chiriquf Province, Panama, 1200-1400 m eleva- tion. It has not yet been collected in Costa Rica. The type collection was flowering in June, but the LUTEYN AND WILBUR: FAMILY #172 ERICACEAE 63 flowers are immature; fruits are unknown. En- dangered. Gonocalyx amplexicaulis is a very distinct spe- cies easily recognizable by its strongly amplexi- caul leaves with sharply acuminate blade apices. Its lianoid habit, leaf size, calyx shape, corolla size, and type of anther dehiscence are similar to those of G. smilacifolius (Grisebach) A. C. Smith. That species is limited to the Caribbean islands of Dominica and Guadeloupe, however, and further- more has petiolate, ovate to ovate-lanceolate leaf blades, longer pedicels, more flowers per inflores- cence, and smaller corollas. Nevertheless, mor- phologically, G. amplexicaulis and G. smilacifol- ius are similar and may be each other's closest relatives. Additional material is needed, especially with mature flowers. Gonocalyx costaricensis Luteyn, Brittonia 28: 40, fig. 2. 1976. Figure 4. Epiphytic shrubs, without lignotuber; mature stems terete, striate, glabrous, reddish; bark thin, cracking in a reticulate pattern, reddish; twigs terete to subterete, often flattened with blunt angles, smooth to minutely striate, glabrous, reddish-brown. Leaves with petioles subterete, flattened adaxially, rugose, ca. 2 mm long and 1 mm diam., glabrous or puberulent; leaf blades coriaceous, ovate-lanceolate or elliptic-lanceolate, 2.5-6 cm long, 1- 1.5 cm wide, base rounded or obtuse, apex long-acu- minate, tip ultimately blunt, glabrous, obscurely 3 (5)- plinerved, midrib and lateral nerves plane to slightly raised on both surfaces, reticulate veinlets raised slightly on both sides but conspicuous only beneath. Inflores- cences axillary, solitary, racemose, 8-16-flowered; ra- chis subterete, smooth, glabrous, 1-1.8 cm long; floral bract persistent, linear-lanceolate or aristate, ca. 1 mm long, ca. 0.5 mm wide, glandular-fimbriate at base, cil- iate apically; pedicels gradually thickening apically, stri- ate, glabrous, 4-10 mm long and 0.5-1 mm diam., sometimes with cartilaginous teeth apically; bracteoles basal, aristate, ca. 1 mm long, ciliate apically. Flowers with calyx glabrous, 6-9.5 mm long, tube cylindric or obconic and spreading apically, slightly expanded as a rim at base but not conspicuously apophysate, conspic- uously striate, 1.5-4.5 mm long, limb cylindric-spread- ing, conspicuously striate, 4.5-5 mm long, lobes broadly triangular, acute, striate, 1-2 mm long, sinuses obtuse; corolla thinly carnose, bistratose, cylindric, 12-18 mm long, 5-7 mm diam., glabrous, red, lobes triangular, 1.2- 1.5 mm long, obtuse to acute; stamens equal, 11-13.5 mm long; filaments distinct at anthesis, marginally pi- lose, 2-4.3 mm long; anthers 9.5-14 mm long, thecae 2-3.5 mm long, the base short-apiculate, tubules ca. 7- 1 1 mm long, dehiscing by oval, subterminal pores, 0.5- 1.6 mm long; style strongly exserted, 14-19 mm long. Berry (only one seen) spherical, to 10 mm diam., striate. Gonocalyx costaricensis is endemic to the low- er montane, wet forest region around Monteverde, Costa Rica (Cordillera de Tilaran), 1100-1600 m elevation. It is known only from that area and from only 20 collections. Flowering specimens have been collected in February-April, June, and October-December; fruiting in May. Endan- gered. Gonocalyx costaricensis is characterized by its flowers in racemes; tiny, aristate, basal bracteoles; terete and conspicuously striate calyx; red corolla, which is 12-18 mm long; and distinct staminal filaments. It is morphologically similar to G. liliae but is distinguished by glabrous or glabrate (not short-pilose) vegetative and floral parts, shorter floral bracts (1 mm vs. 5-20 mm long), and dis- tinct (not connate) staminal filaments. At Monte- verde, G. costaricensis may be easily confused with Vaccinium montever 'dense, which grows in the same general area and has a similar epiphytic habit, leaves, striate calyx limb, and rather broad corolla. Gonocalyx costaricensis, however, nor- mally has flat (not somewhat concave) leaves, to- tally red corollas that are strictly cylindric and much thinner in texture (vs. corollas greenish but suffused with red or purple, broadly cylindric - campanulate in shape and subcarnose), and sta- mens with thin, erect, elongated tubules (vs. tu- bules wide, spreading, and shorter with respect to thecae) (see Luteyn, 2001: 444). The species is visited by hummingbirds (pers. observ.). Gonocalyx liliae Al. Rodr. & J. F. Morales, No- von, in prep. (Hollowell, pers. comm.), fig. 1. 2004. Figure 5. Terrestrial herbs (fide label) 30-50 cm tall, erect or pendent; mature stems glabrate, lightly striate; twigs red- dish-brown to grayish-brown, short- pilose. Leaves al- ternate to rarely subopposite, with petioles 2-4 mm long; leaf blades coriaceous to subcoriaceous, ovate to narrowly ovate, slightly revolute, 1.9-4 cm long, 0.8-2 cm wide, base obtuse, apex acute, margin entire, sparse- ly pilose on both surfaces, the lower usually glandular fimbriate, obscurely plinerved. Inflorescences axillary, terminal or subterminal, solitary, racemose, 4-6-flow- ered; rachis subterete, slightly ribbed, pilose, 1-1.5 cm long; floral bract persistent, foliaceous, ovate, 5-20 mm long, apex acute; pedicels subterete, striate, 8-1 1 mm long, pilose; bracteoles deciduous, basal, linear, 0.4-0.8 mm long. Flowers with calyx ca. 10 mm long, short- pilose, tube campanulate, terete, 4-5 mm long, ca. 4 mm diam., limb campanulate, suberect, 4-5.5 mm long, lobes 1-1.2 mm long, deltate, acute; corolla campanu- late, tube ca. 12 mm long, ca. 3 mm diam., reddish to orangish-red, pilose, lobes deltate, acute, 2-2.5 mm long; stamens equal, slightly shorter than corolla; fila- ments connate, 2-2.5 mm long, sparsely pilose; anthers ca. 9.5 mm long, thecae incurved at base, ca. 2.3 mm long, tubules ca. 7 mm long, dehiscing by terminal, 64 FIELDIANA: BOTANY oblique and introrse pores; style ca. 6 mm long. Berry immature, subglobose, shortly pilose, greenish-red. Gonoclayx liliae is endemic to Costa Rica, where it is found in the pluvial oak forest of the Pacific slopes in the Cordillera de Talamanca, 1600-2000 m. The only known collections are San Jose: Canton de Acosta, Fila Bustamante, Ha- cienda Tiquires, cabeceras Rio Tiquires, J. F. Mo- rales 4279; Canton de Dota, Providencia, A. Rod- riguez 7261, and "Santa Rosa de Copey," Tonduz 12237. Flowering collections have been made in April-May; fruiting between May and June. En- dangered. Among the Costa Rican species of Gonocalyx, G. liliae is easily recognized by its conspicuous pubescence of its vegetative, floral, and fruiting parts; flowers in racemes; terete calyx tube; and connate staminal filaments. It shares similar veg- etative and inflorescence features with G. costar- icensis, but the latter has glabrous parts, incon- spicuous floral bracts, and distinct staminal fila- ments. [The above description and discussion are based on the protologue; the authors have not yet seen the type collection, and the Tonduz collec- tion lacks corollas and stamens.] Gonocalyx megabracteolatus (Wilbur & Luteyn) Luteyn, Brittonia 53: 437. 2001. Macleania me- gabracteolata Wilbur & Luteyn, Ann. Missouri Bot. Card. 68: 163. 1981. Epiphytic shrubs, without lignotuber; mature stems and twigs subterete, conspicuously and bluntly to nar- rowly ridged and grooved, striate, glabrous or nearly so, scattered glandular-fimbriate, the basal portion of each year's growth conspicuously surrounded by a series of persistent, tightly clasping, imbricate, ovate to ellitic, acute to rounded, striate, glabrous perulae 5-20 mm long. Leaves with petioles stout, rugose, 3-6 mm long, densely short-pilose, or puberulous to glabrate; leaf blades coriaceous, elliptic, slightly revolute, (5) 7-10 (12.5) cm long, (2.7) 3-4 (5.7) cm wide, base rounded or broadly tapering, apex acute or more typically acu- minate with the tip 1 cm or more long, margin entire, glabrous above or sparingly to moderately spreading short-pilose or puberulent along basal raised portion of midrib, glabrate beneath, younger leaves with reddish- brown appressed, glandular fimbriae 0.1-0.2 mm long that leave punctate depressions when deciduous, 5-7- plinerved with lateral veins arising at or near base, mid- rib much thickened and prominently raised in basal 1 cm then becoming impressed apically, raised beneath, lateral nerves slightly raised basally then impressed api- cally above and slightly raised beneath, reticulate vein- lets obscure above and slightly raised but obscure be- neath. Inflorescences axillary, borne near ends of branches, sessile, racemose, 4-7-flowered, all parts weakly to moderately reddish-brown appressed glandu- lar-fimbriate with fimbriae 0.1-0.2 mm long; rachis stri- ate or irregularly angled, 3-7 cm long, lower 1.5-2 cm enveloped by persistent, 'ovate to oblong bracts (inflo- rescence), 5-8 mm long; floral bract oblong to spatulate, 9-13 mm long, apex rounded, margin entire; pedicels striate, 5-11 mm long, apically broadened; bracteoles apical or up to 10 mm below apex, elliptic, striate, 13- 16 mm long, ca. 5 mm wide, apex acute. Flowers with the calyx 10-12.5 mm long, moderately to densely glan- dular-fimbriate with reddish-brown fimbriae, tube ± pentagonal, strongly ribbed at base or narrowly winged in lower half continuing apically onto limb, 4-6 mm long, limb striate, campanulate-spreading. 6-8 mm long, lobes striate, deltate, 3-5 mm long, apex acute, sinuses acute; corolla thinly carnose, unistratose. broadly cylin- dric-campanulate. terete, 12-18 mm long, 7-16 mm api- cal ili.nn . glabrous but scattered glandular-fimbriate, pale greenish but suffused with red on surfaces exposed to sun, lobes broadly triangular-deltoid, acute, 2-3 mm long; stamens 10, ca. 10-13.5 mm long; filaments dis- tinct, 3-5 mm long, spreading long-pilose with slender trichomes 0.2-0.4 mm long; connectives smooth; an- thers (7) 8-10 mm long, curving slightly inward, thecae 3-3.5 mm long, the base incurved and shortly and blunt- ly apiculate, tubules 2, (3.5) 5-6.5 mm long, dehiscing by apically oblique, oval clefts ca. 0.8-1 mm long; style 18-19 mm long, glabrous, ± sigmoid. Berry not seen. Gonocalyx megabracteolatus is found in pre- montane cloud forest, 1 100-2050 m elevation. It is endemic to western Panama (several collections from Chiriqui Prov.) and eastern Costa Rica (one collection. San Jos£: Cant6n de Coto Brus, Zona Protectora Las Tablas, Cuanca T6rraba-Sierpe, Sura, 7 km NW of Progreso, Navarro V. & Pi- cado 583). Flowering material has been collected in January, February, April, and October; fruits are unknown. Endangered. Gonocalyx megabrateolatus is characterized by having flowers in racemes, large bracteoles borne at the apex of the pedicel, broadly cylindric-cam- panulate corolla, and distinct staminal filaments. It was first described in the genus Macleania by Wilbur and Luteyn (1981), but its relationships were obscure at that time. After a detailed ex- amination of the stamens from all known material of this taxon, it was transferred to Gonocalyx, the morphological placement being reinforced by mo- lecular data (cf. Luteyn, 2002). That study showed the taxon fell into the same clade and is a sister species of G. costaricensis. We do not, however, feel that G. megabracteolatus is closely related to G. costaricensis at the species level and await a more detailed morphological study of Gonocalyx to decide its placement. Gonocalyx pterocarpus (J. D. Smith) Luteyn, Brittonia 28: 38, fig. 1. 1976. Themistoclesia pterocarpa J. D. Smith, Bot. Gaz. 44: 113. LUTEYN AND WILBUR: FAMILY #172 ERICACEAE 65 1907. Cavendishia pterocarpa (J. D. Smith) A. C. Smith, Contrib. U.S. Natl. Herb. 28: 449. 1932. Figures 2O, 3, and 7. Epiphytic shrubs or sometimes small, stunted trees in elfin forest with branches 3-6 m and bole to 20 cm diam.; mature stems terete to subterete, short puberulent or glabrate, striate, reddish- to blackish-brown; bark thin, cracking longitudinally twigs subterete, bluntly angled, densely puberulent. Leaves with petioles subterete, channeled adaxially, somewhat winged, 1-3 mm long, densely short puberulent; leaf blades coriaceous, ob- ovate, oblong-elliptic or elliptic, 1-2 (3) cm long, 0.5- 1 .5 cm wide, base cuneate and attentuate, apex rounded and slightly emarginate, margin entire or remotely cre- nate apically, scattered hairs on midrib adaxially, ob- scurely 5-plinerved, all nerves slightly raised on both surfaces. Inflorescences axillary, 1-2-flowered, sub- tended by 3-5, ovate, acute, ciliate and fimbriate mar- gined bracts, 0.7-1.5 mm long, 0.5-1 mm wide, not clearly differentiated into inflorescence bracts, floral bracts, or bracteoles; pedicels subterete to bluntly an- gled, slightly thickening and oblique apically, 5-9 (13) mm long, ca. 1 mm diam., glabrous, with few cartilag- inous teeth apically. Flowers with calyx glabrous, 3-4.5 mm long, obprismatic, 5-winged opposite sinuses, tube 1.5-2.5 mm long, limb campanulate, 1.5-2 mm long, lobes triangular, acute, ca. 0.5 mm long, sinuses rounded to flat; corolla thinly carnose when fresh and membra- nous, bistratose, cylindric, narrowed to throat, slightly pentagonal, (14) 16-20 (25) mm long, 3-6 mm diam., glabrous externally but with scattered, short, glandular- fimbriae, crimson when fresh, lobes triangular or oblong, to 3 mm long, apex obtuse, throat and lobes densely white pubescent internally; stamens equal, 14-19 mm long; filaments distinct, 3-5 mm long, glabrous, white; anthers 11-16.5 mm long, thecae 2.5-4 mm long, tu- bules ca. 8.2 mm long, dehiscing by lateral slits 3.5-4 mm long; style 19-23 mm long, sometimes exserted. Berry spherical, 8-10 mm diam., dark blue-black. Gonocalyx pterocarpus is found in premontane cloud forest, 1000-2400 m elevation. It is distrib- uted in Costa Rica (Cordillera Tilaran and Cor- dillera Central) and western Panama (Chiriquf Prov.). Flowering occurs throughout year; fruiting collections have been made in January, April, June, July, November, and December. Rare/In- frequent. Gonocalyx pterocarpus is characterized by its 1-2-flowered inflorescences, obprismatic calyx tube that is narrowly 5-winged to the sinuses, dis- tinct staminal filaments, anther tubules 3-4 times longer than the thecae and dehiscing by lateral slits, and internally densely short-pubescent co- rolla lobes. It is morphologically similar to and can superficially be confused with G. almedae. Gonocalyx pterocarpus differs by having longer pedicels that are glabrous (5-9 [13] mm vs. nor- mally 1-3 mm long, glabrous or puberulent), a shorter calyx (3-4.5 mm not 5-8 [10] mm long), and distinct staminal filaments (not connate). The species is visited by the hummingbird Lampornis calolaema in Costa Rica (pers. observ.; Luteyn, 1998). Lateropora A. C. Smith REFERENCES — R. L. Wilbur & J. L. Luteyn, Ad- ditions to the Ericaceae of Panama. Ann. Missouri Bot. Card. 68: 162-163. 1981. J. L. Luteyn, New species, new records, and neotypification of some Mesoamerican Ericaceae. Brittonia 48: 241-249. 1996 [Lateropora, pp. 246-247]. Terrestrial or epiphytic shrubs. Leaves alternate, pet- iolate, the blade coriaceous, the margin entire, the ve- nation plinerved. Inflorescences axillary, compact, um- belliform or racemose; floral bract persistent; pedicels articulate with calyx, short, stout; bracteoles 2, basal, subopposite, persistent. Flowers 5-merous, without odor; calyx synsepalous, tube campanulate, lobes erect; corolla sympetalous, aestivation valvate, thick-carnose, bistratose, broadly urceolate or shortly campanulate, densely pilose internally; stamens 10, equal, nearly as long as the corolla; filaments distinct, weakly adnate to base of corolla; connectives lacking disintegration tissue or spurs; anthers lacking awns, thecae strongly papillate, tubules 2, distinct, short (vestigial), dehiscing laterally by clefts extending entire length of anther; pollen grains in tetrahedral tetrads, lacking viscin threads; ovary in- ferior; style filiform, glabrous. Fruit a berry; seeds nu- merous. Lateropora consists of three species distributed from eastern Costa Rica to central Panama. The genus is characterized by succulent, dark green corollas and anthers with vestigial to short tubules dehiscing laterally along the entire length of the anther. The genus is unique in its type of anther dehiscence and is in a rather isolated position within the Vaccinieae, although its overall rela- tionships may lie with the green-flowered species of Vaccinium (e.g., V. poasanum, V. costaricense, etc.). One species is currently known to occur in Costa Rica. 66 FIELDIANA: BOTANY Key to the Species of Lateropora la. Calyx lobes 2-3 mm long, 4-5 mm wide basally; corolla lobes ca. 3 mm long; petioles 5-15 mm long; leaves 6-13 cm long, 4-9 cm wide L ovata Ib. Calyx lobes 1-1.5 mm long, 1.8-2.2 mm wide basally; corolla lobes 1-2 mm long; petioles 3-4 mm long; leaves 2.5-3.8 cm long, 1.2-1.8 cm wide L. tubulifera Lateropora ovata A. C. Smith, Contrib. U.S. Natl. Herb. 28: 334. 1932. Symphysia ovata (A. C. Smith) vander Kloet, Taxon 53: 97. 2004. Illustration: A. C. Smith, Contrib. U.S. Natl. Herb. 28: pi. 1. 1932. R. L. Wilbur & J. L. Luteyn, Ann. Missouri Hot. Card. 65: 91, fig. 9. 1978. Figures 2J and 10. Epiphytic or terrestrial shrubs 2-3 m tall; mature stems and twigs subterete, glabrous, striate, thick and widely divergent. Leaves with petioles rugose, 5-15 mm long, 1.5-3 mm ilium., narrowly winged apically, gla- brous; leaf blades thick-coriaceous, ovate to broadly el- liptic, sometimes nearly orbicular, 6-13 cm long, 4-6 (9) cm wide, base cuneate, apex acute to broadly obtuse, bluntly short-acuminate or nearly rounded, glabrate but with appressed, reddish-brown, glandular-nmbriae 0.2- 0.6 mm long, 3-5 (7)-plinerved with inner pair of lateral veins arcuate, arising above base and sometimes subop- posite, midrib thickened and raised in basal V3-V4 then apically impressed above, raised and conspicuous be- neath, inner 1-2 pairs of lateral nerves impressed above and raised beneath, usually conspicuous but sometimes inconspicuous beneath, the outer pair of lateral nerves sometimes raised above but always obscure, reticulate veinlets slightly elevated but often obscure on both sur- faces. Inflorescences axillary, compact umbelliform ra- cemes; rachis stout, 12-16 mm long, glabrous, bracteate at base; floral bract thick, scale-like, broadly ovate, 4-5 mm long, 3.5-4.5 mm wide, apex rounded, margin cil- iate and glandular-fimbriate; pedicels 3-10 mm long, ca. 2 mm in diam., glandular-fimbriate with trichomes 0.1- 0.2 mm long at apex; bracteoles 2, basal to submedial, subopposite, keeled, broadly ovate, 3-4 mm long, 4-5 mm wide, apex rounded, ciliate. Flowers with calyx ca. 6 mm long, sparingly appressed glandular-fimbriate, tube cylindric-campanulate, usually pentagonal and bluntly 5-ribbed opposite sinuses, 3.5-5 mm long, ca. 4 mm diam., limb thick-walled, campanulate, 1-2.5 mm long, glabrous, lobes broadly triangular, acute, 2-3 mm long, 4-5 mm wide at base, erect or incurved, short- ciliate, sinuses concave at anthesis or sometimes tearing and then acute; corolla broadly urceolate or subglobose, 6-8 mm long, 7-8 mm diam., pale greenish-white, sparsely pubescent externally near middle with ap- pressed trichomes 0.4-0.6 mm long, lobes triangular, ca. 3 mm long, 4.5-5 mm wide at base, acute, spreading to reflexed, densely tangled villous internally with white trichomes 0.4-0.7 mm long; stamens 10, ca. 6.7-7 mm long; filaments 3-4 mm long, broadened at base to ca. 1 mm wide, tapering gradually upward to ca. 0.6 mm, glabrous in basal '/„ strikingly pilose apically especially near connective with trichomes 0.5-1 mm long; anthers 3-5 mm long, thecae ca. 2.7-4 mm long, strongly in- curved at base with lower '/, protruding inwardly or even turning upward, tubules submembranous, ca. 0.3-0.7 mm long; style stout, ca. 7 mm long. Berry not seen. Lateropora ovata is found in premontane cloud forest, 1100-2200 m elevation. This little-known species has been collected only from western Chi- riquf Province, Panama (nine collections) and once in the eastern Cordillera de Talamanca, Cos- ta Rica (Lim6n Prov., Cant6n Talamanca, Dtto. Bratsi, Reserva Indfgena Bri Bri, 1.7 km SW of Kivut and 6. 1 km SW of Alto Lari, Schmidt 642). Flowering specimens have been collected in Jan- uary, February, and March; fruits are unknown. Endangered. Lateropora ovata is characterized by its coria- ceous, large, ovate to broadly elliptic, basally cu- neate leaves and compact umbelliform racemes. It may be distinguished from the other two species in the genus by its combination of larger and coarser leaves with longer petioles and longer ca- lyx and corolla lobes (a key to the three species is provided in Wilbur & Luteyn, 1981). More col- lections are needed to understand the range of var- iation in this and all species of Lateropora and to determine their interspecific relationships. Lateropora tubulifera Wilbur & Luteyn, Ann. Missouri Bot. Card. 68: 162. 1981. Symphysia tubulifera (Wilbur & Luteyn) vander Kloet, Taxon 53: 97. 2004. Shrubs 1 .5 m tall, with twigs moderately thick, striate to irregularly angulate, 2-2.5 mm diam., glabrous. Leaves with petioles flattened adaxially, 3-4 mm long, 2-2.5 mm wide, glabrous; leaf blades coriaceous, ellip- tic, 2.5-3.8 cm long, 1.2-1.8 cm wide, base acutely ta- pering, apex acute to obtuse, margin slightly thickened, glabrous or glabrate beneath, 5-plinerved with inner pair of lateral nerves arising from basal 2 cm, midrib con- spicuously thickened and raised in proximal 1-2 cm then impressed apically above and conspicuously raised be- neath, lateral nerves impressed completely or only ba- sally above and conspicuously raised beneath, reticulate nerves slightly impressed, plane or slightly raised above, sometimes obscure, raised beneath. Inflorescences cor- ymbosely racemose, (1) 3-5-flowered, 2-3 cm long overall, circumscribed at base by 3-4, broadly oblong to orbicular, short-ciliate bracts, 1-3 mm long, ca. 1.5- 2.2 mm wide; rachis 1-1.5 cm long, glabrous; floral LUTEYN AND WILBUR: FAMILY #172 ERICACEAE 67 bract ovate to oblong, 1 .5-2.5 mm long, 1 .5-2 mm wide, slightly keeled and short-apiculate, irregularly narrowly hyaline margined, glabrous, short-ciliate; pedicels slight- ly clavate, 10-18 mm long, medially ca. 1 mm diam., glabrous except for inconspicuous fringe of glandular fimbriae borne apically at articulation with trichomes 0.2-0.4 mm long; bracteoles located along basal '/3 of pedicel, appressed, subopposite to alternate, ovate to ob- long, 2-2.6 mm long, 1.5-1.8 mm wide, ciliate, irregu- larly hyaline margined. Flowers with calyx 3-6 mm long. 3-3.2 mm diam., glabrous, tube campanulate, ca. 2.5 mm long, bluntly 10-ribbed in lower V2, limb some- what flaring or at least wider than tube, ca. 2.5-3.5 mm long, lobes erect, deltate, 1-1.5 mm long, 1.8-2.2 mm wide at base, apex acute; corolla urceolate, 3.5-7 mm long, to 7 mm diam., externally glabrous but internally densely pilose to tangled villous in apical [/2 with white trichomes 0.6-1 mm long, pink (fide label), lobes tri- angular, 1-2 mm long, acute, glabrous externally and densely tangled villous internally; stamens ca. 5.2 mm long; filaments 2-2.5 mm long, <0.5 mm wide at base, densely pilose apically with white trichomes 0.5-1.2 mm long; anthers conspicuously granular, 2.5-4.2 mm long, thecae strongly incurved in basal l/4-V3, tubules 0.8-1.2 mm long. Berry not seen. Lateropora tubulifera is known from only three collections, from short, windswept, montane cloud forest in the Fortuna Dam region, Chiriqui Province, Panama, 1800-2120 m elevation. The species has not yet been collected in Costa Rica. Flowering specimens have been collected in Jan- uary, February, and May; fruits are unknown. En- dangered. Lateropora tubulifera is characterized by its an- ther tubules, at least twice as long as those of the other two species, and its small leaf blades. Macleania W. J. Hooker REFERENCES — P. Yeo, Notes on some species of Macleania (Ericaceae). Baileya 15: 45-59. 1967. J. L. Luteyn, A review of and taxonomic realign- ments within the neotropical genus Macleania (Ericaceae: Vaccinieae). BioLlania 6: 455-465. 1997. Epiphytic or terrestrial shrubs sometimes arising from lignotubers. Leaves alternate, petiolate, the blade cori- aceous, the margin entire, the venation pinnate or pli- nerved. Inflorescences axillary (in ours) or terminal, ra- cemose or subfasciculate, few- to many-flowered, rarely flowers solitary; pedicels articulate with calyx (in ours) but this sometimes obscure or rarely continuous; brac- teoles basal to medial. Flowers 5-merous, without odor; calyx synsepalous, tube short-cylindric or campanulate, often angled to conspicuously winged, the wings some- times protruding beyond calyx limb as spurs (not ours), limb erect, spreading; corolla sympetalous, aestivation valvate, subcylindric or elongate urceolate; stamens 10, equal, about !/4 to nearly equaling corolla in overall length; filaments distinct or connate, abaxially attached to anthers near their bases; connectives lacking disinte- gration tissue or spurs; anthers stout, lacking awns, the- cae conspicuously papillate, tubules about as long as thecae, 1 or 2 (then laterally connate but with a septum present, or distinct, rarely completely distinct), rigid, broadly conical, dehiscing by introrse, elongate, distinct or fused clefts; pollen grains in tetrahedral tetrads, lack- ing viscin threads; ovary inferior; style filiform, exsert- ed; stigma truncate. Fruit a berry; seeds numerous, often with a mucilaginous sheath, the embryo sometimes green. Macleania consists of approximately 38 species distributed from Mexico to Peru. The genus is characterized by stamens of equal lengths but !/4-% as long as the corolla; staminal connectives without spurs; anther tubules often short, elon- gate-conical, rigid, and only laterally to totally fused; and ovary inferior. The genus has been di- vided into two very distinct subgenera, each with one species in Costa Rica (Luteyn, 1997). Ma- cleania is related to Psammisia, differing most conspicuously by its staminal connectives that lack spurs (vs. spurs present or absent in Psam- misia} and anther tubules connate (not distinct to the base). Two species are currently known to oc- cur in Costa Rica. The genus is currently being revised by J.L.L. Key to the Species of Macleania la. Anther tubules 1; staminal filaments connate (rarely distinct); calyx sharply angled to winged; corolla angled, totally orange, densely white pilose at throat and lobes internally; berry translucent white (subgen. Macleania) M. insignis Ib. Anther tubules 2; staminal filaments distinct; calyx essentially terete or pentagonal; corolla terete or pentagonal, reddish with white or pinkish throat, glabrous internally; calyx and corolla essentially glabrous; berry blue-black (subgen. Aponema) M. rupestris 68 FIELDIANA: BOTANY Macleania insignis M. Martens & Galeotti, Bull. Acad. Roy. Sci. Bruxelles 9: 531. 1842. Thi- baudia laurifolia M. Martens & Galeotti, Bull. Acad. Roy. Sci. Bruxelles 9: 530. 1842. M. ova- la Klotzsch, Linnaea 23: 20. 1851. M. cordata var. linearifolia J. D. Smith, Bot. Gaz. 16: 12. 1891. M. linearifolia (J. D. Smith) A. C. Smith, Contrib. U.S. Natl. Herb. 28: 368. 1932. M. in- signis var. linearifolia (J. D. Smith) Stand!. & L. O. Williams, Fieldiana, Bot. 31: 172. 1965. M. subracemosa L. O. Williams, Fieldiana, Bot. 31: 172. 