.LLINOIS LIBRARY

URBANA-CHAMPAI'

HIST SUP'

FIELDIANA

Botany

Published by Field Museum of Natural History

WraWlH/JTBgys^

6 1985

FLORA COSTARICENSIS

FAMILY #54 PODOSTEMACEAE

FAMILY #55 PROTEACEAE

FAMILY #56 OLACACEAE

FAMILY #57 OPILIACEAE

FAMILY #58 LORANTHACEAE

FAMILY #59 ARISTOLOCHIACEAE

FAMILY #60a HYDNORACEAE

FAMILY #60b RAFFLESIACEAE

FAMILY #61 BALANOPHORACEAE

FAMILY #62 POLYGONACEAE

FAMILY #63 CHENOPODIACEAE

FAMILY #64 AMARANTHACEAE

FAMILY #65 NYCTAGINACEAE

FAMILY #66 PHYTOLACCACEAE

FAMILY #67 AIZOACEAE

FAMILY #68 PORTULACACEAE

FAMILY #69 BASELLACEAE

FAMILY #70 CARYOPHYLLACEAE

WILLIAM BURGER, Editor

mber 29,

Publication 1350

Families of seed plants known or expected to occur in Costa Rica and adjacent areas,

listed alphabetically and numbered according to the sequence of Engler s Syllabus der

Pflanzenfamilien, edition 11, reworked by L. Diels (1936).

96 Caesalpiniaceae,

see Leguminosae

200 Acanthaceae

136 Actinidiaceae

67 Aizoaceae

11 Alismataceae

64 Amaranthaceae

30 Amaryllidaceae

117 Anacardiaceae

77 Anonaceae

184 Apocynaceae

119 Aquifoliaceae

19 Araceae

166 Araliaceae

4 Araucariaceae

59 Aristolochiaceae

185 Asclepiadaceae

61 Balanophoraceae

127 Balsaminaceae

69 Basellaceae

48 Batidaceae

153 Begoniaceae

74 Berberidaceae

49 Betulaceae

194 Bignoniaceae

145 Bixaceae

133 Bombacaceae

189 Boraginaceae

24 Bromeliaceae

91 Brunelliaceae

38 Burmanniaceae

106 Burseraceae

12 Butomaceae

115 Buxaceae

154 Cactaceae

lpiniacea

Legumin

114 Callitrichaceae

207 Campanulaceae

36 Cannaceae

83 Capparidaceae

203 Caprifoliaceae

151 Caricaceae

138 Caryocaraceae

70 Caryophyllaceae

40 Casuarinaceae

120 Celastraceae

72 Ceratophyllaceae

63 Chenopodiaceae

42 Chloranthaceae

144 Cistaceae

169 Clethraceae

146 Cochlosperraaceae

161 Combretaceae

25 Commelinaceae

208 Compositae

95 Connaraceae

186 Convolvulaceae

116 Coriariaceae

168 Cornaceae

89 Crassulaceae

84 Cruciferae

206 Cucurbitaceae

92 Cunoniaceae

6 Cupressaceae

1 Cycadaceae

18 Cyclanthaceae

16 Cyperaceae

118 Cyrillaceae

112 Dichapetalaceae

135 Dilleniaceae

32 Dioscoreaceae

205 Dipsacaceae

88 Droseraceae

178 Ebenaceae

156 Elaeagnaceae

130 Elaocarpaceae

143 Elatinaceae

172 Ericaceae

23 Eriocaulaceae

102 Erythroxylaceae

113 Euphorbiaceae

96 Fabaceae,

see Leguminosae

50 Fagaceae

148 Flacourtiaceae

82 Fumariaceae,

see Papaveraceae

45 Garryaceae

183 Gentianaceae

99 Geraniaceae

198 Gesneriaceae

7 Gnetaceae

15 Gramineae

142 Guttiferae

29 Haemodoraceae

165 Halorrhagaceae

93 Hamaraelidaceae

81 Hernandiaceae

124 Hippocastanaceae

121 Hippocrateaceae

101 Humiriaceae,

see Linaceae

60a Hydnoraceae

13 Hydrocharitaceae

188 Hydrophyllaceae

142 Hypericaceae,

see Guttiferae

123 Icacinaceae

33 Iridaceae

47 Juglandaceae

27 Juncaceae

97 Krameriaceae

191 Labiatae

43 Lacistemaceae

80 Lauraceae

159 Lecythidaceae

96 Leguminosae

20 Lemnaceae

199 Lentibulariaceae

28 Liliaceae

101 Linaceae

152 Loasaceae

182 Loganiaceae

58 Loranthaceae

157 Lythraceae

76 Magnoliaceae

108 Malpighiaceae

132 Malvaceae

37 Marantaceae

139 Marcgraviaceae

196 Martyniaceae

21 Mayacaceae

163 Melastomataceae

107 Meliaceae

75 Menispermaceae

96 Mimosaceae,

see Leguminosae

79 Monimiaceae

170 Monotropaceae

52 Moraceae

87 Moringaceae

34 Musaceae

46 Myricaceae

78 Myristicaceae

174 Myrsinaceae

162 Myrtaceae

10 Najadaceae

65 Nyctaginaceae

71 Nymphaeafeae

137 Ochnaceae

56 Olacaceae

181 Oleaceae

164 Onagraceae

57 Opiliaceae

39 Orchidaceae

197 Orobanchaceae

98 Oxalidaceae

17 Palmae

82 Papaveraceae

150 Passifloraceae

195 Pedaliaceae

66 Phytolaccaceae

5 Pinaceae

41 Piperaceae

171 Pyrolaceae

201 Plantaginaceae

176 Plumbaginaceae

3 Podocarpaceae

54 Podostemonaceae

187 Polemoniaceae

111 Polygalaceae

62 Polygonaceae

26 Pontederiaceae

68 Portulacaceae

9 Potamogetonaceae

175 Primulaceae

55 Proteaceae

158 Punicaceae

140 Quiinaceae

60b Rafflesiaceae

73 Ranunculaceae

86 Resedaceae

128 Rhamnaceae

160 Rhizophoraceae

94 Rosaceae

202 Rubiaceae

104 Rutaceae

126 Sabiaceae

44 Salicaceae

125 Sapindaceae

177 Sapotaceae

90 Saxifragaceae

193 Scrophulariaceae

105 Simarubaceae

192 Solanaceae

122 Staphyleaceae

134 Sterculiaceae

180 Styracaceae

179 Symplocaceae

2 Taxaceae

141 Theaceae

173 Theophrastaceae

155 Thymelaeaceae

131 Tiliaceae

85 Tovariaceae

109 Trigoniaceae

14 Triuridaceae

100 Tropaeolaceae

149 Turneraceae

8 Typhaceae

51 Ulmaceae

167 Umbelliferae

53 Urticaceae

204 Valerianaceae

31 Velloziaceae

190 Verbenaceae

147 Violaceae

129 Vitaceae

110 Vochysiaceae

22 Xyridaceae

35 Zingiberaceae

103 Zygophyllaceae

FLORA COSTARICENSIS

FAMILY #54

FAMILY #55

FAMILY #56

FAMILY #57

FAMILY #58

FAMILY #59

FAMILY #60a

FAMILY #60b

FAMILY #61

FAMILY #62

FAMILY #63

FAMILY #64

FAMILY #65

FAMILY #66

FAMILY #67

FAMILY #68

FAMILY #69

FAMILY #70

PODOSTEMACEAE

PROTEACEAE

OLACACEAE

OPILIACEAE

LORANTHACEAE

ARISTOLOCHIACEAE

HYDNORACEAE

RAFFLESIACEAE

BALANOPHORACEAE

POLYGONACEAE

CHENOPODIACEAE

AMARANTHACEAE

NYCTAGINACEAE

PHYTOLACCACEAE

AIZOACEAE

PORTULACACEAE

BASELLACEAE

CARYOPHYLLACEAE

FIELDIANA

Botany

Published by Field Museum of Natural History

New Series, No. 13

FLORA COSTARICENSIS

FAMILY #54

FAMILY #55

FAMILY #56

FAMILY #57

FAMILY #58

FAMILY #59

FAMILY #60a

FAMILY #60b

FAMILY #61

FAMILY #62

FAMILY #63

FAMILY #64

FAMILY #65

FAMILY #66

FAMILY #67

FAMILY #68

FAMILY #69

FAMILY #70

PODOSTEMACEAE

PROTEACEAE

OLACACEAE

OPILIACEAE

LORANTHACEAE

ARISTOLOCHIACEAE

HYDNORACEAE

RAFFLESIACEAE

BALANOPHORACEAE

POLYGONACEAE

CHENOPODIACEAE

AMARANTHACEAE

NYCTAGINACEAE

PHYTOLACCACEAE

AIZOACEAE

PORTULACACEAE

BASELLACEAE

CARYOPHYLLACEAE

WILLIAM BURGER, Editor

Curator and Chairman

Department of Botany

Field Museum of Natural History

Accepted for publication October 18, 1982

December 29, 1983

Publication 1350

Library of Congress Catalog Card Number: 78-172358

ISSN 0015-0746

PRINTED IN THE UNITED STATES OF AMERICA

CONTENTS

LIST OF ILLUSTRATIONS vi

ACKNOWLEDGMENTS vii

PODOSTEMACEAE by William Burger (based on P. Van Royen) 1

PROTEACEAE by William Burger 8

OLACACEAE by William Burger 14

OPILIACEAE by William Burger 27

LORANTHACEAE (in a wide sense) by William Burger and Job Kuijt 29

EREMOLEPIDACEAE 31

LORANTHACEAE (in a narrow sense) 33

VISCACEAE 59

ARISTOLOCHIACEAE by Kerry Barringer 79

HYDNORACEAE by Luis D. Gomez 87

RAFFLESIACEAE by Luis D. Gomez 89

BALANOPHORACEAE by Luis D. Gomez 93

POLYGONACEAE by William Burger 99

CHENOPODIACEAE by William Burger 138

AMARANTHACEAE by William Burger 142

NYCTAGINACEAE by William Burger 180

PHYTOLACCACEAE by William Burger 199

AIZOACEAE by William Burger 213

PORTULACACEAE by William Burger 217

BASELLACEAE by William Burger 222

CARYOPHYLLACEAE by Richard Baker and William Burger 227

LIST OF ILLUSTRATIONS

1. Podostemaceae 2

2. Proteaceae 10

3. Olacaceae and Opiliaceae 15

4. Olacaceae: Heisteria 18

5. Loranthaceae (s.s.): Psitticanthus and Gaiadendron 34

6. Loranthaceae (s.s.): Struthanthus and Phthirusa 35

7. Loranthaceae (s.s.): Struthanthus 36

8. Loranthaceae (s.s.): Oryctanthus. Eremolepidaceae and Rafflesiaceae 37

9. Viscaceae: Dendrophthora and Phoradendron 61

10. Viscaceae: Phoradendron, second group 62

11. Viscaceae: Phoradendron, third group 63

12a. Aristolochiaceae: A. maxima and its allies 80

12b. Aristolochiaceae: six species 81

13. Hydnoraceae 88

14. Rafflesiaceae: Bdallophyton 92

15. Balanophoraceae: Langsdorffia and Helosis 94

16. Balanophoraceae: Corynaea 95

17. Polygonaceae: Polygonum 102

18. Polygonaceae: Rumex 103

19. Polygonaceae: climbing species 104

20. Polygonaceae: species with winged fruit 105

21. Polygonaceae: Coccoloba with rounded leaves 108

22. Polygonaceae: Coccoloba with epistipular petioles 109

23. Polygonaceae: Coccoloba with basal petioles 110

24. Polygonaceae: Coccoloba, rarely collected species Ill

25. Amaranthaceae: Achyranthes and Cyathula 146

26. Amaranthaceae: Alternanthera and Gomphrena 149

27. Amaranthaceae: Alternanthera and Pfaffia 152

28. Amaranthaceae: Amaranthus 158

29. Amaranthaceae: Celosia, Chamissoa, and Pleuropetalum 163

30. Amaranthaceae: Iresine 171

31. Nyctaginaceae: Bougainvillea and Mirabilis 182

32. Nyctaginaceae: Boerhavia and Salpianthus 183

33. Nyctaginaceae: Guapira and Pisonia 184

34. Nyctaginaceae: Neea psychotrioides complex 185

35. Nyctaginaceae: Neea amplifolia complex 186

36. Phytolaccaceae: Microtea, Rivina, and Trichostigma 200

37. Phytolaccaceae: Phytolacca 201

38. Phytolaccaceae: Petiveria and Stegnosperma. Chenopodiaceae 202

39. Caryophyllales: succulent species 223

40. Caryophyllales: Basellaceae and Portulacaceae 224

41. Caryophyllales: herbs with narrow opposite leaves 225

42. Caryophyllales: herbs with opposite or whorled leaves 226

ACKNOWLEDGMENTS

This is the fourth issue of the Flora Costaricensis. The first dealt with Piperaceae

(Fieldiana, Bot. 35, 1971), the second dealt with the families Chloranthaceae

through Urticaceae (Fieldiana, Bot. 40, 1977), and the third dealt with Gramineae

by Richard Pohl (Fieldiana, Bot., new series No. 4, 1980). In the work of preparing

both this present volume and in all previous volumes, we have had the generous

cooperation and support of the Museo Nacional de Costa Rica. Use of special

facilities at the Museo for drying, processing, and shipping have been essential to

our field program. Sr. Luis D. Gomez has been especially helpful to the flora

program during the years he served as Director and since that time. Sr. Jorge

Gdmez-Laurito, associated with both the Herbario Nacional at the Museo Na-

cional and the Biology Department of the University, has also been most helpful.

We are especially grateful for the assistance of the National Science Foundation,

Program for Systematic Biology, which has aided this program for many years.

Financial support for fieldwork, assistants, and a postdoctoral position have been

provided by NSF grants DEB 74-08575 and DEB 81-03184.

vii

PODOSTEMACEAE

REFERENCE: P. van Royen, The Podostemaceae of the New World, pt. 1, Meded.

Bot. Mus. Herb. Rijks Univ. Utrecht 107:1-150, 1951. Part 2, Acta Hot. Neerl.

2:1-21. 1953.

Small herbs, growing in fast-moving freshwater rapids and waterfalls, usually attached

to larger rocks (rarely wood), annual or perennial, bisexual, very variable in size and form,

ranging from thalloid and closely attached to rocks to mosslike or with long branched

leaves, sterile plants usually larger and thicker than the fertile, stems usually forming a

broad holdfast and difficult to distinguish from the roots in some genera; lateral stipules

or intrapetiolar stipules sometimes present. Leaves very different in different genera and

species, in 2 or 3 ranks or borne from the edge or center of the thalloid stem, minute and

appressed (in Tristichia) to large and much branched (in Marathrum). Inflorescences of

solitary or fascicled pedicellate flowers, terminal, axillary or extra-axillary on special

shoots or in 2-sided spikelike monochasia, each flower at first enclosed within a thin

spathella or a few leaves or with groups of 10-20 flowers at first enclosed within a

thickened spathe. Flowers bisexual, radially symmetrical or bilaterally symmetrical, very

small (mostly less than 1 cm), borne on prominent pedicels often slightly expanded at the

apex; perianth of 2 to many parts, in 1 or several whorls or on only 1 side, either thin and

petal-like or reduced to small scales, free or united at the base; stamens 1 to many, often

alternate with the perianth parts (tepals) in 1 or 2 whorls, in a whorl or on only 1 side of

the flower, filaments free or united (rarely on an andropodium), anthers opening by 2

longitudinal slits, introrse or extrorse, basifixed or dorsifixed; ovary superior, narrowed at

the base or on a short stalk, sometimes obliquely oriented in the flower, 1- to 3-locular,

placentation axile with 2 to many anatrapous ovules, styles (1) 2 or 3, usually slender. Fruit

a thin-walled capsule, seeds 2 to many, without endosperm, embryo straight.

A family of about 200 species in 43 genera, pantropical but reaching the middle

Atlantic states of the United States and Japan in the Northern Hemisphere and

northeastern Australia in the Southern Hemisphere. This is one of the most

unusual families of angiosperms, both in morphology and in habitat. Because of

their restriction to fast-moving water, these plants are difficult to collect, and our

knowledge of them is poor. Fortunately, we have the excellent monograph by van

Royen covering the American genera and species (see the references above).

Nearly all of this treatment is based on van Royen's work, but it may be that some

of his species of Marathrum were circumscribed too narrowly.

KEY TO THE GENERA OF PODOSTEMACEAE

la Leaves minute (to 4 mm), in 3 imbricate ranks on short erect stems and resembling

bryophytes; young flowers protected by a few leaves, with 3-5 broad perianth parts;

ovary 2- or 3-locular Tristichia

Ib Leaves 1-50 cm long, cuneate to much branched and divided or tripinnate, borne on

flat thalloid stems and resembling algae; young flowers enclosed in a narrow spath-

ella, perianth parts very narrow or minute and scalelike; ovary 1- or 2-locular

Marathrum

MARATHRUM

tenue

10cm

M. utile

FIG. 1. Podostemaceae: species of Marathrum and Tristichia trifaria, lower left, enlarged.

BURGER: FLORA COSTARICENSIS 3

MARATHRUM Humboldt and Bonpland

Small aquatic herbs attached by a broad irregular base to rocks in fast-moving water,

glabrous, bisexual, sterile plants often larger than the fertile, the stem often difficult to

distinguish from roots or leaf bases; an intrapetiolar stipule or ligule often present. Leaves

variously inserted on the thalloid base or distichous, very variable in size and form, from

subentire with a few distal lobes to repeatedly forked or repeatedly pinnate and branch-

like, conforming to the flow of water or prostrate on the rocks, petiole and rachis often

fleshy. Inflorescences of solitary or fascicled flowers arising from between the leaf bases,

each flower at first enclosed within a translucent tubular-clavate spathella which splits

irregularly near the apex, pedicels becoming elongate and often slightly expanded at the

apex; flowers bisexual, radially symmetrical (stamens and perianth on only 1 side in M.

tenue), perianth of 3-25 parts in 1 or several whorls or on only 1 side, lanceolate to linear

or filiform or reduced (0.3 mm) and scalelike at the base of the flower, free or united at the

base, sometimes inserted on the cuplike apex of the pedicel; stamens 2-25, whorled or on

only 1 side, united only at the base, filaments linear to lanceolate, anthers introrse; ovary

superior and 2-locuIar, usually ellipsoid and narrowed at the base, with 6 or 8 longitudinal

ribs, styles 2, free or united near the base. Fruit a capsule, splitting along the longitudinal

ribs to produce 2 equal halves each with 5 ribs, seeds small and numerous.

Marathrum includes about 20 to 25 species, ranging from Mexico and Cuba

through Central America to northwestern South America where the genus inter-

grades with Apinagia (see van Royen, 1951). Different species have leaves that vary

from short, simple, and rhombic or flabellate to much longer and divided or

compound-pinnate, but the flowers and fruit seem quite similar. Standley noted

that the name Pasacarnehas been used for these plants in Costa Rica and that cattle

will wade out into stream rapids to feed on them in the dry season (see Fieldiana,

Bot. 24, pt. 4:402, 1946). Fertile material has only been collected in the dry season,

from early January to middle May.

While appreciative of van Royen's excellent monograph, I believe that some of

his species are separated on rather minor characters. Nevertheless, it seemed

important to provide the user of this flora with the distinctions that van Royen

used to separate species. Accordingly, the key utilizes van Royen's dichotomies

for all the species of Costa Rica and adjacent areas, but the last five species in the

key are probably covered under the expanded description of M. oxycarpum. In his

monograph of 1951, van Royen had already submerged some of the species he

described for the Flora of Panama (Ann. Missouri Bot. Card. 37:126-137, 1950);

see the discussion under M. oxycarpum.

KEY TO THE COSTA RICAN SPECIES OF Marathrum

BASED ON THE KEY OF VAN ROYEN (1951)

la Leaves cuneate or somewhat flabellate, relatively broad (2-30 mm), lobed or pin-

natisect only distally, with distinct reticulate venation when dry M. utile

Ib Leaves with a central narrow (1-4 mm) rachis or much divided from near the base,

often twice pinnatifid and branchlike in appearance, without broadly reticulate

venation 2a

2a Stamens and perianth parts 2 or 3, borne on 1 side of the flower, apex of the pedicel

often becoming asymmetric in fruit with ribs of the fruit decurrent into the pedicel on

1 side; anthers 1 mm long or less; ultimate divisions of the leaf short and very slender,

producing a characteristically "fimbriate" ultimate cluster M. tenue

2b Stamens and perianth 5-10 or more, in whorls around the flower and fruit; apex of the

pedicel not strongly asymmetric; anthers to 2 mm long; distal leaf segments not

usually densely fimbriate with short filiform divisions 3a

3a Pedicel usually becoming broadened and cuplike at the apex 4a

3b Pedicel only slightly enlarged at the apex, never forming a cuplike base for the

flower . . . . 5a

4 FIELDIANA: BOTANY

4a Leaf segments repeatedly forked, with long (0.5-12 mm) and very slender terminal

divisions, never pinnatisect in appearance; rare in Central America

M. foeniculaceum

4b Leaf segments usually in alternate arrangements on the central rachis and giving a

pinnatifid appearance, ultimate division 1-3 mm broad or if slender not usually

exceeding 5 mm in length; common in Central America but may intergrade with M.

oxycarpum M. schiedeanum

5a Fully developed stamens about half as high as the ovary; not recorded from Costa Rica:

M. stenocarpunt (Weddell) van Royen, Panama and Colombia.

5b Fully developed stamens more than half as high as the ovary, usually equalling or

exceeding the ovary in height; all the following species (as separated by van Royen)

seem very similar, and a description is provided only for M. oxycarpum in a wider

sense 6a

6a Styles filiform, widened at the top; sometimes emarginate, 2-3 mm long, cohering

over a long distance; stamens 8-10; ultimate divisions of the leaf up to 3 cm long

(= 3 mm?) and up to 0.3 mm wide M. oxycarpum

6b Styles of different shape, cohering at the base only, up to 2 mm long; stamens 5-9;

ultimate divisions of the leaf 5 mm long or less 7a

7a Largest leaves shorter than 7 cm long 8a

7b Largest leaves up to 30 cm long 9a

8a Leaves repeatedly pinnate, 3-6 cm long; styles filiform, up to 1 mm long; pollen grains

20 x 11 ^.m: M. leptophyllum van Royen, western Panama.

8b Leaves repeatedly pinnate or cuneate and dissected at the top, up to 1 cm long; styles

cylindrical, ca. 1.5 mm long; pollen 15 x 14 /im: M. pusillum van Royen, central

Panama.

9a Leaves without an intrapetiolar stipule at the base M. cheiriferum

9b Leaves with an intrapetiolar stipule at the base M. minutiflorum

Marathrum cheiriferum van Royen, Ann. Missouri Bot. Card. 37:134-135.

1950.

It is very difficult to see the intrapetiolar stipules on many specimens of Mar-

athrum, and to use such a characteristic to separate species seems impractical. In

addition, the separated specimens seem so similar in other aspects of both vege-

tative and floral morphology that one doubts they are different species. Costa

Rican material identified as M. cheiriferum by van Royen is tentatively being placed

under M. oxycarpum in a wide sense (q.v.).

Marathrum foeniculaceum Humboldt and Bonpland, Plantae Aequin. 1:40-41,

t. 11. 1808. Lads foeniculacea (H. & B.) Martius, Nov. Gen. & Sp. 1:6. 1824.

Aquatic herbs to 50 cm long, attached by an irregular base to 2 cm broad; intrapetiolar

stipules present, to 3 mm high. Leaves repeatedly divided or forked (rarely repeatedly

pinnate in early stages), 2.5-50 cm long, petiole flattened, 1-8 cm long, widened at the

base, ultimate divisions usually very slender and without a midvein, to 15 mm long.

Flowers solitary or fascicled, spathella up to 15 mm long, pedicels up to 35 mm long and

expanded at the apex to form a broad shallow cup; perianth parts 5-8, linear, ca. 1 mm

long, arising from the edge of the disklike pedicel apex; stamens 5-8, 4-6 mm long,

anthers ca. 1 mm long; ovary 3-4.5 mm long, 1-1.5 mm in diameter, styles 1-2.5 mm long.

Fruit ca. 4.5 mm long, each valve with prominent ribs.

This species ranges from Belize to Colombia, but has not been collected from

Costa Rica.

Marathrum foeniculaceum is characterized by the pedicels expanded at the apex

to form a shallow "floral" cup and the relatively long slender distal divisions of the

leaves which obscure the pinnate form of the leaf and give the leaf a more alga-like

appearance. This leaf form is quite distinctive and unlike that of other species of

Marathrum in our area.

