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* </<•/• PfiaroenftmiUe*, edition

I I. reuorkedh\ I Dicls (18

156

130

143

172

23

102

113

96

50

I4S

82

45

183

99

198

7

15

142

29

165

93

81

124

121

101

60a

13

188

142

riaeagnaccac

Haucarpaccae.

1 lalmaccae

Erica*

l-.rutcaulaceae

Eupbortriaceae

Fabaci

sec Leguminosae

1 agaccae

1 lacourtiaceae

1 umanaccae.

sec Papaveraceae

Garryaccae

Gentianaceae

Cieraniaccae

Gcsneriaceae

Gnetaceae

Gramineae

Gutti ferae

H aemodoraceae

Halorrhagaceae

Hamamelidaceae

Hernandiaceae

H ippocastanaceae

Hippocrateaceae

Humiriaceae,

see Linaceae

Hydnoraceae

Hydrocharitaceae

Hydrophyllaceae

Hypericaceae.

see Guttiferae^

uraceae

1 59 Lecythidaceae

96 Leguminosae

20 Lemnaceae

1 99 Lentibulariaceae

28 Liliaccae

101 Linaceae

152 Loasaceae

1 82 Logan iaceae

58 Loranthaceae

157 Lythraceae

76 Magnoliaceae

108 Malpighiaceae

132 Malvaceae

37 Marantaceae

139 Marcgnn iaceae

196 Martyniaceae

2 1 Mayacaceac

1 63 Melastomataceac

107 Mel iaceae

75 Mcnispermaceac

96 Mimosaceae.

see Leguminosae

79 Moniiiu

170 Monotropaceae

52 Moraceac

87 Morinij

34 Musaceae

iceae

78 M\nstK

174 Mvrsinaccac

1 h2 '

10 "^

65

71

137

56

181

164

57

39

197

98

17

82

150

195

66

5

41

171

201

176

3

54

187

111

62

26

68

9

175

55

158

140

60b

73

86

128

160

94

202

104

126

44

125

177

90

193

105

192

122

134

180

179

2

141

173

155

131

85

109

14

100

149

8

51

167

204

31

190

147

129

110

22

35

103

Nyctaginaec.ie

Nymphaeaceae

( )chnu>

( )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

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

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.

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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chte und Pflanzengeographie, 105: 449-480.

SMITH, A. C. 1932. The American species of Thibau-

dieae. Contributions from the United States National

Herbarium, 28: 31 1-547, 19 pis.

. 1933. The genera Sphyrospermum and Dister-

igma. Brittonia, 1: 203-232.

-. 1941. Vacciniaceae. In Woodson, R. E., Jr., and

R. W. Schery, Contributions toward a Flora of Pana-

ma. V. Annals of the Missouri Botanical Garden, 28:

438-452.

STANDLEY, P. C. 1920-1926. Trees and shrubs of Mex-

ico. Contributions from the United States National

Herbarium, 23: 1-1721. [Polycodium and Vaccinium

pp. 1101-1103]

LUTEYN AND WILBUR: FAMILY #172 ERICACEAE

103

STANDLEY, P. C., AND L. O. WILLIAMS. 1996. Ericales.

Fl. Guatemala, Fieldiana: Botany, 24(pt. 8): 73-127.

STEVENS, P. F. 1971. A classification of the Ericaceae:

Subfamilies and tribes. Journal of the Linnean Society.

Botany, 64: 1-53.

STEVENS, P. F, J. L. LUTEYN, E. G. H. OLIVER, T. L. BELL,

E. A. BROWN, R. K. CROWDEN, A. S. GEORGE, G. J.

JORDAN, P. LADD, K. LEMSON, C. B. MCLEAN, Y. MEN-

ADUE, J. S. PATE, H. M. STAGE, AND C. M. WEILLER.

2004. Ericaceae, pp. 145-194. In K. Kubitski, ed.,

Families of Flowering Plants, vol. VI.

TAKHTAJAN, A. 1980. Outline of the classification of

flowering plants (Magnoliophyta). Botanical Review,

46: 225-359.

VANDER KLOET, S. P. 1985. On the generic status of Sym-

physia. Taxon, 34: 440-447.

VANDER KLOET, S. P., J. L. BALTZER, J. H. APPLEBY, R.

C. EVANS, AND D. T. STEWART. 2004. A re-examina-

tion of the taxonomic boundaries of Symphysia (Eri-

caceae). Taxon, 53: 91-98.

WALLACE, G. D. 1995. Monotropoideae, in Ericaceae —

Part II. The superior-ovaried genera (Monotropoideae,

Pyroloideae, Rhododendroideae, and Vaccinioideae

p.p.). Flora Neotropica, Monograph, 66: 1-560. [Mon-

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WILBUR, R. L. 1992. A synopsis of the genus Disterigma

(Ericaceae: Vaccinieae) in Mexico and Central Amer-

ica with the description of two previously undescribed

species. Bulletin of the Torrey Botanical Club, 119:

280-288.

WILBUR, R. L., AND J. L. LUTEYN. 1978. Ericaceae. Fl.

Panama. Annals of the Missouri Botanical Garden,

65: 27-144.

. 1981. Additions to the Ericaceae of Panama.

Annals of the Missouri Botanical Garden, 68: 154-

166.

YEO, P. 1967. Notes on some species of Macleania (Er-

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104

Index

The index includes all accepted scientific names (in Roman type), synonyms (italics), as well as

common English names (Roman) and vernacular Spanish names (italics), and scientific terms (Roman)

that may not be familiar to all and that have been defined in the text. 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