1965. Figures 2L, 6, and 7. Epiphytic or terrestrial, infrequently epipetric shrubs 0.5-2 (4) m tall, arising from a lignotuber; mature stems terete, glabrous to pilose, glabrate, often nitid, light to dark brown; bark exfoliating in thin sheetlets, reddish- brown; twigs bluntly angled, glabrous to often densely short-puberulous with white hairs. Leaves with petioles terete or subterete then flattened adaxially, rugose, (1) 2-4 (6) mm long, glabrous or rarely densely short-pu- berulous; leaf blades ovate to lance-ovate, elliptic to el- liptic-ovate, rarely lance-oblong, oblong or suborbicular, (2) 3-8 (12) cm long, (1) 1.5-4 (5.6) cm wide, base rounded, usually subcordate, apex obtuse to acute, gla- brous on both surfaces but usually with brownish, glan- dular fimbriae especially beneath, lower surface also of- ten with a dark, circular, concave gland ca. 0.2 mm diam. on either side of midrib base, pinnately nerved with 3-4 secondary nerves per side, or 3-5 (7)-pliner- ved with lateral nerves originating near base, midrib thickened in basal 0.6-3.5 cm and slightly impressed above and raised beneath or more usually slightly raised on both surfaces, lateral nerves and veinlets usually slightly raised on both surfaces. Inflorescences axillary, sessile, subfasciculate to rarely short-racemose, (1) 2-3 (S)-flowered; rachis (when present) subterete, rugose, 0.6-1 cm long, ca. 1.5-2 mm diam.; floral bract persis- tent, carinate, ovate to triangular-ovate, 1 .2-2 mm long, apex short-acuminate, margin ciliate; pedicels subterete. rugose, (4) 8-16 mm long, glabrous or weakly to mod- erately densely short-pilose, usually also with a few glandular fimbriae apically; bracteoles basal to medial, triangular, 1-3 mm long, with 1-2 circular, dark glands ca. 0.2 mm diam. abaxially at base (the bracteole base sometimes appearing simply darker and thicker in col- or), often minutely ciliate. Flowers with calyx 5.3-8 (10) mm long, glabrous or weakly to moderately densely short-pilose, rarely with a few glandular fimbriae, tube obconic, 3-5 (7) mm long and 3-4 mm diam., glabrous or minutely to moderately appressed short-pubescent, broadly to narrowly but distinctly winged, the wings 0.3-1.8 mm broad, truncate or slightly rounded at base not extended beyond limb apically, limb spreading to erect, 1-3 (4) mm long, conspicuously lobed, lobes nar- rowly triangular to deltate, (0.4) 0.8-2 mm long, mar- gins rarely ciliate, sinuses flat or acute; corolla tubular, slightly expanded at base, contracted at throat, sharply and conspicuously 5-angled, (14) 20-27 (30) mm long, 4-6 (8) mm diam., glabrous to slightly puberulous es- pecially apically externally, throat sparsely to densely villous internally, orange to reddish-orange or rarely pinkish, lobes acute, spreading, 1.5-3 mm long, rarely greenish; stamens 8-13 mm long; filaments connate (rarely distinct) into a tube 3-6 (7) mm long, glabrous; connectives often alternately apically thickened and shouldered; anthers 7-10 mm long, thecae 4-6.5 mm long, tubule I, 2-4 mm long, dehiscing by a broad in- trorse cleft more than half the tubule length; style 22- 31 mm long, exserted 1-2 mm. Berry terete, translucent white. 9-12 mm diam., glabrous. Macleania insignis .is found in premontane cloud forest, (650) 900-2400 m elevation. It is distributed from Mexico to Costa Rica. In Costa Rica it is found in the Cordillera Tilardn and west- ern Cordillera Central. Flowering specimens have been collected January-March; fruiting in May. Locally Common. Macleania insignis is characterized by its small, ovate to elliptic-ovate leaf blades; few-flowered fasciculate inflorescences; and concolorous corol- las. It is the only member of subgen. Macleania found in Costa Rica (see Luteyn, 1997). Plants of some populations, mostly in Nicaragua, in the heavily collected area between Jinotega and Ma- tagalpa, have short-racemose inflorescences that may also be pubescent to some degree ( =F M. sub- racemosa). Frequently these plants may also have staminal filaments that are distinct or only slightly connate. These features are not consistent, how- ever, nor are they correlated in any way; each may be found scattered throughout the Mesoamerican range and may even vary on the same plant. Therefore, we have chosen not to recognize for- mally any other taxon in this group. A. C. Smith (1932) recognized some populations from south- ern Central America (as M. ovata) that were gen- erally differentiated from the more northern M. insignis by longer and broader, deltoid calyx lobes that formed acute basal sinuses with the adjacent lobes. Furthermore, according to Smith, M. insig- nis was considered to have apiculate calyx lobes less than 1.5 mm long that formed a broad, flat sinus between adjacent lobes. There is, however, much greater variability in calycine lobes than was apparent in the relatively few specimens available to Smith, and there is an apparent con- tinuum between the extremes. Hence, there is no way to recognize two taxa based on differences in calyx lobes. Yeo (1967) suggested that M. ova- ta might be best regarded as a geographical sub- species of M. insignis, but he did not make the combination, and we see no reason to do so. The calycine variation does not suggest geographical subspecies since apiculate-lobed plants are found at least as far south as Costa Rica and deltoid, larger-lobed plants are to be found in southern LUTEYN AND WILBUR: FAMILY #172 ERICACEAE 69 Mexico along with the apiculate-lobed plants. Macleania insignis is morphologically most sim- ilar to M. cordifolia Bentham from Ecuador; it differs from the other Costa Rican species of Ma- cleania by the characters mentioned in the key above. The species is visited by hummingbirds in Costa Rica (pers. observ.). Macleania rupestris (Kunth in H.B.K.) A. C. Smith. Phytologia 1: 131. 1935. Thibaudia ru- pestris Kunth in H.B.K., Nov. gen. sp. pi. 3: 270. 1818. Psammisia rupestris (Kunth in H.B.K.) Klotzsch, Linnaea 24: 45. 1851. P. gla- bra Klotzsch, Linnaea 24: 46. 1851. P. costar- icensis Klotzsch, Linnaea 24: 47. 1851. M. tur- rialbana J. D. Smith, Bot. Gaz. 27: 339. 1899. M. glabra (Klotzsch) Hoerold, Bot. Jahrb. Syst. 42: 268. 1909. M. costaricensis (Klotzsch) Hoerold, Bot. Jahrb. Syst. 42: 268. 1909. M. irazuensis S. F. Blake, Proc. Biol. Soc. Wash- ington 35: 121. 1922. M. racemosa Cufod., Arch. Bot. Sist. 9: 199. 1933. Illustration: R. L. Wilbur & J. L. Luteyn, Ann. Missouri Bot. Card. 65: 100, fig. 11. 1978. Figures 2M and 10. Epiphytic or terrestrial shrubs 0.6-2 m tall, often lia- noid, rarely small trees, often with basal lignotubers to 60 cm diam.; mature stems terete to subterete, smooth to furrowed with epidermis cracking longitudinally, dull to nitid, glabrous to rarely moderately puberulent; twigs subterete, bluntly angled, striate, glabrous to densely short-pilose with white hairs to 0.8 mm long. Leaves with petioles rugose, sometimes narrowly winged api- cally, 3-1 1 (14) mm long, 2-4.5 mm diam., usually gla- brous; leaf blades thick-coriaceous, oblong, elliptic, ovate-elliptic to ovate, sometimes spatulate, (2) 4-9.5 (15) cm long, (1.2) 2-5 (7.5) cm wide, base rounded to broadly cuneate, sometimes attenuate, rarely subcordate, apex obtuse or rounded to acute, rarely short-acuminate, usually glabrous but occasionally puberulent or spread- ing short-pubescent on lower surface near base, occa- sionally sparsely punctate above, pinnately nerved with 2-4, arcuate-ascending secondary nerves per side, mid- rib thickened and raised in basal 1-3 cm then slightly impressed apically above and prominently raised be- neath, secondary nerves slightly impressed, raised or plane above and prominently raised beneath, reticulate veinlets slightly raised to plane above and slightly raised beneath. Inflorescences axillary, racemose, (4) 10-20 (40)-flowered, glabrous throughout or occasionally mod- erately spreading short-pubescent; rachis subterete, sometimes sharply angled or ribbed, striate, (0.5) 1-3.5 (13) cm long; floral bract deltate or ovate to oblong, 2- 5 (7) mm long, 2-3 mm wide, often ciliate, sometimes margin glandular-fimbriate; pedicels subterete, angled, striate, rugose, (7) 10-24 mm long; bracteoles nearly basal to medial, ovate to broadly ovate, broadly deltate to oblong, 1-2 (4) mm long, to 3 mm wide, glabrous or often ciliate, apex acute to rounded or short-acuminate. Flowers with calyx 5-8 mm long, glabrous, tube rugose, short-cylindric, 2.5-4 (5) mm long, terete or bluntly an- gled, the base truncate, sometimes striate, limb campan- ulate to spreading, 2-5 mm long, lobes broadly trian- gular to oblong-ovate, 1-2.5 mm long, apiculate, acute to acuminate, sinuses concave; corolla thick-carnose, bistratose, cylindric to somewhat cylindric-urceolate or bottle-shaped, terete or pentagonal, (11) 13-19 (24) mm long, ca. 6-7 mm diam., deep red or pinkish-red at base then paler or yellowish or whitish apically, lobes deltate to oblong, 1.5-3 mm long, usually white, apex acute; stamens 9-16 mm long; filaments distinct, 2-5.5 mm long, glabrous or marginally short-pilose; anthers 8-14 mm long, thecae 4-8 mm long, incurved at base, tubules 2 but usually laterally connate to base, rarely distinct, 3-6.2 mm long; style usually long-exserted. Berry spherical, glabrous, 12-15 mm diam., dark bluish-black. Macleania rupestris is found in lower montane forest; premontane dry and wet forest; montane moist, wet, and rain forest; subalpine rain forest; and humid paramo, (1400) 2000-3400 m eleva- tion. It is broadly distributed from Costa Rica to Peru. Flowering occurs in nearly every month; fruiting specimens in March-July and September. Common. Macleania rupestris is the only member of sub- gen. Aponema in Costa Rica, where it is the third most commonly encountered and collected ericad (following Cavendishia bracteata and Pernettya prostrata}. It is characterized by usually oblong, elliptic, or ovate-elliptic to ovate, sometimes spat- ulate, pinnately nerved leaf blades; racemose in- florescences; inconspicuous floral bracts; corollas about 2 cm long that are red with white apices; staminal filaments distinct; anther tubules two (but laterally connate); and a dark blue-black ber- ry. The extensive morphological variation in M. rupestris is perhaps to be expected in a species as widespread and common as this. Since there are no consistently distinctive populations, we do not consider the variation to be of taxonomic value. Common names include colmillo, muelita, mue- las, and San Juan (Costa Rica) and uva (Panama). In Costa Rica, the hummingbirds Eugenes fulgens spectabilis and Panterpe insignis forage popula- tions of M. rupestris for nectar and probably pol- linate it in the process (pers. observ.; Col well, 1973, as M. glabra). The flower-piercer Diglossa plumbea also visits M. rupestris to rob nectar, which is the reason for the small holes often seen at the base of the corolla. The berry is edible, being sometimes sweet or insipid, and is rarely found in marketplaces; the flowers are said to taste "sour" like Rumex (they contain oxalic acid). 70 FIELDIANA: BOTANY Monotropa Linnaeus REFERENCES — G. D. Wallace, Monotropa. Fl. Neotrop. Monogr. 66: 19-25. 1995. Fleshy, succulent herbs lacking chlorophyll, black- ening upon drying, the shoots arising from perennial root masses; mycorrhizal root parasites. Leaves reduced to bract-like scales, congested. Inflorescences racemose or reduced to a solitary flower (in ours); pedicels re- curved; floral bract and bracteoles present but sometimes similar to perianth. Flowers (3) 4-5 (6)-merous; calyx similar to corolla, the sepals distinct and imbricate (in ours), or apparently absent; corolla cylindrical or cam- panulate, the petals distinct but imbricate, scale-like, saccate at base, tardily deciduous; stamens 8-10 (12); filaments distinct, the outer whorl (opposite petals) shorter than the inner; connectives lacking disintegration tissue or spurs; anthers 1-locular at anthesis, smooth, lacking awns, dehiscing by a single, terminal slit or by ends of a slit which may later open along entire length; pollen grains single, lacking viscin threads; nectaries of 8-10 paired lobes between filament bases; ovary supe- rior, (8) 10-lobed, (4) 5 (6)-locular at base with axile placentation, 1-locular above with 4-5, bilobed parietal placentas; style stout, columnar, thick, fleshy, tapering to ovary; stigma funnel-shaped to discoid, obscurely 4- 6-lobed; nectary present as 8-10 lobes projecting from between filament bases. Fruit a loculicidal capsule, 4- 6-valved, globose to ovoid, 8-10-grooved, erect; seeds numerous, minute, tailed. Monotropa is a genus of two species occurring in circumpolar, boreal, and temperate (= montane in Costa Rica) forests of the United States, Mex- ico, Mesoamerica, and Colombia. One species oc- curs in Costa Rica and adjacent western Panama. Monotropa is readily characterized by its succu- lent herbaceous, achlorophyllous, mycorrhizal habit. The genus was placed in its own family, Monotropaceae, until recently but is now consid- ered part of a more broadly circumscribed Erica- ceae as subfam. Monotropoideae (see above). Monotropa uniflora Linnaeus, Sp. pi. 1: 387. 1753. Hypopitys uniflora (L.) Crantz, Inst. Rei Herb. 2: 476. 1766. M. coccinea Zucc., Flora 15(2): 100. 1832. M. uniflora Linnaeus var. coc- cinea (Zucc.) Domin, Sitzungsber. Konigl. Bohm. Ges. Wiss. Prag 1915: 5. 1915. Illustra- tions: R. L. Wilbur & J. L. Luteyn, Ann. Mis- souri Bot. Card. 65: 103, fig. 12. 1978. G. D. Wallace, Fl. Neotrop. Monogr. 66: 24, fig. 5. 1995. Figures 1A and 15. Terrestrial herbs 5-30 cm tall, arising from a golden- colored, succulent, highly branched (coralline), brittle, root mass. Bracts sessile, elliptic, oblong to lanceolate, 6-16 (20) mm long, (2) 3-5 (10) mm wide, base round- ed, apex acute, margin entire or somewhat erase. Inflo- rescences somewhat fleshy scapes, (5) 13-30 cm long, glabrous. Flowers solitary, nodding at anthesis, the peri- anth more or less 5-merous, cylindric; sepals 4-5, lan- ceolate, 13-16 mm long, apex obtuse or acute, margin erase; petals 5-8, oblong, obovate or spatulate, 8-16 (20) mm long, apex rounded, margin becoming erase apically, sometimes pubescent within; stamens (8) 10 (14), alternately unequal, 7-15 mm long; filaments te- rete, subulate, pubescent, 11-12 mm long, the bases clasped by elongate, slender, paired, nectariferous lobes; anthers ca. 2 mm diam., reniform, appearing peltate, ori- ented ± horizontally at anthesis; ovary 5-6-locular, glo- bose or ovoid; style 2-7 mm long; stigma 2-6 mm diam., funnel-shaped, obscurely 5-lobed. Capsule glo- bose, 7-1 1 mm long, held erect at maturity, the segments persistent; seeds spindle-shaped, ca. 1 mm long. Monotropa uniflora is distributed in boreal and temperate forests of North America, eastern Asia, Japan, Korea, China, and westward in the Hima- layas. In the Neotropics, it is found from Mexico to Colombia, where it forms a mycorrhizal asso- ciation with a fungus in turn associated with the roots of Pinus and/or Quercus. In Costa Rica it is found in the shade of humid Quercus- forest floors in primary cloud forest and bamboo forest, at 1500-2900 m elevation. It has been found in the Cordillera Tilaran and Cordillera de Talamanca. Flowering specimens have been collected in Au- gust through January and March; fruiting speci- mens in August, November, and December. Rare/ Infrequent. Monotropa uniflora is characterized by having inflorescences of solitary, nodding flowers that become erect in fruit. The red-flowered color form (= M. coccinea) is striking in appearance. In Cos- ta Rica and Panama, virtually all plants are red- dish in some part, usually the lower mature stems, but red bracts, calyces, and petals are also known; seemingly no populations are totally white or cream colored. Monotropa uniflora is related to M. hypopitys L., a species also of boreal and tem- perate forests of Europe, North America, Asia, and the Himalayas, with neotropical populations found as far south as Guatemala. Monotropa un- iflora differs from M. hypopitys by its single-flow- ered inflorescences (not several), perianth of se- pals (not of distinct sepals and petals), horizontal anthers dehiscing by slits at bothr ends (not erect and dehiscing only by a single terminal slit), stout style (not slender), elongate and slender nectary lobes (not short and stout), and plants black-dry- ing (not brownish) (see Wallace, 1995; Bidarton- do & Bruns, 2001). LUTEYN AND WILBUR: FAMILY #172 ERICACEAE 71 Orthaea Klotzsch REFERENCES — J. L. Luteyn, Orthaea (Ericaceae- Vaccinieae): New species and redefinition of the genus. Nord. J. Bot. 7: 31-37. 1987. Low, frequently epiphytic shrubs. Leaves alternate, short-petiolate, the blades coriaceous to subcoriaceous, the margin entire, the venation pinnate or plinerved. In- florescences axillary, racemose, subfasciculate or 1-2- flowered; floral bract minute (in ours) to sometimes large and showy, but usually caducous; pedicels articulate with calyx; bracteoles 2, usually basal. Flowers 5-mer- ous, without odor; calyx synsepalous, tube terete or 3- winged (in ours) and sometimes basally apophysate, limb erect to suberect, usually 5-lobed (3-lobed in ours); corolla sympetalous, aestivation valvate, carnose, tubu- lar to subcylindric; stamens (8) 10, alternately unequal, usually about '/, length of corolla; filaments distinct or slightly coherent at base, alternately unequal; connec- tives lacking disintegration tissue (present in some South American species), lacking spurs; anthers equal or alter- nately subequal, lacking awns, thecae smooth, tubules 2, distinct, shorter than thecae, dehiscing by terminal or slightly obliquely introrse pores; pollen grains in tetra- hedral tetrads, lacking viscin threads; ovary inferior; style filiform, about as long as corolla. Fruit a berry; seeds numerous. Orthaea is a neotropical genus of ca. 34 species distributed from Mexico to Bolivia and east to Guyana and Trinidad. The genus is characterized by stamens that are about '/3 the length of the co- rolla and among themselves alternately unequal (the filaments alternately unequal), short anther tu- bules, and an inferior ovary. The genus has not yet been found in Costa Rica. Orthaea is morpholog- ically similar to Cavendishia, both having alter- nately unequal stamens and similar habits. Orthaea differs from Cavendishia most prominently by its usually shorter stamens with regard to the corolla (ca. '/3 the length vs. equaling the corolla length), anthers of basically equal lengths (unequal in Cav- endishia), and usual absence of showy floral bracts (present in Cavendishia). On the basis of gross morphology, Luteyn (1987b) redefined the genus Orthaea, recognizing the new subgenus Lysiclesia, in which our species belongs. Orthaea panamensis (Luteyn & Wilbur) Luteyn, Nord. J. Bot. 7: 33. 1987. Lysiclesia panamen- sis Luteyn & Wilbur, Brittonia 29: 261, fig. 2. 1977. Figure 1H. Lax, somewhat scandent, almost vine-like shrubs; ma- ture stems terete to bluntly angled, smooth, short-pilose, glabrescent, glaucous. Leaves with petioles subterete, flat- tened adaxially, rugose, 2.5-3 mm long, ca. 1 mm diam., short-pilose; leaf blades ovate-elliptic, (2.5) 3-3.5 cm long, (1.2) 1 .5-2 cm wide, base obtuse or rounded, apex abruptly short-acuminate and apiculate, glabrous, obscure- ly 3 (S)-plinerved, midrib nearly plane above but raised beneath, lateral nerves and reticulate veinlets slightly raised and prominent above, obscure beneath. Inflores- cences axillary, racemose, 2-3-flowered; rachis bluntly angled, 1 .5-2.2 cm long, ca. 1 mm diam., short-pilose and also bearing scattered, capitate-glandular trichomes ca. 1 mm long; floral bract lanceolate, ca. 1 .5 mm long, ciliate; pedicels subterete, 15-17 mm long, ca. 1 mm diam., short-pilose and also bearing scattered, capitate-glandular trichomes; bracteoles medial, strongly nerved, ovate-lan- ceolate, 3-4 mm long, ca. 1.5 mm wide, acuminate, cil- iate and also bearing capitate-glandular trichomes. Flow- ers with calyx 36-48 mm long, short-pilose externally and internally, tube ca. 4-6 mm long but obscured by wings, limb short-campanulate, ca. 2 mm long, conspic- uously 3-parted, lobes triangular, acuminate, 25-38 mm long, basally ca. 10-14 mm wide, with margins and lam- ina bearing scattered capitate-glandular trichomes, sinuses acute; corolla membranous, bistratose, cylindric, narrow- ing to throat, ca. 35 mm long, pink, short-pilose externally (except at base) and internally and at throat, lobes flaring, triangular, ca. 1 mm long, obtuse; stamens alternately ca. 10 mm and 12 mm long; filaments alternately ca. 5 mm and 8.5 mm long, distinct, pilose apically; anthers very slightly unequal, alternately ca. 5 mm and 5.5 mm long, thecae 2.5-3 mm long, tubules 2.5-3 mm long, with ter- minal pores or in shorter stamens slightly obliquely in- trorse pores; style ca. 36 mm long, pilose apically. Berry not seen. Orthaea panamensis is found in premontane cloud forests, 900-1200 m elevation. It is endemic to Panama and is known only from a few localities in western and central Panama. It has not yet been collected in Costa Rica. Flowering specimens have been collected in October, December, and Febru- ary; fruit are unknown. Rare/Infrequent Orthaea panamensis is characterized by its calyx limb, which has three very large, bract-like lobes instead of the usual five smaller lobes. This is a striking feature that defines Orthaea subgen. Lysi- clesia (A. C. Smith) Luteyn (Luteyn, 1987b) and immediately allows one to recognize the taxon. Only three species are found in this subgenus, the other two being known only from north-central Co- lombia (Antioquia Dept.). The three species are closely related and possibly even conspecific; more collections of all three species are needed to deter- mine their circumscription and relationships. Pernettya Gaudich. nom. & orth. conserv. REFERENCES — H. O. Sleumer, Taxonomy of the genus Pernettya. Bot. Jahrb. Syst. 105: 449-480. 72 FIELDIANA: BOTANY 1985. D. J. Middleton & C. C. Wilcock, A critical examination of the status of Pernettya as a genus distinct from Gaultheria. Edinburgh J. Bot. 47: 291-301. 1990. J. L. Luteyn, Pernettya. Fl. Neo- trop. Monogr. 66: 365-383. 1995. Terrestrial shrubs. Leaves alternate, petiolate, the blade coriaceous, the margin entire or often subserrate- crenulate with teeth then bristle-tipped with eglandular or glandular-tipped hairs, glabrous or rarely setose be- neath, the venation pinnate. Inflorescences axillary, flowers solitary (ours) or in racemes; floral bract and bracteoles indistinguishable: pedicels articulate with ca- lyx, with 2-many bracts along length nearly to apex. Flowers 5-merous, nodding, without odor; calyx persis- tent, synsepalous but parted nearly to base, lobes imbri- cate at base, normally membranous and not accrescent in fruit, rarely becoming succulent after anthesis but not surrounding fruit; corolla sympetalous, aestivation im- bricate, lobes reflexed, less than '/I0 as long as corolla tube; stamens (8) 10, about V2 as long as corolla; fila- ments distinct, slightly to conspicuously dilated above base, papillate; connectives with disintegration tissue present abaxially at base of awns, lacking spurs; anthers apically 4-awned, the awns sometimes vestigial (in spe- cies from Brazil), thecae minutely papillate at base, de- hiscing at apex by introrse, terminal pores; pollen grains in tetrahedral tetrads, lacking viscin threads; ovary su- perior, 5-locular; style filiform, as long as corolla. Fruit a berry, subglobose to globose, apically depressed, gla- brous to densely short-pubescent; seeds wingless, nu- merous, small. Pernettya consists of about 14 species distrib- uted in Tasmania, New Zealand, and Mexico to Argentina. One species occurs in Costa Rica. Per- nettya is characterized by a superior ovary and baccate fruit. It is closely related to Gaultheria, the primary distinction being the baccate fruit of Pernettya vs. the capsular fruit of Gaultheria (see discussion above under Gaultheria). Pernettya prostrata (Cav.) DC., Prodr. 7: 609. 1839. Andromeda prostrata Cav., Icon. 6: 43, t. 562, f. 2. 1801. P. coriacea Klotzsch, Linnaea 24: 85. 1851. P. setigera Klotzsch, Linnaea 24: 85. 1851. P. congesta Klotzsch in 0rsted, Amer. Centr. 9: 1 863, nom. nud. P. oerstediana Klotzsch in 0rsted, Amer. Centr. 9: 1863, nom. nud. P. ciliaris D. Don ex G. Don van alpina J. D. Smith, Bot. Gaz. 25: 148. 1898. G. alpina (J. D. Smith) Sleumer, Notizbl. Bot. Gait. Ber- lin-Dahlem 13: 207. 1936. Illustrations: R. L. Wilbur & J. L. Luteyn, Ann. Missouri Bot. Card. 65: 106, fig. 13. 1978 (as P. coriacea). J. L. Luteyn, Fl. Neotrop. Monogr. 66: 367, fig. 1C, M, and 374, fig. 3A-E. 1995. Figures IF and 4. Prostrate and stoloniferous, to erect or ascending shrubs, sometimes forming dense tufts or extensive mats (0.1) 0.2-1 (3) m tall; mature stems terete to sub- terete, sometimes sharply angled, glabrous or weakly to densely puberulous or laxly crisped pilose, and also usu- ally moderately to densely strigose or hirsute-setose with basally swollen ferruginous, appressed or spreading eglandular and/or rarely gland-tipped hairs 0.5-3 mm long. Leaves with petioles 1-4 mm long, often canalic- ulate adaxially, glabrous to puberulous above; leaf blades subcoriaceous to coriaceous, narrowly to broadly elliptic, oblong, or ovate-oblong to linear-oblong, (2) 6- 14 (40) mm long, (2) 4-7 (12) mm wide, base rounded, obtuse, attenuate or cuneate, apex obtuse, acute or sub- acuminate, midrib rarely protracted into an achlorophyl- lous mucro, rarely short-mucronate or rounded, margin slightly to conspicuously thickened, usually slightly rev- olute. minutely crenate-serrulate with 6-12 (17) cadu- cous teeth per side, these bristle-tipped or gland-tipped and 0.2-0.3 mm long, to hirsute-setose and eglandular to 2 mm long, blades themselves usually glabrous or rarely weakly short-pilose along the basal half of midrib above, rarely strigose beneath. Inflorescences with 5- 10 ovate, ciliate, appressed, persistent bractlets at base of pedicel and 1-6 undifferentiated bractlets scattered along pedicel, these bractlets (1.5) 2-4 mm long; pedi- cels (1) 4-10 (13) mm long, glabrous or puberulous, sometimes also bearing short-strigose or hirsute-setose, straight or crisped, eglandular or gland-tipped hairs. Flowers with calyx lobes ovate to ovate-lanceolate, 2- 4 mm long, apex acuminate to acute, weakly to densely ciliate, sometimes also basally with short, gland-tipped hairs; corolla cylindric to campanulate-urceolate, (3.5) 5-6 (7) mm long, 2.5-5 mm diam. at base, white to rose tinted, lobes 1-1.5 mm long; stamens with filaments 1.5-3 mm long, glabrous or rarely pilose; ovary gla- brous to densely short-pubescent; style 2-4 mm long, glabrous or rarely short-pilose in lower half. Berry subglobose, 6-16 mm diam., glabrous to densely short- pubescent, dark blue-black, rarely light purple to almost white. Pernettya prostrata is found in montane cloud forest, scrub vegetation, subparamo to pdramo, frequently as a pioneer species in new habitats, (900) 2000-3800 m elevation. It is distributed from Mexico to northwestern Argentina. In Costa Rica it is common at higher elevations of the Cen- tral Valley volcanoes and the Cordillera de Tala- manca. Flowering and fruiting occur throughout the year. Common. Pernettya prostrata is characterized by its usu- ally strigose to setose mature stems, the trichomes having swollen bases, an obtuse to acute (rarely mucronate-tipped) leaf apex, eglandular pubes- cence, glabrous filaments, and essentially Andean distribution. It is morphologically highly variable with regard to habit, leaf size and shape, stem and leaf indumentum, calyx lobe succulence, and fruit color. Also, because of its broad geographical dis- tribution and often isolated populations, many dif- LUTEYN AND WILBUR: FAMILY #172 ERICACEAE 73 ferent names have been assigned to the different populations; Costa Rican populations have tradi- tionally been named P. coriacea. After studying the species from throughout its entire geographi- cal range with abundant herbarium material, Lu- teyn (1995) was not able to see any morphological variation that consistently correlated with geog- raphy, geology, elevation, habitat, habit, etc. Since virtually any character might appear anywhere in the range of the species, P. prostrata was best treated as a polymorphic species of wide distri- bution. In Costa Rica, in the paramo vegetation on Cerro de La Muerte and Cerro Chirripo at 3300-3800 m, one puzzling variant may be seen in high-elevation populations, where plants may have distinctly and persistently mucronate leaf apices, a feature otherwise found only in P. how- ellii Sleumer (Galapagos Isl.) and some of the temperate Chilean/ Argentinean species. However, although distinctive, this character is not even consistent within a population, and some young leaves with mucronate leaf tips may lose this fea- ture with age. Pernettya prostrata is most closely related to P. hirta (Willdenow) Sleumer, which is known only from one small area east of Bogota, Colom- bia. Pernettya prostrata is often confused with Vac- cinium floribundum, both in the field and in her- barium material. However, closer examination of even sterile material shows easily discernible dif- ferences: Pernettya prostrata has usually strigose- setose mature stems with thick trichomes that have swollen bases; the leaf margins are slightly revolute beneath and of the same color as the lam- ina proper; the midrib and lamina are the same color on the lower surface of the leaf; and the ovary is superior with the berry subtended by the calyx lobes, whereas Vaccinium floribundum has glabrous or short-pilose mature stems; the leaf margins are flat, thickened, smooth in texture, and of a conspicuously lighter color than the leaf lam- ina proper; the midrib is also a lighter color than the lamina; and the ovary is inferior with the ber- ry having calyx lobes at the apex. Flowers of P. prostrata are visited by the bee Bombus ephippia- tus in Costa Rica (Luteyn, 1998). The berries are reportedly toxic in quantity (but see Luteyn, 1995). Psammisia Klotzsch REFERENCES — A. C. Smith, The American spe- cies of Thibaudieae. Contrib. U.S. Natl. Herb. 28: 311-547. 