BURGER: FLORA COSTARICENSIS 5

Marathrum minutiflorum Engler, Bot. Jahrb. Syst. 61, Beibl. 138:4. 1927. M.

allenii Woodson, Ann. Missouri Bot. Card. 25:827-828. 1938. M. indifferens van

Royen, loc. cit. 37:132-133. M. minutiflorum forma allenii (Woods.) v. Royen,

Meded. Bot. Mus. Herb. Rijks Univ. Utrecht 107:80. 1951. M. minutiflorum forma

diversifolium v. Royen, loc. cit. 81. M. minutiflorum forma indifferens (v. Royen) v.

Royen, loc. cit. 81. M. minutiflorum forma intermedium v. Royen, loc. cit. 81.

At present, it seems best to interpret material placed under the above names by

van Royen (1951) as robust specimens of M. oxycarpum in a wide sense; see the

description and discussion under M. oxycarpum.

Marathrum oxycarpum Tulasne, Ann. Sci. Nat. Paris, ser. 3, 11:94-95. 1849.

Small- to medium-sized herbs, 2-50 cm long, base with an irregular shape, ca. 1 cm wide

and to 3 cm long; intrapetiolar stipules usually present. Leaves usually 3 x pinnate, 3-20

(50) cm long, petiole terete or compressed, to 12 cm long, pinnae alternate on the rachis,

ascending, ultimate divisions of the leaf spathulate (rarely filiform). Flowers solitary, at

first enclosed in a tubular spathella to 12 mm long, pedicels slightly thickened at the apex

but never forming a cup, to 5 (9?) cm long; perianth parts 5-10, 1-3 (4) mm long, narrowly

lanceolate; stamens 5-10, 2.5-4.5 (6) mm long, anthers 1-2 mm long; ovary 2-4 mm high,

1-2 mm in diameter, with 8 distinct longitudinal ribs, styles 1-3 mm long, slender. Fruit

2.5-5 mm long, the smaller rounded and obtuse at the apex and the longer more ellipsoid,

each valve with 5 prominent ribs.

Aquatic plants of rocky streams growing at or up to 1 m below the water level.

Ranging from near sea level to 1,200 m elevation in Costa Rica and flowering

from January to May. Most collections are from the Pacific slope, but a few

come from the Caribbean watershed. This species, defined in a wider sense,

ranges from Mexico to Colombia.

Marathrum oxycarpum is recognized by its distally slender pedicel, whorled

perianth and stamens, strongly ribbed fruit, and its complex tripinnate leaves.

This species may be polymorphic, with some plants (often collected without

leaves) producing smaller, often stalked round fruit and other plants (usually

leafy) producing larger (3.5-5 mm) sessile ellipsoid fruit. These latter plants look

similar to specimens of M. schiedeanum, with some individuals actually having an

expanded pedicel apex. Some plants have large (2 mm) anthers, but with other

floral characters being typical. The minute perianth parts at the base of the flower

(and alternating with the longer narrow perianth parts) are often conspicuous in

this species. All perianth parts may be deciduous before the fruit is mature.

Marathrum oxycarpum, as here defined, may be something of a catchall, since the

more distinctive entities of the genus are separated in the earlier dichotomies of

the key to species. Nevertheless, the material identified by van Royen as M.

cheiriferum and M. minutiflorum seems to fall into this assemblage very naturally;

all these plants seem to share exactly the same altitudinal range, and their inclu-

sion in one species gives a more evenly distributed pattern of collecting localities

through Central America. Perhaps M. leptophyllum and M. pusillum of Panama

belong here as well (see the key). This larger circumscription of M. oxycarpum also

allows us to identify the small-fruited material lacking well-developed leaves

which may represent late-flowering plants (in respect to local water level) that are

less robust. Another reason for the broader interpretation of this species is that

one does not expect so many similar sympatric species in a genus whose life-style

is so specialized and so dependent on effective dispersal. See the discussion

under M. schiedeanum.

6 FIELDIANA: BOTANY

Marathrum schiedeanum(Cham.) Tulasne, Ann. Sci. Nat. Paris, ser. 3, 11:95.

1849. Lads schiedeanum Chamisso, Linnaea 9:504-505, t. 566. 1835. Marathrum

flexuosum Liebmann, Fdrh. Skand. Naturf. 5:511. 1849. Figure 1.

Aquatic herbs to 50 cm long, base irregular, to 2 cm wide; intrapetiolar stipules usually

present, 1.5 mm long, obtuse. Leaves up to 50 cm long, repeatedly pinnate, the young

leaves sometimes repeatedly divided, petiole terete or slightly flattened, 0.5-13 cm long,

pinnae usually ascending, ultimate divisions of the leaf to ca. 3 mm long and varying from

2-3 mm broad to filiform, acute to obtuse at the apex, with or without a midvein. Flowers

solitary or fascicled, spathella 1-1.5 cm long, pedicels 1-4 (9) cm long, expanded at the

apex and forming a distinctive shallow or deep (2 mm) cup, 2 mm broad at the apex;

perianth parts 5-8, linear to lanceolate, 0.5-1.5 mm long; stamens 5-8, 3.5-4.5 mm long,

anthers ca. 1 mm long; ovary 2-5.5 mm long, 1.5-3 mm in diameter with 8 longitudinal

ribs, styles 1-2 mm long. Fruit to 5.5 mm long and 3 mm thick, ellipsoid, each valve with

5 prominent ribs.

This species is found from near sea level to 1,200 m elevation on the Pacific slope

and ranges from Mexico to southern Costa Rica; it flowers and fruits from January

to May.

Marathrum schiedeanum is characterized by the pedicel expanded at the apex to

form a "floral" cup from which the perianth and stamens arise and the often large

tripinnate leaves with ultimate divisions that may be broad and pinnatifid or

slender and filiform. This species may hybridize or intergrade with M. oxycarpum.

An alternative interpretation is that the cupulate nature of the pedicel apex is not

a character worthy of distinguishing species in this instance; the character seems

variable in some collections and there do appear to be intermediate individual

collections, such as Bunting & Licht 1018 from Nicaragua. It may be that all the

material under both M. oxycarpum in its expanded circumscription and M. schie-

deanum is part of a single polymorphic species. For this reason, it seems best to

place sterile material under the oldest name (M. schiedeanum), but to continue to

use the pedicel apex to distinguish fertile material into two species.

Marathrum tenue Liebmann, Forh. Skand. Naturf. 5:511. 1849. Neolacis myr-

iophylla Weddell, in DC., Prodr. 17:63-64. 1873. Blandowia myriophylla (Wedd.)

Nash, N. Amer. Fl. 22, pt. 1:5. 1905. Apinagia myriophylla (Wedd.) Engler, Natur.

Pflanzenfam. 18a:38. 1930. Figure 1.

Small aquatic herbs to 20 cm long, from a flat or branched base to 1.5 cm high; stipules

absent but the petioles sheathing at the base. Leaves repeatedly pinnate, central rachis to

15 (20) cm long, petiole usually terete and up to 6 cm long, ultimate divisions filiform and

less than 2 mm long and resulting in distinctive densely 3-dimensional pinnae. Flowers

solitary or fascicled, spathellae 3-10 mm long, pedicel to 2.5 cm long, somewhat asym-

metric at the apex or becoming asymmetric; perianth parts 2, 3, or 4, on 1 side of the

flower, lanceolate to filiform, to 1.5 mm long; stamens 2 or 3, 2.5-4.5 mm long, filaments

lanceolate (resembling perianth parts), anthers 1-1.5 mm long; ovary 2-3 mm long, 1-2

mm in diameter, styles ca. 1 mm long. Fruit ca. 3 mm long, strongly ribbed with the ribs

on the side of the flower lacking perianth and stamens often decurrent on the pedicel, each

half of the fruit with 5 prominent ribs or with 3 prominent and 2 less prominent ribs.

Usually submerged aquatic plants growing on rocks in fast water and flowering

in the dry season, January to May. The species is found from near sea level (100

m) to 1,600 m elevation in Costa Rica, but has only rarely been collected below

1,000 m elevation. The species ranges from Mexico to Costa Rica and is to be

expected in western Panama.

Marathrum tenue is distinguished by the small number of stamens and perianth

parts arising on only one side of the flower. Because of this asymmetry, the apex

BURGER: FLORA COSTARICENSIS

of the pedicel is usually enlarged on one side, and the ribs of the fruit are often

decurrent into the pedicel on the opposite side; no other species in our area has

similar flower and fruit. In addition, the leaves with prominent central rachis and

alternate pinnae with numerous (often closely crowded) ultimate divisions are

quite different from typical leaves of the other species in our area.

Marathrum utile Tulasne, Ann. Sci. Nat. (Paris), ser. 3, 11:95. 1849. Figure 1.

Small aquatic herbs, to 35 cm long, base ca. 2 cm long, shoots opposite along the root

but difficult to see. Leaves 3-35 cm long, 1-6 cm broad, often rhombic when young,

becoming spathulate to cuneate, entire to pinnatisect or palmisect distally, the distal lobes

or segments usually few (2-8), broad areas of the leaf with distinctive venation (dichoto-

mous and reticulate), the lamina gradually narrowed to the base and the petiole not clearly

differentiated. Flowers solitary or in fascicles, spathella 2-2.5 cm long, pedicels 1-4 cm

long, expanded distally and forming a short broad cup; perianth parts 5-8, up to 2 cm

long (difficult to distinguish from the filaments); stamens 5-8, 2-5.4 mm long, anthers

1-1.5 mm long; ovary 2-4 mm long, ellipsoid to ovoid, style 1.5-2 mm long. Fruit to 4 mm

long, with conspicuous longitudinal ribs.

Herbs growing in the fast water of streams and known only from the Pacific

slopes of the Cordillera Central and Cordillera de Talamanca between 1,000 and

2,000 m elevation in Costa Rica. The species is known from Costa Rica, Colombia,

and Panama.

Marathrum utile is immediately distinguished by its broad laminae with anas-

tomosing venation and divided only near the apex. The pedicels slightly cupulate

at the apex are also distinctive. Although only collected above 1,000 m in Costa

Rica, this species is known from lower elevations in Colombia.

TRISTICHA Du Petit-Thouars

Bryophyte-like herbs, forming dense mats on rocks in fast-moving water, stems thin and

terete, glabrous, bearing leaves on 3 sides; stipules absent. Leaves alternate and in 3 ranks

or becoming whorled in groups of 3 or 6, subsessile, lamina small and usually imbricate,

entire or divided at the tip, with or without a midvein. Inflorescences absent, the flower

solitary and terminal on the main stems or on short lateral shoots, young flower buds

protected by 2 or 3 distinctly larger overlapping leaves, pedicels long and slender; flowers

small and radially symmetrical, perianth 3-parted, free or united into a 3-lobed tube;

stamen 1, filament slender, anthers introrse with an elongate connective; ovary 3-locular,

with 3 slender erect styles. Fruit a capsule dehiscing by 3 equal valves, each valve with 3

ribs, seeds numerous.

A genus of three species in tropical and subtropical America, Africa, and Asia.

Only one species is known to occur in the Americas. Like other members of the

family, these plants are found only in fast-moving fresh water, and they resemble

mosses and leafy liverworts more than they do flowering plants; even the little

flowers and fruits resemble the stalked capsules of mosses.

Tristicha trifaria (Bory ex Willd.) Sprengl, Syst. Veg., ed. 16, 1:22. 1825. Du-

fourea trifaria Bory ex Willd., Sp. PI., ed. 5:55. 1811. Duforea hypnoides St. Hil.,

Mem. Mus. Hist. Nat. 10:472. 1823. Tristicha hypnoides (St. Hil.) Spreng., Syst.

Veg. 4, pt. 2:10. 1827. T. trifaria (Bory ex Willd.) Tulasne, Arch. Mus. Hist. Nat.

6:179. 1852. Figure 1.

Small herbs, 1-4 (10) cm high, mosslike and firmly attached to rocks, often forming

dense mats, stems often numerous, terete, leafy stems 1-3 cm long; without stipules.

Leaves very small and overlapping on the stem, often somewhat dimorphic, with those on

the upper stems 0.3-1 mm long and ovate, those of the lower stems to 0.4 mm broad and

8 FIELDIANA: BOTANY

2 mm long and spatulate in form, apex of the lamina entire or rarely split into 2-4 acute

lobes, glabrous. Flowers at first enclosed within 2 or 3 larger leaves, borne on a slender

pedicel 3-10 (20) mm long; perianth of 3 free or united petals, 1-2 mm long, perianth lobes

obtuse; stamen 1.5-2.5 mm long, anthers ca. 0.8 mm long; ovary 0.5-1 mm long and ca.

0.8 mm in diameter, ellipsoid, rounded or narrowed at the base, styles ca. 0.5 mm long.

Fruit an ellipsoid thin-walled capsule, ca. 1.5 mm long, with prominent longitudinal ribs.

Aquatic plants attached to rocks in fast-flowing freshwater streams from near

sea level to 1,300 m elevation; flowering and fruiting from January through April

in Costa Rica. The plants also flower in November and December in El Salvador

and Nicaragua where the dry season may begin earlier. They are found on both

the Pacific and Caribbean slopes. The species ranges from Cuba and southern

Mexico to Colombia, Venezuela, Brazil, Uruguay, and northern Argentina.

Tristicha trifariais recognized by its small bryophyte-like stems, with three ranks

of very small leaves attached by holdfasts to rocks in rapidly moving water. The

small flowers and ellipsoid fruit borne on a slender pedicel resemble the stalked

capsule of a moss.

PROTEACEAE

REFERENCE: H. Sleumer, Proteaceae americanae. Bot. Jahrb. Syst. 76:139-211.

1954.

Trees or shrubs (in ours), usually bisexual; stipules absent. Leaves simple or compound,

the juvenile leaves often pinnately compound and the mature foliage often simple, alter-

nate in a spiral to opposite or whorled (sometimes on the same stem), the laminae usually

stiff and drying coriaceous, entire to serrate or pinnately lobed. Inflorescences racemes,

spikes, umbels, or heads, rarely with flowers solitary or paired; flowers radially or bilater-

ally symmetrical, perianth of 1 outer whorl of usually 4 parts, free or united, valvate and

tubular but splitting apart at the apex and usually becoming recurved, sometimes splitting

on only 1 side; stamens opposite the perianth parts and the filament often adnate to them,

the anthers introrse and often exposing the pollen to the air or transferring pollen to the

style prior to pollination, an inner whorl of perianth-like nectar scales of hypogynous disk

elements usually present around the base of the ovary; pistil superior and often short-

stipitate, solitary, 1-carpellate, ovary with 1 loculeand 1 to many ovules, style solitary and

unbranched with a thickened or enlarged stigma. Fruit usually woody, splitting open (a

follicle) or remaining closed (a nut) with 1 to many seeds; seeds flattened or round,

sometimes winged, the cotyledons often unequal, lacking endosperm.

A moderately large family of about 60 genera and a little more than 1,000 species

largely restricted to the warmer portions of the Southern Hemisphere. The family

is best represented in Australia and southern Africa, especially in areas marked by

a long dry season. A number, such as Banksia and Protect, are cultivated in the

subtropics for their spectacular inflorescences. The stiff foliage usually evergreen

and often with both pinnate and simple leaves on different parts of the same plant

help to distinguish these woody plants. The usually narrow-tubular flower buds

with four valvate perianth parts, perianth-opposing stamens, and one-locular

pistils with simple style and stigma further distinguish the family. These plants

may resemble some members of the Loranthaceae which have inferior ovaries and

are often parasitic.

la Introduced plants of gardens and roadsides; mature leaves pinnate or deeply pin-

natifid, alternate along the stem, (simple leaves absent); flower buds bent backward

and somewhat U-shaped Grevillea

Ib Native plants of forest and woodland; mature leaves usually simple, alternate to

opposite or whorled on the stems (pinnately compound leaves present or absent);

flower buds straight 2a

BURGER: FLORA COSTARICENSIS 9

2a Leaves isomorphic, always simple and entire, the tertiary veins becoming prominent

on both surfaces when dry; anthers borne on prominent filaments; fruit round and not

splitting open Panopsis

2b Leaves heteromorphic, immature and flowerless shoots usually with pinnately com-

pound leaves and mature or flowering shoots usually with simple leaves; anthers

sessile or subsessile on the perianth parts; fruit somewhat flattened and splitting open

along the edge Roupala

GREVILLEA R. Brown

Shrubs or trees, bisexual, evergreen. Leaves alternate, simple to pinnately compound or

pinnately divided. Inflorescences usually terminal, racemes or umbels, flowers in pairs

along the rachis, with hairs usually attached in the middle; flowers bisexual, radially or

bilaterally symmetrical, the perianth tube straight and slender or curved, the apical part

of the perianth usually expanded and bearing the sessile stamens, anthers ovate, the

connective not produced beyond the anthers, hypogynous disk usually a single semi-

circular gland on the upper side of the receptacle or rarely completely circular or absent;

ovary usually stipitate, the 2 ovules attached laterally near the middle of the locule wall,

style filiform, often protruding from the slit in the underside of the perianth tube before

the perianth opens completely, stigma on the terminally expanded style. Fruit a follicle,

coriaceous or woody, often with the long style persisting, usually opening along the upper

margin; seeds flat or round, usually bordered by a wing.

A primarily Australian genus of about 190 species. Two species are widely

planted in the tropics and in Central America. They have large (12 to 35 cm)

pinnate or pinnatifid leaves, colorful racemes, and small (2 cm) follicles splitting

open along one side.

la Trees to 20 (30) m tall, often planted along avenues and in parks; leaves pinnate with

pinnatifid pinnae; flowers orange; fruit glabrous G. robusta

Ib Shrubs or small trees to 7 m tall; leaves pinnate or once pinnatifid; flowers red; fruit

villous G. banksii

Grevillea banksii R. Brown, Trans. Linn. Soc. 10:176. 1810.

Tall shrubs or slender trees to 7 m tall, branches and inflorescences reddish brown

tomentulose. Leaves pinnate or pinnatifid, 10-20 cm long, with 3-11 linear or lanceolate

segments, 5-10 cm long, margins entire and revolute. Flowers red to rose red, in compactly

flowered racemes 5-12 cm long, pedicels ca. 5 mm long, perianth tube strongly recurved

below the apex, ca. 1.5 cm long, densely puberulent; fruit ca. 1.5 cm long with a long (4

cm) persisting style.

Usually planted in gardens and near houses.

Grevillea robusta Cunningham, in R. Br., Prot. Nov. 24. Figure 2.

Small and slender to large and robust trees, branches and inflorescences pale grayish to

orange-brown tomentulose. Leaves usually pinnate, 15-30 cm long with ca. 10-20 pin-

natifid leaflets, pinnae or leaflets ca. 5-10 cm long, margins usually lobed and revolute.

Flowers orange, in loosely flowered racemes 10-20 cm long, pedicels ca. 1.5 cm long,

perianth tube strongly recurved at about the middle, 1 cm long (2 cm measured along the

curve); fruit ca. 2 cm long.

Often planted in parks and along streets and avenues.

PANOPSIS Salisbury

Trees, bisexual, the foliage monomorphic. Leaves alternate, subopposite, opposite, or

whorled, simple, entire, drying coriaceous. Inflorescences terminal or axillary, usually

solitary, racemose with many flowers, the flowers arising solitary, 2, or 3 together on the

rachis, with or without bracts; flowers bisexual, radially symmetrical, the perianth cylin-

VGREVILLEA,

irobusta&'t

Juvenile leaves

From a

ROUPALA |I photograph

loranthoides II of the type

10 cm

FIG. 2. Proteaceae: a cultivated species of Grevillea and native species of Roupala and

Panopsis. Note: reduced scale.

10

BURGER: FLORA COSTARICENSIS 11

drical and straight in bud, separating at anthesis into 4 perianth parts (tepals), the tepals

free to the base and becoming recurved; stamens arising from the lower half of the tepals

or from their base, filaments largely free, anthers oblong, connective apiculate; hypogy-

nous disk a thin 4-lobed cup resembling a short perianth tube; ovary subsessile, ovules 2,

pendulous, stigma terminal. Fruit indehiscent with a thick woody pericarp, seeds lacking

wings, rounded.

An American genus of about 11 species ranging from Costa Rica southward to

Peru, Bolivia, and Brazil. The lack of pinnate leaves on younger parts, racemose

flowers with distinct filaments, and the hard fruit that do not split open dis-

tinguish this genus from other Proteaceae.

Panopsis suaveolens (Kl. & Karst. ex Klotzsch) Pittier, Contr. Fl. Venez. 22.

1923. Andripetalum suaveolens Klotzsch & Karsten ex Kl., Linnaea 20:472. 1847.

Panopsis costaricensis Standl., J. Wash. Acad. Sci. 17:164. 1927. Figure 2.

Trees 5-15 (25) m tall, trunks to 80 cm in diameter, with brown bark, leafy internodes

(1) 5-30 mm long, 2-5 mm thick, reddish brown strigulose at the nodes but soon becoming

glabrous, with conspicuous lenticels 0.3-1 mm long, axillary buds densely covered with

lustrous reddish brown hairs. Leaves alternate to opposite, often clustered near the ends

of branchlets, petioles 2-12 mm long, 1-2 mm thick, striate and glabrous; laminae (3) 5-15

(17) cm long, (1.3) 2-6 (9) cm broad, narrowly oblong to elliptic or narrowly to broadly

obovate, bluntly obtuse and rounded at the tip, obtuse to cuneate at the base and some-

what decurrent on the petiole, margins entire or slightly undulate, lamina drying very

stiffly chartaceous to subcoriaceous, smooth and glabrous on both surfaces when mature,

often lustrous, the 4-8 pairs of major secondary veins arising at angles of 30°-65°, arcuate

and weakly connected near the margin, tertiary veins raised and prominent on both

surfaces (dry). Inflorescences axillary or in terminal clusters, always near the ends of

stems, 6-16 cm long, racemes with more than 30 flowers each, rachis with slender as-

cending yellowish brown hairs 0.1-0.5 mm long, pedicels 2-5 mm long, densely ap-

pressed puberulent, perianth 6-7 mm long, sparsely puberulent; filaments 4-5 mm long,

anthers ca. 1.5 mm long with a slender apiculate connective, hypogynous disk forming a

thin cup or tube 0.5-0.7 mm long, enclosing the base of the ovary and mostly glabrous;

ovary ca. 1.5 mm long, densely covered with hirsutulous orange-brown hairs to 0.5 mm

long, style ca. 5 mm long. Fruit indehiscent, a very hard woody nut, 3.5-6 cm long and

becoming 4.5 cm in diameter, ellipsoid to ovoid or subglobose, narrowed at the base and

apex, the surface smooth and glabrous, dark brown with paler markings.

Trees of the very wet forests along the Continental Divide and the Caribbean

slopes between 1,000 and 2,200 m elevation in Costa Rica; flowering from April to

July and fruiting from July to March. The species ranges from central Costa Rica

southward to Colombia, Venezuela, and Ecuador.

Panopsis suaveolens is recognized by its glabrous leaves with prominent tertiary

venation that are alternate, subopposite, or opposite, often on the same stem. The

puberulent racemes with many flowers and hard round fruit that do not open

further distinguish the species. The trees have been called Palo de papa and Papa

in Costa Rica and Aguacatia in Panama.

ROUPALA Aublet

Shrubs or trees, bisexual, the juvenile and sterile shoots usually with pinnately com-

pound leaves and the flowering branches usually with simple foliage. Leaves hetero-

morphic with both pinnately compound and simple leaves, alternate in a spiral to sub-

opposite or opposite, the laminae very stiff, dentate to entire with the pinnate leaves

usually dentate. Inflorescences axillary or terminal, spikes or racemes, flowers often aris-

ing 2 or 3 together on the rachis; flowers bisexual and radially symmetrical, the perianth

cylindrical and straight in bud, separating at anthesis into 4 perianth parts (tepals), the

tepals free to the base and becoming recurved; stamens adnate to about the middle of the

12 FIELDIANA: BOTANY

tepals, the filament almost entirely adnate, anthers linear-oblong, sessile or subsessile,

connective slightly prolonged beyond the thecae, a hypogynous disk or perianth-like

whorl of flat scales or glands present at the base of the pistil; ovary with 2 pendulous

ovules, stigma terminal. Fruit a short-stipitate follicle with thick hard walls, splitting

obliquely into 2 flattened or slightly rounded valves, the seeds winged.