1932 (Psammisia, pp. 384-406). J. L. Luteyn, New species and notes on Neotropical Er- icaceae. Opera Bot. 92: 109-130. 1987a (Psam- misia, pp. 120-121). J. L. Luteyn, Ericaceae. Fl. Ecuador 54: 1-406. 1996 (Psammisia, pp. 164- 209). Epiphytic shrubs. Leaves alternate (in ours) to subop- posite, petiolate, the blades coriaceous, the margin en- tire, the venation plinerved or pinnate. Inflorescences axillary (in ours) or terminal, subfasciculate or race- mose, few- to many-flowered; floral bract small, incon- spicuous; pedicels articulate with calyx; bracteoles 2, usually basal. Flowers 5-merous, without odor; calyx synsepalous, tube terete and rarely winged, limb erect or spreading; corolla sympetalous, aestivation valvate, car- nose, subcylindric, elongate-urceolate or subglobose; stamens 10, equal, '/3 to nearly as long as corolla; fila- ments distinct or connate; connectives lacking disinte- gration tissue, but all laterally or apically 2-spurred, or alternately spurred, or rarely spurs obscure or lacking, when present the spurs either acute and conspicuous or rounded and not very apparent; anthers rigid, lacking awns, thecae conspicuously papillate, tubules 2, distinct, rigid, !/4 to about as long as thecae, dehiscing by elon- gate, introrse clefts; pollen grains in tetrahedral tetrads, lacking viscin threads; ovary inferior; style filiform, ex- serted, glabrous. Fruit a leathery berry, green to green speckled with brown. Psammisia consists of approximately 70 spe- cies distributed from Mexico to Bolivia and east to Guyana, Amazonian Brazil, and Trinidad. The genus is characterized by having stamens of equal lengths but only !/3 to nearly as long as the overall corolla length; their connectives are usually spurred; the anther tubules are often short, elon- gate-conical, rigid, and distinct; and an inferior ovary is present. There are three species in Costa Rica. Psammisia is related to Macleania, differing most conspicuously by its spurred stamens (spurs absent in Macleania) and anther tubules that are distinct to the base (not connate). The genus is currently being revised by J.L.L. 74 FIELDIANA: BOTANY Key to the Species of Psammisia la. Rachis elongate, at anthesis 2-6 cm long, the inflorescence distinctly racemose; bracteoles 3 mm long P. williamsii Ib. Rachis short, at anthesis ca. 0.7-2 cm long or less (elongating in fruit up to 4 cm long), the inflorescence short- racemose but often appearing fasciculate or corymbose by elongation of lower pedicels; bracteoles 2.5 mm long or less 2 2a. Staminal filaments typically connate over entire length; anther connectives spurred; inflorescence parts lacking exudate; pedicels 8-25 mm long P. ramiflora 2b. Staminal filaments distinct or only slightly connate at base; anther connectives lacking spurs; inflo- rescence parts (especially calyx) with white, scale-like or crystalline exudate (most conspicuous when inflorescence young, but usually persistent); pedicels 20-60 (71) mm long P. ulbrichiana Psammisia ramiflora Klotzsch, Linnaea 24: 44. 1851. Psammisia symphystemona J. D. Smith, Bot. Gaz. 20: 291. 1 895. Figures 2N and 13. Terrestrial or usually epiphytic shrubs; mature stems slender, terete, glabrous. Leaves with petioles (3) 5-12 (17) mm long, sometimes narrowly winged apically, gla- brous; leaf blades coriaceous to thick-coriaceous but usually drying membranous, oblong or ovate-oblong, 8- 19 (27) cm long, 3-7 (11) cm wide, base cuneate to rounded, apex caudate-acuminate, glabrous, 5-plinerved, lateral nerves arising slightly above base and ascending near margins, midrib impressed or flat above and prom- inently raised beneath, reticulate veinlets conspicuous and raised on both surfaces. Inflorescences axillary, subfasciculate or short-racemose, 4-15-flowered, exu- dates none; rachis usually < 1 cm long but elongating to 1-4 cm long after anthesis, glabrate; floral bract oblong or triangular, (1) 3-4 mm long, often persistent; pedicels 8-25 mm long, often elongating after fruit; bracteoles supramedial, 1.5-2.5 mm long. Flowers with calyx 5- 8.5 mm long, tube subcylindric or broadly obconic, ca. 2-4 mm long, glabrous, dark green, limb spreading- campanulate at anthesis but becoming erect and cylin- dric after anthesis forming a persistent crown on matur- ing ovary, 3-5.5 mm long, light green, lobes triangular to ovate, ca. 1 .5-2 mm long, apex acute to rounded and short-apiculate, sinuses rounded to sometimes apparent- ly tearing and then acute; corolla parchment thin (thinly fleshy when fresh), unistratose, cylindric throughout, ta- pering to throat, sometimes slightly constricted below middle, 28-35 mm long, 6-10 mm diam. medially and at base but contracted to 4-5 mm diam. at throat, orange to red with a white tip, throat often with appressed glan- dular trichomes 0.1-0.4 mm long, lobes oblong, 2.5-3.5 mm long; stamens 8-10 mm long; filaments 3.5-4.5 mm long, connate for their entire length, lightly coherent near base only, sometimes completely separate, short- pilose adaxially or along margin in apical free part; con- nectives apically and alternately unequally 2-spurred. ciliate with trichomes ca. 0.2 mm long; anthers 5.8-7.7 mm long, thecae 3-5.3 mm long, tubules 2-3.5 mm long. Berry subglobose, 8-15 mm diam., crowned by persistent calyx limb that may elongate to 7 mm long. Psammisia ramiflora is found in lowland rain forest and premontane and montane cloud forest, 200-2200 m elevation. It is distributed in Guate- mala (one collection), Costa Rica, and Panama. Flowering occurs in nearly every month; fruiting specimens in June and December. Common. Psammisia ramiflora is characterized by its short-racemose inflorescence that often appears fasciculate or corymbose because of the elonga- tion of the lower pedicels, spurred anther connec- tives, inflorescence parts without exudate, pedi- cels 8-25 mm long, and proportionally long co- rolla lobes. It is variable in leaf size, texture and venation, corolla color, and degree of connation (or lack thereof) of the Staminal filaments. Typi- cally, in Costa Rica, the leaf blades are subcori- aceous when fresh, but they dry membranous, the leaf venation is strongly 3 (5)-plinerved with the inner pair of lateral nerves distinct and conspic- uous to the apical tip on both surfaces, and the tertiary veinlets are prominent beneath, running perpendicular to the midrib and inner laterals. The leaf veins are also lighter in color than the blade proper when dry. The calyx limb and lobes are usually a distinctly lighter shade of green when fresh as well as a lighter shade of brown when dry. Also, the corolla is red, with a white throat and lobes, and the filaments are usually connate. Psammisia ramiflora is related to P. ulbrichi- ana, a species disjunct from Colombia and Ec- uador, and until now has been consistently con- fused with it. They may be distinguished by the characters mentioned in the key above, with the presence of a white, crystalline exudate being es- pecially noticeable in P. ulbrichiana. The species is visited by the hummingbird Doryfera ludovicae veraguensis and by the genus Phaethornis (pers. observ.; Luteyn, 1998). The flowers are somewhat pungent to the taste (they contain oxalic acid) and are said to give energy when eaten. Psammisia ulbrichiana Hoerold, Bot. Jahrb. Syst. 42: 306. 1909. Figure 13. LUTEYN AND WILBUR: FAMILY #172 ERICACEAE 75 Terrestrial or epiphytic shrubs, sometimes with scan- dent stems. Leaves with petioles 8-19 mm long, gla- brous; leaf blades ovate, elliptic or oblong-lanceolate, (15) 18-40 cm long, 6-13 (17.5) cm wide, base cuneate to rounded, often ± abruptly short-acuminate, apex ta- pering and long-acuminate, glabrous, 5-7-plinerved. In- florescences axillary, short-racemose but conspicuously corymbiform by elongation of lower pedicels, 10-20 (35)-flowered; rachis 0.7-2 cm long, elongating to 4 cm in fruit, glabrous, although often covered with a scaly or crystalline, white exudate when young which may persist to anthesis; floral bract broadly ovate to oblong, 3-3.6 mm long, apex acute, margin deciduously glan- dular-fimbriate; pedicels 20-60 mm long, sometimes elongating to 71 mm post-anthesis; bracteoles medial to supramedial, triangular, ca. 1.8-2.5 mm long, apex acute, margin glandular-fimbriate. Flowers with calyx 6.5-9 mm long, tube campanulate, the base narrowing and rounded or sometimes truncate, 2.5-4 mm long, limb 4-5.5 mm long, lobes ca. 2-4 mm long, broadly ovate, round, apiculate, margin often appearing irregu- larly torn, sinuses acute; corolla cylindric, swollen in basal V3-V2, conspicuously once-constricted below mid- dle (best seen when fresh), (18) 25-38 mm long, ca. 6 mm diam. near base, glandular-fimbriate apically, dark rose-red in basal swollen part and lighter rose-red api- cally, lobes 2-3 mm long, triangular to oblong, acute; stamens 9-15 mm long; filaments distinct or slightly connate at base, 3-6 mm long, sparsely pilose at margins apically; connectives lacking spurs or with short (mi- nute, vestigial) spurs; anthers 10-13 mm long, thecae 4-7 mm long, tubules sometimes laterally connate in basal half, 3-5.5 mm long. Berry slightly depressed- spherical, 13-18 mm diam. Psammisia ulbrichiana is found in premontane cloud forest, 900-1200 m elevation. It is distrib- uted from Costa Rica to Peru. Flowering speci- mens have been collected in July; fruiting collec- tions in August. Rare/Infrequent. Psammisia ulbrichiana is characterized by its large, 5-7-plinerved leaves; short-racemose inflo- rescence that is conspicuously corymbose because of the elongation of the lower pedicels; the con- stricted (submedially) corolla; anther connectives lacking spurs; inflorescence parts with white, scale-like or crystalline exudate (most conspicu- ous when the inflorescence is young, but usually persistent); and pedicels 20-60 (71) mm long. In Costa Rica and Panama, it is morphologically similar to and has been confused with P. ramiflo- ra, as described above. The species is visited by hummingbirds in Ecuador (pers. observ.). Psammisia williamsii A. C. Smith, Ceiba 1: 62. 1950. Figure 13. Epiphytic shrubs 1-1.5 m tall; mature stems terete, glabrous. Leaves with petioles subterete, rugose, 8-11 mm long and ca. 2 mm diam; leaf blades subcoriaceous, narrowly oblong-elliptic, 10-20 cm long, 3-8.5 cm wide, base ± attenuate and decurrent onto petiole, apex acuminate, 5-plinerved, midrib strongly impressed above and prominent beneath, thickened in basal 1-1.5 cm, lateral nerves and reticulate veinlets inconspicuous. Inflorescences axillary, racemose, 5-7-flowered; rachis 2-6 cm long, striate, circumscribed at base by many papery bracts ca. 2 mm long; floral bract oblong-deltate, 4-5 mm long, apex acute; pedicels rugulose, 30-37 mm long; bracteoles basal, ca. 3 mm long. Flowers with ca- lyx ca. 9 mm long and 10 mm apical diam., tube cu- puliform, inconspicuously glandular, ca. 4 mm long and 5 mm diam., limb spreading-erect, slightly longer than tube, lobes deltate, apiculate, ca. 1 mm long, sinuses concave to complanate; corolla carnose, cylindric but slightly swollen below middle, conspicuously and abruptly constricted at throat, ca. 35 mm long, ca. 10- 12 mm diam. near base, red with yellowish throat and lobes, the lobes oblong-deltate, obtuse, 2-3 mm long; stamens 10-11 mm long; filaments distinct, ca. 5 mm long; connective carnose and apically 2-spurred, the spurs spreading, obtuse; anthers 9-10 mm long, thecae ca. 1.5 mm diam., shortly incurved at base, tubules scarcely 2 mm long, much shorter than thecae, dehiscing through oval rims. Berry not seen. Psammisia williamsii is found in premontane cloud forest, 1800-2100 m elevation. It may be endemic to Costa Rica and is known with certain- ty only from the type collection and Luteyn et al. 4526 (both Cartage Province, El Muneco area). A recent collection from Chiriqui Province, Panama (Fortuna Dam site area), Luteyn & Pedraza 15550, may also be this species; that collection has vegetative and inflorescence features includ- ing the color combination of the corolla of P. wil- liamsii, but unfortunately it is only in bud. Flow- ering specimens have been collected (in Costa Rica) only in December; fruits are unknown. En- dangered. Psammisia williamsii is characterized by its ra- cemose inflorescence with an elongate rachis (at anthesis 2-6 cm long) and large, carnose corollas. It is morphologically similar to P. ramiflora and P. ulbrichiana but differs by a longer and race- mose inflorescence with a longer rachis, longer bracteoles and larger flowers, and its corollas, which have a different shape and color combina- tion. These character differences are slight, how- ever, and more collections of P. williamsii are needed before its range of morphology and spe- cific relationships can be determined. The species is visited by hummingbirds (pers. observ.). Satyria Klotzsch REFERENCE — J. L. Luteyn, New species and notes on Neotropical Ericaceae. Opera Bot. 92: 109-130. 1987 (Satyria, pp. 121-122). 76 FIELDIANA: BOTANY Epiphytic or terrestrial shrubs. Leaves alternate, ev- ergreen, petiolate, the blades coriaceous, the margin en- tire, the venation usually plinerved (in ours), rarely pin- nate. Inflorescences axillary, racemose or subfascicula- te, often ramiflorous and then 1 -several arising from a callus-like mound; floral bract small, inconspicuous, usually persistent; pedicels articulate with calyx; brac- teoles 2, usually persistent. Flowers 5-merous, without odor; calyx synsepalous, tube subglobose to shortly campanulate, limb flaring or spreading, lobes (3-4) 5, ovate to triangular or merely apiculate; corolla sympet- alous, aestivation valvate, carnose, cylindric to narrowly vasiform, lobes triangular to narrowly oblong, acute; stamens 10, of alternately unequal lengths, usually about '/, as long as corolla; filaments equal and firmly connate into a tube for more than half their length, mem- branous to thinly coriaceous; connectives lacking dis- integration tissue or spurs; anthers basifixed, firm, alter- nately unequal in length, lacking awns, slightly dimor- phic in shape, the longer ones spreading apically with tips often incurved and decorated in a dendritic pattern and with tubules twisted so dehiscence latrorse, the shorter ones laterally coherent apically, tips straight, of- ten not ornate, tubules straight so dehiscence introrse, thecae subcoriaceous, slightly to not papillate, tubules 2, distinct, not sharply differentiated from thecae, dehisc- ing by subapical pores; pollen grains in tetrahedral tet- rads, lacking viscin threads; ovary inferior; style fili- form, as long as corolla or slightly exserted, glabrous; stigma usually truncate; nectariferous disc usually con- spicuous, annular-pulvinate. Fruit a spherical berry, dark blue-black (in ours), rarely translucent white; seeds numerous. Satyria consists of about 25 species distributed from southern Mexico to Bolivia and east to French Guiana. Four species are currently known to occur in Costa Rica. Satyria is characterized by its numerous, usually ramiflorous inflorescences; alternately unequal stamens (equal filaments, un- equal anthers) that are about '/, as long as the co- rolla; entirely connate staminal filaments; and in- ferior ovary. Morphology as well as molecular data (but see Kron et a!., 2002b) indicate that the genus is related to Cavendishia and Orthaea, two other vaccinioid genera with alternately unequal stamens (each with unequal filaments). Satyria is currently being revised in a Ph.D. study by E. Ann Powell (Wake Forest University). Key to the Species of Satyria la. Inflorescence clearly long-racemose; corolla glabrous or usually puberulent over entire length; co- rolla red, greenish toward apex, often bluntly 5-angled; lowland, primary rain forest to premontane cloud forest 5. panurensis Ib. Inflorescence fasciculate to short-racemose; corolla glabrous or rarely shortly and sparsely puber- ulent apically; corolla red or orange, white or green toward apex, terete to bluntly 5-angled; pre- montane to montane cloud forest 2 2a. Corolla (15) 17-36 mm long; leaves (6) 10-27 (30) cm long; inflorescences 3-11 (30)-flowered, fasciculate to short-racemose 3 2b. Corolla 7-21 mm long; leaves 6.5-18 (25) cm long; inflorescences 6- 1 6-flowered, short-racemose 4 3a. Corolla cylindrical throughout or only somewhat swollen toward the base, red to rose-red becoming white at throat and lobes; rachis 0.5-1.5 (4) cm long 5. warszewiczii 3b. Corolla conspicuously swollen in basal V4-V^ red becoming green at throat and lobes; rachis 0.5- 0.9 cm long S. ventricosa 4a. Leaves conspicuously bullate, the inner pair of lateral nerves arising from about or just below the middle; calyx lobes conspicuous, ovate to triangular, ca. 2-2.3 mm long; corolla 7-1 1 mm long, conspicuously swollen and globose basally then abruptly constricted and cylindrical apically .... 5. species "A" 4b. Leaves flat, not bullate, the inner pair of lateral nerves arising near base; calyx lobes inconspicuous, apiculate, ca. 0.2-0.8 mm long; corolla 10-21 mm long, cylindric and of equal diameter along entire length 5 5a. Leaf blades 4-9 (11) cm wide; calyx lobes triangular-apiculate, ca. 0.2-0.3 mm long; corolla 10- 15 mm long; Guatemala to western Costa Rica ». . S. meiantha 5b. Leaf blades 2-3 cm wide; calyx lobes ovate-apiculate, 0.5-0.8 mm long; corolla 15-21 mm long; western Panama S. species "B" LUTEYN AND WILBUR: FAMILY #172 ERICACEAE 77 Satyria meiantha J. D. Smith, Bot. Gaz. 47: 256. 1909. Figure 15. Usually epiphytic shrubs. leaves with petioles stout, 7-20 mm long, 2-3 mm diam., glabrous; leaf blades el- liptic, lance-ovate, oblong or ovate-oblong, (8) 12-18 (25) cm long. 4-9 (11) cm wide, base usually cuneate to rounded, apex acuminate to acute, glabrous, 3 (S)-pliner- ved with inner pair of lateral nerves arising in basal 15 mm, midrib thickened and raised in basal 10-15 mm above, then apically thinner and becoming slightly im- pressed near apical tip, raised and conspicuous beneath, lateral nerves slightly raised but inconspicuous above be- coming slightly impressed like midrib near apical tip, raised and conspicuous beneath, reticulate veinlets slightly raised but inconspicuous on both surfaces. Inflorescences short-racemose, 6-12-flowered, frequently up to 7 togeth- er, ramiflorous from callus at leafless nodes on older branches (persisting over many years); rachis angled, stri- ate, 5-15 mm long, puberulent, inconspicuously short- glandular-fimbriate; floral bract oblong-ovate, deciduous, <2 mm long, apex acute to acuminate, glabrous or pu- berulent, margin deciduously glandular-fimbriate; pedicels 6-18 mm long, glabrous or minutely puberulous, glan- dular-fimbriate especially apically near articulation; brac- teoles at basal '/, to middle of pedicels, oblong-ovate, ca. 1.5 mm long, apex acuminate, glabrous or puberulent, margin deciduously glandular-fimbriate. Flowers with ca- lyx 3.5-5.5 mm long, glabrous or minutely puberulous, glabrate, with or without glandular fimbriae 0.1-0.2 mm long, tube cylindric-campanulate, 2-2.5 mm long, ca. 2 mm diam., limb erect to spreading, 1 .5-3 mm long, lobes triangular, apiculate, ca. 0.2-0.3 mm long, sinuses flat to rounded; corolla coriaceous, thinly bistratose, tubular but slightly swollen at base and contracted apically, broadly and bluntly pentagonal, rarely slightly zygomorphic, ca. 20 mm long and ca. 6 mm diam. when fresh, but drying 10-15 mm long and 2-4 mm diam., basal % rose-red becoming white in apical '/4, glabrous or densely but mi- nutely puberulous, also typically with inconspicuous red- dish, glandular fimbriae ca. 0.1 mm long, lobes oblong, acute, 1-2 mm long; stamens alternately 6.6-9 mm and 7.6-10 mm long; filaments ca. 4-5 mm long, equal, con- nate into a tube, apically, abaxially and marginally short- pilose onto connectives, also bearing glandular fimbriae apically; anthers alternately 4.2-5.5 mm and 5.2-6.5 mm long, thecae alternately ca. 3-3.5 mm long and 3.5-4.5 mm long, tubules apically alternately straight and slightly flaring ca. 1.6-2.5 mm long, dehiscing by clefts ca. 1-2 mm long. Berry globose, to at least 8 mm diam., turning purple. Satyria meiantha is found in premontane cloud forest, 850-1700 m elevation. It is distributed in Guatemala, Nicaragua, and Costa Rica, where it is limited to the Cordillera Tilaran. Flowering specimens have been collected in January-April, August, and October; fruiting in June, October, and November. Locally Common. Satyria meiantha is characterized by having in- florescences that are axillary or frequently ramiflo- rous, short-racemose, glabrous rachises, glabrous or rarely shortly and sparsely puberulent apically corollas that are red becoming white apically, and a premontane to montane cloud forest habitat. It is morphologically similar to S. warszewiczii, and the small differences given in the key may not stand rigorous analysis; more fieldwork is needed to compare the two. The species is visited by the hummingbird Lampornis calolaema (male) in Cos- ta Rica (pers. observ.; Luteyn, 1998). Satyria panurensis (Benth. ex Meissn.) Benth. & Hook.f. ex Niedenzu, Bot. Jahrb. Syst. 1 1 : 249. 1890. Thibaudia panurensis Benth. ex Meissn. in Martius Fl. Brasil. 7: 125. 1863. 5. panuren- sis (Benth. ex Meissn.) Benth. & Hook.f., Gen. PI. 2: 568. 1876, nom. illeg. Riedelia panuren- sis (Benth. ex Meissn.) Kuntze, Rev. Gen. PI. 2: 384. 1891. 5. elongata A. C. Smith, Contrib. U.S. Natl. Herb. 28: 521. 1932. 5. triloba A. C. Smith, Contrib. U.S. Natl. Herb. 28: 523. 1932. Figures 1G and 15. Epiphytic, lianoid shrubs; mature stems terete, striate, glabrous. Leaves with petioles 4-6 (9) mm long, sub- terete, rugose, weakly puberulent; leaf blades ovate to oblong, 10-13 (22.5) cm long, 3-6 (7) cm wide, base rounded, subcuneate, or subcordate, apex acuminate, glabrous, nitid, 3-5-plinerved with nerves arising from near base and spreading near margins, midrib slightly raised above, prominent beneath, lateral nerves slightly raised to plane above and raised beneath, reticulate vein- lets slightly raised on both surfaces, conspicuous to ob- scure. Inflorescences axillary, racemose, 8-20-flowered; rachis 1 .5-4.5 cm long, subterete, bluntly angled, striate, puberulent; floral bract ovate, 2-3.7 mm long, apex acute-acuminate, glabrous or puberulent at base, margin ciliolate and minutely glandular-fimbriate; pedicels 8-13 mm long, subterete, subrugose, striate, glabrous or pu- berulent; bracteoles nearly base, ovate, 1-2 mm long, apex acute-acuminate, densely puberulent or glabrous, margin sparsely glandular-fimbriate. Flowers with calyx 3-4 mm long, tube subcylindric, sparsely to densely pu- berulous, 1.5-2 mm long, limb 1-2 mm long, spreading- campanulate, often rotate to reflexed, lobes oblong- ovate, 0.5-1 mm long, apiculate, often irregularly divid- ed or fused, the margins often torn or thinner in texture, sinuses acute; corolla thin-carnose, thinly unistratose, tubular but gradually narrowing apically, pentagonal, 16-34 mm long, 3-5 mm diam. near base, glabrous or usually puberulent, dull dark red to scarlet, lobes ca. 1.5-4 mm long, reflexed, oblong, subacute, greenish; stamens alternately 7.5-10 mm and 8.5-11 mm long; filaments 4-6 mm long, equal, glabrous, produced into slender connectives, these minutely pilose with hairs up to 0.2 mm long; anthers alternately 4.8-7 mm and 6-8 mm long, thecae ca. 3.5-4.5 mm long, elongated at base, tubules ca. 1.5-2.7 mm long, the tips lacking ornamen- tation, dehiscing by pores 1.5-2 mm long. Berry 10-15 mm diam., spherical, glabrous, dark blue-black. 78 FIELDIANA: BOTANY Satyria panurensis grows in primary rain forest to premontane cloud forest, 5-850 (1250) m ele- vation (to 1400 m in Nicaragua). It is scattered on the Caribbean slopes of Mesoamerica from south- ern Mexico to western Panama, then it is found along the wet Pacific Coast of Puntarenas Prov- ince, Costa Rica, and Choco Depto., Colombia, and finally crosses the Andes occurring mostly in Amazonian rain forest in an arc from Amapa, Brazil, west through the Guianas and south to Peru; it has also been collected in white-sand sa- vannas in Amazonia. In Costa Rica it is infre- quently collected on the Caribbean slopes includ- ing inundated forest in Lim6n Province, but then it has also been collected a few times on the wet Pacific slopes of the Osa Peninsula. Flowering specimens have been collected in March, June, July, September, and October; fruiting specimens in January, March, August, October, and Novem- ber. Rare/Infrequent. Satyria panurensis is characterized by having axillary and clearly long-racemose inflorescences, puberulent rachis and (usually) corolla, often bluntly pentagonal corolla that is red basally but becoming greenish apically when fresh, and its lowland, primary rain forest distribution. Morpho- logically it is without close relatives, perhaps be- cause of its usual isolation in primary lowland rain forest. The taxa called S. elongata and 5. tri- loba (see A. C. Smith, 1932) are no more than inconsequential variants within this geographical- ly wide-ranging species. Satyria ventricosa Luteyn, Brittonia 53: 440, fig. 1. 2001. Lianoid shrubs climbing to at least 9 m; mature stems terete to subterete, glabrous; twigs subterete, bluntly an- gled, somewhat striate, glabrous. Leaves with petioles subterete, strongly rugose, (6) 9-15 mm long, slightly winged apically due to decurrent leaf blade, glabrous; leaf blades coriaceous, elliptic to lance-elliptic, (15) 19- 27 cm long, (4) 7-14 cm wide, base cuneate, attenuate and shortly decurrent onto petiole to rounded, apex nar- rowing and very short-acuminate, glabrous on both sur- faces but with scattered, minute (ca. 0. 1 mm long), red- dish, glandular fimbriae beneath, 3 (S)-plinerved with inner pair of lateral nerves arising from (1.5) 2-3 cm above base, midrib conspicuously thickened and raised in basal 2-2.5 cm becoming progressively thinner and plane apically above and conspicuously raised beneath, lateral nerves slightly raised basally becoming weakly impressed apically above and raised beneath, reticulate veinlets plane to weakly impressed above and slightly raised beneath. Inflorescences short-racemose, 4-11- flowered, frequently several together, ramiflorous from a callus or at least arising from leafless nodes on older branches (persisting over many years); rachis subterete. sharply to bluntly angled, 0.5-0.9 cm long, moderately densely glandular-fimbriate otherwise glabrous; floral bract continuous with rachis, persistent, strongly spread- ing to reflexed, ovate to elliptic-ovate, 1 .3-2.5 mm long, margin glandular-fimbriate otherwise glabrous; pedicels thin, striate, 19-21 mm long, glabrous; bracteoles nearly basal, ovate-lanceolate, ca. 0.8-2 mm long, apex short- acuminate, margin glandular-fimbriate otherwise gla- brous. Flowers with calyx ca. 2-4.5 mm long, some- times seemingly slightly stipitate, glabrous, tube cylin- dric to bluntly pentagonal, rugose, ca. 1-2.5 mm long, basally apophysate, limb rotate-campanulate, often spreading and reflexed, ca. 1-2 mm long, lobes apicu- late, to ca. 0.5 mm long, sinuses concave; corolla car- nose, bistratose, essentially tubular but bluntly pentag- onal, conspicuously swollen in basal I/.,-1/, (especially noticeable when fresh). 21-36 mm long, 5-J5 mm diam. at base becoming ca. 2-4 mm diam. at throat, glabrous or sparsely short-puberulent in apical half, also bearing few, scattered, glandular fimbriae apically, red at base but becoming green at throat and lobes, lobes erect, ovate, 1.5-3 mm long, apex acute; stamens alternately ca. 8-9 mm and 9-10 mm long; filaments equal, connate into a tube ca. 3-4.5 mm long, glabrous below but cil- iate apically, also abaxial surface of tube glandular-fim- briate; anthers alternately ca. 4.8-6 mm and 5.5-7 mm long, thecae and tubules nearly indistinguishable and about equal in length, tubules apically acuminate, alter- nately somewhat ornate. Berry not seen. Satyria ventricosa is currently endemic to the Continental Divide, premontane cloud forests of central (Veraguas Prov.) and western (Bocas del Toro and Chiriquf Prov.), Panama, 980-1400 m elevation, where it is known from only about six collections. It has not yet been collected in Costa Rica. Flowering specimens have been collected in February and October; fruits are unknown. En- dangered. Satyria ventricosa is characterized by having a cauliflorous, subfasciculate to short-racemose inflo- rescence and by the size, shape, and color patterns of its corolla. It differs from all other species of Satyria in Mesoamerica by its large and ventricose- based corolla (up to 13-15 mm basal diam. when fresh). Both 5. ventricosa and 5. panurensis have corollas that are red at the base and green apically in the throat and lobes, although they differ in sev- eral other features mentioned in the key. Satyria warszewiczii Klotzsch, Linnaea 24: 22. 1851. 5. clonantha Klotzsch, Linnaea 24: 22. 1851. Riedelia warszewiczii (Klotzsch) Kuntze, Rev. Gen. PI. 2: 384. 1891! R. clonantha (Klotzsch) Kuntze, Rev. Gen. PI. 2: 384. 1891. S. ovata A. C. Smith, Contrib. U.S. Natl. Herb. 28: 524. 1932. Illustration: R. L. Wilbur & J. L. Luteyn, Ann. Missouri Bot. Gard. 65: 118, fig. 15. 1978. Figure 13. LUTEYN AND WILBUR: FAMILY #172 ERICACEAE 79 Epiphytic or terrestrial shrubs, sometimes lianoid 1- 2.5 (4) m long; mature stems stout, terete, glabrous, grayish to brown or reddish-brown. Leaves with petioles stout, rugose. (5) 8-22 mm long; leaf blades chartaceous to coriaceous, oblong, ovate-oblong, lance-oblong or el- liptic. (6) 10-25 (29) cm long. (2) 4-10 (14) cm wide, base cuneate, rarely rounded, apex acute to acuminate, glabrous or rarely inconspicuously glandular-fimbriate beneath with trichomes 0.2 mm long or less and drying reddish-brown, 3-(5)-plinerved, midrib and lateral nerves slightly raised in basal '/2-'/, and then apically slightly impressed above, prominently raised beneath, reticulate veinlets slightly raised on both surfaces but usually obscure. Inflorescences short-racemose, 3-10 (30)-flowered, frequently several together, ramiflorous from a callus or at least arising from leafless nodes on older branches (persisting over many years); rachis stri- ate, 5-15 (40) mm long, glabrous; floral bract ovate to triangular, ( 1 .8) 2-3 (4.5) mm long, to 4 mm wide, apic- ulate, inconspicuously ciliate; pedicels terete, often api- cally swollen, 8-20 (25) mm long, glabrous or rarely sparsely puberulent; bracteoles alternate, narrowly tri- angular, 1-1.5 (2) mm long, apex acute, inconspicuously ciliate. Flowers with calyx 6-7.5 mm long, glabrous or deciduously puberulous, tube cylindric to campanulate, 1.5-3 mm long, occasionally slightly enlarged at base, limb thinly coriaceous, 1 .5-4.5 mm long (usually longer and more conspicuous than tube), either spreading-cam- panulate and apically truncate with lobes not obvious to apiculate and at most 1 mm long, or limb spreading apically with lobes oblong-ovate, conspicuous and strongly reflexed, to ca. 2 mm long, sinuses concave; corolla thick-carnose when fresh but often drying thin- membranous, bistratose, cylindric or tapering to slightly flaring throat, occasionally with a somewhat swollen base and apically flaring, (15) 17-35 mm long, mostly (2) 3-4 (6) mm diam., usually bright red in basal 3/4 with apex white, rarely shortly and sparsely puberulent api- cally, lobes oblong, 1-2 mm long, apex acute, white; stamens alternately unequal, 6.5-9.5 mm and 7.5-1 1 mm long; filaments (1.5) 3-5 mm long, the apical free portion weakly ciliate and rarely abaxially bearing a few glandular fimbriae; anthers alternately ca. 3.5-6.5 mm and 4.5-8 mm long, thecae alternately ca. 3.8-5 mm and 4.8-6 mm long, tubules 2.7-4 mm long with oval de- hiscence clefts 1 .8-2.5 mm long. Berry 7-8 mm diam., spherical, blue-black but often glaucous. Satyria warszewiczii is found in premontane and montane cloud forest, (100) 300-2500 m el- evation. It is distributed from southern Mexico to Panama. Flowering occurs in all months; fruiting collections March-June and October. Common. Satyria warszewiczii is characterized by having axillary or frequently ramiflorous inflorescences that are short-racemose or arise from a low, pad- like mound; glabrous rachis; glabrous or rarely shortly and sparsely puberulent apically corolla, which is red becoming white apically; and its pre- montane to montane cloud forest habitat. There is considerable morphological variability in the shape and venation of the leaves and in the size of the flowers in this wide-ranging species. The extremes are striking, but only one species can be recognized, S. clonantha and 5. ovata represent- ing but minor variations. Common names in Costa Rica include colmillo and uva. The species is vis- ited by the hummingbird Lampornis calolaema in Costa Rica (pers. observ.; Luteyn, 1998). Satyria species "A*' Terrestrial or sometimes epiphytic shrubs to 2 m tall; mature stems terete to subterete, striate, epidermis crack- ing longitudinally, glabrous, reddish-brown; twigs sub- terete to complanate, conspicuously bluntly angled, of- ten striate, glabrous, reddish-brown. Leaves with peti- oles subterete, rugose, 5-10 (25) mm long, 1.5-2.5 mm diam., glabrous; leaf blades coriaceous, longitudinally conspicuously bullate, lanceolate to sometimes elliptic- lanceolate, (4) 6.5-10 (13) cm long, (1.5) 2-3.5 (5) cm wide, base cuneate, apex acuminate to short-acuminate, glabrous, deciduously and shortly glandular-fimbriate beneath with usually only blackened bases of trichomes present on mature leaves, 5 (7)-plinerved with inner pair of lateral nerves arising at or a little below the middle, midrib thickened and raised in proximal 3-4 cm espe- cially basal 1 .5 cm then plane to slightly impressed api- cally above and raised beneath, lateral nerves strongly impressed above for entire length and conspicuously raised beneath, reticulate veinlets slightly impressed above but obscure beneath. Inflorescences usually aris- ing from axils of leafy nodes (rarely, when epiphytic and in forest canopy, arising from leafless nodes), short-ra- cemose, (3) 6-10 (14)-flowered, circumscribed at base by a series of few, broadly ovate, acute, marginally glan- dular-fimbriate bracts to ca. 2 mm long and 1-1.5 mm wide; rachis subterete, sharply angled, very congested, ca. 3-6 mm long, glabrous; floral bract narrowly ovate- lanceolate, 2.3-4 mm long, ca. 1 mm wide, apex acu- minate, glabrous to ciliate, margin glandular-fimbriate with reddish-brown trichomes <0.5 mm long; pedicels thin, angulate, abruptly swollen at tip and there often hypocrateriform, 15-17 mm long, glabrous or scattered white-puberulent to short-pilose especially at apex, also bearing few, reddish-brown, glandular fimbriae at artic- ulation; bracteoles located medially or submedially, op- posite to subopposite, ovate, 0.8-2 mm long, apex acu- minate, glabrous to ciliate, margin glandular-fimbriate. Flowers with calyx 4-5 mm long, glabrous or white- puberulent, usually separated from pedicel by a 0.5 mm long, constricted stipe after anthesis, tube subcylindric- campanulate, rugose to muricate, ca. 1.7-2.5 mm long, limb spreading-erect, ca. 2.2-3 mm long, lobes trian- gular to ovate, ca. 2-2.3 mm long, sinuses acute, appar- ently sometimes tearing slightly at base; corolla carnose, bistratose, terete, ca. 7-12.5 mm long overall, conspic- uously swollen and globose in basal '/2 where reddish- orange and 4-8.3 mm diam., abruptly cylindrical in api- cal '/2 where white and ca. 2-4 mm diam., glabrous or white-puberulent externally, glabrous or sparsely white- pilose on lobes internally with translucent hairs to 0.4 mm long, also bearing scattered, reddish-brown, glan- dular fimbriae apically ca. 0. 1 mm long, lobes triangular- ovate, 0.5-1 mm long, apex acute; stamens 10, alter- 80 FIELDIANA: BOTANY TABLE 3. Salient morphological differences between Satyria species "A" and 5. allenii. Satyria species "A" Satyria allenii Petiole length (mm) Petiole diameter (mm) Petiole indumentum Leaf blade base Leaf blade margin Leaf venation Inflorescence bracts Inflorescence indumentum Rachis length (mm) Calyx limb shape Corolla length (mm) Corolla shape 5-10(25) 1.5-2.5 Glabrous Cuneate Flat Inner pair lateral nerves arising from middle; midrib thick and raised in basal 3-4 cm 2 x 1-1.5 mm Glabrous 3-6 Spreading-erect 7-12.5 Basally globose, apically cylindric 6-12 3-4.5 Pubescent Rounded Revolute Inner pair lateral nerves arising from' near base; midrib thick and raised at most in basal 5 mm 5X4 mm Densely short-pilose 12-30 Rotate-reflexed 15-17 Cylindric throughout nately ca. 4.6-5.5 mm and 5.3-6 mm long; filaments ca. 2.6-3.2 mm long, glabrous; connectives sparsely ciliate with translucent hairs ca. 0.3 mm long; anthers alter- nately ca. 2.5-3.2 mm and 3.2-3.8 mm long, thecae al- ternately ca. 1 .5-2 mm and 2-2.5 mm long, tubules ca. 1-1.5 mm long, broadening apically and rounded at apex, basically without ornamentation, the dehiscence clefts about as long as tubules overall. Berry not seen. Satyria species "A" is found in lowland rain forest and premontane cloud forest areas in Costa Rica and Panama, (0) 800-1200 (1400) m ele- vation. In Costa Rica it has been collected only once (Cartago: Turrialba, Parque Nacional Barbil- la, Sendero Cientifico Barthon, E. Mora 7972), where it was apparently collected in lowland rain forest at 0 m elevation (fide label). In Panama (Chiriqui: Fortuna Dam site; Veraguas: Cerro Tute; Code: El Cope; Panama: Cerro Campana and Cerro Jefe), it is locally common when found growing terrestrially along open, disturbed road- sides but is rather rare when growing as an epi- phyte in closed canopy. Flowering specimens have been collected in January, February, March, September, and October; fruits are unknown. Lo- cally Common (in Panama). Satyria species "A" is characterized by its small, proportionally long-petiolate; strongly bul- late, lanceolate to elliptic-lanceolate leaves; short- racemose inflorescences found at leafy or leafless nodes; and short corollas that are conspicuously swollen and globose basally and cylindric apical- ly. The swollen nature of the corolla is not easily seen in dried herbarium specimens. The leaves of Satyria species "A" often dry a dark reddish- brown color. There seems to be a positive corre- lation between plants growing in forest with a dense canopy, the epiphytic habit, and inflores- cences arising from leafless nodes. Plants growing in an open habit are terrestrial and have inflores- cences arising from leafy nodes. However, few collections and localities are known for this spe- cies. Floral pubescence (i.e., glabrous vs. short- pilose pedicels, calyces, and corollas) seems to vary even within one population growing epi- phytically on a single tree (at Cerro Jefe). Satyria species "A" is morphologically most similar to and has until now been confused with S. allenii A. C. Smith, a species of central Panama (Cocl6 and Panamd Prov.). Although superficially similar in overall appearance, these two species differ in many features (see Table 3). Its overall relation- ships, however, await further study. The species is visited by the hummingbird Lampornis calo- laema castaneoventis in Panama (pers. observ.). Satyria species "B" Terrestrial shrubs to 2 m tall; mature stems terete, epidermis cracking longitudinally, glabrous; twigs sub- terete, bluntly angled, glabrous. Leaves with petioles te- rete, rugose, 6-9 mm long, 1.5-3 mm diam., glabrous; leaf blades coriaceous, lanceolate to elliptic-lanceolate, 6.5-8 cm long, 2-3 cm wide, base cuneate, apex bluntly short-acuminate, glabrous, weakly and thinly glandular- fimbriate on lower surface, 3 (S)-plinerved with all ve- nation relatively inconspicuous and inner pair of lateral nerves arising up to 2 cm above base, midrib thickened and raised above in basal 1.5-2 cm then weakly raised or impressed apically, raised and conspicuous beneath, lateral nerves weakly impressed aljove and weakly raised beneath, ± inconspicuous on both surfaces, retic- ulate veinlets obscure above and weakly raised to ob- scure beneath. Inflorescences racemose, 7-16-flowered, apparently typically solitary in leafy or leafless nodes but new inflorescences arising from same nodes in suc- cessive seasons; rachises subterete, strongly angled, 0.7- 2 cm long, glabrous; floral bract ovate- lanceolate, 2-2.5 LUTEYN AND WILBUR: FAMILY #172 ERICACEAE 81 mm long, ca. 1-1.5 mm broad, apex acuminate, margin glandular-rimbriate, glabrous; pedicels subterete, striate, 10-13 mm long. 1-1.5 mm diam.. glabrous; bracteoles subopposite, medial, ovate to lanceolate, 1.2-1.5 mm long, apex acuminate, margin glandular-fimbriate, gla- brous. Flowers with calyx ca. 3-5 mm long, glabrous, tube cylindric, ca. 1.5-2.6 mm long, ca. 2.5-4.2 mm diam., limb campanulate, ca. 1.5-2.4 mm long, lobes broadly ovate, apiculate, ca. 0.5-0.8 mm long, ca. 2.5 mm wide at base, sinuses concave or sometimes tearing and then acute; corolla thin-carnose, bistratose, cylindric throughout but slightly bluntly pentagonal, ca. 15-21 mm long, 4-8.5 mm diam. at base, glabrous, rose-red in basal half and white in apical half, lobes oblong-deltate, ca. 1.5-2 mm long, apex acute; stamens 10, alternately ca. 6-6.8 mm and 7-7.8 mm long; filaments ca. 3-3.8 mm long, glabrous or ciliate along apical free portion; connectives marginally short-puberulent; anthers alter- nately ca. 4-5 mm and 4.8-6 mm long, thecae alter- nately ca. 2.4-2.8 mm and 3.3-3.5 mm long, tubules alternately ca. 1.5-2.3 mm and 1.6-2.5 mm long, the apical margins bluntly erose. Berry immature, ± spher- ical, at least 6 mm diam., green. Satyria species "B" is known only from open, windswept slopes along the edge of premontane cloud forest, at the Fortuna Dam region (Chiriqui Prov.), Panama, 1000-1200 m elevation. It has not yet been collected in Costa Rica. Flowering collec- tions have been made in January and February; immature fruits in mid- January. Endangered. Satyria species "B" is characterized by its overall glabrousness; small lanceolate leaves with relatively inconspicuous nervation in general, only the midrib being obvious; short-racemose in- florescences; and corollas that are cylindric throughout their entire length. Morphologically, it is most similar to S. allenii, found in central Pan- ama, that species being short-pilose overall and with much larger and broader elliptic leaves with prominent venation. Sphyrospermum Poepp. & Endl. REFERENCE — A. C. Smith, The genera Sphyro- spermum and Disterigma. Brittonia 1: 203-232. 1933. Slender, often straggly, epiphytic or terrestrial shrubs with pendent branches. Leaves alternate, sub- sessile to petiolate, the blades coriaceous, sometimes very thickly so, often brittle, the margin entire, the ve- nation obscure. Inflorescences axillary, flowers soli- tary or rarely paired; floral bract minute, persistent; pedicels continuous with calyx, slender, pendent, api- cally swollen; bracteoles 2, basal, deciduous. Flowers 4-5-merous, without odor; calyx synsepalous, tube subglobose or obconic, limb suberect; corolla sympet- alous, aestivation apparently valvate, campanulate, cy- lindric (tubular), to infundibuliform; stamens usually as many or twice as many as corolla lobes, equal or alternately slightly unequal; filaments distinct, usually longer than anther, usually somewhat sigmoid at base; connectives lacking disintegration tissue or spurs; an- thers inserted abaxially near base, membranous, lack- ing awns, thecae minutely papillate, tubules 2, distinct, as long and wide as or longer than thecae, dehiscing by introrse, oval clefts; pollen grains in tetrahedral tet- rads, lacking viscin threads; nectariferous disc annular- pulvinate, but not apparent; ovary inferior, the ovules numerous, the placentation axile but often appearing central or even basal, the partitions soon decaying; style filiform, about as long as corolla, rarely slightly exserted, glabrous; stigma truncate. Fruit a juicy berry, subglobose or ellipsoid, translucent white to lavender or purplish, glabrous or pubescent, when immature dry and brittle, when mature the pericarp thin-membra- nous; seeds elongate, striate, the embryo green and clearly visible through the endosperm. Sphyrospermum is a genus of about 22 species and ranges from southern Mexico through the highlands of Central America and the Andes of South America to Bolivia, east to French Guiana, and in the Caribbean from Haiti to Trinidad. Six species are currently known to occur in Costa Rica. Sphyrospermum is characterized by pedicels continuous with the calyx and usually thin and pendulous, terete calyces, usually small corollas, inferior ovary, thin-walled fruits, and seeds with green embryos. It is related to Vaccinium and Di- ogenesia Sleumer, but details of the relationships are not as yet clear. Key to the Species of Sphyrospermum la. Leaf blades linear 5. linearifolium 1 b. Leaf blades variously shaped but never linear 2 2a. Pedicels 1-4 (8) mm long; corolla dark red or maroon, 7-10 mm long, campanulate; berry de- pressed-globose, quadrangular, translucent pearl-white 5. dissimile 2b. Pedicels 5-25 mm long; corolla white to pinkish (rarely bright red), 2-9 mm long, cylindric to cylindric-urceolate; berry spherical, terete, pale to dark lilac to translucent white 3 3a. Corolla 2-3.5 mm long 4 82 FIELDIANA: BOTANY 3b. Corolla 4-9 mm long 5 4a. Corolla 2-2.3 mm long; leaf blades elliptic, ovate to lance-ovate; pedicels 7-15 mm long; calyx glabrous although minutely glandular-fimbriate with reddish fimbriae S. standleyi 4b. Corolla 3-3.5 mm long; leaf blades suborbicular-elliptic or broadly subovate-elliptic to somewhat obovate; pedicels 5-9 mm long; calyx densely grayish pilose S. ellipticum 5a. Leaf blades suborbicular to oblong-ovate, barely longer than broad, (0.7) 0.9-1.5 (1.8) cm long, the apex rounded or obtuse; flowers usually extending well beyond the leaves; corolla 4-6 mm long; stamens 4 5. buxifolium 5b. Leaf blades ovate to ovate-lanceolate, usually conspicuously longer than broad, (1) 2-3.5 (5) cm long, the apex obtuse to acute, sometimes shortly and bluntly acuminate; flowers rarely extending beyond the leaves; corolla (4) 5-7 (9) mm long; stamens 4-5 or 8-10 S. cordifolium Sphyrospermum buxifolium Poepp. & 1 mil . Nov. Gen. Sp. PI. 1: 4, pi. 8. 1835. Sophoclesia cordifolia Benth. var. oophylla Kuntze, Rev. Gen. PI. 2:384. 1891. Vaccinium pachycardium Standl., Fieldiana, Bot. 18: 883. 1938. Illustrat- ed: J. L. Luteyn, Fl. Ecuador 54, pi. 8. 1996. Figure 3. Epiphytic or terrestrial shrublets mostly 15-40 cm tall; mature stems pendent to ascendent, terete, slender, ribbed, glabrate; twigs subterete, striate and ribbed, moderately short-pilose with white hairs, glabrate. Leaves with petioles terete, rugose, sometimes striate, 1-2 mm long, puberulous to glabrate; leaf blades cori- aceous, suborbicular to oblong-ovate, (7) 9-15 (18) mm long, 8-20 mm wide, base cordate or rounded, apex rounded or obtuse, sparsely and inconspicuously short- pilose on both surfaces, glabrate, also usually inconspic- uously glandular-fimbriate beneath, often on both sur- faces, obscurely 3 (S)-plinerved from base, midrib and lateral nerves usually weakly impressed above and raised beneath but obscure, reticulate veinlets usually totally obscure, rarely slightly raised on both surfaces when visible. Inflorescences axillary, flowers 1 (2), usu- ally extending well beyond leaves; floral bract ovate, ca. 0.5 mm long, apex obtuse, pilose; pedicels subterete, gradually swollen apically, (5) 10-18 mm long, glabrous to sparingly pilose, sometimes scattered glandular-fim- briate; bracteoles nearly basal, linear-aristate, ca. 0.5 mm long, ciliate. Flowers 4-merous; calyx ca. 2-2.8 mm long, tube obconic to subglobose, 1-1.8 mm diam., pi- lose to glabrate, glandular-fimbriate, limb suberect, ca. 0.5-1 mm long, lobes apiculate, ca. 0.8 mm long, with a tuft of hairs at apex or glabrous, sinuses concave; co- rolla submembranous, cylindric-urceolate, terete, 4-6 mm long, 1.5-3 mm diam., sparsely pilose to glabrous, sometimes only tips of lobes pilose, also usually glan- dular-fimbriate, white to pinkish, lobes strongly reflexed, narrowly triangular, 0.8-1.3 mm long, apex obtuse; sta- mens 4, equal, slightly shorter than corolla, ca. 5.4-5.5 mm long; filaments seemingly callose-thickened at base, 1 .8-3 mm long, sparsely pilose or glabrous; anthers 1 .8- 3.5 mm long, thecae smooth, ca. 1-1.5 mm long, round- ed but sometimes minutely mucronate at base, tubules ca. 1.6-2.2 mm long, dehiscing by short clefts ca. 0.4 mm long. Berry globose, succulent, ca. (6) 10 mm diam., pilose or glabrous, translucent, whitish or with a violet or lavender tinge to purple. Sphyrospermum buxifolium occurs in tropical and premontane wet forest and rain forest and montane and elfin forest, 100-1800 (2200) m el- evation. It is distributed from Nicaragua south to Bolivia and east to French Guiana. In Costa Rica it is widespread in all the cordilleras. Flowering occurs in nearly every month; fruiting in February and May-December. Common. Sphyrospermum buxifolium is characterized by its small, suborbicular to oblong-ovate leaf blades that are rounded or obtuse at the apex; 4-merous flowers borne on long petioles that usually extend well beyond the leaves; corollas only 4-6 mm long; and 4 stamens. It is morphologically very similar to 5. cordifolium, with which eventually it may be shown to be conspecific. The main differ- ences between the species are the usually smaller, suborbicular leaf blades of S. buxifolium that ad- ditionally are usually shorter than the flowering pedicels vs. the larger, ovate and usually subcor- date leaf blades of S. cordifolium that usually are equal to or exceed the flowering pedicels. Sphyrospermum cordifolium Benth.. PI. Hartw. 222. 1846. Sophoclesia cordifolia (Benth.) Klotzsch, Linnaea 24: 29. 1851. Sophoclesia flaccida Seem., Bot. Voy. Herald 164. 1854. Sphyrospermum majus Griseb., Fl. Brit. W.I. 143. 1859. Sophoclesia major (Griseb.) Benth. & Hook.f., Gen. PI. 2: 576. 1876. Sophoclesia cordifolia (Benth.) Klotzsch var. normalis O. Kuntze, Rev. Gen. PI. 2: 384. 1891. Vaccinium valerii Standl., Fieldiana, Bot. 18: 884. 1938. Themistoclesia valerii (Standl.) Sleumer, Bot. Jahrb. Syst. 71: 390. 1941. Illustration: R. L. Wilbur & J. L. Luteyn, Ann. Missouri Bot. Gard. 65: 122, fig. 16. 1978. Figures 2V, 3, and 5. Terrestrial or epiphytic shrublets mostly 2-6 dm tall; mature stems pendent, arching, or scandent, subterete, striate, glabrate; twigs subterete, ribbed or striate, dense- ly to moderately spreading puberulent, glabrate. Leaves LUTEYN AND WILBUR: FAMILY #172 ERICACEAE 83 with petioles terete, rugose, 1-2 (3) mm long, glabrous to pilose; leaf blades ovate to ovate-lanceolate, (1)2- 3.5 (5) cm long, (0.8) 1.5-2.5 (3) cm wide, base round- ed, subcordate or broadly obtuse, apex obtuse or round- ed to acute, sometimes shortly but bluntly acuminate, moderately short-pilose with white hairs on both surfac- es, glabrate. usually also appressed glandular-fimbriate beneath with fimbriae ca. 0.1 mm long, 3 (S)-plinerved from base, midrib and lateral nerves often impressed above (sometimes raised) and raised beneath, often all but midrib obscure beneath, reticulate veinlets usually slightly raised but obscure. Inflorescences axillary, flowers 1 (2), sometimes extending beyond leaves; floral bract ovate, ca. 0.3 mm long, apex obtuse, pilose; ped- icels terete, (10) 15-25 mm long, pilose or glabrous, also often with inconspicuous, appressed, glandular fimbriae apically; bracteoles nearly basal, oblong-aristate, ca. 0.4 mm long, pilose. Flowers 4-5-merous; calyx ca. 3 mm long, usually moderately to densely spreading pilose, oc- casionally sparingly so or glabrous, usually also ap- pressed glandular-fimbriate, tube subglobose or thickly fusiform to short-cylindric, 1-1.5 (2) mm long, limb cy- lindric-campanulate, erect, 1-1.6 mm long, lobes deltate, 0.5-0.8 mm long, apex acute, often with a tuft of hairs at the tip, sinuses rounded; corolla membranous, cylin- dric-urceolate, (4) 5-7 (9) mm long, 1.2-3 mm diam., white to bright red, glabrous or puberulous apically, sparsely and inconspicuously glandular-fimbriate apical- ly, lobes spreading, narrowly triangular, ca. 0.6 mm long, acute; stamens usually either 4-5 or 8-10, slightly shorter than corolla, 6.5-8 mm long; filaments slender, 2-5 mm long, pilose or glabrous; anthers 2-4 mm long, thecae ca. 1-1.6 mm long, tubules ca. 1.5-2.3 mm long, dehiscing by clefts to 1.3 mm long. Berry globose to fusiform, to 15 mm diam., pilose or glabrous, translucent white or white with a violet tinge at maturity. Sphyrospermwn cordifolium is found in premon- tane and montane wet and rain forest to elfin forest, 100-2000 (2700) m elevation. It is distributed from southern Mexico, south to Bolivia, east to French Guiana, and onto the island of Hispaniola. In Costa Rica it is common throughout the Cordilleras de Tilaran, Central, and Talamanca. Flowering occurs in nearly every month; fruiting in November-Jan- uary and June— August. Common. Sphyrospermum cordifolium is characterized by its ovate to ovate-lanceolate leaf blades that are usually conspicuously longer than broad with the apex obtuse to acute or sometimes shortly and bluntly acuminate and stamens 4-5 or 8-10. Usu- ally it has pedicels about equaling the leaf length, and so the flowers easily stand out from the leaves that subtend them. Sometimes, however, leaves are much longer than the pedicels, so the flowers are overshadowed by the subtending leaves. Its relationship to 5. buxifolium is mentioned under that taxon. Sphyrospermum dissimile (S. F. Blake) Luteyn, Opera Bot. 92: 126. 1987a. Vaccinium dissimile S. F. Blake, J. Bot. 53: 271. 1915. Disterigma dissimile (S. F. Blake) S. F. Blake, Proc. Biol. Soc. Wash. 35: 120. 1922. Illustration: J. L. Lu- teyn, Fl. Ecuador 54, color plate 5. 