An American genus of about 50 species ranging from southern Mexico south-

ward to Peru, Bolivia, and Rio Grande do Sul, Brazil. Three species have been

reported from Costa Rica. The very stiff often heteromorphic leaves, glabrous

except at the base of the petiole in the leaf axil, and often alternate to subopposite,

help to distinguish these woody plants.

la Inflorescence and flowers sparsely puberulent, perianth parts 7-9 mm long; simple

leaves long-acuminate; fruit ca. 3.5 cm long; trees and shrubs of the seasonally dry

Pacific slope, 40-1,200 m elevation R. montana

Ib Inflorescence and flowers glabrous (in ours); trees of moist cloud forests and evergreen

montane forests 2a

2a Simple leaves on mature shoots with laminae 5-15 cm long and with acuminate

apices; perianth parts 10-14 mm long; fruit ca. 4 cm long; trees of wet evergreen forests

(800) 1,500-2,500 m elevation R. glaberrima

2b Simple leaves on mature shoots with laminae 3-7 cm long and with bluntly obtuse or

rounded apices; perianth parts 7-9 mm long; fruit unknown; habitat unknown but

may be exposed ridges and montane wind gaps R. loranthoides

Roupala glaberrima Pittier, Contr. U.S. Natl. Herb. 18:299. 1917. Figure 2.

Trees 5-20 (30?) m tall, bark gray, leafy internodes 3-30 mm long, 1-5 mm thick, very

minutely puberulent but soon becoming glabrous, grayish. Leaves alternate, subopposite,

or occasionally opposite, heterophyllous and pinnately compound on the young snoots or

simple, rarely pinnate on the flowering shoots, simple leaves with petioles 1-8 cm long,

1-2 mm thick, very minutely puberulent near the base adaxially but glabrous elsewhere,

simple leaves with laminae 5-15 cm long, 2.5-6 cm broad, narrowly ovoid to rhombic or

obovate in outline but very variable (often on the same twig), gradually tapering to an

obtuse to acuminate or caudate-acuminate apex, occasionally rounded or slightly emar-

ginate at the tip, abruptly to gradually narrowed below the middle but decurrent and

forming small winged margins on the petiole, margin entire to undulate or bluntly den-

tate, the laminae drying subcoriaceous, smooth and glabrous on both surfaces, usually

slightly lustrous above, the 2-4 pairs of major secondary veins arising at angles of 15-35

degrees from the proximal half of the midvein, secondary veins slightly raised beneath

(dry), tertiary veins obscure. Inflorescences usually solitary in the axils of persisting leaves,

8-22 cm long, racemose with more than 40 flowers, rachis glabrous and ca. 1 mm thick

(dry), pedicels 3-7 mm long, ca. 0.5 mm thick; perianth parts 10-14 mm long, ca. 1 mm

broad, linear oblanceolate, glabrous, white when flowering; anther ca. 3 mm long on a

filament 1 mm long borne on the upper third of the tepal, hypogynous glands ca. 0.5 mm

long, rounded apically; pistil ca. 10-11 mm long with the ovary 1 mm long and glabrous.

Fruit ca. 4 cm long and 2 cm broad, flattened and asymmetrically obovate, abruptly

narrowed at apex and base, opening distally and along one side, surface smooth and

glabrous; seeds not seen.

Trees of the wet cloud forests along the Continental Divide and adjacent areas

on both the Pacific and Caribbean slopes between (800) 1,500 and 2,500 m ele-

vation; flowering from March to July and fruiting from October to February in

Costa Rica. The species ranges from El Salvador to the western part of the Cor-

dillera de Talamanca. The species is apparently rare east of Volcan Barba.

Roupala glaberrima is recognized by the stiff leaves that are pinnately compound

on juvenile shoots but usually simple and long-petiolate on flowering stems, the

conspicuous racemes, and the lack of pubescence on most parts. The collections

from El Salvador differ from ours in having slightly puberulent inflorescences and

resembling Roupala montana. There are no collections from Honduras or Nica-

BURGER: FLORA COSTARICENSIS 13

ragua. The white flowers are very fragrant in anthesis; their odor resembles that

of vanilla. Common names associated with these plants in Costa Rica are Danto

hediondoand Lora malodora. The latter name refers to the bad odor of the cut bark.

This tree was common in the forests around Zarcero; the westernmost collection

in Costa Rica comes from near Monteverde, Guanacaste.

Roupala loranthoides Meisner, in DC., Prodr. 14:425. 1856. Figure 2.

Probably small trees, the branchlets thick with many short twigs and clustered leaves,

twigs at first reddish brown tomentulose. Leaves alternate or subopposite, pinnate leaves

not seen, petioles 6-15 mm long, 1.2-2.2 mm thick, glabrous except for the very base in

the leaf axil (adaxially); laminae (3) 4-7 cm long, 1.4-4.5 cm broad, broadly elliptic or oval

to slightly obovate, bluntly obtuse to rounded at the apex and often emarginate, abruptly

rounded or narrowed near the base and attenuate to form short wings on the petiole,

margin entire in the few leaves seen and the edge revolute, drying very stiffly coriaceous,

smooth and glabrous on both sides, the 3-5 pairs of major secondary veins arising at

angles of ca. 20-30 degrees, both the secondary and tertiary veins slightly impressed

above, venation obscure beneath. Inflorescence 5-8 cm long, axillary, exceeding the termi-

nal leaves, racemose, glabrous, the flowers borne on thick (1-1.8 mm) pedicels, perianth

ca. 8-10 mm long, less than 1 mm thick in the lower half and ca. 1.5 mm thick near the

apex (dry) before anthesis, the hypogynous scales thin and triangular, very short, stigma

subclavate and obtuse, thicker than the style. Fruit unknown.

A very poorly known species. Type material was destroyed by fire in Vienna,

and only two small leaves and photograph remain (F). The original Friedrichsthal

collection was only labeled Monte Rincon, which may be Volcan Rincon de la

Vieja, Guanacaste, of northwestern Costa Rica. The thick small clustered leaves on

short thick twigs suggest that this is a tree of wind gaps or exposed elfin forest.

Roupala montana Aublet, PI. Guyan. Franc. 1:83. 1775. R. complicata H.B.K.,

Nov. Gen. & Sp. 2:153, t. 119. 1817. R. borealisHemsl, Biol. Centr. Am. Bot. 3:78,

t. 76. 1882. R. panamensis Pittier, Contr. U.S. Natl. Herb. 18:229. 1917. Figure 2.

Shrubs or small trees 2-8 (rarely 18) m tall, leafy internodes 1-30 mm long, 1-6 mm

thick, minutely reddish brown strigulose but quickly becoming glabrescent, becoming

grayish, the axillary buds densely brownish strigulose. Leaves alternate or subopposite,

heteromorphic, simple on the flowering branches, juvenile leaves pinnately compound but

very variable in the number of leaflets and usually coarsely serrate, adult (simple) leaves

with petioles 1.5-6 cm long, 0.7-2.5 mm thick, minutely puberulent near the base ad-

axially; simple leaves with laminae 4-14 cm long, 2-7.5 (9) cm broad, narrowly to broadly

ovate, gradually or abruptly narrowed to the often long-acuminate apex, abruptly nar-

rowed at the obtuse or rounded base, decurrent and forming small winged margins on the

petiole, margins entire or undulate, rarely coarsely serrate as on the juvenile foliage, the

laminae drying coriaceous, smooth and glabrous (in ours), often lustrous above, the 3-6

pairs of major secondary veins arising at angles of 20-50 degrees, secondary and tertiary

veins slightly raised on both surfaces (dry). Inflorescences usually axillary, solitary, 7-15

(18) cm long, racemose with more than 40 flowers, rachis densely puberulent with as-

cending yellowish hairs 0.2-0.3 mm long, pedicel becoming 4-5 mm long densely strigu-

lose; perianth parts 7-9 mm long, 1 mm broad, linear-oblanceolate, puberulent near the

base, anthers 2-3 mm long on filaments ca. 0.5 mm long, hypogynous glands thick, ca. 0.5

mm tall, pistil ca. 6 mm long, ovary 1-1.5 mm long and densely strigose. Fruit 3-4 cm long,

ca. 1.3 cm broad, asymetrically obovate or elliptic and somewhat flattened, the surface

glabrous and often lustrous, opening distally and along 1 side; seeds winged, thin and flat,

ca. 2.4 cm long and 1.2 cm broad.

Trees and shrubs of the seasonally very dry deciduous and seasonally dry

partly deciduous (tropical and premontane dry and moist) forest formations of the

Pacific slope between 40 and 1,200 m elevations, flowering from January to March

14 FIELDIANA: BOTANY

and fruiting from June to October in Costa Rica. The species ranges from

Veracruz, Mexico, to Peru, Bolivia, and Brazil.

Roupala montana is recognized by the stiff leaves that dry grayish and are pin-

nately compound on juvenile shoots, but simple and long-petiolate on flowering

stems, and the long puberulent racemes. Sleumer recognized both the typical

form of this species and var. dentata (R. Br.) Sleumer as occurring in Costa Rica,

but the differences between these appear to be inconsequential.

OLACACEAE

Trees, shrubs, or woody climbers, bisexual (in ours), autotrophic or hemiparasites on

roots of the host, stems with or without spines, mostly glabrous; stipules absent. Leaves

alternate in a spiral or distichous (rarely opposite), simple, petioles often with ridges

continuous with the laminae margins; laminae usually entire and glabrous, venation

pinnate. Inflorescences 1 to several or fasciculate in the leaf axils, paniculate, racemose,

cymose, or umbellate in form, often few-flowered and the flowers sometimes in groups of

3, bracteoles present or absent. Flowers bisexual, or rarely unisexual, radially symmetrical,

sometimes polymorphic, often small and inconspicuous (not so in Ximenia, Chaunochiton,

et al.); calyx united and usually a shallow cup with 3-6 small lobes or teeth, free or united

to the ovary (as in Schoepfia), corolla of 3-6 petals, valvate in bud, free or united, sometimes

puberulent on the inner surface; stamens as many as or 2-3 x as many as the petals,

opposite the petals or petal lobes when of the same number, filaments free or adnate to the

corolla tube, anthers 2-thecous with mostly lateral dehiscence; a disk said to be present at

the base of the ovary but usually obscure in ours, ovary superior or half-inferior (in ours),

1-5 locular (at least at the base) with the partitions often failing to reach the upper part of

the ovary, ovule 1 in each locule, pendulous from the apex of the central placenta, style

1 or none, stigma 1- to 5-lobed. Fruit usually a drupe with fleshy exocarp and single seed,

with endosperm, in some genera the calyx enlarges to form a colorful structure subtending

the fruit.

The Olacaceae are a pantropical family with about 27 genera and 230 species.

The family exhibits some interesting South American- African relationships. The

Olacaceae are characterized by the woody habit, simple alternate leaves usually

articulate at the base, a general absence of pubescence on vegetative parts (but not

in Minquartia), regular flowers with very small calyx and valvate petals or petal

lobes, and pistil with single style and the unusual arrangement of locules. Some

genera are known to be root parasites, while others appear to be normal auto-

trophs; all have green leaves and resemble nonparasitic trees and shrubs. This

family is very closely related to the Opiliaceae, and some authors unite the two.

These families are, in turn, related to the Loranthaceae (in a wide sense) and

Santalaceae (not recorded for Central America). Together, they make up the very

natural order Santalales. This is a unique and rather isolated order of di-

cotyledons, many of which show tendencies or adaptations for parasitism and

reduction of ovule structure. They share many vegetative and reproductive fea-

tures, such as simple articulate entire leaves, absence of stipules, scarcity of

pubescence, flowers often in triads, valvate perianth parts, poorly differentiated

calyx, and pistil with an undivided style. Members of the Olacaceae may be

confused with plants of the Rhamnaceae, Celastraceae, and Opiliaceae.

la Young stems and inflorescences with minute branched hairs; leaves with subparallel

tertiary veins; calyx not becoming expanded in fruit; corolla tube campanulate and ca.

2 mm long; tall trees of evergreen wet forests Minquartia

Ib Young vegetative parts and inflorescences essentially glabrous, hairs unbranched

where present; leaves lacking consistently subparallel tertiary venation 2a

2a Stems often with short thick spines, leaves often clustered on short shoots; petals free

XIMENIA

americana

1:

SCHOEPFIA

schreberi

SCHOEPFIA

vacciniiflora

MINQUARTIA

guianensis

FIG. 3. Olacaceae: Minquartia, Schoepfia, and Ximenia species. Opiliaceae: Agonandra

macrocarpa, upper left.

15

16 FIELDIANA: BOTANY

and with conspicuous hairs within; calyx remaining small in fruit; shrubs and small

trees flowering in the dry season in the Pacific lowlands Ximenia

2b Stems lacking spines, leaves rarely borne on short shoots 3a

3a Calyx very small and becoming almost completely united to the fruit, forming much

of the smooth covering of the fruit and defining a small round distal area at the tip;

petals united to form a short tube; shrubs to large trees in wet and dry areas

Schoepfia

3b Calyx becoming enlarged in fruit and separate from the fruit, a fleshy disk or thin and

broadly rotate structure 4a

4a Common understory shrubs and small trees to 8 m tall; the fruit subtended by a fleshy

red disk (the calyx); flowers very small and fasciculate, petals separate and 1-2 mm

long, glabrous Heisteria

4b Rare trees 10-30 m tall; fruit subtended and partly enclosed in a thin papery calyx

5-10 cm broad; flowers 1-3 cm long, petals united in the lower half and puberulent

within Chaunochiton

CHAUNOCHITON Bentham

Medium-sized to large trees, bisexual, stems glabrous, lacking spines; stipules absent.

Leaves simple and alternate, articulate at the base, petiolate; laminae elliptic and entire,

pinnately veined, glabrous. Inflorescences of congested cymules forming small panicles in

the axils of leaves. Flowers bisexual, radially symmetrical 2-6 cm long but only 2-5 mm

in diameter; calyx small and cuplike, 5-toothed, becoming greatly enlarged and thin in

fruit; petals 5, very narrowly linear-spatulate, valvate in bud, united in the lower half to

form a narrow tube, pilose within for almost their entire length, a disk absent; stamens 5,

opposite the petals and united to their base, filaments long and slender, anthers very small

and clavate; ovary superior, free or united with the calyx only near the base, 2-locular with

1 pendulous ovule in each locule, style very long with globose or 5-lobed stigma. Fruit a

drupe united at the base to the thin broadly expanded calyx, the calyx broadly cam-

panulate to rotate and chartaceous, exocarp fleshy with a crustaceous endocarp.

Chaunochiton is a very poorly known genus ranging from Costa Rica to the

Amazonian Basin. The genus is readily recognized in fruit with an expanded

rotate papery thin calyx that may be 10 cm broad. The slender flowers resemble

those of Psittacanthus (Loranthaceae), but the long narrow petals are united near

the base and are puberulent within.

Chaunochiton kappleri(Sagot ex Engler) Ducke, Arch. Jard. Bot. Rio de Janeiro

3:41. 1922. Heisteria kappleri Sagot ex Engler, in Martius, Fl. Brasil. 12, pt. 2:14.

1872. Figure 4.

Medium-sized to tall trees 12-30 m high, bisexual, leafy internodes 0.5-4 cm long, 1-4

mm thick, ashy gray in color, glabrous. Leaves alternate in a spiral, articulate at the base,

petioles 6-12 mm long, with lateral ridges continuous with the lamina margins; laminae

ca. 3-6 cm long and 2-3.5 cm broad, ovate-elliptic to ovate-oblong or broadly elliptic,

obtuse or bluntly acute at the apex, obtuse or rounded at the base, decurrent on the

petiole, margin entire, the laminae drying thin-chartaceous, smooth and glabrous above

and below, with 4-7 pairs of major secondary veins, the lower secondaries often more

strongly ascending than the upper. Inflorescences not seen. Flowers not seen, but if C.

breviflorum Ducke is synonymous with this species, then the calyx should be ca. 1 mm long

and the corolla ca. 1 cm long with the petals ca. 0.5 mm broad at the base and 1 mm broad

at the thickened apex. Fruit a drupe 6-10 mm long and 5-10 mm in diameter, with a

somewhat muricate surface when dry, the fruit is subtended by the expanded calyx which

may be 8-12 cm broad, subentire or weakly 5-lobed, drying pale brown and mem-

branaceous, apparently rotate or slightly campanulate in form, the fruiting pedicel 15-20

mm long.

Plants of lowland evergreen wet forest formations and known from only a single

collection in North America. Fruit and leafless stems were collected by Pittier

BURGER: FLORA COSTARICENSIS 17

(11948) in the Diquis basin at ca. 600 m altitude on March 15th, 1898. The species

is otherwise only known from Amazonian Brazil and the Guianas.

Chaunochiton kappleri is recognized by the fruit being subtended by a thin broad

calycular development and the slender flowers with the petals puberulent within.

The small leaves clearly articulate at the base, lack of pubescence, weakly five-

lobed calyx, and small ovary are all characteristic of Olacaceae. The collection from

Costa Rica is said to have been a small slender tree 12-15 m high and 20-25 cm

diameter. Its local name is said to be Manglillo. Botanists visiting the General

Valley, Osa Peninsula, and Golfo Dulce region should keep this unusual plant in

mind; the species and the genus are very poorly known.

HEISTERIA Jacquin

Shrubs or small trees, rarely woody lianas as in H. scandens, not known to be parasitic,

bisexual, the young stems with longitudinal ridges, glabrous. Leaves alternate and often

distichous, articulate at the base, petioles usually with 2 adaxial ridges continuous with

the lamina margins; lamina entire, glabrous, with laticiferous ducts. Inflorescences of few

to many flowers in axillary fascicles, often forming a raised mound of tissue with sessile

scarious bracts irregularly arranged over the surface. Flowers bisexual, very small, pale

green to white or yellowish, pedicellate to subsessile, glabrous on the outside, calyx

usually broadly cup-shaped, subentire to 5- or 6-toothed or -lobed, free from the ovary,

becoming enlarged and thickened in fruit; petals usually 5 (6), valvate in bud, free or united

at the very base, glabrous or puberulent within; stamens 10 or 12 (rarely 5 or 6), free or

united to the base of the petals, antipetalous stamens often shorter than the antisepalous,

anthers rounded and minute; ovary superior, rounded to conical, with 3 locules in the

lower half, 1 pendulous ovule in each locule, style short or none and with 3 poorly

differentiated stigmas. Fruit a drupe with thin fleshy exocarp that is often black and borne

on the expanded brightly colored calyx, the fruiting calyx annular, broadly cupulate or

becoming reflexed, fleshy in texture and red to purple in color, the embryo small in the top

of the endosperm.

The genus, with probably fewer than 20 species, ranges from southern Mexico

and the Lesser Antilles to southern Brazil and Bolivia. Two species are native to

West and Central Africa. Species of Heisteriaare easily recognized in fruit because

of the expanded, slightly fleshy calyx. The usually black fruit and bright red calyx

produce a striking contrast, undoubtedly an adaptation for seed dispersal. The

very small flowers arising directly from an axillary base are difficult to see and rare

in collections. The American species are seriously in need of revision, and our

own species represent some difficult unresolved problems. This is particularly

true regarding H. acuminata, H. macrophylla, and H. costaricensis. A number of

collections appear to be intermediate between these taxa, and all three may

prove to be part of a polymorphic complex. All our species appear to be plants

of the forest shade.

la Woody lianas (perhaps beginning as shrubs); laminae occasionally with the lowest

pair of secondary veins quite prominent and almost triplinerved; petals puberulent

within; fruit red at maturity and subtended by a small (10-15 mm) calyx; 0-800 m

elevation H. scandens

Ib Shrubs or trees; laminae never with the basal pair of secondary veins very prominent;

fruiting calyx often more than 12 mm broad 2a

2a Petals puberulent within; fruit white at maturity, fruiting calyx conspicuously and

deeply 5-lobed; leaves lustrous above and subcoriaceous when dry with the margins

recurved, short-acuminate at the apex; 0-800 m elevation H. concinna

2b Petals glabrous within; fruit becoming black in late stages, fruiting calyx shallowly or

obscurely 5-lobed; leaves not usually lustrous or with the margins recurved when dry,

chartaceous . . . . 3a

HEISTERIA

scandens

CHAUNOCHITONVfruitm9

flowers of

H. acuminata

FIG. 4. Olacaceae: Costa Rican species of Heistcria and Chaunochiton.

18

BURGER: FLORA COSTARICENSIS 19

3a Laminae usually abruptly narrowed at both ends; petiole deeply and narrowly cana-

liculate above (adaxially); fruit becoming black only in late stages, the fruiting calyx

often reflexed; 0-300 m elevation H. cyanocarpa

3b Laminae usually gradually narrowed to the apex; petiole with the adaxial ridges

forming a V-shaped or broadly U-shaped sulcus above, rarely narrowly canaliculate

above; 0-1,700 m elevation. (Note: The following 3 species are differentiated here

on the bases of leaf form and habitat. Plants with intermediate characteristics are

not uncommon, and it may be that these 3 taxa are part of a single polymorphic

complex.) 4a

4a Laminae linear-lanceolate, widest below the middle and often narrowed abruptly at

the base; 0-700 m elevation H. costaricensis

4b Laminae not linear-lanceolate 5a

5a Plants rarely found below 500 m elevation; laminae rarely more than 15 cm long, not

usually tapering gradually to both base and apex, often drying pale in color; under-

story plants of montane and seasonally dry evergreen moist forest formations; fruit

ellipsoid-oblong H. acuminata

5b Plants rarely found above 500 m elevation; laminae usually more than 15 cm long,

often tapering gradually to both base and apex, drying dark in color; plants of wet

forest formations; fruit globose H. tnacrophylla

Heisteria acuminata (H. & B.) Engler, in Martius, Fl. Brasil. 12(2):14, nota. 1872;

and in Flora 56:135. 1873. Rhaptostylum acuminatum Humboldt and Bonpland,

Plant. Aequinox. 2:129, 5:125, 1809, and in H.B.K., Nov. Gen. & Sp. PI. 5, t. 621,

emended illustration. 1824. Figure 4.

Shrubs or small treelets 0.7-3 m tall, glabrous, the leafy internodes 0.3-4 cm long, 1-4

mm thick, with 2 or 4 distinct longitudinal ridges, becoming terete and often drying pale

in color. Leaves obscurely articulate at the base, petioles 4-8 (11) mm long, the 2 longi-

tudinal adaxial ridges separate or forming a V-shaped sulcus above; laminae 6-14 (23) cm

long, 2-5 (7) cm broad, elliptic to very narrowly elliptic or elliptic-oblong (rarely lanceolate

or ovate elliptic), usually widest near the middle, tapering gradually to the short- or

long-acuminate apex, obtuse to acute at the base, margin entire or undulate when dry, the

laminae drying thin- to stiff-chartaceous, glabrous and smooth above and below, the 5-10

pairs of major secondary veins usually quite distinct from the tertiary veins. Inflorescences

in the axils of leaves, or less often in the axils of fallen leaves and undeveloped leaves,

usually with only 1 or 2 flowers in anthesis at a node at one time, pedicels 2-4 mm long;

flowers 2-3 mm long, calyx prominently 5- toothed, 1.5-2 mm broad, petals ca. 2 mm long

and 1 mm broad, glabrous within; stamens 10 and clearly of 2 lengths with the longer

alternipetalous, filaments to 0.4 mm broad, anthers 0.2-0.3 mm broad, pistil ca. 1.2 mm

long and 1 mm broad at the base. Fruit ca. 8-10 mm long and 6-8 mm in diameter, globose

to ellipsoid, soon becoming black, longitudinally ribbed when dry, the fruiting calyx 12-20

(30) mm broad, inconspicuously 5-lobed, bright red, fruiting pedicels ca. 7 mm long.

Plants of forest understory in moist evergreen montane or, less often, lowland

formations; collected with fruit from October to April, but probably flowering

throughout the year. The plants are most often encountered between 600 and

1,600 m elevation in Costa Rica. The species ranges from Chiapas, Mexico, to

Colombia (see below).

Heisteria acuminata is recognized by its small- to medium-sized, usually elliptic

leaves that taper gradually to an acuminate apex, glabrous flowers, weakly lobed

fruiting calyx that is broader than the fruit, and its frequent occurrence in cloud

forests on the Pacific slope. These plants are not known from below 600 m on the

Pacific slope of Costa Rica and adjacent Panama. The few lowland collections of

the Caribbean slope appear to be restricted to the moist (not the wet) forest

formations as diagrammed by Tosi in the Mapa Ecologico. Our highland material

is very similar to that of Guatemala and Chiapas, but I have seen no comparable

highland material from Colombia. Much of the Central American material has

been referred to as H. tnacrophylla or misidentified as H. longipes in the past.