1996. Fig- ure 3. Terrestrial, or more commonly, creeping epiphytic, vi- ney shrublets interwoven amongst bryophytes along ground or high in trees; mature stems wiry, perhaps to 1 m long, 1-1.5 mm diam.; twigs densely spreading pu- berulent, glabrate. Leaves with petioles 1-2 mm long, puberulent; leaf blades coriaceous, ovate, lance-ovate or narrowly to broadly elliptic, 1-3 (4) cm long, 0.8-1.5 (2) cm wide, base rounded to cuneate, apex acute, gla- brous above and beneath, also often sparsely to moder- ately appressed glandular-fimbriate beneath with fimbri- ae 0.1-0.2 mm long. Inflorescences axillary, flowers solitary; floral bract broadly reniform, ca. 0.3-0.4 mm long, apex rounded, glabrous; pedicels 1-2 (4) mm long, weakly hispidulous; bracteoles basal to medial, ovate to broadly oblong, apex obtuse to broadly rounded, 0.8-1 mm long, puberulent. Flowers 4-merous; calyx ca. 2.2- 2.8 mm long, tube campanulate, 0.8-1.2 mm long, densely hirsutulous with hyaline to golden-yellowish hairs 0.1-0.2 mm long, limb erect-spreading, ca. 1.3 mm long, sparsely hirsutulous, lobes broadly deltate, 0.7-0.8 mm long, acute or obtuse, moderately to densely hirsu- tulous; corolla campanulate, 7-10 mm long, 6-9 mm diam., dark red or maroon, sparsely and inconspicuously glandular-fimbriate with fimbriae 0.1-0.2 mm long, lobes erect to spreading, deltate, 1.5-2 mm long, ca. 3.5 mm wide at base, acute; stamens 8, ca. 4.3-5.8 mm long, black to brownish-red; filaments 1.3-4 mm long, sparsely to densely pilose; anthers 2-3.3 mm long, the- cae ca. 1.7 mm long, papillose, tubules 1.6-2.5 mm long, dehiscing by introrse clefts 0.5-0.7 mm long. Ber- ry depressed-globose, bluntly quadrangular, 0.7-1 cm diam., translucent pearl-white. Sphyrospermum dissimile occurs in tropical wet forest to premontane and montane cloud forest, 400-1900 m elevation. The species is geograph- ically widespread from Costa Rica to Ecuador, but it is rarely collected and easily overlooked be- cause it grows among mosses at ground level or high in the canopies of old, moss-covered trees and because of its dark, dull (not brightly colored) red corollas. In Costa Rica it is found in the Cor- dillera Central, especially along the Atlantic slopes. Flowering occurs in nearly every month; fruiting specimens have been collected only in July. Rare/Infrequent. Sphyrospermum dissimile is characterized by its lianoid, thin-stemmed habit; short pedicels; and dark red, campanulate corollas. There has been much controversy over the generic placement of this species. Blake (1915) originally described it in Vaccinium, but he soon transferred it (without 84 FIELDIANA: BOTANY comment) to Disterigma (Blake, 1922). A. C. Smith (1933), followed by Sleumer (1936), moved it back into Vaccinium, mainly because its pedicels and bracteoles were not of the "typical" Disterigma type. There it remained, although Wil- bur and Luteyn (1978) stated that it did not "rest comfortably" in either genus. In 1982, Luteyn collected the first mature fruits in Ecuador and then again in Colombia in 1984, and in 1987 he transferred the species to Sphyrospermum based on the translucent, whitish, thin, membranous pericarp of the mature fruits and the green em- bryos in the mature seeds — characters that are used to distinguish the genus Sphyrospermum (see Luteyn, 1987a, for more details). [Note: R.L.W. favors continued placement of this species within Vaccinium as V. dissimile; J.L.L. maintains it in Sphyrospermum for the reasons given in the dis- cussion above.] Plants of this species are said to be cultivated at "Lankester's Garden" (at Finca Silvestre, C6ncavas de Cartago). Sphyrospermum ellipticum Sleumer, Feddes Re- pert. Spec. Nov. Regni Veg. 41: 121. 1936. Fig- ures 5 and 7. Epiphytic shrubs; mature stems thin, elongate, terete, glabrous; twigs short-puberulent. Leaves with petioles ca. 2 mm long; leaf blades subcoriaceous, suborbicular- elliptic or broadly subovate-elliptic to somewhat ob- ovate, (1.5) 2.8-6 cm long, 2.3-3 cm wide, base rounded or slightly cordate, apex rounded or nearly so, obscurely 3-5-plinerved from base, midrib and lateral nerves slightly raised beneath, reticulate veinlets obscure. Inflo- rescences axillary, flowers solitary; floral bract not seen; pedicels terete, striate, 5-9 mm long, sparsely short-pi- lose, glabrous; bracteoles nearly basal to located medi- ally, aristate, 0.3-1 mm long. Flowers 4-merous; calyx ca. 2 mm long, tube subglobose, abruptly contracted at apex, ca. 2 mm diam., densely grayish pilose, limb cam- panulate, sparsely pilose, lobes deltate, ca. 0.4 mm long, acute, sparsely pilose, sinuses rounded; corolla subcy- lindric, somewhat inflated at middle, 3-3.5 mm long, ca. 1.5 mm diam., scattered short-pubescent and fimbriate, white to pink, lobes ca. 0.5 mm long; stamens 4, ca. 2.8 mm long; filaments ca. 1.6 mm long, subglabrous; an- thers ca. 1 .2 mm long, thecae ca. 0.6 mm long, tubules ca. 0.6 mm long. Berry spherical, pilose, ca. 10 mm diam., translucent white to lavender. Sphyrospermum ellipticum is found in primary rain forest and cloud forest, 30-1000 m elevation. It is distributed from Costa Rica to Ecuador but is extremely rare and poorly collected. The only known collections of the species in Costa Rica in- clude Parque Nacional Corcovado, Kernan & Phil- lips 5/6; Parque Nacional Guanacaste, Estaci6n San Ramon, Morales et al. 2545; Osa Peninsula, Raven 2/657; Osa Peninsula, vie. Jalaca Station, Allen 5307; locality unknown, McAlpin s.n. Flow- ering specimens have been collected April-June; fruiting in April-August. Endangered. Sphyrospermum ellipticum is characterized by its combination of usually elliptic leaf blades with rounded bases and apices, short pedicels, dense and grayish calyx pubescence, and short corollas. It is morphologically similar .to and may be confused with S. standleyi; the differences between these two species are given in the key. Many more col- lections of both are needed in order to determine their circumscription and exact relationships. Sphyrospermum linearifolium Al. Rodr. & J. F Morales in Morales & Rodriguez, Novon 15: 335, fig. 1. 2005. Epiphytic, pendent shrubs; mature stems thin, glabra- te, longitudinally striate; twigs reddish-brown to grayish- brown, shortly and sparsely puberulent. Leaves with pet- ioles 1-2 mm long; leaf blades coriaceous to subcoria- ceous, linear, usually revolute, 3.5-14.4 cm long, 0.2- 0.8 cm wide, base narrowly obtuse, apex narrowly acute, glabrous above, shortly and sparsely glandular-fimbriate beneath, obscurely 2-3-plinerved, midrib sparsely pilose above. Inflorescences axillary, flowers solitary; floral bract not seen; pedicels 1 1-33 mm long, glabrous; brac- teoles deciduous, linear, ca. 0.7 mm long. Flowers with calyx tube subglobose, terete to subterete, ca. 1.2 mm diam., glabrous, limb suberect, 0.8-1 mm long, lobes deltate, 0.2-0.6 mm long, acute, glabrous; corolla ur- ceolate, ca. 5-6 mm long, ca. 3 mm diam., pink to whit- ish-pink, glabrous, lobes triangular, ca. 0.7 mm long, acute; stamens 5, slightly shorter than corolla; filaments ca. 1.8 mm long, glabrous to sparsely villose; anthers ca. 2.3 mm long, tubules dehiscing- by oval pores. Berry spherical to subglobose, 5-8 mm diam.. glabrous, whit- ish to whitish-maroon. Sphyrospermum linearifolium is endemic to Costa Rica, where it has been collected on the Caribbean slopes of the Cordillera Central, at vol- canoes Barva and Turrialba, 700-1500 m eleva- tion. Only three collections are known: Lim6n: Altos de Pascua, cabeceras Quebrada Linda, 1 April 2001, Morales 7823 and 1 1 July 2003, Mo- rales & Callejas 9426, and Heredia: Parque Na- cional Braulio Carrillo, falda Atldntica del Volcan Barva, 28 March 2002, Morales 9772. Flowering specimens have been collected in March and April; fruiting collections in Match, April, and July. Endangered. Sphyrospermum linearifolium is characterized by its linear leaves with a usually revolute margin. It is probably most closely related to 5. cordifol- ium but is easily differentiated by leaf morphol- LUTEYN AND WILBUR: FAMILY #172 ERICACEAE 85 ogy. More collections are needed of this rare and distinctive species. [The above description and discussion are based on the protologue; we have not seen the type collection] Sphyrospermum standleyi A. C. Smith, Britton- ia 1: 210. 1933. Figure 5. Straggly, epiphytic or terrestrial shrubs 0.4-0.6 (1) m tall; mature stems subrugose, cinerous. Leaves with pet- ioles subterete, 1.5-2 mm long; leaf blades subcoria- ceous, the surfaces wrinkled, elliptic, ovate to lance- ovate, 2-6.5 cm long, 1.5-3 cm wide, base rounded or obtuse, apex obtuse, 3 (5)-plinerved, midrib impressed to plane (apically) above, lateral nerves impressed to plane but sometimes slightly raised apically, mostly ob- scure with only the inner 2 conspicuous above, reticulate veinlets obscure above, all nerves raised and conspicu- ous beneath. Inflorescences of axillary or ramiflorous, solitary or paired flowers; floral bract not seen; pedicels 7-15 mm long, swollen apically; bracteoles located along basal '/3 of pedicel, subopposite, aristate, ca. 0.2- 0.3 mm long. Flowers 4-merous; calyx 1-2.8 mm long, 1 .5 mm diam., glabrous but minutely glandular-fimbriate with fimbriae 0.1 mm long, reddish, tube ca. 1-2 mm long, limb suberect, ca. 0.8 mm long, lobes apiculate, ca. 0.2 mm long; corolla cylindrical, 2-2.3 mm long, ca. 1 mm diam., white to pinkish, lobes deltate, acute, ca. 0.3 mm long; stamens 4, slightly shorter than co- rolla; filaments ca. 1 .2 mm long; anthers ca. 1 mm long, tubules as long as thecae. Berry subglobose, 3-8 mm diam., translucent white. Sphyrospermum standleyi is endemic to the premontane forest, 900-1500 m elevation in Cos- ta Rica, from the Cordillera de Tilaran (at Mon- teverde, Alajuela Prov.) and the Cordillera de Tal- amanca (at Orosi, Cartago Prov. and in the Canton de Talamanca, Limon Prov.). This species is known from only five collections and is poorly understood. It is not found in Panama as previ- ously reported in the Flora of Panama (Wilbur & Luteyn, 1978: 124), the specimen cited there hav- ing been misidentified. Flowering and fruiting specimens have been collected in March and June only. Endangered. Sphyrospermum standleyi is characterized by its small corolla (2-2.3 mm long), elliptic or ovate to lance-ovate leaf blades, pedicels that are 7-15 mm long, and glabrous calyx, although it may bear mi- nute, reddish, glandular-fimbriae. The species' re- lationships are obscure at this time, but it is prob- ably related to S. ellipticum (see discussion above). Themistoclesia Klotzsch REFERENCE — J. L. Luteyn, New species, new records, and neotypification of some Mesoameri- can Ericaceae. Brittonia 48: 241-249. 1996. [Themistoclesia, pp. 247-248] Epiphytic shrubs. Leaves alternate, usually distichous, petiolate, the blades subcoriaceous to coriaceous, occa- sionally thick-fleshy, the margin entire, the venation ob- scurely plinerved or rarely pinnate. Inflorescences axil- lary, fasciculate, racemose or flowers solitary (in ours); pedicels continuous with calyx (possibly articulate in T. alatd); bracteoles 2, basal, inconspicuous. Flowers (4) 5- merous, without odor; calyx synsepalous, tube obconic or short-cylindric, bluntly angled or rarely strongly winged, limb erect to slightly spreading, lobes minute to well de- veloped; corolla sympetalous, aestivation valvate, ovoid to cylindric, terete or angled but not winged, the lobes usually apiculate, ca. '/IQ-'/S overall corolla length, not strongly reflexed; stamens 4-5 or 8-10, equal or alter- nately slightly unequal, equal to or more than '/2 as long as corolla; filaments distinct; connectives lacking disin- tegration tissue or spurs; anthers lacking awns, thecae smooth, tubules 2 and distinct, or fused into 1 (T. pen- tandra), as wide as and nearly equal to or about two times longer than thecae, these smooth, dehiscing by short, in- trorse clefts or pores; pollen grains in tetrahedral tetrads, lacking viscin threads; ovary inferior; style filiform, in- cluded, glabrous. Fruit a berry; seeds covered by a trans- lucent, mucilaginous sheath, the embryo green. Themistoclesia is a genus of about 25 species distributed from Costa Rica to Bolivia. Four spe- cies are currently known to occur in Costa Rica. It is characterized by pedicels that are continuous with the calyx; an angled to rarely winged calyx; usually small and distichous leaves (in Costa Rica); terete and small corollas; dehiscence by short, terminal or oblique pores; seeds that are covered by a translucent, mucilaginous sheath: and green embryos. It is related to Sphyrosper- mum and Diogenesia Sleumer, but taxonomic re- visions of all three genera are needed before their relationships can be clarified. Key to the Species of Themistoclesia la. Leaves spirally arranged along stems; leaf blades flat, apically acute to obtuse; calyx tube distinctly 4-5- winged; stamens 5 (6, 8 or 10); anther tubule 1 T. pentandra 86 FIELDIANA: BOTANY Ib. Leaves distichously arranged along stems; leaf blades often concave, apically acute or acuminate; calyx tube bluntly 5-angled, not winged; stamens (8) 10; anther tubules 2 2 2a. Corolla 3-4.8 mm long; leaf blades less than 2 cm long, base narrowly obtuse to rounded, often appearing acute due to slightly involute margin T. smithiana 2b. Corolla 5-7 mm long; leaf blades more than 2 cm long, base rounded, subtruncate or subcordate 3 3a. Bracteoles ovate to hemispheric, completely concealing the glabrous pedicels; corolla cylindric, short-pilose throughout; stamens ca. 5 mm long, the filaments short-pilose, the thecae bearing finger- like appendage at base : T. costaricensis 3b. Bracteoles awl-shaped, never concealing the hispidulous pedicels; corolla urceolate, hispidulous apically; stamens 2.5-2.7 mm long, the filaments glabrous, the thecae lacking basal appendage . . T. horquetensis Themistoclesia costaricensis Luteyn & Wilbur, Brittonia 29: 265. 1977. Figure 5. Pendent, epiphytic shrubs; mature stems to 0.5 m long, terete, minutely striate, tan to reddish-brown, his- pidulous with reddish-brown trichomes 0.6-0.8 mm long, glabrate with age. Leaves distichous, with petioles subterete, densely puberulous to short-pilose, 1-1.5 mm long; leaf blades flat or often concave, ovate, (1.6) 2- 2.5 cm long, (0.8) 1-2 cm wide, base rounded or sub- cordate, apex acute or short-acuminate, hispidulous above especially near margin, glabrous beneath, brown above and olive-green or light brown beneath, obscurely 5-plinerved, midrib slightly impressed basally, raised apically above, slightly impressed but inconspicuous be- neath, lateral nerves and reticulate veinlets slightly raised but inconspicuous above, totally obscure beneath. Inflorescences axillary, flowers solitary, with 4, minute, basal bracts; floral bract ovate, ca. 1 .5 mm long, ciliate; pedicels completely concealed by bracteoles, terete, 1- 2 mm long, glabrous; bracteoles basal, ovate to hemi- spheric, ca. 1 mm long, apically hispidulous. Flowers 5-merous; calyx ca. 3.2-4 mm long, glaucous, copiously short-pilose with translucent or reddish tinged trichomes 1-1.2 mm long, tube obconic, bluntly 5-angled, ca. 3 mm long, limb 1-1.5 mm long, campanulate, lobes apic- ulate, obscure; corolla cylindric, narrowed at throat, 5- 6.2 mm long, ca. 2.2 mm diam. near base, short-pilose throughout, white, lobes ca. 1 mm long; stamens 1 0, ca. 5 mm long, equal; filaments 2.8-3.2 mm long, short- pilose over entire length; anthers ca. 2.5-2.8 mm long, thecae ca. 1.2 mm long, with a minute, 0.1 mm long, finger-like appendage protruding from base, tubules 2, distinct, ca. 1.6 mm long, slightly longer than thecae, dehiscing by narrow clefts ca. 1.3 mm long; style 5-6 mm long. Berry spherical, ca. 9 mm diam., short-pilose, light bluish or lilac. Themistoclesia costaricensis is found in pre- montane cloud forest, 1300-1850 m elevation, and is distributed in Costa Rica, adjacent Panama (Chiriqui Prov.), then again along the Panama/Co- lombia border region and from there south to cen- tral Colombia (Valle Dept.). In Costa Rica it is known only from the type locality: Cartago: prop- erty of ICE hydroelectric plant, 15-20 km beyond suspension bridge over the Rio Grande de Orosi, Luteyn el al. 4407. Flowering specimens have been collected in January, March, April, June, July, and December; fruiting in July (only known from one Panamanian collection). Endangered. Themistoclesia costaricensis is characterized by its distichous leaves with flat or concave leaf blades that are greater than 2 cm long and have an acute or short-acuminate apex, bluntly 5-an- gled calyx, ovate to hemispheric bracteoles that completely conceal the glabrous pedicels, cylin- dric and short-pilose corollas, and 10 stamens with the filaments short-pilose and the thecae ap- pendiculate at base. Morphologically, it is similar to T. horquetensis; the rather minor differences that distinguish the two are given in the key. Both species blend in well with their environment be- cause of their straggly, wispy habit with few, sol- itary, tiny greenish-white flowers that are partially hidden by the leaves. Therefore, they are rarely noticed in the field and are poorly collected. Themistoclesia horquetensis Luteyn & Wilbur, Brittonia 29: 266. 1977. Epiphytic shrubs ca. 5 cm tall; mature stems subterete or bluntly angled, minutely striate but becoming smooth, glaucous, pale reddish-brown, hispidulous with translu- cent trichomes 0.2-0.3 mm long, glabrous with age. Leaves distichous, with petioles subterete, flattened adaxially, 2-3 mm long, ca. 1 mm diam., hispidulous with translucent trichomes 0.2-0.3 mm long; leaf blades ovate, somewhat concave, (2.5) 3-4.5 cm long, 1.5-2.5 cm wide, base broadly rounded or subtruncate, some- times involute, apex short-acuminate, apiculate, obscure- ly short-pilose above, especially nearjnargin, glabrous beneath, 3 (S)-plinerved, nerves obscure, slightly im- pressed above, only midrib slightly raised beneath. In- florescences axillary, flowers solitary (rarely 2), with 2- 4, minute, basal bracts; floral bract ovate, 0.3-0.5 mm long, ciliate; pedicels angled, striate, glaucous, 1.5-2.5 mm long, hispidulous; bracteoles supramedial, awl- shaped, ca. 0.5 mm long, ciliate. Flowers 5-merous; ca- LUTEYN AND WILBUR: FAMILY #172 ERICACEAE 87 lyx 3-4.3 mm long, glaucous, sparsely hispidulous with translucent hairs 0.2 mm long, tube obconic, bluntly 5- angled, 2-3 mm long, limb broadly campanulate, 1.5- 1 .7 mm long, lobes apiculate; corolla urceolate, strongly constricted at throat, 5-7 mm long, ca. 2.2 mm diam. near base, glabrous below but upper part of throat and lobes hispidulous, white to pale greenish-white, lobes ca. 0.5-1 mm long; stamens 10, ca. 2.5-2.7 mm long; fil- aments 1.5-2 mm long, glabrous; anthers 1-1.7 mm long inserted near base, thecae ca. 0.7-0.9 mm long, lacking basal appendages, tubules 2, distinct, ca. 0.8 mm long, equaling thecae, dehiscing by broad clefts ca. 0.7 mm long; style ca. 3.5 mm long. Berry ca. 5 mm diam. Themistoclesia horquetensis is found in pre- montane cloud forest, 1200-2750 m elevation. It is endemic to Costa Rica and adjacent Panama (Chiriquf Prov.). In Costa Rica it has been col- lected only twice in Limon Province (Valle de Sil- encio, Davidse et al. 28705, and Caribbean slope between Rio Terbi and Rib Sini, Davidse et al. 29060). Flowering specimens have been collected in January-March, July, September, and Decem- ber; fruiting in January. Endangered. Themistoclesia horquetensis is characterized by its distichous leaves with blades that are greater than 2 cm long and with short-acuminate and apiculate apices, bluntly 5-angled clayx, awl- shaped bracteoles that never conceal the hispidu- lous pedicels, urceolate and apically hispidulous corollas, and 10 stamens with the glabrous fila- ments. Morphologically, it is similar to T. costar- icensis and is discussed with that species. Themistoclesia pentandra Sleumer, Bot. Jahrb. Syst. 71: 392. 1941. Figures 2S and 5. Epiphytic, often slender and straggly shrubs 0.3-0.6 (1.5) m tall, arising from lignotubers; mature stems as- cendent, terete, striate, fissured, grayish to brown or red- dish-brown, moderately to densely spreading short-pu- bescent or short-pilose with hyaline trichomes 0.2-0.6 mm long, glabrate with age. Leaves spirally arranged, with petioles broadly canaliculate adaxially, (1.5) 2-4 mm long, short-pilose; leaf blades elliptic to oblong-el- liptic, flat, (1.5) 2-3.5 (4.5) cm long, (0.8) 1.2-2 (2.5) cm wide, base cuneate, less commonly rounded, apex acute to obtuse, moderately but inconspicuously spread- ing short-pilose to puberulous on both surfaces when young but becoming glabrate or more sparsely pubes- cent, lower surface also inconspicuously moderately glandular-fimbriate and with 1-2 pairs of circular, con- cave, blackish glands ca. 0.2 mm diam. at base, obscure- ly 3-5-plinerved, midrib and lateral nerves slightly raised above and beneath, reticulate veins obscure on both surfaces. Inflorescences axillary, flowers solitary or rarely paired, with 2 or 4, minute, triangular bracts at base, < 1 .5 mm long; pedicels terete, 1 .5-2.5 mm long, puberulous; bracteoles appressed, lanceolate, 1.2-1.5 mm long, puberulous. Flowers 4-5-merous; calyx 5-7 mm long, tube 3.4-4 mm long, conspicuously 4-5- winged, sparingly to moderately beset with minute, glandular trichomes, the wings 1-2 mm wide, pale greenish, conspicuously ciliate with hyaline trichomes 0.2-0.4 mm long, these trichomes also occurring else- where on surface, limb 0.8-1 mm long, lobes minutely apiculate; corolla cylindric to somewhat cylindric-ur- ceolate, narrowly winged over entire length, sharply quadrangular or pentagonal, 3-4.5 mm long, 2-2.5 mm diam. at base, moderately hyaline short-pilose through- out both internally and externally, also bearing scattered, tiny glandular fimbriae, light salmon-orange to orangish- red with pale yellow to cream tips, lobes erect or slightly spreading and exposing tips of stamens, narrowly tri- angular, 1-1.5 mm long, acute, minutely mucronate; sta- mens 5 (6, 8, or 10), 3.2-3.5 mm long, equal or nearly so, about as long as corolla; filaments ca. 1.5 mm long, glabrous or sparingly ciliate, broadened at base, thecae 1-1.2 mm long, tubule 1, conical, 1.5-1.8 mm long, de- hiscing by a single, oblique pore, ca. 0.8-1 mm long; stigma truncate; style 3-3.5 mm long, about as long as corolla. Berry ± ellipsoid-quadrangular, ca. 8 mm long, ca. 10 mm diam., violet to blue-black. Themistoclesia pentandra is found in primary and premontane cloud forests, 700-1800 m ele- vation. It is endemic to Costa Rica and Panama. In Costa Rica, it has been collected fewer than 1 2 times, all from the Caribbean slopes of the cor- dilleras de Tilaran, Central, and Talamanca. Flow- ering specimens have been collected in January, March, April, and July; fruiting in March and April. Rare/Infrequent. Themistoclesia pentandra is characterized by having leaves spirally arranged along the stems, leaf blade apices that are acute to obtuse, a strong- ly 4-5- winged calyx tube, 5 (6, 8, or 10) stamens, and a single anther tubule. While the original de- scription and indeed the specific epithet indicate 5 stamens, most recent collections seen have 8 or 10, although many herbarium specimens lack co- rollas and stamens. It is possible that more than one species is included under this binomial. The interspecific relationships of T. pentandra are un- clear at this time, although it is morphologically very similar to T. alata Luteyn, a species known from the Panama/Colombia border region to NW Ecuador. Both species have strongly winged calyx tubes and single anther tubules. Themistoclesia alata is itself an anomalous species in the genus, seemingly having the unique character of pedicels articulated with the calyces (Luteyn, 1996a). Themistoclesia smithiana (Standl.) Sleumer, Bot. Jahrb. Syst. 71: 392. 1941. Vaccinium smithian- um Standl., Fieldiana, Bot. 18: 833. 1938. Il- lustration: R. L. Wilbur & J. L. Luteyn, Ann. 88 FIELDIANA: BOTANY Missouri Bot. Card. 65: 129, fig. 17. 1978. Fig- ures 2T and 3. Pendent to spreading, epiphytic shrublets mostly 0.2- 0.3 (0.5) m tall; mature stems terete, slender, brown or reddish-brown to grayish, densely to moderately short- pilose with hyaline trichomes 0.2-0.5 mm long, matur- ing to glabrate. Leaves distichous, with petioles terete, densely hispidulous, 0.5-1.5 mm long; leaf blades thick- ly coriaceous, concave, lanceolate, (7) 10-15 (18) mm long, (3) 6-8 mm wide, base narrowly obtuse to round- ed, or sometimes strongly involute thus appearing acute, apex acute to acuminate, often deciduously apiculate, inconspicuously but copiously short-pilose above when young, becoming glabrate, trichomes slender, spreading, ca. 0.2 mm long, beneath glabrous except for reddish- brown, minute, glandular trichomes 0. 1 mm long or less, venation obscure above except sunken midrib, complete- ly obscured beneath. Inflorescences axillary, flowers solitary, with several basal bracts; floral bract ovate, ca. 0.8 mm long, short-pilose; pedicels terete, short-pilose, 1-1.2 mm long; bracteoles linear to linear-oblong, 0.4- 0.6 mm long, reddish-brown, short-pilose. Flowers 5- merous; calyx ca. 5 mm long, copiously short-pilose with hyaline trichomes 0.3-0.6 mm long, tube obconic, terete, ca. 2.5-3 mm long, limb ca. 2 mm long, cam- panulate-rotate, lobes triangular, 1.2-1.5 mm long, con- spicuous, acute to acuminate; corolla terete or obscurely 5-angled, urceolate, broadly ovoid, abruptly and con- spicuously constricted at throat, 3-4.8 mm long, 2.5-3 mm diam. near base, 0.8-1 mm diam. at throat, glabrous in basal 4/5 but short-pilose apically, pale yellow to pale greenish-white or white, lobes linear-oblong, 0.5-0.8 mm long, strongly flaring and reflexed; stamens 8 or 10, equal, ca. 2.5-3.5 mm long; filaments slender, ca. 1.3- 2 mm long, terete, glabrous, attached medially; anthers ca. 1.5-2 mm long, thecae 0.6-1 mm long, straight, tu- bules 2, 0.8-1.3 mm long, dehiscing by short, oval clefts ca. 0.6 mm long; style ca. 3 mm long. Berry ± spher- ical, inconspicuously angled, 5-7 mm long, ca. 4 mm diam., bluish. Themistoclesia smithiana is found in montane cloud forest, 1 100-2000 m elevation. It is endem- ic to the cordilleras Central and Talamanca of Costa Rica and adjacent western Panama. Flow- ering specimens have been collected in Decem- ber-January and July-August; fruiting in July and December. Rare/Infrequent. Themistoclesia smithiana is characterized by its distichously arranged leaves with small (<2 cm long), concave blades that are narrow and involute at the base and acute or acuminate at the apex, bluntly 5-angled calyx tube, small (3-4.8 mm long) corolla that is sharply constricted at the throat, and 8 or 10 stamens. An unusual popula- tion exists in Limon Province (Parque Nacional Cordillera de Talamanca, Canton de Talamanca, Herrera 5915 and 5434), which has very long and narrow, lanceolate, acuminate leaf blades 15-22 mm long and ca. 5 mm wide. The narrowness of the leaf blades is accentuated because the margin is strongly revolute over its entire length; all other features fall within the range of the species. The interspecific relationships of T. smithiana await a revision of the entire genus. Thibaudia Ruiz & Pav. ex J. St.-Hil. Terrestrial or epiphytic shrubs. Leaves alternate, pet- iolate, the blades coriaceous to subcoriaceous, the mar- gin entire (in ours) or crenulate, the venation pinnate or plinerved (in ours). Inflorescences axillary or terminal, glomerulate, subfasciculate, racemose or paniculate (in ours), few- to many-flowered; floral bract small, incon- spicuous; pedicels articulate with calyx (in ours) or rare- ly continuous; bracteoles 2. submedial. Flowers 5-mer- ous; calyx synsepalous, tube short-cylindric or campan- ulate, occasionally narrowly 5-angled or winged, limb erect to slightly spreading apically; corolla sympetalous, aestivation valvate, subcylindric, occasionally 5-angu- late, rarely weakly to strongly winged, rarely slightly zygomorphic, the lobes valvate; stamens 10. equal, of- ten nearly as long as corolla; filaments equal, distinct or connate (in ours); connectives lacking disintegration tis- sue or spurs; anthers equal, lacking awns, thecae smooth or minutely papillate, tubules 2, distinct (in ours) or rare- ly partially connate, about as long and wide as thecae or rarely twice as long, dehiscing by elongate, introrse clefts; pollen grains in tetrahedral tetrads, lacking viscin threads; ovary inferior; style filiform. Fruit a berry; seeds numerous. Thibaudia is a genus with about 70 species dis- tributed from Honduras south to Bolivia and east to Suriname. One species occurs in Costa Rica. It is characterized by small floral bracts, pedicels usually articulate with the calyx, and stamens of similar morphology and equal in length with co- rolla. It is closely related to Cavendishia and per- haps Anthopterus. Thibaudia costaricensis Hoerold, Bot. Jahrb. Syst. 42: 311. 1909. Psammisia rhododelphis K. Schum. ex Werckle, Bol. Fomento Costa Rica 1: 934. 191 1, nom. nud. Illustration: R. L. Wilbur & J. L. Luteyn, Ann. Missouri Bot. Card. 65: 131, fig. 18. 1978. Figures 2P and 12. Usually epiphytic shrubs; mature stems elongate to 2 m long, terete, cinereous, glabrous. Leaves with petioles glabrous, 2-3 (4) mm long; leaf blades chartaceous or thin-coriaceous, oblong, lanceolate-oblong to oblong-el- liptic, (10) 12-24 cm long, (1.7) 3.5-5 (7.5) cm wide, base cuneate and often short-attenuate, apex caudate acuminate, apparently glabrous or with inconspicuous, glandular-fimbriate trichomes on both surfaces, 3-5-pli- nerved, lateral nerves originating near base, midrib and inner pair of laterals strong and conspicuous for entire length, midrib impressed above and prominently raised LUTEYN AND WILBUR: FAMILY #172 ERICACEAE 89 beneath, reticulate veinlets slightly raised on both sur- faces. Inflorescences axillary, ramiflorous, often at leaf- less nodes, usually on branches of previous season, usu- ally paniculate, sometimes racemose then usually with 2-3 (6) racemes per node, 1 5-30-flowered, numerous; rachis slender, 2-5 cm long, laxly and sparsely puberu- lous or glabrous; floral bract deciduous, sometimes ca- ducous, ovate to broadly ovate, 1-1.5 mm long, long- acuminate, margin fimbriate and sometimes also ciliate; pedicels articulate with calyx, subterete, glabrous or pu- berulous, 10-16 mm long, apically slightly swollen, pinkish to rose; bracteoles submedial, deciduous, ovate- lanceolate, 0.8-1.2 mm long, long-acuminate, margin fimbriate and sometimes also ciliate. Flowers with calyx ca. 3-4 mm long, tube campanulate, rugose, ca. 1.7-2 mm long, ca. 2.5 mm diam., sparsely puberulous or gla- brous, pinkish to roseate, limb 1-2 mm long, erect or spreading, lobes 0.5-1 mm long, acute, sometimes car- tilaginous; corolla actinomorphic, membranous, bistra- tose, cylindric, slightly contracted at base and throat, 9- 10 mm long, 3-3.5 mm diam., translucent white, lobes 1-1.5 mm long; stamens ca. 7-9.2 mm long, equal or almost so; filaments membranous, firmly connate into a glabrous tube 2-3 mm long; anthers 5.5-7.2 mm long, thecae 2-3.5 mm long, tubules distinct, 3-3.7 mm long, dehiscing by elongate, oval clefts about '/2 as long as tubule; stigma broadly peltate, ca. 1 mm diam.; style ca. 10-12 mm long, exserted 1-2 mm at maturity, glabrous. Berry spherical, 7-8 mm diam., blue-black. Thibaudia costaricensis is found in cloud for- est, 700-2200 m elevation. It is distributed in Honduras, Costa Rica, and western Panama (Chi- riqui Prov.). Flowering specimens have been col- lected in February-June, October, and December; fruiting in May and June. Widespread. Thibaudia costaricensis is characterized by its paniculate inflorescence (unique among Costa Ri- can ericads), long-lanceolate leaf blades, and small, translucent white corollas. This distinctive and relatively uniform species is the only repre- sentative of the genus in Central America. Sterile collections with long and narrow leaf blades may be confused with Satyria warzsiwiczii. Thibaudia costaricensis is morphologically most similar to T. archeri A. C. Smith (Colombia) and T. albiflo- ra A. C. Smith (Ecuador), the three species having in common paniculate inflorescences; T. costari- censis has smaller leaves and corollas and less robust inflorescences. Utleya Wilbur & Luteyn in Luteyn & Wilbur REFERENCE — J. L. Luteyn & R. L. Wilbur, New genera and species of Ericaceae ( Vaccinieae) from Costa Rica and Panama. Brittonia 29: 255-276, fig. 4. 1977. [Utleya, pp. 267-270] Epiphytic shrubs; stems winged. Leaves alternate, petiolate, the blades coriaceous, the margin entire or cre- nate, the venation pinnate. Inflorescences axillary, ra- cemose, few-flowered; floral bract small, inconspicuous; pedicels articulate with calyx; bracteoles 2, basal. Flow- ers 5-merous, without odor; calyx synsepalous, broadly and conspicuously 5-winged from base to apex, the wings alternating with lobes, tube obconic, limb erect, spreading; corolla sympetalous, aestivation valvate, ur- ceolate, conspicuously 5-spurred, the spurs solid, arising from apical part of corolla tube, vertically oriented, op- posite corolla lobes; stamens 10, equal, nearly as long as corolla; filaments distinct; connectives lacking disin- tegration tissue or spurs; anthers lacking awns, thecae slightly papillate, tubules distinct, about as wide as the- cae, dehiscing introrsely although somewhat latrorsely by elongate clefts over entire length and somewhat onto theca; pollen grains in tetrahedral tetrads, lacking viscin threads; ovary inferior, 5-locular; style filiform; stigma truncate. Fruit unknown, but presumably a berry. Utleya is a monotypic genus, endemic to cen- tral Costa Rica. It is characterized by having winged stems, pedicels articulate with the calyx, calyx wings that alternate with the lobes, fleshy corollas with spurs opposite the lobes, and sta- mens that dehisce by introrse or slightly latrorse, oval slits that extend the entire length of the tu- bules and slightly onto the thecae. Its relationships are uncertain at this time but may lie with Them- istoclesia, although in that genus the pedicels are continuous with the calyx; dehiscence is through terminal pores or short, oblique clefts; and the seeds are covered by a translucent, mucilaginous sheath. [Note: mature fruits are unknown in Utle- ya, although the immature seeds, at least, lack the mucilaginous sheath of Themistoclesia.] The na- ture of the corolla spurs in Utleya is of some in- terest; these spurs are very conspicuous, fleshy, and solid (not hollow) and are situated in the ex- treme apical part of the corolla opposite the lobes. Specimens of Macleania pentaptera Hoerold may sometimes bear apiculate spurs directly on the tip of the corolla lobes, but they are of a very differ- ent type and are in no way comparable to the pro- nounced spurs of Utleya. Utleya costaricensis Wilbur & Luteyn in Luteyn & Wilbur, Brittonia 29: 267, fig. 4. 1977. Fig- ures 2R and 10. Epiphytic shrubs 1 m tall; mature stems glabrous, subterete and usually with narrow, corky wings; twigs glabrous, sharply angled, 3-5-winged with wings to 1 mm broad, reddish-brown; axillary buds with outer pair 90 FIELDIANA: BOTANY of scales subulate, appearing stipular, 2-2.5 mm long. Leaves with petioles 4-12 mm long, sharply angled, canaliculate or carinate adaxially, carinate beneath, gla- brous but with scattered glandular fimbriae; leaf blades broadly elliptic to ovate-elliptic, 2.5-5 (6) cm long, ( 1 ) 1.5-3.5 cm wide, base cuneate and decurrent onto pet- iole, apex short-acuminate, acute or obtuse, margin en- tire or apically crenate. glabrous on both sides but with scattered, reddish, glandular rimbriae, pinnately nerved, midrib very prominent above and raised in the basal '/3-'/2 but plane to slightly impressed apically. carinate beneath, secondary nerves arcuate and anastomosing near margin, slightly impressed above but raised be- neath, reticulate veinlets slightly raised on both sides. Inflorescences racemose, 3-4 flowered; rachis glabrous, 3-6 mm long, sharply angled, with 2-4 tiny (<1 mm long), ovate bracts at base; floral bract ovate, ca. 1 mm long; pedicels subterete, striate, angled, 12-14 mm long, ca. 0.5 mm diam., glabrous but with scattered glandular rimbriae; bracteoles ovate, 1-1.5 mm long, apex acute, margin glandular-fimbriate. Flowers with calyx 5.5-8 mm long, 7-9 mm wide across apical tip including wings, broadly 5-winged, the wings 2-3 mm wide, al- ternate with and produced beyond lobes, glabrous but glandular-fimbriate at base, tube obconic and 2-2.5 mm long, limb spreading and ca. 2 mm long, lobes ca. 0.5 mm long, apiculate: corolla fleshy, urceolate, strongly constricted at throat, pentagonal, ca. 4 mm long. ca. 6 mm diam. including spurs, light pink, glabrous exter- nally but densely pilose internally at throat, the spurs ovate-obtuse, laterally compressed, about twice as long as lobes, ca. 1.5 mm broad at widest point, the lobes slightly reflexed at anthesis, triangular, ca. 1 mm long, white; stamens ca. 3 mm long; filaments distinct, ca. 2 mm long, densely long-pilose in apical 'A; anthers ca. 1.5-1.8 mm long, thecae strongly incurved at base. ca. 1 mm long, tubules ca. 0.5-0.75 mm long; style ca. 2 mm long, included, glabrous. Berry not seen. Utleya costaricensis is endemic to the premon- tane forest, 1500-1800 m elevation. It is known only from the Caribbean slope of central Costa Rica (Cartago Prov., Orosi), from which it has been collected fewer than 12 times. Flowering specimens have been collected in May-July and December; mature fruits are unknown. Rare/In- frequent. Vaccinium Linnaeus REFERENCES — H. O. Sleumer, Die Arten der Gattung Vaccinium L. in Zentral- und Sudameri- ka. Notizbl. Bot. Gart. Berlin-Dahlem 13: 111- 140. 1936. S. P. Vander Kloet, J. L. Baltzer, J. H. Appleby, R. C. Evans, & D. T. Stewart, A re- examination of the taxonomic boundaries of .Sym- physia (Ericaceae). Taxon 53: 91-98. 2004. Shrubs or rarely Irees, often rhi/omatous; axillary buds with outer pair of scales sometimes elongate and appearing stipular. Leaves alternate to pseudo-opposite, persistent, petiolate. the blades with margin entire or ser- rate, the venation pinnate or plinerved. Inflorescences axillary, racemose, or rarely I -2-flowered; pedicels ar- ticulate (in ours) or rarely continuous with calyx; brac- teoles 2. basal. Flowers 4-5-merous, without odor; ca- lyx synsepalous, tube cylindric to globose, lobes rarely obsolete; corolla sympetalous, aestivation imbricate or valvate. cylindric. urceolate, or campanulate, white to pink, greenish, or yellowish-green; stamens 8 or 10, equal, about equaling corolla; filaments equal, distinct or weakly connate at base, equal to or longer or shorter than anthers; connectives lacking disintegration tissue, with or without spurs: anthers equal, lacking awns, the- cae smooth or minutely papillate, tubules 2. distinct, about as wide as thecae. dehiscing introrsely (sometimes obliquely), latrorsely, or terminally by pores or clefts; pollen grains in tetrahedral tetrads, lacking viscin strands: ovary inferior, 4-5 (falsely 10)-locular; style fi- liform, included and about as long as corolla, or rarely slightly exserted. glabrous; stigma small, simple or somewhat capitate. Fruit a spherical berry, crowned by the persistent calyx lobes and capped by the conspicuous nectariferous disc; seeds numerous, sometimes with mu- cilaginous sheath. Vaccinium is a genus with approximately 450 species distributed in the Northern Hemisphere, Mesoamerica, the West Indies, South America. East Africa and Madagascar, Malesia, and Pacif- ica. Ten species are known from or are to be ex- pected in Costa Rica. In Costa Rica, the genus is characterized by having pedicels articulate with the calyx, corollas of united petals, stamens of equal morphology and about equaling the corolla in length, inferior and 4-5 (sometimes falsely 10)- locular ovaries, and baccate fruit. Vaccinium is morphologically heterogeneous and not monophy- letic. In Latin America, Vaccinium is morpholog- ically most similar to Disterigma, the two being easily distinguished by the lack of calyx-clasping bracteoles at the apex of the pedicels in Vaccin- ium, although see Vander Kloet (1985) and Van- der Kloet et al. (2004) for a different opinion. Key to Species of Vaccinium la. Leaf blades with (indistinctly) crenate to crenate-serrate margins, 1-4.7 cm long, (0.3) 1-2 (2.3) cm wide; corolla white to pinkish or red, membranous, unistratose, the aestivation imbricate 2 Ib. Leaf blades with margins entire, (2) 3-8 (20) cm long, (0.8) 1-5 (8.5) cm wide; corolla usually LUTEYN AND WILBUR: FAMILY #172 ERICACEAE 91 yellowish- to creamy-green (rarely suffused with purplish), fleshy (membranous in V. almedae), bistratose, the aestivation valvate 4 2a. Anther connectives spurred V. consanguineum 2b. Anther connectives lacking spurs 3 3a. Corolla appressed- to spreading-puberulous externally and moderately to densely pilose internally; pedicels 6-10 mm long; berry pilose, not glaucous V. talamancense 3b. Corolla glabrous; pedicels 3-4 (6) mm long; berry glabrous, sometimes glaucous V. flohbundum 4a. Calyx limb with 5 saccate spurs opposite and at base of each calyx lobe V. luteynii 4b. Calyx limb terete and lacking spurs 5 5a. Calyx (4) 7-16 mm long, the tube always shorter than the limb, calyx lobes (1.8) 2-5 (6) mm long 6 5b. Calyx 2-4.5 mm long, the tube longer than the limb, except V. furfuraceum, calyx lobes <1.5 mm long 9 6a. Leaf blades conspicuously bullate, the base ± clasping, the apex rounded; corolla cylindric-urce- olate, proportionally less prominent than calyx V. bocatorense 6b. Leaf blades flat (sometimes weakly bullate in V. jefense, sometimes concave in V. monteverdense), the base not clasping, the apex acute to acuminate; corolla broadly cylindric to cylindric-campan- ulate, proportionally more prominent than calyx 7 7a. Corolla 7-10 mm long, internally densely floccose; calyx lobes smooth; anther tubules ca. 1 mm long, glabrous, dehiscing introrsely to terminal V. floccosum 7b. Corolla 10-18 mm long, internally glabrous; calyx lobes conspicuously striate; anther tubules ca. 3.5-5 mm long, long-pilose along inner surfaces, dehiscing latrorsely 8 8a. Leaf blades often slightly concave, (2.4) 3.4-5.2 (8) cm long, the venation relatively inconspicuous; bracteoles 0.8-1.5 mm long; corolla 13-18 mm long, glabrous V. monteverdense 8b. Leaf blades flat but often bullate, (4.5) 8-12 (20) cm long, the venation strong, conspicuous; bracteoles 5-8 mm long; corolla 10-13.5 mm long, hirsute and glandular-fimbriate externally especially on surface of lobes V. jefense 9a. Corolla cylindric to urceolate, 5—8 mm long; staminal filaments about equaling or slightly shorter than anthers, the anther tubules <2 mm long 10 9b. Corolla cylindric, cylindric-campanulate, to cylindric-funnelform, (5) 8-12 mm long (if corolla <8 mm long then color white and the mouth flaring to ca. 5 mm diam., V. almedae); staminal filaments conspicuously shorter than anthers, the anther tubules 3 mm long or longer 11 lOa. Leaf blades acute to rounded at base; calyx lobes 1.2-1.8 (2) mm long V. costaricense lOb. Leaf blades rounded to more typically cordate and amplexicaul at base; calyx lobes 0.5-1.2 mm long V. orosiense lla. Petioles 7-15 mm long; axillary buds with outer pair of scales long- aristate, to 10 mm long; corolla narrowly cylindric to cylindric- campanulate, glabrous, pale green to white, 5-9 mm long, 5-6 mm diam.; anther tubules dehiscing by truncate, terminal pores V. almedae 1 Ib. Petioles 1-4 mm long; axillary buds with outer pair of scales narrowly lanceolate, to 4.5 mm long; corolla broadly cylindric to cylindric-campanulate, glabrous to densely scurfily glandular-squamose externally, green to pale yellowish-green or brownish-lilac (fide label) in V. furfuraceum), 8-12 mm long, 6-10 mm diam.; anther tubules dehiscing by introrse pores or latrorse clefts 12 12a. Leaves and floral parts generally densely fimbriate; leaf blades bullate, basally rounded, the retic- ulate veinlets conspicuously raised beneath; corolla squamosely glandular-fimbriate externally; sta- mens dehiscing by latrorse clefts, the tubules short-pilose along inner surfaces, not much longer than thecae V. furfuraceum 12b. Leaves and floral parts without fimbriae, but leaf blades moderately fimbriate especially beneath; leaf blades flat, basally acute, broadly cuneate or acuminate, the reticulate veinlets usually obscure beneath; corolla essentially glabrous; stamens dehiscing by introrse pores, the tubules glabrous, conspicuously longer than thecae V. poasanum 92 FIELDIANA: BOTANY Vaccinium almedae Wilbur & Luteyn, Sida 21(3): 1607, fig. 1. 2005. Epiphytic shrubs 1-2 m tall; mature stems terete, stri- ate, dull to nitid, glabrous; twigs subterele to complan- ate. bluntly angled to furrowed, glabrous; axillary buds with outer pair of scales subulate, long-aristate to 10 mm long, appearing stipular. Leaves with petioles flattened or shallowly canaliculate adaxially, rugose, 7-15 mm long, 2-3.5 mm diam., glabrous; leaf blades coriaceous, elliptic. 6-18 cm long, 3-7.5 (9) cm wide, base rounded to short-acuminate, sometimes appearing involute in basal V2 cm, apex acute to acuminate, margin entire, gla- brous on both surfaces, 3-5-plinerved. midvein raised in basal '/,-'/, then slightly impressed apically above, lateral nerves ± impressed above but raised beneath, reticulate veinlets slightly impressed above and raised beneath but there indistinct. Inflorescences axillary or ramiflorous. fasciculate to very short-racemose, (2) 4-10-flowered, when ramiflorous the inflorescence apparently usually subtended by a narrowly ovate, acuminate bract to 8 mm long; rachis (when present) to ca. 2 mm long, congested; floral bract small, ovate, 0.6-1.5 mm long, apex long- acuminate, margin ciliate and glandular-fimbriate; pedi- cels slender, 5-10 mm long, 0.2-0.3 mm diam., gla- brous; bracteoles basal, ovate-deltate, 0.5-0.8 mm long, apex long-acuminate, margin ciliate and glandular-fim- briate. Flowers 5-merous; calyx ca. 3-4 mm long, gla- brous, tube cylindric-obconic, ca. 1.5-2.2 mm long, limb thin, broadly flaring to rotate, often reflexed at anthesis, 1.5-1.8 mm long, lobes minute, ca. 0.2-0.4 mm long, sinuses flat to concave; corolla aestivation apparently valvate, membranous, thinly bistratose, often inflating in bud, apparently cylindric to cylindric-campanulate at an- thesis, 5-9 mm long, 2.5-4 mm wide at base but broad- ening apically to ca. 5-6 mm diam., glabrous both ex- ternally and internally, pale green to white with lobes bordered red (fide label), lobes narrowly triangular to deltate, ca. 2.5 mm long, apex acute; stamens 10, 7-8 mm long; filaments 1-2.5 mm long, distinct (possibly united in basal 0.5 mm), ciliate, conspicuously shorter than anthers; connectives lacking spurs, glabrous; an- thers 6-7 mm long, thecae ca. 1.4-2.2 mm long, in- curved and apiculate at extreme base only, tubules dis- tinct, ca. 3-4.8 mm long, glabrous, dehiscing by truncate to slightly flaring, terminal pores; style slightly exserted. Berry immature. 3-4 mm diam.. glabrous. Vaccinium almedae is known from three col- lections made in premontane, wet forest near the Fortuna Dam and Cerro Colorado sites in Chiriquf Province, Panama, 1 150-1500 m elevation. It has not yet been collected in Costa Rica. Flowering specimens have been collected in January; im- mature fruits in late January and April. Endan- gered. Vaccinium almedae is characteri/.ed by its long- aristate, axillary bud scales that appear stipular; short-racemose to fasciculate inflorescences that are sometimes ramiflorous, rotate to flaring calyx limb; minute calyx lobes; pale green to white co- rolla; staminal filaments that are conspicuously shorter than the anthers; and anther tubules that are conspicuously longer than the thecae and that dehisce by truncate to slightly flaring, terminal pores. Its relationships are unclear at this time. Vaccinium bocatorense Wilbur. Sida 1 1 : 441. 1986 | as hocatorensis]. Epiphytic, scandent trtelet; mature stems ± terete, furrowed, epidermis cracking longitudinally; axillary buds with outer pair of scales narrowly lanceolate, 7-9 mm long, abruptly long-acuminate, appearing stipular. Leaves apparently subopposite, with petioles thick, stout, rugose. 2-3 mm long. 3-4 mm diam., short-pu- berulent adaxially. ± glabrous abaxially; leaf blades co- riaceous, ovate to ovate-elliptic. 4-6.5 cm long. 3.5-5 cm wide, base rounded and slightly cordate, often some- what clasping, apex ± rounded, margin entire, glabrous above and moderately but inconspicuously ercct-short- pilose beneath with widely spaced, slender, hyaline tri- chomes 0.2-0.3 mm long, apparently lacking glandular fimbriae, 5 (7)-plinerved with 2-3 pairs of lateral nerves arising within 5-10 mm of base, midrib and lateral nerves impressed above and raised beneath, reticulate veinlets obscure. Inflorescences congregated apically and sometimes appearing terminal, racemose to subco- rymbose, 6-12-flowered; rachis 2-5 cm long, ± terete, moderately and inconspicuously short-pilose; floral bract persistent, lanceolate to lance-ovate. 3-6 mm long, apex acute to acuminate, sparingly puberulent to short-pilose externally; pedicels terete, striate. 10-15 (20) mm long, ca. 1 mm diam., eglandular, moderately to densely pu- berulent with erect hyaline trichomes; bracteoles borne in basal '/„ lanceolate to lance-oblong. 4-6 mm long, 0.6-1.5 (1.8) mm wide, adaxially glabrous and abaxially sparingly puberulent to short-pilose, margin ciliate. Flowers 5-merous; calyx rugose. 7-10 mm long, mod- erately to densely white-pubcrulent with erect hyaline trichomes and also inconspicuously and moderately fer- rugineous, glandular-fimbriate. "purplish-red." tube broadly cylindric. 3.5-4 mm long, fimbriatc and glan- dular-pustulate throughout, limb cylindric-campanulate, 4.5-5.5 mm long, smooth, lobes narrowly triangular to lanceolate, 4-5 mm long. ca. 2-3 mm wide at base, acute, spreading-recurved or reflexed; corolla aestiva- tion valvate, thick and fleshy, bistratose. urceolate to cy- lindric-urceolate, constricted at throat, 7-9 mm long, 6- 7 mm diam., externally glabrous, internally glabrous or nearly so in basal half but increasingly tangled white- pilose apically. lobes broadly deltoid, ca. 2-2.5 mm long. ca. 2.5 mm wide at base, livid red (fide label) externally, exceedingly densely matted-tomcntose inter- nally; stamens 10. ca. 5-6.2 mm long; filaments weakly connate at base. ca. 2.5-3 mm long, ca. 0.5 mm wide at middle, moderately to densely sericeous at margin and on both surfaces especially on connective abaxially. slightly shorter than anthers; connectives lacking spurs; anthers ca. 3-4 mm long, thecae ca. 2.5 mm long, strongly incurved at base, tubules distinct, I -1. 5 mm long, glabrous, dehiscing by obliquely introrse pores; style included, ca. 7 mm long. Berry not seen. Vaccinium bocatorense is known only from the LUTEYN AND WILBUR: FAMILY #172 ERICACEAE 93 type collection made in Bocas del Toro Province, Panama, along the heavily forested headwaters of the Rio Colubre, near the Costa Rica border, 2400-2550 m elevation (L D. Gomez et al. 22368, DUKE, MO). It has not yet been collected within the political boundaries of Costa Rica. The type collection was flowering in March 1984; fruits are unknown. Endangered. Vaccinium hocatorense is characterized by its subopposite leaves with stout, rugose petioles, bullate leaf blades with the apices rounded and the bases rounded to subcordate and often some- what clasping; plinerved venation with the retic- ulate veinlets obscure beneath; elongate calyx with proportionately long lobes; thick and fleshy corolla that is urceolate to cylindric-urceolate and conspicuously constricted at the throat; and sta- mens with introrse dehiscence and glabrous tu- bules. The corolla is proportionally small and in- conspicuous with respect to the calyx. Vaccinium bocatorense keys closest to V. floccosum and V. monteverdense because of their long calyces and calyx lobes, but its overall relationships within the genus are uncertain. The type collection was vis- ited by Bombus ephippiatus (fide label data). Vaccinium consanguineum Klotzsch, Linnaea 24: 64. 1861. V. irazuense Sleumer, Notizbl. Bot. Gart. Berlin-Dahlem 12: 138. 1934. V. consanguineum Klotzsch var. irazuense (Sleu- mer) Sleumer Notizbl. Bot. Gart. Berlin-Dah- lem 13: 126. 1936. Figures 2W, 4, and 8. Shrubs or small trees (0.2) 1-3 (10) m tall; mature stems often compact and densely bushy, stiff, rigid, gla- brous to densely puberulent, the bole (when present) up to 30 cm dbh; axillary buds with outer pair of scales broadly ovate, 1.2-2.5 mm long, apex acute to obtuse, glabrous to short-pilose, ciliate, inconspicuous, not ap- pearing stipular. Leaves with petioles 1.5-3 mm long, often broadly canaliculate adaxially, glabrous or puber- ulent; leaf blades narrowly to broadly elliptic, occasion- ally oblong, (1) 1.5-3 (5.5) cm long, 0.5-1.2 (2) cm wide, usually 2-3 times as long as wide, base acute, apex usually tapering and acute, margin conspicuously callose-thickened and lighter in color than blade proper, shallowly serrulate with teeth mostly (1) 2-4 mm apart, each tooth often tipped with a darkish, glandular callos- ity, glabrous or with scattered glandular trichomes be- neath and moderately puberulous along midrib, often with bluish tint, pinnately nerved with 4-7 secondary nerves per side, all nerves slightly raised on both sur- faces and lighter color than blade proper, secondary nerves and reticulate veinlets often inconspicuous above but usually conspicuous beneath. Inflorescences race- mose, 5-IO-flowered; rachis sharply angled, 1-4 cm long, puberulent; floral bract ovate to oblong, 2-5 mm long, scarious, slightly puberulent or ciliate; pedicels 1- 2 mm long, puberulent; bracteoles subopposite, ovate to oblong, 1.5-3 mm long, scarious, ciliate but otherwise glabrous, entire to sparingly irregularly and minutely toothed. Flowers 4-merous; calyx glabrous, 3-4 mm long, tube campanulate, 1-2.3 mm long, limb 1.3-2.2 mm long, smooth, lobes deltate to broadly ovate, 0.8- 1.5 mm long, apex acute to acuminate, ciliate but oth- erwise glabrous; corolla aestivation imbricate, membra- nous, unistratose, cylindric, 5-7.5 mm long, 2.7-4.5 mm diam., glabrous, creamy white with tinges of red or pink, lobes oblong, 1-1.7 mm long; stamens 8, 4.4-5.7 mm long; filaments distinct, 3-4 mm long, ciliate to pilose; connectives 2-spurred, the spurs originating near apical tip, ascendent, 0.2-0.6 (1) mm long, rarely vestigial; an- thers 2.8-4 mm long, thecae 1.5-1.8 mm long, tubules distinct, 1 .2-2 mm long, glabrous, dehiscing by introrse clefts 0.5-0.7 mm long; style included. Berry 5-6 mm diam., reddish to blue-black, often glaucous. Vaccinium consanguineum is found in montane cloud forest, primary Quercus forest and open pa- ramo with Chusquea, often rooted in ash near vol- canic summits, and bogs, (1300) 1650-3400 (3800) m elevation. It is endemic to Costa Rica and adjacent western Panama. In Costa Rica, it occurs as both a shrub and a tree. [Note: This species was included by Standley (1920-1926, pp. 1101-1 102) as a member of the woody flora of Mexico, but its presence north of Costa Rica was questioned by Sleumer (1936), and the spe- cies was not included by Standley and Williams (1966) in the Flora of Guatemala. The species is known to us only from the mountains of Costa Rica and western Panama.] Flowering and fruiting occur throughout the year, with the preponderance of flowering between January and July. Common. Vaccinium consanguineum is characterized by its pinnately nerved and serrulate-margined leaf blades, racemose inflorescences with small white to pink flowers, spurred connectives (these rarely vestigial), and glaucous berries. The species varies from small, densely branched shrubs to large trees to 10 m tall. Leaf size also varies great- ly with trees often having leaves at the higher end of the range. The small-leaved populations of V. consanguineum look especially like V. floribun- dum. Both species have spurred (or vestigially spurred) anthers, although the Costa Rican popu- lations of V. floribundum lack spurs. Nevertheless, V. floribundum would appear to be its closest rel- ative. The species is visited by bees (pers. ob- serv.). The fruits may be sweet, with good flavor (seemingly the blue-black ones), or more tart and used to make jams (seemingly the red-fruited ones). Common names in Costa Rica include ar- ray an (arrallan) and madrono. 94 FIELDIANA: BOTANY Vaccinium costaricense Wilbur & Luteyn in Lu- teyn & Wilbur, Brittonia 29: 270, fig. 5. 1977. Symphysia costaricensis (Wilbur & Luteyn) vander Kloet, Taxon 53: 97. 