20 FIELDIANA: BOTANY

The original collection of Heisteria acuminata is from Popayan, Colombia, at

about 2,000 m elevation. The leaf shape and venation and floral details illustrated

for this species agree well with Central American highland material. In a recent

letter, Dr. Sleumer points out that the fruit of H. acuminata is larger and ellipsoid-

oblong, whereas the fruit of H. macrophylla is smaller and globose. He thus inter-

prets the species in a way that has H. acuminata reaching its northern limit in Costa

Rica, whereas H. macrocarpa reaches its southern limit in Costa Rica. This species

may also intergrade with H. costaricensis, and it is closely related to H. cyanocarpa:

see the discussions under those species and H. macrophylla.

Heisteria concinnaStandley, Publ. Field Columbian Mus., Bot. Ser. 8:137. 1930.

Figure 4.

Small- to medium-sized trees 3-10 (20) m tall, the branches often said to be drooping,

leafy internodes (0.5) 1-4 (7) cm long, 1-3 (4) mm thick, yellowish or pale brown with 2

often obscure longitudinal ridges, becoming minutely striate and yellowish or pale brown

when dry, glabrous. Leaves distichous, often clearly articulate only in age, petioles 7-20

mm long, 1-2 mm thick, often bent distally and thickened beneath, the adaxial petiolar

ridges usually forming a narrow (0.5 mm) sulcus above; laminae (3) 5-15 cm long, 2.5-6

(7) cm broad, elliptic to broadly elliptic to ovate-elliptic, tapering gradually or abruptly to

the acuminate apex, obtuse or somewhat rounded at the base, margins entire but be-

coming undulate and recurved when dry, the laminae drying subcoriaceous or very stiffly

chartaceous and lustrous above, smooth on both surfaces, the 5-8 pairs of major secondary

veins often difficult to distinguish from the tertiary venation. Inflorescences in the axils of

current or fallen leaves, pedicels 3-6 mm long; flowers white, flower buds ca. 2-3 mm long

before anthesis, calyx ca. 1 mm high and 2-3 mm broad, petals ca. 2 mm long, puberulent

within, anthers 0.2-0.3 mm long, ovary conical, 1.5 mm long. Fruit 10-14 mm long and

8-10 mm thick (dry), ellipsoid to globose, becoming white, fruiting calyx 2-4 cm in

diameter with usually 5 shallow or relatively pronounced rounded lobes, rose red to dark

red in color.

This species is found in moist (partly deciduous) and wet evergreen forest

formations between sea level and about 800 m elevation on both the Caribbean

and Pacific slopes in Costa Rica; fruiting collections have been made from Sep-

tember through March. The species probably ranges from the Caribbean side of

Honduras (Standley 55292) to eastern Panama.

Heisteria concinna is distinguished by its stiff lustrous leaves with unusual pet-

ioles, puberulent petals, and the mature white fruit subtended by a bright red

five-lobed calyx that is 2 to 4 cm broad. Most of our collections come from season-

ally dry evergreen forests on the Pacific slopes, but none come from the seasonally

very dry and largely deciduous formations of lowland Guanacaste. This species is

apparently very closely related to H. cocinnea Jacquin of the Lesser Antilles.

Heisteria costaricensis Donnell Smith, Bot. Gaz. 19:254. 1894. Figure 4.

Shrubs or slender treelets 1.5-2.5 (3) m tall, leafy internodes 0.3-4 cm long, 0.7-3 (5) mm

thick, with 2 or 4 longitudinal ridges 0.1-0.3 mm high, the stems becoming smooth and

terete. Leaves alternate and distichous, indistinctly articulate at the base, petioles 4-8 (13)

mm long, the longitudinal adaxial ridges usually forming a shallow groove ca. 1 mm wide

above, the petiole occasionally thickened and bent; laminae 12-22 (30) cm long, 1.5-3.5

(4.5) cm broad, narrowly lanceolate to linear-lanceolate, broadest below the middle (rarely

near the middle), tapering very gradually to the acute or acuminate apex, narrowed more

abruptly to the acute or slightly rounded base, margin entire or slightly undulate, the

laminae drying chartaceous to stiffly chartaceous, smooth on both surfaces, the 7-37 pairs

of major secondary veins usually easy to distinguish from the tertiary. Inflorescences small

BURGER: FLORA COSTARICENSIS 21

axillary fascicles with apparently only a few flowers at any one time; flower buds ca. 1.5

mm long and subglobose, mature flower with 5-toothed calyx, petals ca. 2 mm long and

1 mm broad, glabrous on both surfaces; stamens 10, filaments liguliform, anthers 0.2-0.3

mm long; pistil conical, ca. 1 mm long and 1 mm in diameter. Fruit globose or ovoid 6-8

mm long, becoming black, vertically ribbed when dry, fruiting calyx 12-25 mm broad,

obscurely or shallowly 5-lobed, bright red, the fruiting pedicels 3-9 mm long.

Plants of wet and moist evergreen forest formations on both the Caribbean and

Pacific slopes from sea level to 700 (1,000) m elevation, but not as yet collected from

below 500 m on the Caribbean side; probably flowering and fruiting throughout

the year. This species is known only from Costa Rica and Panama.

Heisteria costaricensis is immediately recognized by its long linear-lanceolate

leaves, widest below the middle and usually abruptly narrowed near the base.

Characters of flower and fruit are probably the same as those of H. acuminata and

H. macrophylla. These three species are separated here primarily on their different

leaf shape, but they may prove to be members of a single polymorphic species.

Collections from around San Ramon, Alajuela, appear to be intermediate between

typical H. acuminata and H. costaricensis.

Heisteria cyanocarpa Poeppig, Nov. Gen. Sp. 3:35, t. 241, 1845 (fide Sleumer in

herb.). H. longipes Standley, J. Wash. Acad. Sci. 17:8. 1927. Figure 4.

Shrubs or small- to medium-sized trees (1.5) 3-12 (20) m tall, leafy internodes 0.5-3 cm

long, 1-3 mm thick, with 2 or 4 longitudinal ridges, becoming terete, glabrous. Leaves

alternate, usually distichous, articulate at the base, petioles 4-12 mm long, 1-1.5 mm

thick, the 2 adaxial longitudinal ridges forming a very narrow sulcus above, becoming

longitudinally striate and thickened abaxially throughout the length of the petiole; lami-

nae (4) 6-14 cm long, 2-6 (7) cm broad, elliptic to ovate-elliptic, obovate, or oblong,

gradually or abruptly narrowed to the short acuminate apex, abruptly narrowed at the

obtuse (rarely acute) base, margin entire or undulate when dry, laminae drying stiffly

chartaceous, smooth on both surfaces, with 5-9 pairs of major secondary veins but the

secondaries often difficult to distinguish from the tertiaries. Inflorescences in the axils of

current or recently fallen leaves, with 5-10 flowers but only 1 or 2 flowering at a node at

one time, flower buds 1.5-2 mm long, pedicels 1-4 mm long; flowers with a 5-lobed calyx,

petals 1.5-2 mm long, glabrous within; anthers 10, borne at the edge of the disk, filament

ca. 1.2 mm long and 0.5 mm broad, narrowed below the minute (0.2 mm) anther, ovary

1-1.3 mm broad near the base, ca. 1.2 mm long, abruptly narrowed above, stigma ob-

scurely lobed. Fruit 8-12 mm long, 6-8 mm in diameter, ellipsoid, apparently remaining

green for a time but becoming black, fruiting calyx (1) 1.5-2 (2.5) cm broad, bright red,

often reflexed, usually bluntly 5-lobed.

Plants of evergreen and partly deciduous wet and moist forest formations from

sea level to about 300 m elevation on both the Caribbean and Pacific slopes;

flowering collections have been made from December through July, and fruiting

collections have been made from January to September. The species, as presently

understood, ranges from southwestern Costa Rica and the Bocas del Toro region

of Panama to Colombia.

Heisteria cyanocarpa is recognized by the laminae usually being abruptly nar-

rowed at both ends, the petiole narrowly canaliculate above, and the five-lobed

fruiting calyx subtending a green fruit that becomes black only at maturity. The

species appears to be restricted to the lowlands (below 300 m) in our area, but

collections from Colombia which appear to be this species come from as high as

1,500 m. Rather similar looking plants from above 500 m elevation on the Pacific

slope and along the Caribbean slope are here placed under H. acuminata. In a

recent letter (June 1980), Dr. Sleumer suggests that this species might be made

part of H. acuminata in a very wide sense.

FIELDIANA: BOTANY

Heisteria macrophylla Oersted, Vidensk. Meddel. Dansk Naturhist. Foren.

Kjoebenhavn 1856:40. 1857, sensu stricto. H. latifoliaStandley, ]. Wash. Acad. Sci.

17:8. 1927.

Shrubs or small treelets (0.5) 1.5-3 (7) m tall, glabrous, the leafy internodes 0.3-6 cm

long, 1-5 mm thick, with 2 or 4 distinct longitudinal ridges, becoming terete in age, often

drying pale in color. Leaves obscurely articulate at the base, petioles 3-18 mm long, ca. 3

mm thick, the 2 adaxial longitudinal ridges separate or forming a broad V-shaped sulcus

above; laminae 15-36 cm long, 5-10 (17) cm broad, very narrowly elliptic to broadly elliptic

or ovate-elliptic, tapering very gradually to the long-acuminate apex, tapering gradually

to the acute or obtuse base, occasionally slightly rounded at the petiole in larger laminae,

margin entire (undulate when dry), the laminae drying thin-chartaceous and often dark

in color, glabrous and smooth on both surfaces, the 9-17 pairs of major secondary veins

usually quite distinct from the tertiary veins. Inflorescences in the axils of leaves or fallen

leaves, pedicels 1-3 mm long; flowers 2-3 mm long, calyx 5- (rarely 6-) toothed, petals 5

(rarely 4), ca. 2 mm long and 1 mm broad, glabrous; stamens 10, filaments ca. 0.5 mm

broad, 0.7-1.5 mm long, anthers ca. 0.5 mm broad, pistil conical ca. 1 mm broad at the

base. Fruit ca. 8-10 mm long and 6-9 mm in diameter, globose to ovoid or ellipsoid, soon

becoming black, longitudinally ribbed when dry, the fruiting calyx 16-30 mm broad,

inconspicuously 5-lobed, bright red, the fruiting pedicel ca. 7 mm long.

Plants of the wet evergreen forest formations of the Caribbean lowlands and

adjacent slopes between sea level and about 300 m elevation; fruiting collections

have been made from August to October and January to May. The species, as here

delimited, ranges from southern Nicaragua to eastern Panama.

Heisteria macrophylla (in a narrow sense) is recognized by its larger leaves usually

widest near the middle and gradually tapering to both apex and base and the wet

forest habitat on the Caribbean slope. The large thin leaves are often rhombic in

outline. There are collections (Molina et al. 17339, Pittier & Tonduz 9174, Donnell

Smith 6462, and Standley 37332) which appear to be intermediate between typical

H. macrophylla and H. acuminata. Further study may show that H. macrophylla

is only an unusual lowland form of H. acuminata in a wide sense (q.v.); see

Figure 4.

Heisteria scandens Ducke, Arch. Jard. Bot. Rio de Janeiro 4:9. 1925. H. fatoensis

Standl., Publ. Field Columbian Mus., Bot. Ser. 8:137. 1930. H. eurycarpa Standl.,

loc. cit. 11:148. 1936. Figure 4.

Woody lianas climbing to the tops of trees as much as 30 m tall, perhaps beginning as

small shrubs 3-4 m tall, leafy internodes 0.6-4 (6) cm long, 0.7-2.5 (5) mm thick, the 2

longitudinal ridges absent or inconspicuous, the stems soon becoming terete, longi-

tudinally minutely striate and pale gray to dark brown when dry, the axillary cushion-like

inflorescence bases often conspicuous on the leafless stems. Leaves usually distichous

with the laminae in a single plane, articulate at the base, glabrous, petioles 5-12 (20) mm

long, 0.6-1.3 mm thick, the 2 adaxial ridges forming a shallow groove above; laminae (4)

5-13 cm long, (1.5) 3-6 cm broad, ovate-oblong to elliptic-oblong or broadly elliptic,

abruptly narrowed to the short (0.5-1 cm) acuminate apex, obtuse to abruptly rounded at

the base, margin entire or slightly undulate, the laminae drying thin chartaceous or

chartaceous, smooth on both surfaces and often somewhat lustrous above, venation with

3-5 pairs of major secondary veins, occasionally with the lowest pair of veins quite

prominent and the lamina almost triplinerved. Flowers fasciculate from a broad minutely

bracteate mound in the axils of leaves or fallen leaves, each axil producing up to 30 (50)

flowers but only a few seen in anthesis in our material, pedicels 3-6 mm long, ca. 0.3 mm

thick, flowers 2-3 mm long, glabrous on the outside, calyx 1-2 mm broad, with 5 short (0.5

mm) teeth, petals 1.5-2 mm long, minutely puberulent near the apex within; stamens 10,

filaments 1-1.5 mm long, anthers 0.2-0.3 mm long; ovary conical and ca. 1 mm broad at

the base, with shallow longitudinal sulci. Fruit 9-14 mm long, 8-12 mm in diameter,

globose to ellipsoid, becoming red at maturity and longitudinally ribbed when dry, fruit-

BURGER: FLORA COSTARICENSIS 23

ing pedicel ca. 1 cm long, fruiting calyx 10-15 mm broad with 5 shallow lobes, the calyx

smaller than the fruit and often reflexed.

This species is known from only two collections in Costa Rica, both from the

Golfo Dulce region: Allen 5838 at 75 m altitude and Lent 447 at about 650 m. Fruit

have been collected in February and March and in July and August in Costa Rica

and Panama. The species, as presently known, ranges from the Golfo Dulce area

of Costa Rica (on the Pacific side) and the province of Bocas del Toro in Panama

(on the Caribbean side) southward to Peru and Amazonian Brazil.

Heisteria scandens is immediately distinguished by its climbing habit, apparently

unique in the genus. The smooth terete branches with well-spaced leaves, un-

usual leaf venation, petals puberulent within, and the fruit apparently red at

maturity with relatively small fruiting calyx further distinguish these plants. This

species may be closely related to H. acuta Engler of Brazil.

MINQUARTIA Aublet

Trees, often large, apparently autotrophic, bisexual, the young branches rusty tomen-

tulous with branched hairs, the older stems sometimes perforated, lacking spines. Leaves

alternate, not clearly articulate at the base, petiolate; laminae elliptic and acuminate,

entire, chartaceous to coriaceous, glabrous above but minutely puberulent beneath, with

resin canals and lactiferous ducts. Inflorescences spicate, solitary in the leaf axils, puber-

ulent, with small bracts subtending the subsessile or short-pedunculate cymelike groups

of flowers; flowers bisexual, calyx cuplike and 5-toothed, persistent but not enlarging in

fruit, petals connate into a bell-shaped or campanulate tube, puberulent to villous within,

the 5 (6) lobes valvate in bud; stamens 10 (12) in 2 whorls, adnate to the lower part of the

corolla tube at 2 levels, sessile or on short filaments, the antepetalous anthers longer than

the antesepalous; ovary with 3-5 locules, stigma 3- to 5-lobed. Fruit with a somewhat

fleshy exocarp and crustaceous endocarp.

A genus of perhaps three species, ranging from southern Central America to

the Amazonian Basin. The genus stands out among our representatives of the

family because of the unusual pubescence. Plants with immature inflorescences

may be mistaken for species of Euphorbiaceae or Malvales.

Minquartia guianensis Aublet, Hist. PI. Guyan. Franc., Suppl. 4, t. 370. 1775.

Figure 3.

Trees 10-35 m tall, developing low buttresses, sapwood yellow with dark red or dark

brown heartwood, bisexual, leafy internodes 0.7-3 cm long, 1.4-3 mm thick, at first

slightly angled but soon becoming terete, very minutely (0.05 mm) brownish puberulent

but soon becoming glabrous, smooth and brownish; stipules none but the young folded

leaves stipule-like, often ca. 1 cm long and 1.5 mm thick and tapering to a narrow apex.

Leaves alternate and distichous, petioles 8-30 mm long, 1-2.8 mm thick, with 2 longi-

tudinal adaxial ridges continuous with the margins of the lamina, the ridges expanded and

slightly vaginate just below the apex of the petiole; laminae 8-16 (22) cm long, 3-7 (11) cm

broad, elliptic-oblong to ovate-oblong, abruptly short (1-2 cm) acuminate at the base,

margin entire but often slightly undulate, drying chartaceous and slightly lustrous above,

smooth and glabrous above and below, venation pinnate with 3-14 pairs of major second-

ary veins, arcuate ascending only near the margin and never loop-connected, tertiary

venation often subparallel between the secondaries. Inflorescence solitary in the leaf axils,

spikelike with alternate groups of usually 3 flowers subtended by a bract, rachis with a

covering of minute scurfy or branched reddish brown hairs, ca. 5 cm long at anthesis and

to 15 cm long in fruit. Flowers apparently bisexual, short pedicelled, calyx broadly cupulate

with 5 very short teeth, ca. 1 mm long and 1.5-2 mm broad, reddish brown puberulent,

corolla a campanulate tube ca. 2 mm long with 5 recurved lobes, puberulent, stamens 10,

borne on the tube on very short (0.5 mm) filaments, anthers broader than long; ovary with

24 FIELDIANA: BOTANY

an unusual pubescence forming ca. 10 small pocket-like depressions on the upper (distal)

sides, style with 5 distinct finger-like stigmas. Fruit becoming 3 cm long and 2 cm thick,

black and fleshy, edible but with an acrid taste, borne on a short (5 mm) thick (2 mm)

pedicel.

Trees of the lowland evergreen wet forest formations below 500 m elevation in

Costa Rica; mature fruit have been collected in July, but mature flowers have not

been collected in Central America. The species ranges from southern Nicaragua

to Ecuador and Amazonian Brazil.

Minquartia guianensis is recognized by the tall habit, simple alternate distichous

glabrous leaves with unusual petioles and subparallel tertiary venation, solitary

spikelike reddish brown inflorescences, and small densely puberulent flowers.

These trees are known by the names Manu, Crillo, Nispero, Negro, Cricamola, and

"Black Manwood" in Costa Rica and Panama. The species is largely represented

by sterile collections in herbaria; the flower descriptions are based on South

American material. The wood is very hard and very durable, resisting both ter-

mites and fungal decay (see Biotropica 8:71-95, 1976).

SCHOEPFIA Schreber

Shrubs or trees, reported to be root parasites, bisexual, without spines, the vegetative

parts glabrous; stipules none. Leaves alternate, clearly articulate at the base, petiolate, the

laminae entire and stiffly chartaceous to coriaceous, glabrous, venation pinnate. In-

florescences solitary to several or fasciculate in the leaf axils, racemose or spicate with

usually 3 (2-5) flowers closely clustered or short-pedicellate at the end of a short peduncle

(and often resembling the triads of Loranthaceae), bract subtending the ovary of each

flower a shallow usually 3-lobed cup, glabrous or puberulent. Flowers small, calyx united

with the ovary and with an obscurely 3- to 6-lobed or entire distal margin, glabrous,

usually green, corolla united to form a tube, the corolla lobes usually 5 (3-6), short and

valvate in bud, the corolla glabrous on the outside and usually puberulent near the stamen

attachment within; stamens epipetalous, high on the corolla tube, anthers sessile or sub-

sessile, small, about as wide as long; ovary half-inferior, the upper surface often minutely

puberulent, style 1 with a prominent rounded or 3-lobed stigma, base of the ovary imper-

fectly 3-locular, ovules 3, lacking an integument. Fruit a drupe enclosed by the smooth-

walled calyx except near the apex where a circular area of different texture is demarcated

by the calycular rim, seed 1 and apparently erect, embryo small at the top of the endo-

sperm.

A genus of about 35 species in tropical Asia and ranging in the New World from

Mexico through Central America and the West Indies to Bolivia and southern

Brazil. The genus is distinguished by the simple stiff alternate leaves articulate at

the base, short few-flowered axillary inflorescences, tubular corolla with epi-

petalous stamens opposite the valvate corolla lobes, half-inferior ovary with

poorly differentiated adnate calyx, and unusual fruit with circular apical area of

different texture than the smooth lower surface.

la Trees to 20 m tall in the wet evergreen forests of the Caribbean lowlands; leaves 3-6

cm broad with strongly ascending basal secondary veins S. species A

Ib Shrubs or trees rarely more than 10 m tall in deciduous and partly deciduous forests

of the Pacific slope in Costa Rica; leaves 1-4 cm broad 2a

2a Corolla tube campanulate, 2-5 mm long and 2-3 mm in diameter; flowers usually

sessile in groups of 3; leaves ovate to lanceolate on pale grayish white stems; 0-1,000

m altitude S. schreberi

2b Corolla tube cylindrical, 4-7 mm long and 2-3 mm in diameter; flowers often short-

pedicellate; leaves usually elliptic on brownish stems; 800-1,800 m altitude

S. vaccinii flora

BURGER: FLORA COSTARICENSIS 25

Schoepfia schreberi J. F. Gmel., Syst. Veg. 2:376. 1791. S. americana Willd., Sp.

PL 1:9%. 1798. S. arborescens Roem. & Schult., Syst. 5:160. 1819. Figure 3.

Trees 4-8 (rarely 25) m tall, without spines, leafy internodes 0.5-2.5 cm long, 0.7-3 mm

thick, at first somewhat angular or ridged but soon becoming terete, becoming pale

grayish white and smooth, then dark in later years. Leaves alternate and distichous,

clearly articulate at the base; petiole differentiated from the lamina, ca. 4-8 mm long, with

lateral ridges continuous with the lamina margins, not sulcate above; lamina 3-8 cm long,

1.5-3.5 cm broad, ovate, ovate-elliptic to narrowly elliptic or lanceolate, acute to acu-

minate at the apex, acute to obtuse at the base and with the margins attenuate on the

petiole, margins entire and often undulate, lamina drying very stiffly chartaceous or

subcoriaceous, smooth and glabrous above and below, venation pinnate with 2 or 3

(sometimes to 5) pairs of major secondary veins, the basal secondaries often strongly

ascending, prominent beneath. Inflorescences fasciculate in the leaf axils, the flowers

usually borne in groups of 3 (triads) on short (2-6 mm) thin (0.4 mm) glabrous peduncles,

flowers sessile within broad (1.5-2 mm) shallow (0.5-1 mm) cuplike bracts with irregular

and partly puberulent margins, the bracts closely approximate at the apex of the peduncle.

Flowers 3-5 mm long, calyx glabrous and obscurely 5-lobed, or entire, corolla tube cam-

panulate, 2-4 (5) mm long, 2-3 mm in diameter, the corolla lobes 1-2 mm long and

recurved, glabrous on the outside but puberulent around the anther attachment within,

stamens sessile high on the corolla tube, anthers ca. 0.6 mm long; top of the half-inferior

ovary minutely grayish puberulent, style ca. 1 mm long with a prominent flattened

disklike stigma ca. 0.5 mm broad. Fruit globose to ellipsoid, ca. 7 mm long and 6 mm in

diameter, greenish (immature?) with a red apex, the glabrous lower ovary and calyculus

defining a small circular distal minutely puberulent area at the top of the fruit ca. 2 mm

broad and yellowish in color.

Trees of deciduous and partly deciduous forest formations on the Pacific slope

of Costa Rica between sea level and 1,000 m elevation; flowering material has been

collected from December to February and in June and July in Costa Rica. The

species, as presently understood, ranges from southern Florida and Mexico to

northern South America and the West Indies.

Schoepfia schreberi is recognized by its glabrous alternate stiff little leaves that are

clearly articulate on pale-colored twigs, the triads or diads of flowers arising from

small mounds of tissue in the axils of leaves, and the small bracteate flowers with

campanulate corolla tube, epipetalous sessile anthers, and internal pubescence.

Schoepfia vacciniiflora Planch, ex Hemsley, Diag. PL Mex. 5. 1878. Figure 3.

Shrubs or small trees 2-8 (12) m tall, without spines, leafy internodes 0.3-3 cm long,

0.6-4 mm thick, strongly 2-ridged or angled in early stages but becoming terete, pale

brown to pale gray. Leaves alternate and distichous, clearly articulate at the base, petioles

weakly differentiated from the lamina, ca. 2-5 mm long, with lateral margins continuous

with the lamina margins, flat above (adaxially); laminae (2) 3-8 (10.5) cm long, 1-3 (4.5)

cm broad, elliptic to elliptic-oblong or ovate-elliptic, acute to short-acuminate at the apex,

acute to obtuse at the base and somewhat attenuate at the petiole, margins entire or

slightly undulate, drying stiffly chartaceous to subcoriaceous, smooth and glabrous on

both surfaces, venation pinnate with 3-6 pairs of major secondary veins, the lower pairs

of secondaries often strongly ascending (in ours), sometimes loop-connected near the

margin. Inflorescences usually 1 or 2 in the axils of leaves, the peduncle branched above

the 1st flower or the flowers subsessile and clustered at the end of the peduncle, flowers

2 to 4 per inflorescence (most often 3), the flowers usually on short pedicels with shallow

cupulate bracts beneath the base, the bracts 1.5-2 mm broad, generally 3-lobed, glabrous.

Flowers 6-8 mm long, calyx fused with the ovary and with an entire or obscurely 5-Iobed

distal margin, corolla tube (4) 5-7 mm long, 2-3 mm in diameter, cylindrical, corolla lobes

1-1.5 mm long, slightly reflexed at anthesis, glabrous on the outside, puberulent near the

stamen attachment within, anthers sessile high on the corolla tube, style ca. 2 mm long

with stigma 1 mm broad. Fruit ellipsoid, ca. 10 mm long and 6 mm in diameter, glabrous,

26 FIELDIANA: BOTANY

longitudinally striate when dry, glabrous except for a sharply defined circular distal area

1.5-2 mm in diameter at the apex.

Plants of the partly deciduous forest formations on the Pacific slope between 800

and 1,200 (1,800) m elevation in Costa Rica; apparently flowering from June to

January. The species ranges from Guatemala to Costa Rica.

Schoepfia vacciniiflora is characterized by the stiff distichous leaves articulate at

the base, small usually pedicellate flowers with tubular corolla, and ellipsoid fruit

with circular distal area of different texture around the apex. This species is quite

easily distinguished from S. schreberiin Guatemala where the plants differ consis-

tently in flower form and altitudinal range. These distinctions are not so clear-cut

in Costa Rica where we have very few collections that are clearly referable to S.

vacciniiflora: Brenes 3554, 5367, 5661, 20334, 21051, all from near San Ramon, Ala-

juela province. Two high-altitude collections from 1,700 to 1,800 m altitude with

shorter less cylindrical corolla tubes are tentatively placed here: A. Jimenez 289 and

M. Valeria 1657 from San Jose province.

Schoepfia species A. Figure 3.

Trees ca. 20 m tall, leafy internodes 1-3 cm long, 1.5-3 mm thick, 2-ridged in early

stages, becoming terete and grayish brown. Leaves distichous, articulate at the base,

petioles 3-6 mm long, with 2 adaxial ridges becoming lateral in the distal half of the petiole

and continuous with the margins of the lamina, sulcate above at the base; laminae 5-11

cm long, 3-6 cm broad, ovate to ovate-oblong, acute to short-acuminate at the apex,

rounded and truncate or occasionally obtuse at the base, margins entire and revolute when

dry, laminae drying stiffly chartaceous, smooth and glabrous above and below, venation

pinnate with 3-5 pairs of major secondary veins, the lowest 2-4 secondary veins strongly

arcuate ascending. Inflorescences 1 or 2 in the axils of leaves, the peduncles becoming 6-12

mm long and bearing usually 3 subsessile or shortly pedicellate flowers distally. Flowers

not known. Fruit ellipsoid, ca. 12-14 mm long and 9-12 mm in diameter, red but drying

dark brown and lustrous with numerous longitudinal shallow grooves, a circular area of

different texture ca. 4 mm wide delimited by the calycular rim near the apex of the fruit.

Plants of the lowland Caribbean rain forest formations. The species is known

only from a single fruiting collection made in March, near the Rio Kama-Rio

Escondido in Bluefields, Nicaragua, by Proctor, Jones, & Facey (26972).

Schoepfia species A is recognized by the stiff ovate leaves articulate at the base

and with ascending basal secondary veins, the characteristic fruit on short pedun-

cles, and the lowland Caribbean forest habitat. This species resembles S. obliq-

uifolia Turcz. of Venezuela and Brazil, but without flowers it is difficult to say

where its true affinity lies. The very distinctive fruit and articulate leaves leave no

doubt that this tree is placed in the correct genus.

XIMENIA Linnaeus

Shrubs and small trees, hemiparasitic on roots of other plants, bisexual, stems with thick

spines produced by modified axillary branchlets, glabrous or puberulent. Leaves alternate,

articulate at the base, often borne in fascicles from short shoots, deciduous; laminae ovate

to elliptic, entire, pinnately veined. Inflorescences axillary, 1 to several and fasciculate,

usually a few- to many-flowered bracteate cyme or umbel, flowers sometimes solitary.

Flowers bisexual, regular, hypogynous, medium-sized, whitish; calyx small with 4 or 5

small teeth or lobes, not expanding in fruit; corolla of 4 or 5 free petals, valvate in bud, with

conspicuous hairs on the inner surface; stamens twice as many as the petals, filaments

usually narrow, anthers linear to oblong, 2-thecous, dehiscing longitudinally; a disk

absent; ovary superior, 3- or 5-chambered throughout the height of the ovary, ovules

pendulous from the central axis, with 1 integument, style narrow, stigma simple or

slightly lobed. Fruit a globose to ellipsoid drupe, yellow to deep purple, often with a juicy

edible pulp, seed with a small embryo at the apex of the fleshy endosperm.

BURGER: FLORA COSTARICENSIS 27

A circumtropical genus of eight species of which only one is known from

Central America. The presence of spines, articulate simple leaves often borne on

short shoots, the free petals densely hairy within, and the ovules pendulous from

a central column defining three or four separate locules help distinguish these

plants. The genus is named in honor of Francisco Ximenez, who, in 1615, pub-

lished an important work on the plants and animals of Mexico.

Ximenia americana L., Sp. PL 1193. 1753. Figure 3.

Shrubs and small trees to 7 m tall, branches often with thick spines formed by modified

axillary branches, leafy internodes (0) 0.5-4 cm long, 1.5-6 mm thick, with longitudinal

ridges in very early stages but soon becoming terete, pale grayish and lenticellate in age,

short shoots with clustered leaves often present. Leaves alternate, usually in a spiral,

clearly articulate at the base, petioles 3-8 mm long, with 2 longitudinal adaxial ridges

forming a narrow sulcus above that is minutely puberulent within; laminae (1.5) 3-8 (11)

cm long, (1) 1.5-4 (5) cm broad, elliptic to ovate-elliptic, acute to obtuse at the apex and

slightly retuse with a small mucro at the tip, acute to obtuse at the base, margins entire

and decurrent on the petiole, the laminae drying stiffly chartaceous but usually very brittle

and often dark in color, smooth and glabrous above and below, the 3-6 pairs of major

secondary veins becoming obscure in the distal half of the lamina. Inflorescences 1 to

several in the axils of leaves or fallen leaves, of 3-12 flowers in cymose, umbellate or

racemose arrangements, 1-3 cm long, the flowers often in groups of 3, primary peduncles

7-15 mm long, pedicels 2-6 mm long, subtended by minute (0.5 mm) deciduous bracts;

flowers ca. 5-10 mm long, glabrous on the outside, white or cream-colored but usually

drying dark, calyx forming a very shallow (0.5 mm) broad (1-1.5 mm) cup with 4 (3) blunt

teeth or lobes, the margin erose; petals usually 4, liguliform, 6-8 mm long, 1-1.5 mm

broad, valvate in bud, and becoming reflexed, with dense tomentulous hairs 1-2 mm long

on the inner surface, these white but turning reddish brown; stamens 8, equal, glabrous,

filaments 3-5 mm long, anthers linear, 3-4 mm long, ca. 0.7 mm broad, ovary 3-4 mm long

and 1-1.5 mm in diameter, glabrous, style 3-4 mm long, stigma simple. Fruit a yellow-

orange drupe, 2-3 cm long, 1.5-2.5 cm in diameter, ellipsoid, edible but with an acid taste.

Plants of seasonally dry deciduous and semideciduous forest formations of the

Pacific slope below 300 m elevation in Costa Rica; flowering material has been

collected from January to April, and fruit has been collected from March to May.

The species is pantropical.

Ximenia americana is recognized by the pale grayish branches often bearing thick

spines and leaves clustered on short shoots, small leaves with a minutely mu-

cronate tip and clearly articulate base, flowers with long hairs on the interior of

the petals, and the acid-tasting fruit. The small stature, deciduous leaves, and

restriction to the seasonally dry Pacific lowlands are further distinctions. The

indigenous name Tsu-kra has been recorded from the Boruca area (Pittier 11983).

The Central American representatives of this species have been placed in variety

americana by DeFilipps (Trans. Illinois State Acad. Sci. 62:350-358, 1969).

OPILIACEAE

Trees, shrubs, or climbers, bisexual or unisexual, generally glabrous; stipules absent.

Leaves alternate in a spiral, simple and entire. Inflorescences spikes, racemes, umbels, or

panicles, at first enclosed in imbricate bud scales; flowers bisexual or unisexual, radially

symmetrical, small; sepals united and minutely 4- or 5-lobed or reduced to a rim and not

apparent, petals (or tepals if the sepals are not apparent) 4 or 5, free and valvate in bud;

stamens as many as the petals and opposite them, filaments free, anthers 2-thecous, a disk

of 4 or 5 lobes or separate glands present outside the stamens and alternate with them;

ovary superior or united with the disk and half-inferior, 1-locular with 1 ovule pendulous

from a central basal placenta or erect from the base of the locule, style short or none, stigma

1. Fruit a drupe, seed with endosperm.

28 FIELDIANA: BOTANY

A family of seven or eight genera and about 50 to 60 species, mostly of African

and Asian tropics. The plants are probably all root parasites. The family is closely

related to the Santalaceae and Olacaceae and has been considered a part of

the Olacaceae by some workers. The Opiliaceae differ from the Olacaceae in the

structure of the disk, the usual reduction of the calyx, the lack of chambers at the

base of the ovary, and the lack of lignified cells in the leaves. This small family

resembles Olacaceae and Celastraceae in general appearance. The leaves often

turn a dull or dark yellowish green on drying.

AGONANDRA Miers

REFERENCES: L. O. Williams, The Agonandras of Mexico and Central America.

Ciencia (Mexico) 25:227-228. 1966. P. C. Standley, The North American species

of Agonandra. ]. Wash. Acad. Sci. 10:505-508. 1920.

Small trees, shrubs, or somewhat vinelike, usually dioecious, the bark generally corky

and drying pale gray, often with short shoots; stipules none. Leaves alternate and articu-

late at the base, petiolate, the laminae generally small and ovate to elliptic, entire. In-

florescences axillary, racemes at first subtended by decussate and caducous bracts (bud

scales); flowers small and unisexual, pale yellowish to greenish in color, calyx subentire or

minutely 5- or 4-lobed, petals usually 4 (5), free, valvate in bud, usually caducous in the

female flowers; stamens opposite the petals, filaments slender and equal, (stamens absent

in the female flower, a pistillode usually present in the male flower), disk present and

annular or of separate erect glands alternating with the stamens; pistil 1, superior or with

the disk adnate near the base, style absent and with a single sessile stigma. Fruit a drupe

with fleshy rind and stiff endocarp.

An American genus of about 20 species ranging from Mexico to Argentina and

most often found in seasonally very dry habitats. The genus is recognized by the

pale grayish bark, small alternate leaves distinctly articulate at the base, and the

small unisexual flowers in a racemose inflorescence at first enclosed in several

series of decussately imbricate bud scales.

Agonandra macrocarpaL. O. Williams, Ciencia (Mexico) 24:227. 1966. Figure 3.

Shrubs or small trees to 7 m tall (also said to be vinelike), unisexual, leafy internodes

0.1-2 cm long, 0.7-2.5 mm thick, green and strongly 2- or 3-ridged in early stages, older

stems with gray bark and becoming longitudinally furrowed when dry, glabrous. Leaves

alternate in a spiral, petiole 4-10 mm long, ca. 1-2 mm broad with winglike margins

continuous with the margin of the lamina, slightly expanded at the articulate base; lami-

nae 2-5.5 cm long, 1-3 cm broad, elliptic to ovate-elliptic, obtuse to bluntly acute at the

apex, rounded at the tip, acute to obtuse at the base, margin entire or undulate (dry) and

decurrent on the petiole, the lamina drying stiffly chartaceous or subcoriaceous, glabrous

and smooth above and below, venation pinnate with 3 or 4 pairs of major secondary veins.

Inflorescences originating from a bud covered with 4 distichous series of imbricate bud

scales (bracts) 1-2 mm broad at the base. Male racemes less than 3 cm long, the rachis

essentially glabrous, with 3 or 4 ridges and 0.3-0.5 mm thick, pedicels ca. 2 mm long, the

male flowers ca. 3 mm long with reflexed tepals (the calyx obscure), glands of the disk ca.

0.5 mm long, filaments 1.5-3 mm long, anthers 0.5 mm long. Female racemes less than 1

cm long, pedicels ca. 2 mm long, tepals caducous, a yellowish annular disk ca. 1.5 mm

broad conspicuous beneath the ovary, pistil ca. 1.2 mm long, ovary ca. 0.6 mm thick. Fruit

globose or slightly ellipsoid, ca. 2 cm long (dry), the outer fleshy exocarp drying to form

a smooth coriaceous yellowish brown surface, endocarp hard and yellowish, ca. 16 mm in

diameter.

Plants of the seasonally very dry deciduous (tropical dry and tropical dry-moist

zone transition) forest formations below 200 m elevation in northwestern Costa

Rica; flowers have been collected in January and mature fruit in May. The species

BURGER: FLORA COSTARICENSIS 29

is known only from Central Honduras and the provinces of Guanacaste and

northern Puntarenas in Costa Rica.

Agonandra macrocarpa is recognized by the small alternate leaves articulate at the

base, unusual inflorescence buds, minute unisexual flowers in small racemes, and

distinctive fruit. The seasonally very dry habitat, grayish corky stems, and gla-

brous parts further distinguish this species, which resembles some of our species

of Olacaceae. This species is apparently quite rare, with only three collections

known from Costa Rica: Brenes 22481 near Los Loros (Puntarenas), and Daubentnire

755 and Opler 508, both from near Canas. Agonandra obtusifolia Standley of north-

ern Central America and Mexico has minutely puberulent stems, whereas A.

brasiliensis Benth. & Hooker has longer finely puberulent inflorescences and

smaller fruit. Our species is closely related to A. racemosa (DC.) Standley of Mexico

and northern Central America, which has smaller fruit. The genus is poorly

represented in collections, and species designations can only be regarded as first

approximations.

LORANTHACEAE sensu lato

by William Burger and Job Kuijt

REFERENCES: B. A. Barlow and D. Wiens, The classification of the generic segre-

gates of Phrygilanthus(=Notanthera) of the Loranthaceae. Brittonia 25:26-39. 1973.

Job Kuijt, The Biology of Parasitic Flowering Plants. Univ. Calif. Press, 1-246.

1969. J. Kuijt, Mutual affinities of the Santalalean families. Brittonia 20:136-147.

1968. J. Kuijt, A revision of the Loranthaceae of Costa Rica. Bot. Jahrb. Syst.

83:250-326. 1964. J. Kuijt, Commentary on the mistletoes of Panama. Ann. Mis-

souri Bot. Card. 65:736-763. 1978. C. T. Rizzini, in Woodson and Schery, Lo-

ranthaceae, Flora of Panama. Ann. Missouri Bot. Card. 47:263-290. 1960.

Shrubs or small trees, unisexual or bisexual, usually leafy hemiparasites with chloro-

phyll, erect or scandent, parasitic on the aerial stems of the host or rarely terrestrial and

root parasites (as in Gaiadendron) , usually parasitic on the stems of dicotyledonous trees

and shrubs, attaching to the host by means of a haustorium (that developed from the

radical of the seed), sometimes with additional haustoria on epicortical roots, stems some-

times with thickened or articulate nodes. Leaves present or reduced to small scale leaves

(squamate) and apparently absent, usually opposite or subopposite, rarely whorled (some

Psittacanthus) or rarely alternate throughout (Antidaphne, Cladoclea), simple, glabrous, often

articulate at the base, laminae usually very stiff and drying brittle, with entire or undulate

margins, venation pinnate to palmate. Inflorescence basically an unbranched axis with

sessile or pedunculate flowers in groups of 2 (diads), 3 (triads), or solitary (monads), the

flower groups borne in a spicate, racemose, or paniculate arrangement, the flowers emerg-

ing from depressions in the rachis in some genera, bud scales enclosing young in-

florescences in some genera, bracts subtending the flowers or flower groups (monads,

diads, triads, etc.); flowers bisexual or unisexual, radially symmetrical or nearly so, peri-

anth present (absent only in male Antidaphne), floral morphology quite variable (see under

segregated families below), stamens usually as many as the perianth parts and opposite

them, pistil 1 and usually inferior, locule 1 or none, the ovule solitary and often not clearly

differentiated, style and stigma 1. Fruit a 1-seeded berry with a layer of viscous tissue,

often dispersed by birds, endosperm present in all but Psittacanthus, the endosperm with

or without chlorophyll, cotyledons 2 or rarely as many as 12 (in species of Psittacanthus).

While obviously related, the plants included here represent a wide range of

floral morphology and appear to include several different parallel lineages with

unusual adaptation to the parasitic life-style. The reduction of the ovarian cavity

and simplification of tissues of the ovule are probably associated with the evo-

lution of a fruit that germinates on the host and develops parasitizing roots.

30 FIELDIANA: BOTANY

Recent students of these plants are agreed that they are better treated as several

smaller families (see Kuijt, 1968, referred to above).

The plants included in this alliance (Loranthaceae in a wide sense) are usually

easy to recognize because of their shrubby epiphyte-resembling habit and para-

sitic attachment to the aerial stems of the host. The very stiff, usually opposite

leaves with entire or undulate margins and lack of pubescence are also character-

istic. Gaiadendron, a root parasite often found as a terrestrial tree, superficially

resembles our native species of Proteaceae, as do some of our larger species of

Psittacanthus. Cladocolea and Antidaphnehave consistently alternate leaves, whereas

the apparently naked flowers of Antidaphne are quite unique.

The genera of the Loranthaceae in a wide sense (sensu lato) are here placed

under the following families and in the following order: Eremolepidaceae, Lo-

ranthaceae in a narrow sense (sensu stricto), and Viscaceae. The following key

treats all these plants. Additional keys will be found under Loranthaceae (sensu

stricto) and Viscaceae. The figures of Eremolepidacae and Loranthaceae (sensu

stricto) are under the latter family, while the illustrations of Viscaceae are with the

keys to that family.

KEY TO THE LORANTHACEAE IN A WIDE SENSE IN COSTA RICA

Genera and species without family designation

will be found under the Loranthaceae in a narrow sense

la Leaves always alternate on all the stems, foliage leaves present 2a

Ib Leaves opposite, whorled, or occasionally alternate with some opposite or sub-

opposite leaves present, or the leaves reduced to opposing pairs of scale leaves and

foliage leaves absent 3a

2a Flowers apparently bisexual with conspicuous male and female parts in the

same flower (but the stamens or the pistil nonfunctional); perianth of 4 con-

spicuous petals, style at least 4 mm long; spikes without distal leaflike

parts Cladocolea

2b Flowers obviously unisexual, possessing organs of only 1 sex, petals absent in

the male flower and scalelike (2) in the female, style less than 1 mm long; female

spikes often with small distal leaves; (Eremolepidaceae) Antidaphne

3a Flowers solitary in axillary positions on main stem, or borne singly on spikes and

racemes, not found in diads, triads, or in longitudinal or whorled series 4a

3b Flowers borne in diads or triads on spikes and racemes or in longitudinal or whorled

arrangements on thick spikelike axes 5a

4a Inflorescence a raceme, the individual flowers pedicellate, the pedicel support-

ing a bract and 2 fused bracteoles beneath the flower, foliage leaves present

Struthanthus panamensis

4b Inflorescence of 1 to several flowers borne in the leaf axils; normal foliage leaves

absent, scale leaves present. Doubtfully present in Panama and not recorded

from Costa Rica Ixocactus

5a Flowers obviously borne in diads (groups of 2) or triads (groups of 3) and pedun-

culate or sessile on relatively thin (1-3 mm) axes, the diads and triads clearly

separated from each other 6a

5b Flowers borne in longitudinal ranks (or occasionally in whorls or forming rounded

flat groupings) on relatively thick (2-5 mm) spikelike axes, the flowers rarely in

clearly isolated groups of 2 or 3, the flowers often partly sunken into the axis of the

inflorescence lOa

6a Flowers more than 1 cm long, perianth bright red, orange, or yellow, bisexual,

the flowers occasionally smaller and subtended by leaflike bracts in Gaia-

dendron 7a

6b Flowers less than 1 cm long, perianth dull yellow to dark red, bisexual or

unisexual, never subtended by small leaflike bracts 8a

BURGER: FLORA COSTARICENSIS 31

7a Flowers golden yellow, subtended by green leafy bracts 3-20 mm long;

terrestrial or growing on branches of trees Gaiadendron

7b Flowers at least partly bright red or orange, subtended by a cupulate pedicel

apex and without leaflike bracts; parasites of aerial stems Psittacanthus

8a Flowers bisexual, both the pistil and anthers functional, perianth dark red or

dull yellow (in ours), inflorescences axillary and racemose or terminal and

paniculate Phthirusa

8b Flowers functionally unisexual, organs of both sexes may be present but either

the stamens or the pistil not functional, the male flower buds usually thickened

distally with the fertile anthers, the female flowers with tubular flower buds and

the anthers absent or reduced, perianth yellowish to dull white 9a

9a Filaments thin or the anthers subsessile; inflorescence always a simple racemose

or spikelike axis with sessile or pedunculate diads and triads of flowers

Stmthanthus

9b Filaments thick, the upper ones with lateral depressions; inflorescence often a

paniculate arrangement at the ends of twigs Phthirusa adunca

lOa Perianth parts usually 6, the flowers in 4 vertical (longitudinal) ranks on the thick

inflorescence axis, with 2 minute bracteoles subtending each flower and usually

found within the floral depression; spikes without distinct internodes; foliage leaves

present Oryctanthus

lOb Perianth parts usually 3, the flowers variously arranged on the spike, the individual

flowers not subtended by minute bracteoles; spikes with distinct internodes defined

by opposite pairs of scale leaves (also called bracts); (Viscaceae) lla

lla Anthers unilocular; plants with or without foliage leaves; flowers in single or

multiple longitudinal ranks (series) above each scale leaf (bract), the arrangement of

flowers on the internode often somewhat irregular; from 1,500-3,500 m in Costa

Rica Dendrophthora

lib Anthers bilocular; plants with foliage leaves; flowers always in multiple longitudinal

ranks above each scale leaf, flowers usually in regular ranks on the internode; from

sea level to 3,000 m in Costa Rica Phoradendron

EREMOLEPIDACEAE

REFERENCE: Job Kuijt, Mutual affinities of Santalalean families. Brittonia

20:136-147. 1968.

Shrubs, epiphyte-like hemiparasites, unisexual (or bisexual in some species of Ere-

molepis), epicortical roots produced in young stages (or possibly lacking in Eubrachion).

Leaves alternate and simple but reduced to scalelike squamae in Eubrachion, petiolate, the

lamina stiff with entire margin. Inflorescence spicate or racemose, at first enclosed in a

conelike bud of spirally arranged imbricate and scarious bud scales, the female in-

florescence often continuing with new vegetative growth distally in Antidaphne; flowers

very small and unisexual, sessile or the male pedicellate in Antidaphne, with a single

perianth whorl or apparently naked, the male flowers 2- to 4-parted, a perianth absent in

Antidaphne, the stamens opposite the perianth parts in Eremolepis and Eubrachion, filaments

present and free, anthers 2-thecous and dehiscing longitudinally; the female flowers 2- or

3-parted, ovary inferior (apparently half-inferior in Antidaphne and with 2 or 3 minute

deciduous perianth parts borne halfway up the pistil), locule 1, style 1, stigma capitate.

Fruit a small 1-seeded sessile berry, the endosperm with some chlorophyll in Antidaphne,

embryo with 2 cotyledons, viscous layer of the berry formed between the vascular bundles

of the perianth and the bundles of the ovary.

A small family of three genera ranging from southern Mexico and the largest

Caribbean Islands through tropical South America. The Eremolepidaceae are rep-

resented in our area by a single species of Antidaphne. These plants are easily

recognized because of their hemiparasitic habit on the aerial stems of host plants,

stiff alternate leaves, inflorescences emerging from buds with imbricate and alter-

nate bud scales, and the very small, naked or virtually naked unisexual flowers.