2004. Figures 6 and 8. Terrestrial or epiphytic shrubs 1-3 (4) m tall; mature stems grayish to reddish-brown, glabrous; axillary buds with outer pair of scales sometimes narrowly lanceolate, to ca. 4 mm long, abruptly long-acuminate, appearing stipular. Leaves with petioles 1-2 mm long, glabrous; leaf blades coriaceous, sometimes slightly bullate, ovate or lanceolate to elliptic, (2.5) 4-8 (11) cm long, (1.5) 2.5-4.5 (6.2) cm wide, base acutely tapering to rounded, rarely narrowly auriculate, apex acute to shortly and bluntly acuminate, margin entire, slightly thickened and perhaps appearing slightly revolute, glabrous but mod- erately and inconspicuously appressed glandular-fimbri- ate on both surfaces, 3-5 (7)-plinerved and also some- what weakly arcuately veined, basal part of midrib as well as lamina around base colored reddish when fresh and reddish-brown when dry, inner lateral nerves arising in basal 2 cm, midrib thickened and raised above in basal 2 cm then slightly impressed apically, lateral nerves slightly impressed above, reticulate veinlets slightly raised above, all nerves raised beneath but the reticulate veinlets often inconspicuous. Inflorescences often superficially appearing terminal, in umbelliform racemes, 10-15-flowered; rachis 1-2.5 (3) cm long, gla- brous to sparingly and minutely puberulent; floral bract persistent, scale-like, ovate-lanceolate, lanceolate or tri- angular, 1-3.5 mm long, apex long-acuminate, glabrous to puberulent, minutely ciliate; pedicels 10-30 mm long, glabrous to inconspicuously puberulous, often also glan- dular-fimbriate with trichomes ca. 0.1 mm long at artic- ulation; bracteoles located along lower '/3 of pedicel, subopposite, appressed, persistent, lance-ovate to lan- ceolate, 0.8-3 mm long, glabrous to indistinctly ciliate. Flowers 5-merous; calyx 3.5-5 (5.8) mm long, glabrous to densely, minutely puberulent, tube campanulate, nar- rowed and rounded at base, ca. 2-2.5 (4) mm long, limb spreading-campanulate, equal to or more typically short- er than tube, 1 .5-2 (2.5) mm long, smooth, lobes trian- gular, 1.2-1.8 (2) mm long, apex acute, sinuses rounded to obtuse at anthesis but often tearing and then appearing acute; corolla aestivation valvate, fleshy, bistratose, broadly cylindric to somewhat urceolate, 5-8 mm long and 4-7 mm diam. (ca. 8 mm long and 9 mm diam. when fresh), pale greenish-yellow, externally glabrous and internally moderately to densely pilose especially in throat and on lobes with whitish trichomes, lobes erect to spreading or somewhat reflexed, triangular, acute, 1- 2 mm long; stamens 10, ca. 6 mm long; filaments 3-4 mm long, slightly adherent to base of corolla tube, ciliate to short-pilose; connectives short-pilose, lacking spurs; anthers ca. 3-4 mm long, thecae ca. 1.8-2 mm long, markedly incurved and apiculate at base, tubules dis- tinct, 1-2 mm long, glabrous, dehiscing by introrse, slightly oblique or slightly flaring, terminal pores; style about equaling corolla, included or slightly exserted. Berry spherical, 11-16 mm diam., dark maroon to blackish-maroon, or purple. Vaccinium costaricense grows in premontane cloud forest, (700) 1480-2000 m elevation. It is found on the Caribbean watershed of the centra! volcanic cordillera between the northeastern slopes of Volcan Barva (Heredia Prov.) and the northwestern slopes of Volcan Irazii (San Jos6 Prov.) in Costa Rica and Bocas del Toro and Chi- riquf provinces in western Panama. Flowering specimens have been collected in December and March; fruiting in March, July, and December. Rare/Infrequent. Vaccinium costaricense is characterized by its combination of acutely tapering to rounded leaf blade bases, a calyx tube that is longer than the limb, corollas that are pilose internally at the throat, and short anther tubules. There is some notable morphological variation: nearly all the plants from intermediate elevations (1480-2000 m) of the Cordillera Central of Costa Rica have calyces densely puberulent, whereas all specimens from lower elevations (700-800 m) are glabrous. The plants from the Fortuna Dam region of Chi- riquf Province, Panama, have leaf blades often ba- sally acutely tapering and calyces always gla- brous. Two collections from the CentralCordillera (Heredia Prov., Boyle et al. 1095, Wilbur & Lu- teyn 18576) have leaf blades narrowly auriculate, as is also seen in two collections of V. poasanum mentioned below. Vaccinium costaricense has a longer rachis; shorter calyx limb and lobes; smooth, not striate, calyx limb and lobes; and lon- ger anther tubules than its morphologically closest relative V. floccosum, from western Chiriqui Prov- ince, Panama. These differences in the calyx may be influenced by where and how the plants grow, that is, epiphytes in the closed forest vs. terrestrial in exposed, windswept areas (pers. observ.). Vac- cinium costaricense differs from V. orosiense by acutely tapering to rounded leaf blade bases (not cordate and amplexicaul) and shorter calyx lobes (at least in the Costa Rican populations). The characters that distinguish these three species of Vaccinium are admittedly few, and more collec- tions are needed to help clarify their morphologies and relationships. Vaccinium floccosum (L. O. Williams) Wilbur & Luteyn in Luteyn & Wilbur, Brittonia 29: 272. 1977. Hornemannia floccosa L. O. Williams, Brittonia 18: 248. 1966. Symphysia floccosa (L. O. Williams) L. O. Williams, Phytologia 24: 158. 1972. Terrestrial or epiphytic shrubs 1-4 m tall; mature stems slender, glabrous; axillary buds with outer pair of LUTEYN AND WILBUR: FAMILY #172 ERICACEAE 95 scales narrowly lanceolate, 2-5 mm long, long-acumi- nate, glabrous, appearing stipular. Leaves alternate to pseudo-opposite, with petioles 2-4 (6) mm long, slightly winged apically, glabrous; leaf blades coriaceous, ovate to lanceolate or elliptic, (3) 5-9 (1 1) cm long, (1.5) 3- 5 (6.5) cm wide, base long-cuneate to rounded or some- times short-attenuate, apex acute to acuminate, margin entire, slightly thickened, glabrous to moderately ap- pressed glandular-fimbriate above and especially be- neath, indistinctly 3-5 (7)-plinerved and also somewhat arcuately veined with inner lateral nerves arising ca. 1 cm from base, midrib thickened and raised in basal 1 cm then apically weakly impressed above, conspicuous- ly raised beneath, lateral nerves plane to weakly im- pressed above and weakly raised beneath, reticulate nerves weakly raised but obscure on both surfaces. In- florescences loose, umbelliform racemes; rachis gla- brous to sparingly puberulent, 2-10 (20) mm long; floral bract persistent, ovate to lance-ovate or lanceolate, 1- 3.3 mm long, glabrous to sparingly short-ciliate; pedicels thin, 10-20 (25) mm long, glabrous to sparsely puber- ulous. often glandular-fimbriate with trichomes 0.1-0.2 mm long at articulation; bracteoles persistent, medial, subopposite, ovate to lance-ovate, 1-3.2 mm long, gla- brous to inconspicuously ciliate. Flowers 5-merous; ca- lyx 4-7.8 mm long, glabrous, tube campanulate, 2-3 mm long, slightly papillose, base rounded, limb spread- ing-erect, 2-4.8 mm long, longer than the tube, com- monly conspicuously striate (rarely smooth), lobes broadly triangular, 1.8-3.5 mm long, smooth, acute, si- nuses acute; corolla aestivation valvate, fleshy, bistra- tose, broadly cylindric-campanulate, slightly narrowed toward base, widest at mouth, 7-10 mm long, 4-5 mm diam., externally glabrous to moderately appressed spic- ulate, internally densely floccose with a tangle of white, villous trichomes, pale greenish often suffused with red, lobes erect to spreading or reflexed, triangular, 2-2.2 mm long, acute, densely villous-floccose internally; sta- mens 10, ca. 5.2 mm long; filaments slightly adherent to base of corolla, ca. 3-3.2 mm long, ciliate at apical tips; connectives lacking spurs; anthers 3-3.5 mm long, thecae 2.2-2.5 mm long, strongly incurved at base, tu- bules distinct, ca. 1-1.2 mm long, glabrous, dehiscing by slightly introrse to perfectly terminal pores. Berry spherical, 8-10 mm diam., dark purplish. Vaccinium floccosum is endemic to elfin and premontane cloud forests of eastern Costa Rica and western Panama, 1000-2500 m elevation. The species has been collected twice in Costa Rica (Limon Prov.: Caribbean slopes, Canton Tal- amanca, Dtto. Bratsi, Reserva Indigena Talaman- ca, Kivut, J. Bittner 1494 and G. Herrera 5191). Flowering specimens have been collected in Jan- uary-October; fruiting in May-December. Rare/ Infrequent Vaccinium floccosum is characterized by its al- ternate to pseudo-opposite leaves with flat blades, entire margins and plinerved venation, conspicu- ously striate calyx limbs that are longer than the tubes, greenish corollas that are internally densely floccose apically, and anther tubules that are short, ca. 1 mm long, glabrous, and dehiscene introrsely or by perfectly terminal pores. In Panama, leaf morphology is variable with populations from the Boquete region showing leaves pseudo-opposite, coriaceous and inconspicuously petiolate with blades ovate to ovate-lanceolate, apices acumi- nate, and bases rounded or short-attenuate, where- as populations from the Fortuna Dam area show leaves distinctly alternate, thinner coriaceous and more conspicuously petiolate with blades more lance-elliptic, apices longer acuminate, and bases tapering and cuneate. All leaf blades with rounded bases are also pseudo-opposite, whereas leaf blades with cuneate leaf bases may be either dis- tinctly alternate or pseudo-opposite. The synony- my of Vaccinium floccosum, as listed above, in- dicates something of its anomalous characteris- tics. The relationships of this species are as close to the species centered about V. poasanum as they are to the largely West Indian species usually treated as Symphysia racemosa (Vahl) Stearn [= Hornemannia racemosa Vahl., but V. racemosum (Vahl) Wilbur & Luteyn]. The inclusion of all these species within Vaccinium does not signifi- cantly expand the limits of that extremely diverse genus and still seems to be the best treatment for the complex until a worldwide revision of the ge- neric limits is undertaken. Vander Kloet (1985), whose study was based largely on the West Indian "5. racemosa," argued against such an extension of the generic limits of Vaccinium claiming that such an expansion "is tantamount to making the genus a dumping ground for any Vaccinieae of uncertain affinity." In Vander Kloet's experience "the character unique to the genus "Vaccinium" is the absence of hypanthium enlargement prior to anthesis." The species is visited by humming- birds (Luteyn 3756). Vaccinium floribundum Kunth in H.B.K., Nov. Gen. Sp. 3: 266, pi. 251. 1818. Illustrated: J. L. Luteyn, Fl. Ecuador 54: pi. 4. 1996. Figures 3 and 4. Terrestrial shrubs to dwarf, procumbent or scandent subshrubs 0.2-0.8 (1.2) m tall, sometimes arising from lignotubers to 10 cm diam.; mature stems terete, rugose, sometimes "warty" from swollen nodes of fallen leaves, glabrous; twigs subterete to angled, or bluntly angled, striate to ribbed, glabrous to densely short, white pilose or cinereous; axillary buds with outer pair of scales el- liptic-ovate, ca. 1-2 mm long, apex acute or acuminate to rounded, glabrous to puberulent, inconspicuous not appearing stipular. Leaves with petioles subterete, flat- tened adaxially, 2-3 (4) mm long, short-pilose; leaf blades coriaceous, elliptic, ovate to ovate-lanceolate, 96 FIELDIANA: BOTANY 0.4-1.5 (2.5) cm long, 0.3-1.2 cm wide, base cuneate to rounded, sometimes short-attenuate, apex rounded to acute, margin minutely crenate-serrate with each tooth terminating in a short, glandular hair, glabrous to mi- nutely puberulent above or along margins or midrib ba- sally, usually minutely glandular-fimbriate beneath, mid- rib and other nerves and margins often lighter color be- neath, pinnately nerved with 3-5 secondary nerves per side but these usually obscure, reticulate veinlets slightly raised on both surfaces or impressed above and raised beneath. Inflorescences racemose or rarely weakly pa- niculate, 6-10-flowered; rachis subterete, angled, striate, 1-2 (4.5) cm long, glabrous to puberulent; floral bract deciduous, membranous, broadly elliptic to ovate or ob- long, 2.5-4.5 mm long, apex acute to rounded, margin usually curved inward, minutely appressed puberulent on abaxial surface to nearly glabrous, minutely ciliate and occasionally glandularly fimbriate apically; pedicels subterete, angled, striate, 3-4 (6) mm long, glabrous to short-pilose; bracteoles nearly basal, similar to floral bract, nearly half as long as pedicel at anthesis, 2.5-4.8 mm long. Flowers 4-5-merous (sometimes on same plant); calyx ca. 3-4 mm long, tube campanulate to short-cylindric, 1.5-1.8 mm long, glabrous to densely short-pilose, green to purplish, often heavily suffused with pink to red, limb erect-spreading, 0.8-1 mm long, smooth, glabrous, lobes triangular-ovate, acute or rarely obtuse, 1-1.5 mm long, glabrous, sinuses acute; corolla aestivation imbricate, membranous, unistratose, cylin- dric, 5-6.5 mm long, glabrous, white to pinkish, lobes ovate-oblong, ca. 1.5 mm long, obtuse; stamens 8 or 10, 3.8-6 mm long; filaments distinct, 2-2.5 mm long, densely short-pilose; connectives lacking spurs; anthers 2.5-3.5 mm long, thecae 1-1.5 mm long, tubules dis- tinct, 1.5-2 mm long, glabrous, dehiscing by obliquely introrse pores; style about as long as corolla. Berry spherical, 4-6 mm diam., blue-black, glabrous, some- times glaucous. Vaccinium floribundum is distributed in high montane cloud forest, subparamo thickets, and grass paramo, from Costa Rica to northern Ar- gentina. In Costa Rica, it is rare and has been collected only at 2700-3740 m elevation in the Cordillera de Talamanca at Cerro Chirripo, the Sakira summit of the Buenavista massif, Cerro Sabila of the Cerro de la Muerte area, Cerro de las Vueltas, Cerro Echandi along the Costa Rica/ Panama border, and Cerro Kamuk. In Costa Rica, flowering specimens have been collected in Jan- uary, March, July, and August; fruiting collections in July-September and December. Rare/Infre- quent (in Costa Rica and adjacent Panama). Vaccinium floribundum is characterized by its small leaves the blades of which are pinnately nerved and minutely crenate-serrate margined, ra- cemose inflorescences with small white to pink flowers, spurless connectives, and glaucous ber- ries. In Costa Rica the stamens lack spurs, where- as populations in South America have at least ves- tigial spurs. Vaccinium floribundum is most close- ly related to V. consanguineum, as mentioned above. The differences between V. floribundum and Pernettya prostrata are discussed with Per- nettya. Vaccinium furfuraceum Wilbur & Luteyn, Sida 21(3): 1609, fig. 2. 2005. Figure 8. Much-branched, epiphytic liana; mature stems bluntly angled, densely canescent; axillary buds with outer pair of scales ovate, 0.7-1.2 mm long, short-acuminate, densely canescent, not appearing stipular. Leaves with petioles thin, ca. 2-3 mm long, 1-1.2 mm diam., spread- ing pilose; leaf blades coriaceous, ± bullate, ovate, (2.5) 4-6 cm long, (1.2) 1.8-2.5 cm wide, base rounded, apex short-acuminate with the tip itself bluntly acute, margin entire, glabrous above or nearly so except midrib dense- ly canescent at base becoming less dense to glabrate apically and moderately pilose along principal veins, moderately to densely spreading pilose to hirsute be- neath along all nerves with hyaline trichomes up to 0.7 mm long, also with dense (especially noticeable when young), persistently short, thick (squamose), glandular fimbriae, ca. 0.1-0.2 mm long on both nerves and sur- faces, weakly 5-plinerved with inner pair of lateral nerves arising ca. 0.8 cm above base, midrib and lateral nerves impressed above and raised beneath, reticulate veinlets raised on both surfaces. Inflorescences congre- gated apically, sometimes appearing terminal, ± short- racemose but with rachis often contracted and flowers appearing corymbose, 3-10 ( 1 2)-flowered, all parts (in- cluding calyx and corolla) densely and squamosely glan- dular-fimbriate with appressed, tan, tawny to grayish, thickened trichomes 0.1-0.3 mm long, also sparingly to moderately short-pilose with hyaline spreading tri- chomes 0.3-0.6 mm long; rachis (2) 4-8 mm long; floral bract persistent, narrowly lanceolate to oblong-ovate, 2.5-2.6 mm long, ca. 1.2 mm wide, apex acute, glan- dular-fimbriate on lamina and at margin; pedicels cylin- dric, striate, (6) 10-19 mm long, extending to ca. 25 mm long after anthesis; bracteoles near base, lanceolate, ca. 2.8 mm long, 1.6 mm wide, apex long-acuminate, weakly ciliate, weakly glandular-fimbriate marginally. Flowers 5-merous; calyx 3-4.5 mm long, tube cylindric, ca. 1.5 mm long, base inconspicuously bluntly apophy- sate, limb flaring or spreading, ca. 3 mm long, smooth, lobes broadly ovate, short-acuminate, ca. 1-1.2 mm long, tearing at sinuses with age, sinuses acute; corolla aestivation valvate, thin-carnose when fresh but mem- branous when dry, bistratose, cylindric-campanulate, broadening toward throat, 8-12 mm long, ca. 8 mm diam. across throat, brownish-lilac fide label ("cafe" lila"), lobes ovate, 2.5-3 mm long, acute, glabrous with- in; stamens 10, 7-9.5 mm long; filaments distinct, 2-3 mm long, densely short-pilose in apical \ especially at connective, conspicuously shorter than anthers; connec- tives lacking spurs; anthers ca. 7 mm long, thecae ca. 2-2.2 mm long, incurved and slightly apiculate at base, tubules distinct, 3-5 mm long, curving inward apically, short-pilose along inner surfaces near base, dehiscing by latrorse clefts ca. 1-2 mm long; style located to one side, slightly curved apically, slightly exserted; nectariferous disc cupular. Berry not seen. LUTEYN AND WILBUR: FAMILY #172 ERICACEAE 97 Vaccinium furfuraceum is endemic to Costa Rica and is known only from the type collection from the southern slope of the Talamanca Valley, near the border with Panama (Limon: Cant6n de Talamanca, between Rio Sukut and Rio Carbri, Muragubishi, 700 m, G. Herrera 3286). The type collection was flowering in July; fruits are un- known. Endangered. Vaccinium furfuraceum is characterized by its generally densely glandular-fimbriate leaf blades and floral parts; thin-petiolate leaves with blades that are bullate, apically short-acuminate, and ba- sally rounded; plinerved venation with the retic- ulate veinlets conspicuously raised beneath; short calyx lobes; thin and membranous corollas that are cylindric-campanulate; and staminal tubules that are pubescent along the inner surfaces and dehiscent latrorsely. The corollas are proportion- ally conspicuous with respect to the calyx. Its morphological relationships lie near V. montev- erdense and V. jefense, with which it has in com- mon pilose inner surfaces of the staminal tubules and latrorse dehiscence. It also shares densely fimbriate leaves and inflorescence parts with V. jefense, perhaps its closest relative. Vaccinium furfuraceum differs most conspicuously from V. jefense by its possession of an overall shorter ca- lyx (3-4.5 mm long vs. 6-10 mm) and calyx lobes (1-1.2 mm long vs. 4.5-6 mm), inconspic- uously apophysate (not 10-ribbed) calyx tube, and nonstriate calyx limb and lobes. Vaccinium jefense Luteyn & Wilbur, Brittonia 29: 272. 1977. Symphysia jefensis (Luteyn & Wilbur) vander Kloet, Taxon 53: 97. 2004. Epiphyte or arching terrestrial shrubs, sometimes lia- noid to 2 m tall; mature stems subterete, glabrous; twigs subterete, glabrous to densely hirsute apically; axillary buds with outer pair of scales narrowly lanceolate, 2.5- 6 mm long, long-acuminate, hirsute, appearing stipular. Leaves alternate, with petioles terete, rugose, 5-9 mm long, 2-3.5 mm diam., hirsute to glabrate; leaf blades coriaceous, elliptic or ovate-elliptic, (4.5) 8-12 (20) cm long, (2) 3-7 (8.5) cm wide, base rounded and cordate to subcordate, apex acuminate to short-acuminate, mar- gin entire, glabrous above and sparsely to moderately hirsute beneath along principal veins, lower surface also bearing numerous, minute, glandular fimbriae, 5-7 (9)- plinerved (or sometimes weakly pinnately veined with 3 pairs of secondary nerves), midrib conspicuously thickened and raised in basal 1-2 (3) cm then apically impressed above and raised beneath, lateral nerves im- pressed above and raised beneath (i.e., leaves somewhat bullate), reticulate veinlets strongly impressed to slightly raised and conspicuous to more usually inconspicuous above but raised and conspicuous beneath. Inflores- cences axillary, rarely appearing terminal, racemose, sometimes appearing subcorymbose, (6) 8-17-flowered, sometimes 2-3 arising from same node; rachis subterete, 1-1.5 cm long, glabrous or sparsely hirsute, densely glandular-fimbriate; floral bract linear-lanceolate, (2.5) 3-9 mm long, ca. 1 mm wide, conspicuously nerved, hirsute, glandular-fimbriate; pedicels terete, striate, (6) 14-26 mm long, ca. 1 mm diam., sparsely hirsute, densely glandular-fimbriate; bracteoles linear-lanceolate, (2.5) 5-8 mm long, ca. 0.5 mm wide, hirsute, glandular- fimbriate. Flowers 5-merous; calyx 6-10 mm long, sparsely to moderately hirsute, sparsely to densely glan- dular-fimbriate, tube cylindric, conspicuously bluntly and roundedly 10-ribbed or fluted, 2-4 mm long, limb campanulate, 4-6 mm long, lobes oblong to ovate, con- spicuously nerved, (2) 4.5-6 mm long, apex acute, often basally slightly imbricate, sinuses acute; corolla aesti- vation valvate, thin-carnose, weakly bistratose, cylin- dric-campanulate, 10-13.5 mm long, 5.5-10 mm diam., sparsely hirsute and glandular-fimbriate externally es- pecially on lobes, green to pale translucent green, lobes triangular-deltate, 2-4 mm long and wide; stamens 10, ca. 8-10 mm long; filaments distinct, ca. 2.5-4 mm long, conspicuously broad at base, margin densely se- riceous; connective lacking spurs, densely sericeous; an- thers distinct, ca. 7-8 mm long, thecae ca. 3-3.7 mm long, strongly incurved at base, tubules distinct, ca. 3.5- 5 mm long, longer than thecae, curved inward, sparsely long-pilose along inner surface, dehiscing by latrorse clefts ca. 1.5-2 mm long; style included, 9-10 mm long. Berry apparently spherical, ca. 12 mm diam, purple. • Vaccinium jefense is found in premontane and montane rain forest, elfin forest, and dense, moss- draped thickets, from western Chiriqui Province eastward through Panama Province, Panama, (400) 750-1400 m elevation. It has not yet been collected in Costa Rica. Flowering specimens have been collected in January-July and sporad- ically throughout the rest of the year; fruiting col- lections throughout most of the year. Rare/Infre- quent (in Panama). Vaccinium jefense is characterized by its weak- ly bullate leaf blades that are basally rounded and subcordate to cordate, conspicuous leaf nervation, glandular-fimbriate (sometimes densely) inflores- cences, long and narrow bracteoles, elongate and conspicuously striate calyx limb and lobes, sparsely hirsute corollas (especially on the lobes), and anther tubules that are sparsely long-pilose along the inner surfaces and dehisce by latrorse clefts. It resembles several other green-flowered species of Vaccinium, and its morphological re- lationships are mentioned above in the discussion of V. furfuraceum. Vaccinium luteynii Wilbur in Wilbur & Luteyn, Sida21(3): 1611, fig. 3. 2005. Terrestrial or epiphytic shrubs to 4 m tall; mature 98 FIELDIANA: BOTANY stems terete, epidermis cracking longitudinally, gla- brous; twigs subterete to complanate, striate, glabrous; axillary buds with outer pair of scales lanceolate, 2.4-3 mm long, long-acuminate, appearing stipular. Leaves subopposite, with petioles stout, 1-5 mm long, glabrous; leaf blades coriaceous, ovate to oblong-ovate or ovate- elliptic, ca. 5-15 cm long, 3-6 cm wide, base rounded to almost subclasping, apex acute to somewhat acumi- nate, margin entire, slightly thickened and revolute, gla- brous or nearly so on both surfaces, 3-5 (7)-plinerved with lateral nerves arising from basal V4-V3 of midrib, midrib basally much thickened and weakly raised in proximal '/» abaxially, then thinner and weakly im- pressed becoming plane to slightly raised near apex, lat- eral nerves weakly impressed to plane basally abaxially but soon slightly raised apically, reticulate veinlets raised abaxially, all venation somewhat raised beneath. Inflorescences axillary (often appearing terminal), cor- ymbose racemes, 10-13-flowered; rachis bluntly angled, 1 .5-4.5 cm long, glabrous to sparingly and minutely pu- berulent; floral bract persistent, scale-like, triangular or lanceolate, 2-2.5 mm long, midrib prominent, glabrous but often minutely ciliate; pedicels irregularly angulate or ridged, 20-40 mm long, ca. 0.6-1 mm diam., gla- brous; bracteoles persistent, basal, subopposite, lanceo- late to narrowly triangular, 1.2-2 mm long, glabrous or minutely ciliate. Flowers 5-merous; calyx subglobose, 4-4.2 mm long, glabrous, tube subcylindric, deeply and bluntly rugose, ca. 1 mm long, limb ca. 3 mm long, smooth above but broadly and saccately spurred oppo- site and below each lobe, the spurs solid and sometimes slightly and broadly bilobed, calyx lobes triangular, 0.6- 1 mm long, acute, glabrous; corolla aestivation valvate, thick and somewhat fleshy, bistratose, broadly urceolate to urceolate-campanulate, 5-8 mm long, ca. 6 mm diam. just beneath throat and 3.5-4 mm diam. at base (ca. 10 mm long and 8 mm broadest diam. when fresh), green, externally glabrous and internally densely pilose with whitish trichomes especially at throat and on lobes, lobes triangular, ca. 3 mm long, acute; stamens 10, shorter than corolla, ca. 6.5 mm long; filaments 3.5-4 mm long, long-pilose in apical '/2; connectives lacking spurs; an- thers ca. 3.5 mm long, thecae ca. 2-2.5 mm long, lower '/3 strongly incurved upward, tubules distinct, stout, ca. 1 mm long, glabrous, dehiscing by introrse, strikingly oblique pores; style included. Berry not seen. Vaccinium luteynii is endemic to the montane rain forest of western Panama, along the Conti- nental Divide between Bocas del Toro and Chi- riquf provinces, 700-950 m elevation. The species is to be expected in Costa Rica. Flowering spec- imens have been collected in March and October; fruits are unknown. Endangered. Vaccinium luteynii is characterized by its sub- opposite leaves; glabrous leaf blades with rounded to almost subclasping bases; saccately 5-spurred calyx limb; thick, fleshy, broadly urceolate to ur- ceolate-campanulate corollas that are internally densely pilose at the throat and onto the lobes; and glabrous stamens that dehisce introrsely. The interspecific relationships of V. luteynii are uncer- tain at this time, although it is morphologically similar to V. bocatorense and V. floccosum. Vaccinium monteverdense Wilbur & Luteyn in Luteyn & Wilbur, Brittonia 53: 442, fig. 2. 