32 FIELDIANA: BOTANY

ANTIDAPHNE Poeppig and Endlicher

Shrubs, hemiparasites on aerial woody branches, unisexual, stems and leaves glabrous.

Leaves alternate in a spiral, simple and entire, epidermal cells sclerified except around the

stomata. Inflorescences at first enclosed in overlapping small bracts (resembling the scar-

ious bud scales of Fagaceae and other temperate plants), spicate or racemose, the female

inflorescences often with smaller than normal leaves distally and the inflorescence axis

sometimes continuing as a woody stem, the bracts (bud scales) deciduous and subtending

1 to several small flowers. Male flowers without a perianth, pedicellate, the stamens borne

from a central disklike cushion, stamens usually 2-4, filaments slender, thecae 2 and borne

on opposite sides of the connective, dehiscing laterally. Female flowers with 2 or 3 usually

caducous, minute, triangular tepals halfway up the ovary, pistil with a short distinct style

bearing a broad and flattened crescent-like stigma. Fruit a small 1-seeded berry; endo-

sperm pale green.

A genus of only a few species ranging in montane forests from southern Mexico,

Guatemala, and Costa Rica through northern South America to Peru, Bolivia, and

Brazil.

Antidaphne viscoidea Poeppig & Endlicher, Nov. Gen. & Sp. PI. 2:70, t. 199.

1838. Figure 8.

Epiphyte-like hemiparasitic shrubs 30-120 cm in diameter, unisexual, green to olive

green, leafy internodes 5-20 mm long, 1-3.5 mm thick, smooth and glabrous, becoming

longitudinally striate and grayish when dry. Leaves alternate in a 2/5 spiral, often articu-

late at the base, petioles 1-4 mm long, broad and with lateral wings continuous with the

lamina margins; laminae 1.4-8 cm long, 0.7-6 cm broad, obovate or occasionally elliptic

to ovate, rounded to bluntly obtuse at the apex, gradually narrowed to the attenuate or

subcuneate base, margin entire and of tissue differentiated from the leaf epidermis, the

laminae drying coriaceous, smooth and glabrous, venation varying from subpalmate with

dichotomizing primary veins to pinnately veined with 1-4 pairs of major secondary veins,

the major veins slightly raised on both surfaces when dry, obscurely punctate. In-

florescences at first resembling small axillary shoots enclosed by several series of caducous

pale brown bracts resembling scarious bud scales, the inflorescences 1 to several per node,

unisexual, the female often terminating in small leaves. Male inflorescences 5-15 mm long

(only rarely terminating in leaves as in Standley 43340 at US), racemose with 4-12 nodes,

each node with usually 1 (2, 3) naked flowers; male flowers 2-3 mm long (including the

short pedicels), lacking perianth, stamens 2-4 and arising from beneath the edge of a

central disklike cushion, filaments ca. 1 mm long, anthers dehiscing laterally with longi-

tudinal slits. Female inflorescences spicate, at first ca. 5-10 mm long but soon becoming

woody and with small leaves distally, with ca. 10 nodes, the lower nodes with caducous

bracts (bud scales) and the upper with small leaves, with 1-3 flowers per node; female

flowers less than 1 mm long, sessile, lacking perianth-like whorls but with 2 or 3 deciduous

scalelike perianth parts attached halfway up the ovary, style very short, stigma crestlike

and flattened. Fruit a berry ca. 3 mm long, ovoid, sessile on a woody rachis 2-5 cm long

and terminating with leaves 1-5 cm long, the infructescence sometimes continuing to

grow beyond the distal leaves and producing new axillary shoots.

Parasites of trees in moist and wet montane evergreen forest formations be-

tween (700) 1,000 and 2,000 m elevation; probably flowering throughout the year,

but collected with flower or fruit primarily from December to April. The species

ranges from southern Mexico to Amazonian Brazil, Peru, and Bolivia.

Antidaphne viscoidea is distinguished from our other shrubby and epiphyte-like

parasites by the alternate leaves with variable venation, young shoots and in-

florescences enclosed by overlapping caducous bud scales giving a conelike ap-

pearance, and the flowers lacking a perianth in the male or with a very minute

scalelike perianth in the female.

BURGER: FLORA COSTARICENSIS 33

LORANTHACEAE sensu stricto

Shrubs or small trees, leafy hemiparasites with chlorophyll, bisexual or the flowers

functionally unisexual and the plants unisexual (dioecious), parasitic on aerial stems and

resembling epiphytes or a few genera parasitic on roots and terrestrial or epiphytic and

parasitizing the roots of epiphytes (as in Gaiadendron), attaching to the host by means of

roots provided with haustoria, stems sometimes with thickened nodes, glabrous or with

a scurfy epidermis. Leaves usually opposite and decussate, sometimes whorled, sub-

opposite and alternate leaves occasionally present, simple, often articulate at the base,

laminae usually very stiff and drying brittle, with entire or undulate margins, glabrous in

ours. Inflorescences usually with an unbranched axis (in ours) with sessile or pedunculate

flowers in decussate groups of 3 (triads), 2 (diads), or solitary (monads), the flowers

emerging from depressions in the rachis in Oryctanthus; flowers bisexual or functionally

unisexual (and then the plants always unisexual), the perianth generally interpreted as

representing 2 whorls, the outer whorl a calyculus and the inner a well-defined whorl of

petals, the calyculus with small teeth or with only a thin rim (at the distal periphery of the

ovary) continuous with and of the same texture as the ovary wall, inner perianth of 6 (7)

petals, valvate in bud, longer than broad, often brightly colored, stamens equal in number

and opposite the petals, borne on the petals, filaments present or the anthers subsessile

on the petals, dehiscing by longitudinal slits, pollen usually 3-lobed; pistil with inferior

ovary, locule 1 or ovary solid, ovules replaced by a central papilla in the locule, style 1 with

capitate stigma. Fruit a 1-seeded berry, endosperm compound and lacking chlorophyll,

cotyledons 2 or rarely as many as 12 (in Psittacanthus); seed surrounded by viscid tissue in

all but Gaiadendron.

The Loranthaceae (in a narrow sense) are a largely pantropical family of about

65 genera and 900 species. None of the New World genera occur in the Old World.

The definition of genera in the family is often very unsatisfactory, especially

regarding South American representatives. Many neotropical species of this fam-

ily were at one time assigned to the genus Phrygilanthus which has been shown

to be a rather heterogeneous assemblage by Barlow & Wiens (1973), who have

distributed all species over several other genera. Additionally, these authors have

pointed out that Phrygilanthus is nomenclaturally illegitimate. One of the main

tasks ahead is the evaluation of the resultant genera in terms of various mor-

phologic and other features. Another extremely difficult complex is present in

Struthanthus and Phthirusa; few botanists, if any, have followed Baehni and Mac-

bride's union of these two genera into one (Struthanthus), but no one has pro-

posed a satisfactory separation. A special case is the remarkable Phrygilanthus

panamensis Rizzini, occurring in Chiriqui Province, Panama. It was placed in

Struthanthus by Barlow & Wiens (1973), a practice here accepted as expedient but

not final (see under Struthanthus).

Among these parasitic plants, the genus Oryctanthus, while very distinctive,

may be difficult to distinguish from the Viscaceae. Collections of Oryctanthus often

resemble species of Phoradendron or Dendrophthora (Viscaceae) because of the soli-

tary flowers emerging from the rachis and the fruiting rachis with longitudinal

ranks of fruit or depressions left by the fruit. In contrast to Central American

species of the last two genera, where series or groups of unisexual flowers are

present above each inflorescence bract, Oryctanthus has a solitary flower in the axil

of each bract, and all flowers are perfect and accompanied by two minute lateral

bracteoles. Oryctanthus almost invariably has several epicortical roots at its base,

organs which are unknown in any Viscaceae.

PSITTACANTHUS

nodosus

PSITTACANTHUS Nsfl ramiflorus

GAIADENDRON

punctatum

FIG. 5. Loranthaceae (s.s.): species of Psittacanthus and Gaiadendron.

34

• 10 cm

STRUTHANTHUS

panamensis

S. woodsonii

leptostachyus

PHTHIRUSA

retroflexa

\

FIG. 6. Loranthaceae (s.s.): species of Struthanthus, Phthirusa, and Cladocolea.

35

S. rotundatus

5 mm

STRUTHANTHUS

oersted ii

FIG. 7. Loranthaceae (s.s.): additional Costa Rican species of Struthanthus.

36

ANTIDAPHNE

viscoidea

A stem of Casearia

with old flowers

of the parasite

Apodanthes caseariae

ORYCTANTHUS

alveolatus

O. spicatus

ORYCTANTHUS

occidentalis

FIG. 8. Eremolepidaceae: Antidaphne, upper left. Rafflesiaceae: old flowers of Apodanthes

on the stem of a Casearia (Flacourtiaceae), upper right. Loranthaceae (s.s.): four species of

Oryctanthus.

37

38 FIELDIANA: BOTANY

KEY TO THE COSTA RICAN GENERA OF LORANTHACEAE IN A NARROW SENSE

la Flowers unisexual (aborted stamens or styles sometimes present) 2a

Ib Flowers bisexual (perfect) 4a

2a Phyllotaxy alternate throughout; flowers mostly single in the axis of squamate

inflorescence bracts, the latter sometimes caducous 3a

2b Phyllotaxy decussate; flowers in paired triads Struthanthus

3a Flowers with clearly recognizable aborted organs of the opposite sex; spikes

never with terminal leafy organs; petals conspicuous, always 4; style at least

4 mm long Cladocolea

3b Flowers lacking aborted organs of the opposite sex; female spikes normally

with small, distal leaves which may expand into full size after flowering;

petals lacking in male, less than 0.5 mm in female, 2 or 3 in number; style less

than 0.5 mm long Antidaphne

4a Inflorescence a spike or raceme with decussate single flowers 5a

4b Inflorescence a spike or raceme with decussate triads or diads 6a

5a Flowers less than 4 mm long, sessile or even sunken in inflorescence axis, flanked

by minute separate bracteoles within the rachis cavity; pollen with 3 circular

depressions on each hemisphere Oryctanthus

5b Flowers more than 6 mm long, sessile in a cup made up of primary bract and 2

conspicuous bracteoles, this compound structure elevated on a 1-mm long pedi-

cel; pollen lacking depressions Struthanthus panamensis

6a Flowers less than 4 mm long; epicortical roots from base of plant Phthirusa

6b Flowers longer than 4 mm; basal roots absent or resembling those of normal terrestrial

plants 7a

7a Floral bracts green and foliose, flowers golden yellow; plants either of true epi-

phytic habit or terrestrial; fruit with abundant, grooved endosperm

Gaiadendron

7b Floral bracts never bright green and foliose; flowers always with some red; plants

parasitic on tree branches, lacking any epicortical or other roots; fruit lacking

endosperm Psittacanthus

CLADOCOLEA Van Tieghem

REFERENCE: Job Kuijt, The genus Cladocolea. ]. Arnold Arbor. 56:265-335. 1975.

Erect or scandent shrubs, hemiparasitic on aerial stems, bisexual or unisexual, lateral

branches and inflorescences often emerging in a pseudoendogenous fashion, glabrous or

puberulent in early stages; epicortical roots absent or present, on stems or at the base of

the plant. Leaves opposite and decussate, alternate, or irregularly arranged, simple and

entire, sometimes reduced to scales. Inflorescences usually a determinate spike, a capitu-

lum, dichasium, or raceme, in some species the terminal flower is lost and in a few the

inflorescence is reduced to a solitary flower with a pair of bracts, inflorescences usually

axillary to leaves or fallen leaves; lateral flowers borne in the axils of scalelike or leaflike

bracts but without bracteoles. Flowers bisexual or functionally unisexual with organs of

the nonfunctional sex often well developed, mostly green to yellow in color, sessile or

pedicellate, 4-, 5-, or 6-parted, stamens borne on the petals, monomorphic or dimorphic

within the flower, anthers 4-thecous, pollen rounded-triangular, style straight or often

geniculate or curved distally. Fruit a 1-seeded berry with endosperm, embryo di-

cotyledonous, elongate or globose, the haustorial disk weakly developed.

The genus Cladocolea, with 23 species, is centered in Mexico north of the Isth-

mus of Tehuantepec. A single species ranges to Panama in a (presently) disjunct

distribution, and four species occur in tropical South America. The genus is

distinguished from closely allied plants by the terminal floret (determinate in-

florescence) and by the lack of bracts (bracteoles may be present). Most of the

species of the genus have bisexual flowers, but our species has functionally uni-

sexual flowers that may appear to be bisexual. The genus Cladocolea is closely

related to Struthanthus, especially species such as S. orbicularis and S. polystachyus.

BURGER: FLORA COSTARICENSIS 39

Cladocolea oligantha(Standl. & Steyerm.) Kuijt, J. Arnold Arbor. 56:317. 1975.

Struthanthus oliganthus Standley & Steyermark, Publ. Field Mus. Nat. Hist., Bot.

Ser. 23:154-155. 1944. Figure 6.

Small shrubs, branch parasites, unisexual, few branched with relatively straight stems,

terete, becoming dark brown and often with transverse lenticels; no basal or stem roots

seen. Leaves alternate, gradually narrowed at the base and the petiole (not clearly differ-

entiated) ca. 2-4 mm long and 0.5-2 mm thick; laminae 2-3.5 (4.5) cm long, 0.8-1 (1.6) cm

broad, oblanceolate or occasionally lanceolate, apex acute or rounded, attenuate at the

base, venation usually obscure, pinnate or subpalmate. Inflorescences of 2 kinds: (1) the

primary inflorescences borne on the current year's growth are mostly solitary in the axil

of a leaf with usually simple 3-flowered dichasia borne on peduncles 4-5 (10) mm long and

1 mm thick with the flowers borne in 1 plane, and (2) the secondary inflorescences (very

similar to the primary but borne on the year-old growth) with 4-6 flowers borne in a

congested spiral at the apex of a short peduncle and having a single terminal flower, the

secondary inflorescence subtended by 1 or 2 secondary leaves that resemble primary

foliage leaves but developed later. Flowers functionally unisexual, apparently bisexual but

the anthers of female flowers nonfunctional and the male flowers with a prominent style

but lacking a functional stigma, male and female flowers of about the same size and form,

ca. 5 mm long and 2 mm thick in bud before anthesis, said to be reddish, 4-parted, anthers

1.5 mm long and subsessile halfway up the petals, 4-thecous with a projecting connective,

style at least 4 mm long, stigma inconspicuous. Fruit red becoming black, ovoid, ca. 7 mm

long and 5 mm in diameter, smooth, embryo nearly 4 mm long, with 2 cotyledons.

Epiphyte-like hemiparasites found between 800 and 2,000 m elevation in Gua-

temala and Mexico and from near sea level in Panama. The species ranges from

central Mexico to central Panama, but has not been collected in El Salvador,

Honduras, Nicaragua, Costa Rica, or the westernmost provinces of Panama. The

species is usually found on trees of the Burseraceae, but is also recorded on

Leguminosae.

Cladocolea oligantha is recognized by the shrubby parasitic habit (usually on trees

of the Burseraceae), the small oblanceolate alternate leaves, the small in-

florescences with three to six congested flowers at the apex of the peduncle, and

the flowers that appear to be bisexual but are functionally unisexual.

GAIADENDRON G. Don

Shrubs or trees to 14 m tall, terrestrial or epiphytic, hemiparasites on the roots of shrubs

or herbs. Leaves opposite or subopposite, petiolate, usually dark green, thick, minutely

punctate beneath. Inflorescences raceme-like panicles with a single main axis bearing

usually opposite triads of flower, the triads short-pedunculate, the flowers each sub-

tended by a bract, larger bracts leaflike; flowers bisexual and regular, narrow and tubular

before anthesis, ovary inferior with a short calyculus, the epigynous perianth parts (petals)

6 or 7, free and often recurving, bearing the adnate stamens, anthers dorsifixed and

versatile, dehiscing longitudinally. Fruit a berry, subtended by the persisting bracts.

A very small genus ranging from the mountains of Nicaragua southward to

Peru. These plants resemble some of our native Proteaceae as well as species of

Psittacanthus and Struthanthus (q.v.).

Gaiadendron punctatum (R. & P.) G. Don, Gen. Syst. 3:431. 1834. Loranthus

punctatus Ruiz & Pavon, Fl. Peruv. & Chil. 3:47, pi. 277a. 1802. G. poasense Donn.

Sm., Bot. Gaz. 56:61. 1913. Figure 5.

Shrubs or small trees, hemiparasites, epiphytic or if terrestrial often on disturbed sites,

bisexual, leafy internodes 5-40 mm long, 1.5-6 mm thick, glabrous, longitudinally ridged

and grayish in age, terete. Leaves opposite or subopposite, petioles4-10 mm long, 1-2 mm

40 FIELDIANA: BOTANY

thick, sulcate above, revolute margin of the lamina base continuous with the margins of

the petiole; laminae 3-8 (10) cm long, 1.3-4 (4.7) cm broad, elliptic or somewhat obovate,

usually obtuse at the apex and the tip bluntly rounded, acute to obtuse at the base, margin

entire and revolute (especially near the petiole), the laminae drying subcoriaceous, smooth

on both surfaces and often lustrous above, glabrous, the 6-16 pairs of major secondary

veins often obscure on both surfaces or slightly impressed above, small dark punctate dots

usually visible on the lower surface. Inflorescences 1-3 in the axils of leaves or several

terminal (and occasionally causing a forked growth pattern), usually near the ends of

twigs, 4-8 (14) cm long, raceme-like panicles with a single prime rachis and up to 16 triads

often in opposite pairs, the triads on peduncles 1-3 mm long with 2 shorter (3-8 mm)

bracts subtending the lateral flowers and a larger (5-20 mm) bract subtending the median

flower. Flowers bisexual, perianth of 6 (7) parts, 7-12 (20) mm long, narrow and hooked

apically, golden yellow to orange-yellow, recurved after anthesis, filaments borne halfway

up the perianth, free portion variable in length, anthers (0.5) 0.7-1.5 (2) mm long, ver-

satile; ovary inferior, ca. 3-4 mm long with a short (0.3 mm) flared calyculus, ca. 1 mm

thick, style to 15 mm long, stigma simple. Fruit often in triads subtended by the persisting

bracts and on peduncles to 10 mm long, orange berries 7-10 mm long, ellipsoid or slightly

obovoid (dry), apically blunt, endosperm with as many lobes (in cross section) as the

flower had perianth parts, embryo with 2 (3) cotyledons.

Plants of the evergreen montane forest formation between (800) 1,600 and 2,800

(3,000) m elevation; flowering throughout the year, but collected most often be-

tween October and April in Costa Rica. The species ranges from Nicaragua south-

ward to Peru.

Gaiadendron punctatum is recognized by its narrow flower with showy orange

and recurving perianth parts, flowers usually borne in threes on an unbranched

raceme-like axis, conspicuous floral bracts (especially those of the central flowers),

and lustrous leaves. The plants and their flowers resemble our native species of

Proteaceae, some of which occur in the same habitats. Plants of this species are

parasites of herbs and small trees. These plants can be both true epiphytes and

true parasites by being epiphytic and parasitizing other epiphytes, but not the

supporting tree (see Kuijt, On the ecology and parasitism of the Costa Rican tree

mistletoe, Gaiadendron punctatum, Canad. J. Bot. 41:927-938, 1963).

ORYCTANTHUS Eichler

Nomen Conservandum Propositum

REFERENCE: J. Kuijt, Revision of the genus Oryctanthus (Loranthaceae). Bot.

Jahrb. Syst. 95:478-534, 1976.

Small shrubs, hemiparasites of aerial stems, bisexual, young stems terete or with longi-

tudinal ridges, often with contiguous small lenticels or with a scurfy reddish brown

surface; attached to the host by a primary haustorium and a small number of epicortical

roots from the base of the plant only, or rarely such roots absent. Leaves opposite or

subopposite, rarely with a few alternate leaves on a stem, symmetrical, sessile or petiolate,

edge of the lamina often with differentiated tissue, venation palmate to pinnate, stellate

fiber bundles of lamina often visible when leaves dry. Inflorescences short to long spikes,

1 to several in the axils of leaves or on squamate (leafless) terminal shoots and forming

compound inflorescences (as in some O. alveolatus) , the flowers in 4 longitudinal (vertical)

ranks and emerging from depressions within the rachis, the depressions (pits) subtended

by a floral bract that is often largely united with the thickened rachis and obscure, 2 minute

lateral prophylls (bracteoles) present on each side (proximal) within the floral depression;

flower small, sessile, bisexual (in ours) solitary in the depressions in the rachis, calyculus

subentire, perianth of 6 free parts (petals), stamens adnate to the lower half of the perianth

parts, filaments very short, anthers very small and rounded but with variably prominent

connective, 4- or 2-thecous, dehiscing longitudinally; both petals and stamens dimorphic;

pollen globose, triangular-rounded with 3 deep circular depressions on each hemisphere,

BURGER: FLORA COSTARICENSIS 41

style slender, stigma capitate. Fruit a 1-seeded berry, ovoid to cylindrical, truncate at the

apex with the calyculus prominent and often with a persistent circular perianth scar.

A genus of about 10 species ranging from southern Mexico to Brazil and Bolivia.

Our species are found only in evergreen or partially deciduous life zones. The

emergence of solitary flowers from depressions in the inflorescence axes in early

stages is an unusual feature in the Loranthaceae (s.s.), but is superficially similar

to the situation in Phoradendron and Dendrophthora of the Viscaceae (q.v.). The

genus also possesses anatomical peculiarities (see Kuijt, Notes on the anatomy of

the genus Oryctanthus, Canad. J. Bot. 39:1809-1816, 1961).

la Young stems flattened or with longitudinal ridges, at least when young 2a

Ib Young stems terete 3a

2a Leaves petiolate, laminae generally small (1.8-7 cm) and elliptic; stems with

longitudinal ridges with rough reddish brown surface; spikes to 2.5 cm long,

flowers inclined forward (toward the apex of the spike); 500-1,800 m elevation

O. spicatus

2b Leaves sessile or subsessile, laminae usually broadly ovate, 4-14 cm long; stems

lacking scurfy reddish brown ridges; spikes 2-12 cm long, flowers emerging at

right angles to the spike; 0-1,000 m elevation O. cordifolius

3a Spikes always axillary to foliage leaves; fruits and flowers emerging at right angles to

the rachis; fruit usually constricted around the center; 0-1,400 m elevation

O. occidentalis

3b Spikes axillary to foliage leaves but also often on squamate (leafless) terminal shoots

resembling panicles; fruits and flowers inclined forwardly on the rachis (toward the

apex of the spike); fruit ovoid; 0-800 (1,200) m elevation O. alveolatus

Oryctanthus alveolatus (H.B.K.) Kuijt, Bot. Jahrb. Syst. 95:504. 1976. Loranthus

alveolatus H.B.K., Nov. Gen. & Sp., Quarto text, 3:444. 1820. L. amplexicaulis

H.B.K., loc. cit. 445, non DC., Prodr. 4:305-306. 1830. Oryctanthus botryostachys

Eichl., Fl. Brasil. 5, pt. 2:89-90. 1868. Figure 8.

Small shrubs, usually less than 1 m in diameter, leafy internodes (1) 2-8 cm long, 1.3-4

mm thick, bark flaking off or rough and scurfy on young stems and inflorescences, reddish

brown, terete. Leaves opposite or subopposite, petioles 0-4 mm long, 1-2 mm broad, flat

or slightly sulcate above, scurfy margins of the lamina continuous with the upper edges

of the petiole; laminae 2-8 (10) cm long, 1.5-5 (9) cm broad, narrowly to broadly ovate or

suborbicular in the larger leaves, bluntly obtuse to rounded at the apex, obtuse to rounded

at the base, margin entire and occasionally undulate (dry) with a distinct edge of differ-

entiated slightly roughened or scurfy reddish brown tissue, the laminae drying stiffly

chartaceous to subcoriaceous, smooth and glabrous, venation palmate to subpalmate with

3-7 primary veins, the major veins slightly raised above and usually obscure below.

Inflorescences 1 or a few in the axils of leaves or at nodes in long terminal squamate shoots

that resemble a racemose panicle, spikes 1-4 (6) cm long, peduncle less than 3 mm long,

rachis 1.5-3 mm thick, terete, reddish brown with rough or scurfy texture; flowers emerg-

ing from deep depressions in the rachis, inclined forwardly at an angle of ca. 45°, alternat-

ing in 4 vertical ranks, perianth 1-1.5 mm long, green on the outside and reddish within.

Fruit becoming 5 mm long and 3.5 mm thick, ovate, lower half white, upper half dark

green, truncated at the apex, the mature fruit usually angled forwardly as in the flowers.

Epiphyte-like hemiparasites of evergreen wet lowland forest formations usually

below 1,000 m elevation; probably flowering throughout the year, but collected

only between January and August. The species ranges from central Costa Rica on

both the Caribbean and Pacific coastal plains to Brazil and Bolivia.

Oryctanthus alveolatus is recognized by the sessile to short-petiolate leaves, the

scurfy reddish brown young stems that are always terete, and the relatively short

spikes that are often found in opposite groups or pairs on terminal leafless stems

that resemble open compound inflorescences. In Costa Rica, O. alveolatus seems to

42 FIELDIANA: BOTANY

fall rather clearly into two categories corresponding to what has been known as

O. amplexicaulis and O. botryostachys. The former has nearly orbicular sessile leaves

and a terminal compound inflorescence; the latter bears more ovate leaves with

short, stout petioles, and seems to produce axillary inflorescences only. In South

America, however, no such distinction can be maintained, and the present con-

servative arrangement is therefore considered advisable for Costa Rica.

Oryctanthus cordifolius (Presl) Urban, Bot. Jahrb. Syst. 24:30. 1898. Viscum

cordifolium Presl, Epim. Bot. 253. 1849. Figure 8.

Large shrubs, leafy internodes 3-12 cm long, 1.8-6 mm thick, with a roughened or

minutely striate surface in early stages (dry), with 2 (4) longitudinal ridges and somewhat

flattened but soon becoming terete. Leaves opposite, sessile or subsessile with the petiole

obscured by the basal lobes; laminae 4-14 cm long, 3-12 cm broad, narrowly to broadly

ovate or suborbicular, bluntly obtuse to rounded and emarginate at the apex, truncate to

rounded and cordate at the base, margins entire and somewhat revolute, tissue of the edge

differentiated, usually brownish and smooth, the laminae drying very stiffly chartaceous

to subcoriaceous, smooth and glabrous, dark green and lustrous in life, venation palmate

or subpalmate with 5 or 7 primary veins (occasionally pinnate), the major veins arcuate

ascending, prominent above and somewhat obscure beneath. Inflorescences 1-3 in the

axils of leaves, 2-12 cm long, peduncles 4-20 mm long, brownish and smooth; flowers

sessile in depressions of the spike at right angles or slightly angled toward the apex, the

4 vertical ranks of closely spaced flowers often producing an alternating position in flowers

of adjacent ranks, perianth ca. 2.5 mm long, red, ovary usually included in the depression

at anthesis. Fruit becoming 5 mm long and 3 mm thick, ovoid to somewhat thickened

above and obovoid, becoming dark maroon or black, smooth or occasionally muricate near

the truncated apex.

Epiphyte-like hemiparasites of evergreen wet forest formations between sea

level and 1,000 m elevation on both the Caribbean and Pacific sides of Costa Rica;

probably flowering throughout the year, but collected only from February to April

and in August and September. The species ranges from southern Mexico to

Colombia.

Oryctanthus cordifolius is recognized by its strictly opposite sessile and subcor-

date leaves, young stems with longitudinal ridges, fruit becoming over 5 mm long

(3 mm dry), and the spikes often becoming more than 8 cm long. This species is

very similar to O. occidentalis, and occasionally, collections seem to be intermediate

between the two. However, this may be due more to the variability within popu-

lations of O. cordifolius than to a close relationship.

Oryctanthus occidentalis (L.) Eichler, in Martius, Fl. Brasil. 5, 2:89. 1868. Lo-

ranthus occidentalis L., Amoen. Acad. 5:396. 1760. Figure 8.

Small shrubs ca. 1 m long, leafy internodes (1) 2-7 (10) cm long, (0.7) 1.2-3 (5) mm thick,

terete, with a scurfy reddish brown surface when young and often with many lenticels;

epicortical roots slender. Leaves opposite or subopposite, petioles (0) 2-6 mm long, 1.4-4

mm broad, somewhat sulcate above with wings formed by the decurrent lamina margins;

laminae 3-9 (16) cm long, 2-8 (14) cm broad, elliptic to elliptic-ovate or ovate, very broadly

ovate in the unusually large leaves, bluntly obtuse to rounded at the apex, obtuse to

rounded (truncate to subcordate in the very large laminae) at the base, margin entire and

with differentiated rough brownish tissue along the edge, slightly revolute near the pet-

iole, laminae drying very stiffly chartaceous to coriaceous, smooth and glabrous, the 1-3

pairs of major secondary veins arising from the lower V* of the midvein, arcuate ascending,

usually slightly raised above and obscure beneath. Inflorescences axillary to the current

foliage, 2-6 spikes per leaf axil, 8-30 mm long and elongating in fruit to as much as 10 (15)

cm, peduncles 2-15 (25) mm long, 0.8-1.6 mm thick, scurfy brown; flowers usually

emerging at right angles to the rachis, closely approximate in 4 vertical ranks of circular

BURGER: FLORA COSTARICENSIS 43

depressions, perianth 1-2 mm long, yellowish orange to red, anthers 4- (rarely 2-) celled.

Fruit 2-3 mm long, cylindrical and abruptly truncate at the apex, ca. 1.7 mm in diameter

(dry), yellowish at the base, green above but with the calycular ring sometimes becoming

thickened and red (in ours), closely spaced circular depressions left by the falling fruit.

Epiphyte-like hemiparasites of evergreen moist and very wet forest formations

from sea level to 1,000 (1,400) m elevation; flowering throughout the year, but

collected most often in January and February and from May through August. The

species ranges from Costa Rica to Colombia with a morphologically distinct pop-

ulation on the island of Jamaica (the type locality).

Oryctanthus occidentalis is recognized by its stiff strictly opposite leaves on terete

stems, small bisexual flowers emerging in perpendicular fashion from deep de-

pressions in the axis of the spike, and fruit somewhat constricted around the

middle. The leaves vary greatly in different plants and occasionally on the same

plant. The larger leaves (over 8 cm long and 6 cm broad) are not found associated

with the smaller (5- by 2-cm) elliptic leaves, but many collections contain inter-

mediate sizes.

Within Costa Rica, considerable differences seem to exist between individuals

from the Atlantic and Pacific slopes. Collections from the General Valley-Golfo

Dulce area have longer petioles and peduncles, larger leaves, a much more prolific

production of spikes, and a fruit which lacks the tubercular distal surface (Kuijt,

1964). Some plants from this region are capable of originating as many as four or

five shoots from a single root (as in Kuijt 2440).

Oryctanthus spicatus Qacq.) Eichler, in Martius, Fl. Brasil. 5, pt. 2:89. 1868.

Loranthus spicatus Jacq., Enum. PL Carib. 18. 1760. Struthanthus guatemalensis

Standl., Field Mus. Nat. Hist., Bot. Ser. 17:237. 1937. Oryctanthus guatemalensis

(Standl.) Standl. & Steyerm., Publ. Field Mus. Nat. Hist., Bot. Ser. 23:40. 1944.

Figure 8.

Small shrubs, ca. 50 cm in diameter but occasionally with stems 1 m long, leafy inter-

nodes 1-4 (6) cm long, 1.5-4 mm thick, with 2 or 4 longitudinal scurfy reddish brown

ridges, often smooth between the ridges or entirely scurfy on the very young stems, often

somewhat flattened but becoming terete in age; epicortical roots usually 3. Leaves opposite

or subopposite (occasionally alternate), petioles (2) 4-8 mm long, 0.7-1.8 mm thick,

slightly sulcate above with ridges continuous with the lamina margins, the lower side with

a longitudinal scurfy ridge continuous with the midvein; laminae 1.8-6 (8) cm long,

0.8-2.5 (3.5) cm broad, elliptic to oblong or slightly obovate, bluntly obtuse to rounded and

emarginate at the apex, acute to subtruncate at the base, margin entire and becoming

somewhat revolute, edge with differentiated brown and slightly roughened tissue, the

laminae drying stiffly chartaceous to subcoriaceous, smooth and glabrous but the midvein

often raised and with scurfy tissue beneath (abaxially), venation pinnate with 2-4 pairs of

major secondary veins arising from the lower half of the midvein and arcuate ascending,

raised or obscure on either surface. Inflorescences 1 or 3 in the axils of current foliage,

0.4-2.5 cm long, peduncles 2-7 mm long, rachis ca. 1.4 mm thick (dry) with up to 24

flowers; the flowers emerging from shallow depressions in the rachis at an acute angle to

the apex, perianth ca. 1.5 mm long, dark wine red. Fruit to 5 mm long and 3.5 mm thick

(dry), ovoid to somewhat cylindrical, truncated at the apex, smooth.

Epiphyte-like hemiparasites of evergreen or partly deciduous forest formations

between 500 and 1,800 m elevation; apparently flowering throughout the year.

The species ranges from Guatemala to Peru and Venezuela, where it intergrades

with the South American O. florulentus (Rich.) Urban. It is remarkable that the

species has not yet been collected from either Nicaragua or Panama.

Oryctanthus spicatus is recognized by the generally smaller leaves on young

stems with longitudinal scurfy ridges, the small spikes with relatively few flowers,

44 FIELDIANA: BOTANY

and the restriction to middle elevation habitats. The few flowers obscure the

four-ranked nature of the flowers on the spike, leaving the old spike terete rather

than four-angled as in fully mature fruiting spikes in some of the other species.

PHTHIRUSA Eichler

Nomen Conservandum Propositum

Erect or scandent shrubs, usually bisexual, hemiparasitic on aerial stems; rooting only

from the base of the plant (in our species) or also from the stems. Leaves opposite and

becoming articulate at the base (in ours), subopposite or rarely alternate. Inflorescences

usually solitary in the leaf axils, on leafless terminal or lateral shoots in a few species

(South America), imbricate bud scales lacking in early stages, racemose or spicate to

paniculate in superficial appearance, an indeterminate thyrse of decussate dichasia (triads)

in ours, the flowers borne in triads (dichasia) or rarely in monads, each triad subtended

by 1 primary and 2 secondary bracts, the lateral flowers associated with 3rd- and 4th-order

bractlets, bracts and bractlets often connate to form a cup at the base of the flowers, triads

sessile or very short pedunculate; flowers usually bisexual, calyculus thin and entire,

petals usually 6 (4), stamens of 2 lengths, usually borne on the base of the perianth parts,

filaments of 2 lengths, the longer often scalloped by the pressure of adjacent anthers on

the shorter filaments, anthers dehiscing longitudinally, said to produce pollen grains of a

single type: triangular in shape with finely granulose exine and 3 pores, ovary with or

without a locule, with only 1 functional megaspore (in ours). Fruit a 1 -seeded berry,

endosperm without chlorophyll.

A genus of about 30 species ranging from southern Mexico and the West Indies

southward to Brazil. The genus is poorly defined and may not be distinct from

Struthanthus. One of our species seems to be self-pollinating (see Kuijt &

Weberling, The flowers of Phthirusa pyrifolia, Ber. Deutsch. Bot. Ges. 85:467-480,

1972).

In 1830 Martius erected the genus Phthirusa on the basis of a single species now

placed in Cladocolea (Kuijt, 1975). In order to reduce nomenclatural confusion, it

has been proposed that Phthirusa in the sense of Eichler (1868) be conserved over

the original sense of Martius (see Taxon 24:389, 1975). Phthirusa pittieri Krause,

based on a type from Colon, Panama, has proven to be Struthanthus leptostachyus.

KEY TO THE SPECIES OF PHTHIRUSA IN COSTA RICA AND PANAMA

la Inflorescence always simple and usually axillary (a racemose unbranched axis with

pedunculate triads); flowers bisexual and less than 4 mm long; wine red to dark red

or maroon; 0-1,200 m elevation and widespread P. pyrifolia

Ib Inflorescence often terminal and panicle-like, simple racemose inflorescences may also

be present; plants and flowers said to be unisexual, flowers 4-5 mm long and yellow-

ish; 0-1,800 m in Panama P. retroflexa

Phthirusa pyrifolia (H.B.K.) Eichler, Fl. Brasil. 5(2):36. 1868. Loranthus pyrifolius

H.B.K., Nov. Gen. & Sp. 3:441. 1820. Figure 6.

Shrubs or vinelike, to 1 m long, often pendant, hemiparasitic epiphytes, bisexual, leafy

internodes 1-7 cm long, 1.6-5 mm thick, thicker below the nodes, glabrous but often

forming brownish flakes on drying, usually with 2 longitudinal ridges continuous with

the petioles of the next distal node; epicortical roots arising only from the base of the plant.

Leaves opposite, subopposite, or occasionally alternate, dark green and lustrous, petioles

4-20 mm long, 0.8-3 mm thick, distinct, sulcate above with adaxial ridges continuous with

the lamina margins; laminae 4-14 cm long, 2-8 cm broad, elliptic to oblong, ovate or

lanceolate, rounded to obtuse or less often acute at the apex, often mucronate with a small

(1-2 mm) narrow tip but this readily breaking off when dry, subtruncate to obtuse at the

base, margin entire and the edge drying roughened brownish, differentiated from the

BURGER: FLORA COSTARICENSIS 45

epidermis, the laminae drying subcoriaceous, glabrous and smooth, venation pinnate

with the 4-6 pairs of major secondary veins arising throughout the length of the midvein,

the secondaries often abruptly arcuate ascending about halfway between the midvein and

margin in larger leaves. Inflorescences solitary or paired in the leaf axils or from leafless

terminal shoots and paniculate, usually spicate and 4-12 cm long, the triads of flowers

subsessile or on very short (1-2 mm) peduncles, bracteate apically beneath the flowers,

triads usually in opposite pairs 3-12 mm distant on the rachis, rachis ca. 0.8 mm thick and

with minute scaly surface when dry; flowers sessile, bisexual, perianth parts ca. 1.5 mm

long, wine red or maroon, ovary 1-1.5 mm long and ca. 1 mm thick, without a locule,

calyculus small (0.3 mm) and slightly flaring, style simple. Fruit a blunt ellipsoidal berry,

bright orange-red with yellowish apex and dark purple base, ca. 8 mm long and 5 mm thick

but 3-5 mm long and 1.5-3 mm thick when dry, smooth and with a minutely (0.05 mm)

reticulate surface, the triads often projecting obliquely downward (toward the base of the

spike).

A common and widespread hemiparasite on tree branches in evergreen and

partly deciduous forest formations between sea level and about 1,200 m elevation;

apparently absent from the deciduous (tropical dry) forests of lowland Gua-

nacaste. Flowering throughout the year in Costa Rica, but collected most often

between January and August. The species ranges from southern Mexico to the

West Indies and northern South America.

Phthirusa pyrifolia is recognized by its ridged stems, opposite and subopposite

(alternate) leaves often on the same stem, pinnate venation, subsessile triads of

very small flowers well spaced along the slender spikelike inflorescence, and fruit

with unusual minutely reticulate surface. These plants resemble species of Stru-

thanthus, but differ in having very small apparently bisexual dark red flowers and

in the fact that epicortical roots are not produced from the stem, but only from the

cotyledonary zone.

Phthirusa retroflexa (R. & P.) Kuijt, Brittonia 32:521. 1980. Loranthus retroflexus

Ruiz & Pavon, Fl. Peruv. 3:49-50, t. 279b. 1802. L. adu ncus Meyer, Prim. Fl. Esseq.

149. 1818. L. paniculatusH.B.K., Nov. Gen. & Sp. 3:422. 1820. L. theobwmaeWilld.

ex R. & S., Syst. Veg. 7:132. 1829. Struthanthus retroflexus (R. & P.) Blume, in

Schult, Syst. Veg. 7:1731. 1830. Phthirusa paniculate (H.B.K.) Macbr., Field Mus.

Nat. Hist., Bot. Ser. 11:17. 1931. P. adunca (Meyer) Maguire, Bull. Torrey Bot. Club

75:301. 1948. Figure 6.

Shrubs, the stems often scandent, unisexual, leafy internodes 0.5-4.5 (8) cm long, 1.1-4

(5.5) mm thick, terete and smooth, drying pale brown or grayish; roots borne on the stems.

Leaves opposite, subopposite, or alternate, clearly articulate at the base, petioles well

defined, 8-23 mm long, 1-2 mm broad, adaxial groove formed by ridges continuous with

the lamina margins; laminae 3.8-8 cm long, 2.5-5 cm broad, ovate to broadly elliptic or

broadly oblong, acute to short-acuminate at the apex (emarginate only where damaged

during development?), obtuse to rounded at the base, margin entire, stiffly chartaceous

and drying dark and brittle, glabrous, venation pinnate with 4-7 pairs of major secondary

veins arising throughout the length of the midvein, often irregular in arrangement and

sometimes obscure on both surfaces. Inflorescences an unbranched solitary raceme of

triads in the axils of leaves or more often a compound panicle at the ends of stems (formed

by the failure of distal leaves to develop), triads (occasionally diads and monads) borne on

peduncles 1-3 mm long, small triangular bracts ca. 1-2 mm long borne at the apex of the

peduncle beneath each flower and often deciduous; flowers unisexual(?), rather similar in

size and form, perianth ca. 4-5 mm long, pale yellow, anthers ca. 1.5 mm long; ovary ca.

1.5mm long and 1.5 mm broad at the apex, narrowed at the base. Fruit tol cm long, oblong

and pedicellate, orange.

Hemiparasites of both evergreen and deciduous forest formations from sea level

to 1,800 m elevation. Probably flowering throughout the year, but collected most

46 FIELDIANA: BOTANY

often between April and July. The species ranges from westernmost Panama to

Peru and Paraguay (fide Rizzini).

Phthirusa retroflexa is recognized by its epiphyte-like parasitic habit, small leaves

narrowed at the apex (and quite variable as regards arrangement on the stem),

triads of yellow flowers, and the very unusual terminal panicle-like inflorescences

in addition to simpler axillary inflorescences. Although not yet recorded from

Costa Rica, this species is likely to occur near the border with Panama.

PSITTACANTHUS Martius

Hemiparasitic shrubs, parasitizing the aerial parts of trees and shrubs, often large,

bisexual, leafy stems strongly ridged to terete; epicortical roots absent, the haustorium

often becoming large. Leaves opposite, whorled, or very variable in some species and

ranging from alternate to whorled (often on the same stem), symmetrical to asymmetric,

articulate at the base, simple and entire, glabrous. Inflorescences axillary or terminal, 1 to

several per leaf axil, fasciculate at older leafless nodes in some species, primary peduncles

bearing 1-3 secondary peduncles subtending the triads or diads of flowers, floral pedicels

expanded at the apex to form a deep or shallow cup, the cup usually becoming asym-

metrically expanded on 1 side, entire; flowers 2-10 cm long, bisexual, brightly colored and

usually reddish or orange near the base grading into a yellowish apex, the calyculus little

differentiated from the ovary, thin, entire or minutely denticulate, perianth forming a

slender tube before anthesis, thickened distally around the anthers and in some species

expanded just above the base, the 6 perianth parts separating at anthesis, filaments mostly

free, of 2 lengths with the anthers superposed at 2 levels of 3 just beneath the apex of the

perianth, anthers dorsifixed and versatile, slender, style long and slender with a simple

slightly thickened stigma. Fruit a smooth bluish black berry.

An American genus of uncertain size that has not been revised since Eichler 's

work in the Flora Brasiliensis more than 100 years ago. The species are largely

continental and range from Southern Mexico to Argentina. The very colorful

perianth and the numerous inflorescences of some species often make the plants

very conspicuous. The pedicels are expanded to form a dishlike base beneath the

flowers, probably representing a fusion of bracts and bracteoles. The plants may

be confused with our species of Gaiadendron, but other members of the Lo-

ranthaceae in a wide sense are quite different in having much smaller flowers.

la Larger leaves asymmetric and with the midvein curved to 1 side; leafy stems strongly

4-angled with 4 longitudinal ribs or wings; inflorescences terminal, flowers usually in

triads 2a

Ib Leaves symmetrical with the midvein usually straight; stems rarely 3- or 4-angled,

soon becoming terete; inflorescences axillary or borne on older wood, flowers usually

in diads 3a

2a Larger laminae asymmetrically ovate; flowers to 9 cm long; evergreen montane

forests, 1,700-3,300 m elevation P. schiedeanus

2b Larger laminae asymmetrically lanceolate or falcate; flowers to 5 cm long; decid-

uous lowlands P. calyculatus

3a Leaves usually in whorls of 4 and separated by long (3-12 cm) internodes; flowers 4-6

cm long on small axillary inflorescences, anthers subsessile with slender hairs from the

base; evergreen forests, 500-1,000 m elevation P. nodosus

3b Leaves alternate, opposite or in whorls of 3 (very rarely 4), leafy internodes 1-8 cm

long; flowers 3-4 cm long; anthers on long filaments, without hairs 4a

4a Laminae ovate to suborbicular or elliptic, to 14 cm long and 10 cm broad; inflorescences

often on older stems more than 1 cm thick, primary peduncles less than 5 mm long;

(1,000) 1,500-2,500 m elevation P. scheryi

4b Laminae usually elliptic-oblong, to 8 cm long and 4 cm broad; inflorescences from leaf

axils or on older stems not usually more than 1 cm thick, primary peduncles 4-14 mm

long; 500-1,800 (2,200) m elevation P. ramiflorus

BURGER: FLORA COSTARICENSIS 47

Psittacanthus calyculatus(DC) G. Don, Gen. Syst. 3:415. 1834. Loranthus calyc-

ulatusDC., Coll. Mem. pi. 10. 1830. P. chrismarii ; Urban, Bot. Jahrb. Syst. 24:331.

1897. Figure 5.

Shrubs, branch parasites with the branches often long and trailing, leafy internodes 1-6

cm long, 1.7-7 mm thick, 4-angled in cross section with 4 longitudinal ribs, green but

becoming gray. Leaves opposite or subopposite (occasionally alternate), decussate, dark

green, asymmetric and usually somewhat curved, petioles 0-8 mm long, gradually merg-

ing with the lamina, 1-2.5 mm broad with wings continuous with lamina margins; lami-

nae 4-15 (20) cm long, (1) 1.4-3.5 (5) cm broad, lanceolate to falcate or narrowly ovate,

usually with the midvein curved to 1 side, tapering gradually to the acute or blunt apex,

acute to cuneate or attenuate at the base, margin entire or undulate and decurrent on the

petiole, the edge slightly differentiated, the laminae drying coriaceous, smooth and gla-

brous, venation subpalmate or pinnate with 2-4 pairs of major secondary veins arising

from near the base of the lamina, strongly ascending, usually obscure. Inflorescences

terminal with the lowest units axillary to a pair of foliage leaves, or less often axillary,

paniculate, 5-12 cm long, usually with a basal peduncle 1-2 cm long and 1 or 2 secondary

peduncles, peduncles of the triads (diads) 4-16 mm long, pedicels of the flowers 6-20 mm

long, 0.6-1.2 mm thick, flared near the apex to form a somewhat asymmetric cup 3 mm

broad enclosing the base of the ovary; flowers ca. 4 cm long before anthesis, slender but

dilated at base and apex, incurved, ovary ca. 3-4 mm long including calyculus and 2.5-3

mm thick, the calyculus very slightly flared, 3-4.5 mm broad, subentire with several small

notches and differentiated margin, perianth parts 3-4 cm long, ca. 1-2 mm broad, orange

to red, filaments subequal, differing by ca. 4 mm in length, anthers 3 mm long, dorsifixed

and versatile, stigma ca. 3.5 cm long, very slender, stigma slightly thickened. Fruit black

and ellipsoid, becoming 9 mm long and 5 mm in diameter.

Colorfully flowered hemiparasites of the seasonally very dry deciduous forest

formations of Guanacaste and the Nicoya peninsula; probably flowering through-

out the year, but only collected from September to March in Costa Rica. The

species ranges from southern Mexico to northern South America.

Psittacanthus calyculatus is recognized by the medium-sized bright orange-red

flowers, curved narrow leaves that are opposite to almost alternate on the strongly

angled gray stems, and the seasonally very dry lowland habitat. Psittacanthus

mayanus Standl. & Steyerm. of northern Central America with smaller sym-

metrical leaves may prove to be no more than an unusual form of this species.

Psittacanthus nodosus(Desr.) G. Don, Gen. Syst. 3:417. 1834. Loranthus nodosus

Desr., in Lamarck, Encycl. 3:601. 1809. Aethanthus nodosus (Desr.) Engler, in En-

gler & Prantl, Nat. Pflanzenfam., Nachtr. 1:136. 1897. Figure 5.