2001. Epiphytic, possibly rarely terrestrial shrubs; mature stems pendent, 1-2.5 m long, arising from a lignotuber, terete, somewhat rugose, blackish-brown; twigs subter- ete, striate, glabrous; axillary buds with outer pair of scales lanceolate, 1.5-3 mm long, apex acuminate, gla- brous to short-pilose at tip only, sometimes ciliate, ap- pearing stipular but not conspicuous. Leaves with peti- oles subterete, narrowly winged often over entire length, rugose, 4-8 mm long, glabrous; leaf blades thick-cori- aceous, weakly bullate, noticeably concave, ovate-lan- ceolate, (2.4) 3.4-8 cm long, (0.8) 1.2-2 (2.7) cm wide, base rounded or obtuse, usually shortly attenuate and narrowly decurrent onto petiole for 2-3 mm, apex blunt- ly and somewhat abruptly acuminate to short-acuminate, margin entire, glabrous, base of lamina beneath with two circular, concave, blackish glands ca. 0.2 mm diam., one on either side of midrib, these sometimes each bearing the base of a single glandular trichome, 3 (S)-plinerved with usually only midrib prominent, midrib weakly im- pressed above and raised beneath, lateral nerves and re- ticulate veinlets generally obscure, usually plane to weakly impressed or weakly raised above (or inner pair of lateral nerves impressed and prominent in basal half) and weakly raised beneath. Inflorescences racemose, 7- 9 (16)-flowered; rachis subterete to bluntly angled, stri- ate, 2-3 (4.7) cm long, glabrous; floral bract minute, deciduous, ovate, 0.8-1.5 mm long, apex acuminate, glabrous; pedicels subterete, striate, 14-19 mm long, glabrous but surrounded by stout, blackish, glandular fimbriae at articulation; bracteoles basal, subopposite, similar to floral bract. Flowers 5-merous; calyx 10-16 mm long, glabrous, tube cylindric to obconical, 4.5-8.5 mm long, striate to slightly ribbed, limb spreading to campanulate, 5.5-7.5 mm long, conspicuously striate, lobes broadly ovate, acute, 2.5-3.5 mm long, apparently tearing with age and the margins callose-thickened, si- nuses sharply acute; corolla aestivation valvate, thinly fleshy, bistratose, broadly cylindric-campanulate, 13-18 mm long, 9-15 mm diam. at throat, green suffused with maroon to reddish-maroon or purplish apically in throat and lobes, glabrous, lobes strongly reflexed, deltate, ca. 3-3.5 mm long, acute; stamens 10, appressed to corolla, ca. 11.5-12 mm long; filaments distinct although slight- ly adherent to base of corolla, ca. 4.7-5 mm long, mar- ginally long-pilose; connectives lacking spurs; anthers ca. 8-8.4 mm long, thecae ca. 3.2-3.4 mm long, with a minute, blunt apiculus at base, tubules distinct, slightly spreading, ca. 4.8-5 mm long, long-pilose along inner surfaces, dehiscing by short, slightly latrorse clefts ca. 1.4-1.5 mm long; style about equaling corolla. Berry unknown, but immature fruit spherical and at least 8 mm diam. with the limb still conspicuous. Vaccinium monteverdense is found in premon- tane cloud forest and lower montane wet forest, 790-1700 m elevation. It is endemic to the Cor- LUTEYN AND WILBUR: FAMILY #172 ERICACEAE 99 dillera de Tilaran (Monteverde region, Puntarenas Prov.) of Costa Rica, where it has been collected 10 times. Flowering specimens have been col- lected in January-May and October; immature fruiting plants in May. Endangered. Vaccinium monteverdense is characterized by having leaf blades with two circular, concave, blackish glands ca. 0.2 mm diam., one on either side of midrib on lower surface; conspicuously striate calyx limb; cylindric-campanulate corolla that is mostly green but suffused with maroon or reddish-maroon apically; stamens that are adher- ent to the base of the corolla; relatively long fil- aments; and anthers with pilose tubules that de- hisce latrorsely. Furthermore, its leaves are often slightly bullate and concave. At Monteverde, V. monteverdense may be confused with Gonocalyx costaricensis, which grows in the same general area, because of its striate calyx limb, somewhat broad corolla, and similar leaves; but G. costari- censis normally has flat leaves that lack the cir- cular basal glands, totally red corollas that are strictly cylindric and much thinner in texture, and stamens with thin, elongated tubules. The rela- tionships of V. monteverdense are uncertain at this time. Vaccinium orosiense Wilbur & Luteyn in Luteyn & Wilbur, Brittonia 29: 275. 1977. Symphysia orosiensis (Wilbur & Luteyn) vander Kloet, Taxon 53: 97. 2004. Figure 6. Terrestrial or epiphytic shrubs 1-2 m tall, sometimes scandent; mature stems terete to subterete, sometimes complanate, epidermis cracking longitudinally, glabrous; twigs subterete, striate, glabrous; axillary buds with out- er pair of scales narrowly lanceolate, 2.5-3.5 mm long, abruptly long-acuminate, appearing stipular. Leaves ap- pearing subopposite especially on young twigs, clasping to amplexicaul, subsessile with petioles ca. 1 mm long, glabrous; leaf blades coriaceous, ovate to ovate-lanceo- late, (2) 4-9 (14) cm long, (1.5) 3-5 (7) cm wide, base rounded to cordate, apex acute, bluntly acute to shortly and bluntly acuminate, margin entire and slightly thick- ened, glabrous to sparsely and inconspicuously ap- pressed, glandular-fimbriate, 5-7 (9)-plinerved with in- ner lateral nerves arising up to 3 cm above base, midrib thick and raised above in basal '/, then weakly raised to plane apically, conspicuously raised beneath, lateral nerves weakly raised on both surfaces but conspicuous beneath, reticulate veinlets weakly raised, conspicuous and of a slightly lighter color on both surfaces. Inflo- rescences umbelliform racemes, often superficially ap- pearing terminal; rachis 1-1.5 cm long, glabrous; floral bract persistent, scale-like, triangular to ovate- lanceolate, 1-2.2 mm long, glabrous to minutely ciliate; pedicels 15-25 mm long, glabrous; bracteoles located along basal V4 of pedicel, subopposite, appressed, persistent, deltate to lance-ovate, 0.8-1.5 mm long, glabrous to inconspic- uously ciliate. Flowers 5-merous; calyx 2-4 mm long, glabrous, tube campanulate to urceolate, 2-3 mm long, limb 0.5-1 mm long, smooth, lobes triangular, 0.5-1.2 mm long, acute; corolla aestivation valvate, fleshy, bis- tratose, cylindric to urceolate, 5-8 mm long, 5-7 mm diam., yellowish-green often suffused with maroon, ex- ternally glabrous but internally moderately to densely pilose with white trichomes in upper half and especially on lobes, lobes spreading or reflexed, triangular, 1-1.2 mm long, acute; stamens 10, ca. 6 mm long; filaments weakly adherent to base of corolla, ca. 3 mm long, vil- lous; connectives lacking spurs; anthers ca. 4 mm long, thecae 1.5-2.3 mm long, incurved and apiculate at base, tubules distinct, 1.5-2 mm long, ca. 0.5 mm diam., gla- brous, dehiscing by slightly introrse, oblique pores; style about equaling corolla. Berry spherical, ca. 12 mm diam., "morada" in color (fide label). Vaccinium orosiense is found in premontane cloud forest, 1 130-1800 m elevation. It is endem- ic to the Caribbean slope of Costa Rica (Cartago, Parque Nacional Tapanti) and adjacent western Panama (Chiriqui Prov., Fortuna Dam area). It is rare in Costa Rica and has been collected only six times. Flowering specimens have been collected in May and December-January; fruiting in Janu- ary, June, and December. Rare/Infrequent. Vaccinium orosiense is characterized by its am- plexicaul leaves with rounded or cordate blade ba- ses, short calyx lobes, corollas that are moderately to densely pilose internally at the throat, and short anther tubules. When fresh, the basal region of the leaf blades is red, as is also the case in V. costar- icense, V. floccosum, and V. luteynii but usually not in V. poasanum. The relationships of V. oro- siense are mentioned under the discussion of V. costaricense. Vaccinium poasanum J. D. Smith, Bot. Gaz. 24: 395. 1897. Symphysia poasana (J. D. Smith) vander Kloet, Taxon 53: 97. 2004. Figures 2X and 8. Terrestrial or epiphytic shrubs or small trees ( 1 ) 2-5 (7) m tall; mature stems and twigs glabrous, bole (when present) up to 30 cm dbh; axillary buds with outer pair of scales narrowly lanceolate, (1.7) 2-4.8 mm long, abruptly long-acuminate, appearing stipular. Leaves with petioles 1-4 mm long, glabrous to hirsutulous; leaf blades coriaceous, ovate, lanceolate or elliptic to ± ob- ovate, 3-8 (10) cm long, (1) 2.5-4 (6) cm wide, base acute to broadly cuneate, very rarely auriculate, apex acute to acuminate, margin entire, glabrous or rarely short-puberulent basally, sparingly to moderately ap- pressed glandular-fimbriate above and especially be- neath, indistinctly 3-5-plinerved and also somewhat ar- cuately veined, midrib thickened and slightly raised in the proximal 1 cm above and then plane to slightly im- pressed apically, lateral nerves weakly impressed above, 100 FIELDIANA: BOTANY all nerves raised beneath but reticulate veinlets some- times obscure. Inflorescences umbelliform racemes, (4) 7-13-flowered; rachis 0.5-2 cm long, glabrous to spar- ingly puberulent; floral bract persistent, lance-ovate to lanceolate, 1.5-2.5 mm long, ciliate; pedicels slender, 7- 15 mm long, sparingly puberulent, often with glandular- fimbriate trichomes 0.1-0.2 mm long at articulation; bracteoles submedial, subopposite, lance-ovate to lan- ceolate, 1-1.5 mm long, ciliate, often persistent. Flow- ers 5-merous; calyx 3.5-5 mm long, glabrous to densely puberulent, tube short-campanulate to turbinate, 2-3.5 mm long and in diam., limb wide-spreading, 1-2 mm long, smooth, lobes broadly and shallowly deltate, 0.2- 1 mm long, apiculate, sinuses concave or flat; corolla aestivation valvate, thinly fleshy, bistratose, broadly cy- lindric, 8-12 mm long, 6-10 mm diam., glabrous, pale yellowish or greenish-white and occasionally tinged with bright rose or pink, lobes erect to slightly recurved, broadly oblong, 2-3.5 mm long, acute to obtuse; sta- mens 10, 7.5-8.5 mm long; filaments distinct, 2-2.5 mm long, broadened at base, slightly adherent to base of co- rolla, ciliate, conspicuously shorter than anthers; con- nectives lacking spurs; anthers 6-7 mm long, thecae 1 .8-2.2 mm long, incurved at base, tubules distinct, 3.5- 5.5 mm long, glabrous, dehiscing by slightly introrse, oblique pores; style longer than corolla, slightly exsert- ed. Berry subglobose, 7-10 mm diam., dark purple. Vaccinium poasanum is found in montane cloud forest to subparamo, (900) 1500-3000 m elevation. It is broadly distributed from Guate- mala to Panama. In Costa Rica, it is common in the cordilleras Central, Talamanca, and Tilaran. Flowering specimens have been collected in De- cember-July; fruiting in January-April and July- September. Common. Vaccinium poasanum is characterized by its leaves with acute to broadly cuneate blade bases and acute to acuminate apices, apiculate calyx lobes, broadly cylindric corollas, staminal fila- ments that are conspicuously shorter than the overall anthers, and elongate anther tubules. Throughout the extensive range of this species (Guatemala to Panama), the shape of the leaf base is very stable, that is, narrowing and acute to broadly cuneate. Two recent collections, however, from the Cordillera Tilardn (Monteverde area), Luteyn et al. 15234 and 15409, have leaf bases that are auriculate; otherwise, the characters are those of V. poasanum. This extremely rare variant of leaf base shape, uncommon in Ericaceae as a whole, would, under normal circumstances, prob- ably tempt us to recognize it formally were it not for the fact that it has also been noticed in two collections of V. costaricense from the Cordillera Central (mentioned above). Like many other of the green-flowered species of Vaccinium, the in- terspecific relationships of V. poasanum are un- certain. It is, however, morphologically similar to another unpublished new species endemic to cen- tral Panama (Cerro Gampana, Panama Prov.) that differs by having shorter and narrower leaf blades (2-4.7 X 1.2-2.4 cm vs. 3-10 X [1] 2.5-6 cm) with rounded bases (vs. bases acute, broadly cu- neate or acuminate), 1-3-flowered racemes (vs. [4] 7-13-flowered), puberulous to short-pilose pedicels and rachises (vs. glabrous to sparingly puberulent), a cylindric calyx tube that is striate and bluntly 5-10-ribbed when fresh (vs. short- campanulate to turbinate, terete, and smooth), 4- 5 mm long calyx lobes (vs. 0.2-1 mm long), an- ther thecae that nearly equal the anther tubules (vs. conspicuously shorter), and latrorse dehis- cence by means of elongate clefts (not slightly introrse by oblique pores). Vaccinium poasanum is visited by the hummingbird Selasphorus scin- tilla in Costa Rica (pers. observ.; Luteyn, 1998). Vaccinium talamancense (Wilbur & Luteyn) Lu- teyn, Brittonia 53: 444. 2001. Macleania lala- mancensis Wilbur & Luteyn in Luteyn & Wil- bur, Brittonia 29: 263, fig. 3. 1977. Terrestrial shrubs 0.5-1 m tall, without lignotubers; mature stems terete, striate, glabrous; twigs subterete, bluntly and broadly complanate, glabrous or puberulent to densely short-pilose, glabrate; axillary buds with out- er pair of scales lanceolate, ovate to broadly ovate, 3-6 mm long, up to 3 mm wide at base, long-acuminate, short-pilose or ciliate, appearing stipular but not con- spicuous. Leaves imbricate, with petioles thick, rugose, 2-3 mm long, flattened or broadly canaliculate adaxially and slightly winged throughout, glabrous or moderately puberulous to short-pilose; leaf blades coriaceous, ob- ovate to more typically oblong-elliptic, 2.5-4.7 cm long, 1.2-1.8 (2.3) cm wide, base rounded, apex acute or rounded but with a 1-3 mm long apiculus, margin in- distinctly and minutely serrulate, margin a lighter color and seemingly thinner than lamina proper, essentially glabrous above except midrib finely whitish-appressed- pilosulous near base and margin inconspicuously ciliate especially basally, beneath indistinctly and sparingly whitish-appressed-pilosulous at base and also with 1-2 reddish, circular glands, entire surface beneath also bear- ing appressed, reddish-brown, glandular fimbriae ca. 0.2 mm long, pinnately nerved with 3-4 arcuate-ascending secondary nerves per side, midrib slightly thickened and raised basally but impressed apically, secondary nerves impressed above, all nerves prominent and raised be- neath and of a lighter color than lamina proper. Inflo- rescences racemose, 10-15-flowered; rachis strongly and sharply angled, 1-2 (4) cm long, softly short, white pilose; floral bract appressed, linear to lanceolate, 3-4 mm long, puberulent; pedicels 6-10 mm long, softly pi- losulous or puberulous, bearing a series of glandular fim- briae apically at articulation; bracteoles basal, appressed, linear to narrowly lanceolate, 2-3 mm long, puberulent, margin glandular-fimbriate. Flowers 5-merous; calyx 4.5-5 mm long, tube campanulate to cylindric, ca. 2.5- LUTEYN AND WILBUR: FAMILY #172 ERICACEAE 101 3 mm long and diam., moderately puberulous with whit- ish trichomes ca. 0.1-0.2 mm long, limb slightly flaring to campanulate-spreading, 2-2.8 mm long, smooth, lobes triangular, 1.2-1.8 mm long, acute, puberulous ex- ternally and internally, sinuses concave; corolla aesti- vation apparently valvate, membranous (weakly mem- branous when fresh), cylindric, terete, 8-13 mm long, 4-6 mm diam., appressed- to spreading-puberulous ex- ternally with moderately abundant hyaline trichomes ca. 0.4 mm long, moderately to densely pilose with hyaline trichomes internally, pink to red although lobes some- times pale pink to whitish, lobes strongly reflexed, tri- angular, 1.5-2 mm long, densely pilosulous both exter- nally and internally; stamens 10, alternately slightly un- equal, ca. 7-8.8 mm and 7.5 mm long; filaments distinct, alternately slightly unequal, ca. 3.5-4.7 and 4 mm long, glabrous basally but margin with appressed, short-pilose, hyaline trichomes apically, inconspicuously shorter than anthers; connectives lacking spurs; anthers 4.5-5.5 mm long, thecae 1.8-2.8 mm long, strongly incurved at base, tubules distinct, 2.5-3 mm long, glabrous, dehiscing by introrsely oblique pores ca. 0.6-1.4 mm long; style ex- serted. Berry spherical, 8-13 mm diam, short-pilose, blue-black. Vaccinium talamancense is endemic to the Quercus forest and secondary, rocky scrub in the open paramo of Cerro Chirripo, 2590-3460 m el- evation. Flowering plants have been collected in January- April; fruiting plants in March, Septem- ber, and December. Endangered. Vaccinium talamancense is characterized by its leaves with blade margins indistinctly and mi- nutely serrulate, red to pink corollas that are ap- pressed- to spreading-puberulous externally and moderately to densely pilose internally, anther connectives that lack spurs, and a pilose berry that is not glaucous. Its interspecific relationships are uncertain at this time. Vaccinium talamancense was originally described in Macleania by Wilbur and Luteyn (in Luteyn & Wilbur, 1977) but with more complete material and fieldwork was sub- sequently shown clearly to belong in Vaccinium (Luteyn, 2001). The species is visited by hum- mingbirds (pers. observ.). 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Page numbers of illustrations are in boldface. The text does not include new names or combinations. Acari acrocarpus moss Agaricales Andromeda mexicana Andromeda prostrata anonillo Anthopterus 19, 21, 22 subgen. Anthopterus 22, 23 subgen. Gonandra 22 Anthopterus costaricensis 4, 22, 23 Anthopterus pterotus 23 Anthopterus revolutus 4, 6, 19, 22, 23 Anthopterus schultzeae ants 4 apophysate 1 apophysis arbutoid ectomycorrhizal associa- tions Arctostaphylos sect. Comarostaphylis subgen. Comarostaphylis Arctostaphylos arbutoides var. costaricensis Arctostaphylos costaricensis Arctostaphylos cratericola Arctostaphylos myrtifolia var. foliis-oblongo-ellipticus var. foliis-ovalibus Arctostaphylos rigida arrallan [sic] arraydn 30 Ascidae Azalea indica Befaria 24 Bejaria 1, 2, 8, 23, 24 Bejaria aestuans 2 5, 8, 20, 24 Bejaria glauca 24 Bejarieae bistratose 1, 20 Boletales Bombus ephippiatus Cavendishia 10, 11, 13, 14, 15, 16, 18, 19, 20, 21, 24, 25, 26, 29, 30, 35 sect. Callista 25, 29, 31, 36 sect. Cavendishia 25 sect. Engleriana 25 sect. Quereme 25, 36 sen Cavendishiae 25, 29, 30, 32 ser. Englerianae 25 ser. Imbricatae 25, 29, 32, 35 ser. Lactiviscidae 25, 33, 34, 38 Cavendishia albopicata 38 Cavendishia arizonensis 38 Cavendishia atroviolacea 4, 15, 16, 28, 29, 31 var. atroviolacea 28 var. folsomii 29 Cavendishia aurantiaca 36 Cavendishia axillaris 15, 26, 29, 35 Cavendishia bracteata 4, 14, 15, 18, 26, 29, 30, 33, 35, 38 Cavendishia bullata 31 Cavendishia callista 4, 15, 18, 28, 29, 31 Cavendishia calycina 26, 31, 32, 35 Cavendishia capitulata 11, 26, 30, 32, 33 Cavendishia chiriquiensis 13, 33 var. bullata 27, 33, 34 var. chiriquiensis 27, 33 Cavendishia ciliata 16, 27, 34 Cavendishia complectens 15, 26, 29, 34, 35 subsp. complectens var. complectens 34, 35 var. cylindrica 35 var. striata 35 Cavendishia confertiflora 18, 26, 35 Cavendishia costaricensis 30 Cavendishia crassifolia 30 Cavendishia davidsei 10, 11, 26, 35, 36, 37 Cavendishia endresii 13, 27, 30, 33, 36 Cavendishia fortunensis 5, 27, 38 Cavendishia gaultherioides 29 Cavendishia gomezii 26, 36, 37, 38 Cavendishia glutinosa 36 Cavendishia graebneriana Cavendishia herrerae 4, 16, 27 Cavendishia hoffmannii 30 Cavendishia lactiviscida 13, 27 Cavendishia latifolia 30 Cavendishia laurifolia Cavendishia limonensis 4, 13, 26 Cavendishia linearifolia 4, 19, 27 Cavendishia longiflora 31 Cavendishia luteynii 15, 26 Cavendishia megabracteata var. attenuata var. megabracteata 27 Cavendishia melastomoides 13, 27 var. albiflora 28 var. coloradensis 28 var. melastomoides 28 Cavendishia osaensis 4, 16, 27 Cavendishia panamensis 27, 33, 38 Cavendishia pilosa 32 Cavendishia pterocarpa Cavendishia pseudostenophylla 27, 38 Cavendishia pubescens 26, 30 Cavendishia quercina 10, 26, 37 Cavendishia quereme 13, 18, 26, 29, 37, 38 Cavendishia ruiz-teranii Cavendishia skutchii 30 Cavendishia smithii 30 Cavendishia stenophylla Cavendishia subfasciculata 27 Cavendishia talamancensis 10, 26, 36,37 Cavendishia tuerckheimii Cavendishia veraguensis Cavendishia violacea 36 Cavendishia warszewiczii 30 Cavendishia wercklei 28, 31, 36, 38 Cavendishia zamorensis 35 Chimaphila 1,2, 19 Chimaphila acuminata Chimaphila acuta Chimaphila corymbosa Chimaphila dasystemma Chimaphila guatemalensis Chimaphila maculata 4, 5, 19, 20 Chimaphila mexicana Chimaphila occidentalis Chimaphila umbellata subsp. domingensis subsp. umbellata var. mexicana Chupalon Chupalon complectens 34 Chupalon crassifolia 30 Chupalon endresii 36 Chupalon latifolium 30 Chupalon melastomoides Chupalon quereme Chupalon veraguense Chupalon warszewiczii 30 Chusquea 28, 30 colmillo 30, 38 colmillo de perro 3 1 Comarostaphylis 1, 5, 14, 30 Comarostaphylis .arbutoides 4, 14, 20 subsp. arbutoides subsp. costaricensis Comarostaphylis costaricensis Comarostaphylis chiriquensis Comarostaphylis discolor 5 subsp. discolor 5 INDEX 105 Comarostaphylis sleumeri connective Didonica 21 Didonica crassiflora Didonica pendula 4, 6 Didonica subsessilis Diglossa plumbea 30 disintegration tissue 1 Disterigma 8, 11, 21 Disterigma agathosmoides Disterigma dissimile Disterigma fortunense Disterigma hammelii Disterigma humboldtii 6, 7 Disterigma luteynii Disterigma panamense Disterigma pilosum 8 Disterigma trimerum Disterigma utleyorum 4, 11 Disterigma weberbaueri Doryfera ludovicae Doryfera ludovicae veraguensis el macho 29 Empetraceae 3 Epacridaceae 3 Erica 2 Ericoideae 24 Eugenes fulgens spectabilis 30 fimbriae (adj., fimbriate) Gaultheria 1, 2, 12, 20 Gaultheria barbata Galtheria chiriquensis Gaultheria ciliata Gaultheria cordifolia Gaultheria donnellii Gaultheria erecta 5, 12 Gaultheria glandulifera Gaultheria gracilis 12 var. intermedia Gaultheria hapalotricha Gaultheria hartwegiana Gaultheria hirsuta Gaultheria hirtiflora Gaultheria longipes Gaultheria odorata var. costaricensis Gaultheria parvifolia Gaultheria poasana Gaultheria rigida Gaultheria santanderensis Gaultheria setosa Gaultheria setulosa Gonocalyx 7, 8, 9, 11, 21 Gonocalyx almedae 7 Gonocalyx amplexicaulis Gonocalyx costaricensis 8 Gonocalyx liliae 9 Gonocalyx megabracteolatus Gonocalyx pterocarpus 6, 7, 11 Gonocalyx smilacifolius Hornemmania floccose Hornemmania racemosa Hymenoptera hybridization 20 hypanthium 1 Hypopitys uniflora Lampornis calolaema 30, 35 Lampornis calolaema castaneoven- tis Lampornis castaneoventris calolae- mus 33 Lampornis castaneoventris casta- neoventris 33 Lateropora 14, 21 Lateropora ovata 4, 6, 14 Lateropora tubulifera lignotuber 1 limb 1 lobes 1 macho 29 Macleania 10, 11, 14, 21 subgen. Aponema subgen. Macleania Macleania alpicola Macleania cordata var. linearifolia Macleania cordifolia Macleania costaricensis Macleania glabra Macleania insignis 6, 10, 11 var. linearifolia Macleania irazuensis Macleania linearifolia Macleania megabracteolata Macleania nitida Macleania ovata Macleania pentaptera Macleania racemosa Macleania rupestris 2, 4, 6, 14 Macleania subracemosa Macleania talamancensis Macleania turrialbana madrono Mesostigmata Monotropa 1, 2, 19 Monotropaceae 3 Monotropoideae Monotropa coccinea var. mexicana var. nicaraguensis Monotropa hypopitys Monotropa uniflora 4, 5, 19, 20 var. coccinea muelas muelita Mycerinus Orthaea 21, 25 subgen. Lysiclesia Orthaea panamensis 2, 5 oxalic acid Panterpe insignis 30 Pernettya 1, 2, 8 Pernettya buxifolia Pernettya ciliata Pernettya ciliaris var. alpina Pernettya coriacea Pernettya hirta Pernettya howellii Pernettya oerstediana Pernettya prostrata 4, 5, 8, 20 Pernettya setigera Phaethornis Pinus poison Polyboea crassifolia 30 Polyboea quereme Polyboea veraguensis Polyboea warszewiczii Proclesia warszewiczii 30 Psammisia 1, 17, 21 Psammisia alpicola Psammisia costaricensis Psammisia glabra Psammisia ramiflora 4, 6, 17 Psammisia rupestris Psammisia symphystemona Psammisia albrichianna 17 Psammisia williamsii 17 Pyrolaceae 3 Quercus 28, 30, 36 Rhinoseius 3 Rhinoseius richardsoni Rhododendron 2 Rhododendron indicum Rhododendron simsii 2, 20 Riedelia clonantha Riedelia panurensis Riedelia warszewiczii Rumex Russulales San Juan San Miguel 30 Satyria 2, 17, 21 Satyria allenii Satyria clonantha Satyria elongata Satyria meiantha 19 Satyria ovata Satyria panurensis 5, 19 Satyria species "A" 2, 4 Satyria species "B" 2 Satyria triloba Satyria ventricosa Satyria warszewiczii 4, 17 Selasphorus scintilla Siphon a mini Socratesia melastomoides Sophoclesia cordifolia var. normalis var. oophylla Sophoclesia flaccida Sophoclesia major Sphyrospermum 2, 7, 9, 11, 21 Sphyrospermum buxifolium 7 106 FIELDIANA: BOTANY Sphyrospermum cordifolium 6, 7, 9 Sphyrospermum dissimile 4, 7 Sphyrospermum ellipticum 4, 9, 11 Sphyrospermum linearifolium Sphyrospermum majus Sphyrospermum standleyi 9 Symphysia Symphysia costaricensis Symphysia floccose Symphysia jefensis Symphysia orosiensis Symphysia ovata Symphysia poasana Symphysia racemosa Symphysia tubulifera theca (pi., -ae) 1 Themistoclesia 2, 4, 7, 9, 21, 22 Themistoclesia alata Themistoclesia costaricensis 4, 9 Themistoclesia horquetensis 4 Themistoclesia pentandra 4, 6, 9, 21 Themistoclesia pterocarpa Themistoclesia revoluta 23 Themistoclesia smithiana 6, 7 Themistoclesia valehi Thibaudia 16, 22, 25 Thibaudia albiflora Thibaudia archeri Thibaudia bracteata 29 Thibaudia costaricensis 6, 16, 2 1 Thibaudia crassifolia 30 Thibaudia floribunda Thibaudia laurifolia Thibaudia panurensis Thibaudia quereme Thibaudia rupestris Tropicoseius tube tubule 2 unistratose 1, 20 Utleya 14 Utleya costaricensis 4, 6, 14, 2 1 uva Vacciniaceae 24 Vaccinieae 1, 3, 25 Vaccinioideae 1 Vaccinium 1, 2, 7, 8, 10, 12, 21 sect. Oreades Vaccinium almedae Vaccinium bocatense Vaccinium consanguineum 4, 6. 8, 12 var. irazuense Vaccinium costaricense 4, 10, 12 Vaccinium dissimile Vaccinium floccosum 4 Vaccinium floribundum 2, 7, 8 Vaccinium furfuraceum 4, 12 Vaccinium irazuense Vaccinium jefense Vaccinium luteynii Vaccinium monteverdense Vaccinium orosiense 4, 10 Vaccinium pachycardium Vaccinium poasanum 4, 6, 12 Vaccinium santafeense Vaccinium smithianum Vaccinium talamancense Vaccinium valerii INDEX 107 Families of seed plants known or expected to occur in Costa Rica and adjacent areas numbered according to the sequence ol Hnuler's .SY//»v dcr Pfianzenfamilien, edition I 1. reuorked b\ I Diels (1936). 1 Cyvadacoac 2 laxaceac 3 Podocaip 4 Araucariacoao 5 I'inacoao 6 C'uprossacoao 7 (Jnetaceae 1 vphaceae 9 Potaimuiotonaco.ic 10 Najadaceao 1 1 Alisiuatacoao \2 Butomaceae 13 Hydrocharitaccac 14 Triuridaccac 15 (iramincae Id Cypcracoae 17 Palmae 1 S C 'y clanthaceae 19 Araccac 20 l.oinnacoao 2 1 Mayacaceac 22 Xyridaccae 23 Erioeaulaeoae 24 Bromeliaceae 25 Commelinaceae 2<> I'ontedenaceac 27 Juncaceae 28 Liliaceae 29 Haemodoraccac 30 Amaryllidacoao 31 VfoQoziaceae 32 Dioscorcaccac 33 Iridaceae 34 Musaceac 35 Zingibcraceac 36 C'annaccac 37 Maiantaccac 38 Bunnanniaceae 39 Orehidaceac 40 Casuarinaccac 41 Piperaceae 42 Chluranthaceac 43 LaciMcMiaceae 44 Salicacoao 45 ( iarryacoao 46 Myricaccac 47 Juglandaccac 48 Batidacoac 49 Betulaccao 50 Fagacoac 51 Ulmacoac 52 Moraceae 53 Urticaccac 54 Podostemonaccac 55 Protcacoac 56 Olacaccac 57 Opiliaccac 5S I.oranthaccac 59 Aristolochiaccac 6()a Hydnoraccao 60h Ratllcsiaccac 61 Balxnophoiaceae 62 PolygotiaccaL- 63 Chenopodiaccae 64 AmkraotfaaceM 65 Nyctaginaceae 66 Phytdlaccaccac 67 Ai/oaccac 68 PotttdMCaOCM 69 Bascllaccac 70 C'ar\i>pliyllaccac 71 Nymphacacoao 72 C'cratKphyllaccac 73 Kanuiiculaccac 74 Hcrhcridaccac 75 Mcnispcrniaccac 76 Mugnoliacoac 77 Annonacoao 78 Myristicacoac 79 Monimiaccao SO I auraccac 81 llornandiacoao S2 Papa\'oraccac, incl. Fumanaccae S < I 'apparidaccac 84 C'rucil'crac 85 To\ariaccac S(> Kcscdaccac S7 Mni'iniiacoac 88 Droseraceae Nl> ( 'rassiilacoac 90 Saxifragaccac 91 Brunelliaccac uaoniaceae 93 Ilaniamclidaceae 94 Rosaceac 95 C'onnaraceae 96 Leguminosae 97 Knuneriaceae 98 Oxalidacoao 99 Geraniaccao 100 Tropaeolaceao 101 Linaceac. incl. Humiriaceae 102 F.rythroxylaceae 103 Zygopbyllaceae 104 Rutaccao 105 Simaruhaccac 106 Burscraccac 107 Meliaccac ION Malpighiaccac 109 'IVigoiiiacoac 110 Vochysiaccac 1 1 1 Polygalaccac 1 12 Dichapetalaceae 1 1 3 Euphorbiaceae 114 C'allitrichacoac 115 Huxaccau 1 16 Coriariaccac 1 17 Anacardiacoao 118 Cyrillaccac I I1' Ai|iiit'oliaccac 120 Cclastraccac 121 Hippocrateaceae 122 Staphylcaccac 123 Icacinaccac 124 Hippocaslanaccac 125 Sapindacoac 1 26 Sahiaccao 127 Balsaminaccac 12K Khaiiin I2l> N'ilaccao 130 Ivlaoocarpaccac 131 Tiliaccae 132 Malvaceae 133 Bornhacaccac 134 Slcrculiaccac : iillctiiaccac 1 Vi \ctiniiliaceae 137 Ochiiaccae I 5S 139 140 141 142 143 144 145 146 147 I4S 149 150 151 152 153 154 155 156 157 158 159 160 161 162 163 164 165 166 167 168 169 170 171 172 173 174 175 176 177 178 179 ISO 181 182 183 1X4 185 186 187 188 189 190 191 192 193 I93a 194 195 196 197 198 199 200 201 202 203 204 205 206 207 208 Caryoca :.i\ i.icoac Quiinaceac I ho.u Gutdfierae, incl. Hyper Elatinai ( 'islaccac Bixaceac ispennaceac Vtolaceae llacourliaccac [\irnermoeae I'assitli': ( 'aricaccac I iiasaccac Hcgoniacoae ( atiaecac I hymclacaccac 1-lacagnaccac tceae Pumcaceae l.ccyihidaccac Rhizophoraccac ( 'ombrctaccae Myrtaccac Mclaslonialaccac ( )iiat!raccao llaU>rrliai:accac Araliaccae Uinbcllircrac C'ornaccac Clcthraccac Monotropaceae Pyrolaccac Ericaceae Theophrastaceae Myrsinaceae Primulaccac Plumhaginacoac Sapotaceac Ebenaceae Symplocaccac Styracaceae ( )lcaccac I.oganiaccac (icntianaccac Apocynaccac •Nsfk-piadacoac ( 'niiMiK nla. Polcmoniaccae I Isilrophy -llaccac Uoraginaccac Veibenaceae I abialac Solanaceae Sciiipluilanaccac Schlegeliaccac liignoniaccac Pedaliaceae Martyiiiaccac ( )mbanchaceac ( iL-Mlcriaccac l.ciitibulan.K Acanthi Planlaiiinaccac Rubiaceae Caprifoliaceae Valerian Dipsaca Cucurbit lai Compontae 30112066638583 Field Museum of Natural History Roosevelt Road at Lake Shore Drive Chicago, Illinois 60605-2496 Telephone: (3 1 2) 922-9410