Shrubs to ca. 1 m tall, leafy internodes (1) 3-12 cm long, 2.3-7 mm thick, terete, pale

grayish with narrow longitudinal lenticels, the nodes conspicuously thickened. Leaves

almost always in whorls of 4 and producing a striking verticellate appearance, sym-

metrical, articulate at the base, petioles 0-10 mm long, the adaxial ridges continuous with

the lamina margins, ca. 2 mm broad; laminae 5-10 (12) cm long, 2-4.5 (7) cm broad, elliptic

to obovate, ovate, or ovate-elliptic, bluntly acute to rounded at the apex, gradually taper-

ing to the acute, obtuse, or cuneate base, margin entire and becoming recurved, decurrent

on the petiole, the laminae drying stiffly coriaceous, glabrous and smooth but with a rough

appearance, venation pinnate with 2 or 3 pairs of major secondary veins arising from the

lower half of the midvein, secondary veins strongly ascending and usually obscure be-

neath. Inflorescences axillary, 1-6 (rarely more) per node, ca. 6-10 cm long including the

flowers, peduncles and pedicels together usually less than 2 cm long, primary peduncle

2-5 mm long, 1 mm thick (dry), peduncles of the diads about the same length, pedicels

ca. 5 mm long and flared at the apex to form a shallow asymmetric cup 3 mm broad; flowers

4-6 cm long, slender and straight before anthesis, ovary and calyculus ca. 4 mm long and

2 mm thick, perianth ca. 4.5 cm long, the tube 2-4 mm thick before anthesis, yellow

48 FIELDIANA: BOTANY

apically and brilliant red toward the base, filaments apparently equal, anthers ca. 3 mm

long, stigma slightly thickened. Fruit to ca. 12 mm long and 8 mm in diameter, green

becoming black (dry), epidermal cells apparent (0.1 mm) on the lustrous surface.

Unusual branch parasites of premontane wet forest formations between about

500 and 1,000 m elevation; probably flowering throughout the year. The species

ranges from northern Costa Rica to Ecuador.

Psittacanthus nodosusis recognized by its large colorful flowers, round stems with

distant nodes of four whorled leaves, and relatively small inflorescences. Material

of this species has been mixed with Struthanthus costaricensis under the numbers

11458 and 11611 (Tonduz and Herb. Inst. physico-geogr. nat. costar.). Thus, we

believe, the locality of Santa Maria at 1,450 m elevation may be in error and have

listed the altitudinal range as lower. This species is known only from the following

collections: Lent 2563 Rio Naranjo west of Volcan Tenorio, Poveda 965 Pavones de

Turrialba, Standley & Valeria 46816 near Pejivalle, Cartago, Tonduz 11458 near Tuis,

Cartago.

Psittacanthus ramiflorus (DC.) G. Don, Gen. Syst. 3:415. 1834. Loranthus rami-

florus DC., Prodr. 4:308. 1830. P. allenii Woodson & Schery, Ann. Missouri Bot.

Card. 27:309. 1940. P. lateriflorus Woodson & Schery, loc. cit. Figure 5.

Shrubs, branch parasites, leafy internodes 0.1-3 (5) cm long, 2-6 mm thick, terete or

angled and with longitudinal ridges in early stages but becoming terete in age, smooth and

gray green but becoming rough and dark gray in age. Leaves opposite, subopposite, in

whorls of 3, or alternate (often on the same stem), symmetrical, articulate at the base,

petioles 2-12 mm long, 1-2 mm broad, the lateral wings continuous with the lamina

margins; laminae 3-8 cm long, 1.3-4 cm broad, elliptic-oblong to narrowly elliptic, ovate-

elliptic, ovate, elliptic, or obovate-elliptic, bluntly obtuse to rounded at the apex, tapering

gradually or abruptly at the acute to obtuse or cuneate base, margin entire and decurrent

on the petiole, becoming revolute, the laminae drying subcoriaceous, smooth and gla-

brous, venation pinnate, the 3 or 4 pairs of major secondary veins arising from throughout

the length of the midvein. Inflorescences in leaf axils or at leafless nodes on stems usually

less than 1 cm thick, 5-8 cm long, primary peduncle 4-14 mm long, peduncles of the diads

3-6 mm long, pedicels ca. 5 mm long, 0.5-0.8 mm thick and expanded at the apex to form

a flattened shallow cup 2-3 mm broad; flowers 3-4 cm long, ovary ca. 3 mm long and 2

mm in diameter, calyculus thin and entire (subentire), perianth parts ca. 3-3.5 cm long at

anthesis, forming a slender tube before anthesis, thickened at the apex and at the base,

small projections ("ligule") at the apex of the basally expanded portion of the perianth

parts on the inner surface, filaments of 2 lengths, anthers 2-3 mm long, stigma simple and

very slightly thickened. Fruit not seen.

Epiphyte-like hemiparasites of trees and shrubs in premontane and lower mon-

tane wet forest formations between 600 and 1,800 (2,200) m elevation on the Pacific

watershed; collected with flowers from April to December. The species, in our

area, ranges from the Sierra de Tilaran in Costa Rica to Central Panama.

Psittacanthus ramiflorus is recognized by the usually small symmetrical leaves

alternate to whorled on rounded stems and the 3-4 cm long flowers on short

inflorescences that occur in leaf axils or on slender leafless stems. This species is

very closely related to P. scheryi (q.v.), and that species may be no more than a

large-leaved form of P. ramiflorus. Psittacanthus dilatatus A. C. Smith of Colombia

appears to be closely related to P. ramiflorus.

Psittacanthus scheryi Woodson, Ann. Missouri Bot. Card. 28:426. 1941. Fig-

ure 5.

Shrubs 1-2.5 m tall, leafy internodes 1-8 cm long, 3-11 mm thick, terete and relatively

BURGER: FLORA COSTARICENSIS 49

smooth, becoming rough and longitudinally striate with narrow lenticels in age. Leaves

variable in position, alternate to subopposite, opposite, or in whorls of 3 (4), symmetrical,

articulate at the base, petioles 4-14 mm long, 1.3-4 mm broad with ridges continuous with

the lamina margins; laminae 4.5-15 (18) cm long, (2) 3-10 (12) cm broad, ovate to elliptic-

ovate or occasionally narrowly elliptic or suborbicular, obtuse and blunt or rounded at the

apex, abruptly or gradually narrowed at the base, margin entire, often becoming revolute

and undulate, the laminae drying very stiffly coriaceous and often pale in color, smooth

and glabrous, venation pinnate with the 3-6 pairs of major secondary veins arising from

throughout the length of the midvein, usually obscure beneath. Inflorescences fasciculate

in the axils of leaves or fallen leaves, often on older stems over 1 cm thick, to 5 cm long

including flowers, primary peduncles usually less than 5 mm long, peduncles of the diads

(very rarely triads) 3-6 mm long, pedicels 3-7 mm long, ca. 1 mm thick (dry), flared at the

apex and forming an asymmetric cup 3-4 mm wide at its broadest; flowers ca. 3-4 cm long,

ovary ca. 3 mm long and 2 mm in diameter, flared slightly at the short (0.7 mm) thin entire

calyculus, perianth parts ca. 3 cm long, slender and thickened near the apex, abruptly

expanded for ca. 3 mm above the base, a slight projection ("ligule") at the apex of the

basally expanded portion of the perianth parts on the inner surface, orange or reddish near

the base and yellow-orange apically, filaments of 2 lengths and differing by ca. 4 mm,

anthers 2-3 mm long, stigma simple, very slightly thickened. Fruit ca. 12 mm long and 8

mm in diameter, ellipsoid, black and borne on red pedicels, calyculus very short on the

truncate apex.

Epiphyte-like hemiparasites of wet evergreen montane (premontane wet to

montane rain) forest formations between (1,000) 1,500 and 2,500 m elevation in

Costa Rica; flowering throughout the year. The species, as presently understood,

ranges from the Cordillera Central de Costa Rica to western Panama.

Psittacanthus scheryiis recognized by its fasciculate inflorescences often on older

stems, slender medium-sized flowers apically yellowish, and often large thick

leaves that vary from alternate to opposite or whorls of three. This species is

related to P. dilatatus of Colombia with narrower obovate leaves, and it may be no

more than a large-leaved form of P. ramiflorus in a wider sense. The specimens that

have been placed here are being kept separate from P. ramiflorus because their

overall appearance is quite different, and the distinctions given in the key seem

to be working well with newly collected specimens.

Psittacanthus schiedeanus (Schlecht. & Cham.) Blume, in Schult., Syst. Veg.

7:1730. 1830. Loranthus schiedeanus Schlecht. & Cham., Linnaea 5:172. 1830. Fig-

ure 5.

Shrubs, to 3 m long, often high up in tall trees and conspicuous because of the bright

red-orange inflorescences, leafy internodes 0.5-10 cm long, 2-9 mm thick, glabrous,

4-angIed with 4 prominent longitudinal wings, nodes somewhat flattened; haustorial

connections becoming 35 cm in diameter. Leaves opposite or subopposite, usually bluish

green, petioles 3-20 mm long, distinct, 1-2.5 mm thick, inconspicuously winged; laminae

4-26 cm long, 2-10 cm broad, narrowly ovate (rarely lanceolate) to broadly and inequi-

laterally ovate, the larger laminae often very asymmetric in the lower half with the midvein

curved, gradually narrowed to an acute or acuminate apex, narrowed to abruptly rounded

at the often unequal base, margin entire and decurrent on the petiole, edge differentiated

from the epidermal tissue, the laminae drying subcoriaceous to coriaceous, smooth and

glabrous, venation pinnate with the 3-5 pairs of major secondary veins usually obscure.

Inflorescence axillary or terminal at the ends of twigs (and resulting in forked growth),

leafless, 10-20 cm long including the flowers, of a condensed axis 1-3 cm long bearing

8-14 triads in opposite or subopposite pairs, paniculate in general form, peduncles of the

triads 12-35 mm long, bracteate at the apex by adnation, bracts broad based and ca. 2 mm

long, pedicels of the flowers 10-20 (30) mm long, flaring into a cupulate apex enclosing the

flower base, peduncles, pedicels, and exposed areas of buds becoming bright red or

orange; flowers 8-9 cm long before anthesis, ovary ca. 4 mm long and 5 mm thick, crowned

by a short (3 mm) tubular calyculus continuous with the ovary, subentire and minutely

50 FIELDIANA: BOTANY

3-toothed, perianth becoming 9 cm long and only 2-4 mm broad, linear, slightly thickened

below the apex and with an acuminate tip, brilliant orange, becoming recurved, decid-

uous, filaments 4-6 cm long, dimorphic and differing by ca. 5 mm in length, anthers

dorsifixed and versatile, ca. 6 mm long, style ca. 7.5 cm long, stigma slightly thickened.

Fruit a blackish berry 1.5-2 cm long, ca. 1 cm thick, ellipsoid, on pedicels to 3 cm long;

embryo with 6-12 awl-shaped cotyledons triangular in cross section.

Large epiphyte-like parasites, conspicuous when in flower in evergreen mon-

tane forests between about 1,700 and 3,300 m elevation; flowering from December

to August. This species is especially common on the central Volcanos; it ranges

from central Mexico to western Panama.

Psittacanthus schiedeanus is recognized by its large and colorful slender flowers,

the ridged stems, asymmetric larger leaves, and montane habitat. The number of

cotyledons is unusual. The cotyledons persist for some time, becoming raised and

separated from each other by the rapid expansion of the haustorial body. It is

probably this species that is responsible for the majority of "rosas de palo" or "flares

de palo" which are sold as curiosities in Mexico and Guatemala. These represent

placenta-like host malformations from which the parasite has fallen away and

form an intricate, fluted pattern of a radial arrangement (Kuijt, 1964).

STRUTHANTHUS Martius

Nomen Conservandum Propositum

Shrubs, epiphyte-like hemiparasites of trees, often unisexual, glabrous, stems erect,

pendent, clambering, or climbing by means of twisted (grasping) petioles, twining termi-

nal shoots, or attaching roots; epicortical roots from base and stems. Leaves opposite or

subopposite (often on the same stem), articulate at the base (the leaves of new young

shoots often very different from the leaves of mature flowering stems), lamina margins

usually decurrent on the petiole and usually with differentiated tissue on the edge,

venation pinnate. Inflorescences 1 to several (9) in the axils of leaves, ours with an un-

branched axis of sessile or pedunculate triads (monads in S. panamensis) to produce spicate

or racemose panicles, peduncle of the triads associated with bracts that are usually cadu-

cous, central flower of the triad usually sessile, the laterals often becoming short pedi-

cellate, floral bracts very small and usually caducous, the flowers generally small (less than

10 mm) with denticulate or entire calyculus and free petals; male flowers with the distal

part of the bud expanded to include the anthers, petals valvate in bud, usually 6, becoming

reflexed at anthesis, stamens epipetalous and slightly dimorphic, the anthers borne at 2

superposed levels on adjacent petals, filaments usually very short, anthers dehiscing

longitudinally, pollen said to be of 2 types (functional and rugose, sterile and smooth),

nonfunctional style slender, stigma absent; female flowers often smaller than the male,

buds narrowed toward the apex, the petals not usually becoming reflexed, epipetalous

staminodes usually small and slender, style simple and equaling the petals, with longi-

tudinal ridges, stigma often capitate. Fruit a berry, the calyculus not developing, rounded

at the apex, seed with white endosperm.

Struthanthus is a neotropical genus of probably more than 60 species. The genus

has never been monographed nor has its relationship with closely similar genera

been properly defined. The recent segregation of Cladocolea (Kuijt, 1975) has re-

solved some issues, but the problem of distinguishing Phthirusa remains. The

species of Struthanthus also present problems; e.g., they are often not easily iden-

tifiable. This is partly due to similarity of flowers and unisexuality of the plants,

but the variation of floral and vegetative characters within each species adds to the

problem. An extreme example is seen in young plants of S. leptostachyus, the

leaves of which may be up to 11 cm long and 0.5 cm wide. These long, linear,

fleshy, pendent leaves are gradually replaced by the broad thin organs of the

mature plant (Kuijt, 1964).

BURGER: FLORA COSTA RICENSIS 51

KEY TO THE SPECIES OF Struthanthus

la Inflorescences at first enclosed by 4 ranks of decussate imbricate bud scales, these

broadly triangular and scarious, mostly deciduous; flowers bisexual or functionally

unisexual and similar in form in the 2 sexes 2a

Ib Inflorescences not at first enclosed in an imbricate series of decussate bud scales;

flowers in triads or monads with abaxial bracts that may persist or be caducous, but

not thin and scarious; flowers unisexual with the male flower buds distinctly broader

near the apex and enclosing the functional anthers, the female flower buds with

a narrow tubular perianth and with the nonfunctional anthers small or undevel-

oped 3a

2a Flowers in monads, subtended by 3 persisting and conspicuous bracts on a short

thick peduncle; leaves to ca. 10 cm long, often short-acuminate; rare plants

known only from ca. 1,800 m elevation in Chiriqui province, Panama

5. panamensis

2b Flowers usually in triads (groups of 3), sessile on the axis of the inflorescence,

the subtending bracts quickly deciduous; leaves to 20 cm long (in ours); ever-

green and partly deciduous vegetation, 0-2,000 m 5. leptostachi/ns

3a Inflorescence very short, rarely more than 2 cm long, rarely with more than 3 pairs

of triads; laminae usually less than 8 cm long and often rounded or blunt at the apex;

stems terete and often becoming reddish lenticellate, 0-2,100 m S. oerstedii

3b Inflorescences usually more than 2 cm long, rarely with less than 4 pairs of

triads 4a

4a Inflorescences with conspicuous (3 mm) persisting bracts beneath the triads and

obscuring the base of the ovaries; apex of the ovary (edge of the calyculus) 2-2.7 mm

broad, anthers 1.2-1.6 mm long on short filaments; fruit becoming 10 mm long and

6 mm thick; laminae rounded or emarginate apically and usually less than 7 cm long;

1,200-2,000 m in western Panama S. rotundatus

4b Inflorescences with the bracts at the apex of the peduncle less than 3 mm long

or deciduous before the flowers open; flattened apex of the ovary less than 2 mm

broad 5a

5a Leaves often grasping stems with twisting petioles, laminae usually less than 8 cm

long, usually rounded apically; male perianth 4.5-6 mm long, anthers 0.8-1.3 mm

long and usually subsessile; fruit becoming 10 mm long, cylindrical and bluish on

lenticellate peduncles; 0-1,200 m and widespread S. orbicularis

5b Leaves lacking twisting petioles, but the leaves sometimes curved and the stem

sometimes twining; fruit usually less than 7 mm long and the fruiting peduncle

without lenticels 6a

6a Some stems twining and with the leaves early deciduous, laminae attenuate at the

base and often obovate, rounded to short-acuminate at the apex, the secondary veins

prominent on dried laminae; flowers sessile or subsessile, male perianth 4.5-5.5 mm

long, anthers 1-1.2 mm long; female perianth 2.5-3.5 mm long; fruit becoming

pedunculate, berries orange-red and becoming constricted around the middle, the

central berry (of a triad) often projecting obliquely downward/backward (toward the

inflorescence base); (100) 500-1,800 m S. costaricensis

6b Tips of some stems not quickly losing their leaves and twining; fruit not constricted

centrally or projecting backward; leaves often relatively thin and drying stiffly char-

taceous, often with acuminate apices 7a

7a Laminae narrowly ovate-elliptic to lanceolate, long-acuminate at the apex, secondary

veins strongly ascending and usually drying dark and prominent; male perianth ca.

4.5 mm long with anthers 1.2 mm long; female perianth 3-4 mm long; ovary ca. 1

mm thick; 0-500 m elevation S. woodsonii

To Only the smaller leaves lanceolate, the venation often obscure; plants of higher

altitudes, 1,000-3,000 m 8a

8a Stems usually terete, laminae acute to short-acuminate; male flowers 6-8 mm long,

anthers 1-1.6 mm long; female flowers 4-6 mm long; fruit to 7 mm long; areas around

Volcan Irazu and Volcan Turrialba S. marginatus

8b Stems usually 4-angled, laminae often long-acuminate; male flowers 4-5 mm long,

anthers ca. 0.8 mm long; female flowers 3-4.5 mm long; fruit to 5 mm long (dry);

Central Volcanic Highlands to the Cordillera de Talamanca S. quercicola

52 FIELDIANA: BOTANY

Struthanthus costaricensis Standley, Publ. Field Columbian Mus., Bot. Ser.

4:202, 1929. Figure 7.

Shrubs, branch parasites often forming pendent masses, tips of stems often twining

about supporting branches, leafy internodes 2-12 cm long, 1.2-5 (8) mm thick, terete or

obscurely angled (flattened) at first, becoming terete with small lenticels and longi-

tudinally striate, gray; epicortical roots often produced on stems contacting other objects.

Leaves opposite, subopposite, or alternate, recurved when young but not prehensile, often

early deciduous when near the apex of a twining stem, articulate at the base, petioles

10-20 mm long, somewhat vaginate near the base with winglike margins continuous with

the laminae margins; laminae 3-9 (14) cm long, 2-5 (9) cm broad, broadly elliptic to

obovate or spatulate (rarely suborbicular), obtuse to short-acuminate (rarely rounded) at

the apex, gradually tapering to the attenuate base or occasionally obtuse to truncate,

margin entire and slightly revolute (dry), decurrent on the petiole, the laminae drying

subcoriaceous to coriaceous, venation pinnate with 4-7 pairs of major secondary veins

arising mostly from the lower half of the midvein and ascending at angles of ca. 45°.

Inflorescences 1-3 (occasionally to 9 in the males) in the axils of leaves, the spicate panicles

2-6 cm long, triads sessile or subsessile, alternate to opposite and 3-15 mm distant along

the slender (0.5-1.7 mm) ridged rachis, ca. 8 triads on female inflorescences, more in the

male, flowers sessile or the lateral females with very short pedicels, floral bracts minute

and caducous, male flowers 5-7 mm long, ovary and calyculus 1-1.5 mm long, narrowed

at the base and flared at the calyculus to 2 mm broad, perianth ca. 4 mm long before

anthesis, becoming reflexed, stamens with short (0.5 mm) filaments, anthers 0.8-1.2 mm

long; female flowers ca. 4-5 mm long, petals ca. 3 mm long with minute (0.3 mm) slender

epipetalous staminodes, style ca. 3 mm long with capitate stigma. Fruit ellipsoid to cylin-

drical, becoming ca. 5 mm long and 3 mm thick, slightly constricted in the middle, red to

orange, the lateral fruit of a triad on pedicels becoming up to 3 mm long.

Epiphyte-like hemiparasites of moist evergreen or partly deciduous forest for-

mations between 500 and 1,800 m elevation on both the Caribbean and Pacific

slopes of Costa Rica; flowering collections have been made between November

and July. The species is found only in Costa Rica and western Panama.

Struthanthus costaricensis is recognized by the presence of stems with small-

leaved twining tips, subopposite leaves on terete stems, stiff laminae with atten-

uate base (decurrent on the petioles), and relatively small red-orange fruit slightly

constricted in the middle. This species is closely related to S. orbicularis, but that

species lacks the twining stems and has prehensile (grasping) petioles, angled

stems, and larger bluish fruit on lenticellate pedicels.

Struthanthus leptostachyus (H.B.K.) G. Don, Gen. Hist. Dichlam. PI. 3:411.

1834. Loranthus leptostachyusH.B.K., Nov. Gen. Sp. 3:440. 1818. Peristethium lepto-

stachyum (H.B.K.) Van Tieghem, Bull. Soc. Bot. France 42:175. 1895. Phthirusa

pittieri Krause, Feddes Repert. 15:441-442. 1919. Struthanthus polystachyus sensu

auctores. Figure 6.

Large shrubs, with horizontal or scandent branches over 2 m long, leafy internodes 2-9

cm long, 1.5-8 mm thick, raised brownish lenticels often present on older and thicker

stems; epicortical roots produced from the base and from the stems, those from the base

occasionally becoming over 30 cm long. Leaves opposite or subopposite (occasionally a few

alternate leaves present), articulate at the base, petioles 10-25 mm long, 1.4-4.5 mm thick,

slightly sulcate above with adaxial ridges continuous with the lamina margins; laminae

5-16 (20) cm long, 3-8 (12) cm broad, broadly ovate-lanceolate to ovate, elliptic-ovate, or

elliptic-oblong, usually gradually narrowed to the acuminate apex, rounded and sub-

truncate to obtuse at the base, margins entire and becoming revolute, tissue of the edge

slightly differentiated and smooth, the laminae drying stiffly chartaceous, glabrous and

smooth, venation pinnate with 5-10 pairs of major secondary veins arising throughout the

length of the midvein, secondary veins slightly raised above and often obscure beneath.

Inflorescences solitary or 2-4 in the axils of leaves or fallen leaves, unisexual, at first

BURGER: FLORA COSTARICENSIS 53

enclosed in pale scarious keeled bracts (scales) ca. 3 mm long, these caducous with

expansion of the spike, the spikes becoming 2-8 cm long, with ca. 14-18 opposite triads

of flowers or with solitary flowers near the apex of the spike, triads subtended by the bract-

(scale-) scar, rachis ca. 1 mm thick and longitudinally striate when dry, glabrous but

developing lenticels in fruit; flowers 4-6 mm long, unisexual, the female somewhat more

slender than the male, ovary 1-1.5 mm long with thin subentire calyculus, perianth parts

3-4 mm long, male flowers with anthers sessile on the perianth, subequally attached, 1

mm long. Fruit becoming 6 mm long and 4 mm thick, truncated at base and apex or

somewhat ellipsoid.

Epiphyte-like hemiparasites of wet evergreen or partly deciduous forest for-

mations from sea level to 2,000 m elevation on both the Caribbean and Pacific

slopes; probably flowering throughout the year, but not collected in September,

October, and November in our area. The northernmost collection of the species

comes from near Tilaran, Guanacaste, while the southernmost are reported to be

from Peru.

Struthanthus leptostachyus is recognized by its large size (for a parasite), epi-

cortical roots arising from various areas on the stems, leaves with acuminate

apices, small unisexual yellow flowers in sessile triads, and the spikes at first

enclosed in imbricate bracts resembling bud scales. These scales and the rather

large leaves distinguish this species from most of our other species of Stru-

thanthus. This species resembles Phthirusa pyrifolia in a superficial way.

Struthanthus marginatus (Desr.) Blume, in Schult., Syst. Veg. 7, pt. 2:1731.

1830. Loranthus marginatus Desr., in Lam., Encycl. 3:596. 1789. Figure 7.

Shrubs or climbers, branch parasites, often covering the crowns of small trees, leafy

internodes 2-5 cm long, 1.3-3.5 (5) mm thick, pale brown and smooth, becoming grayish

with small round lenticels, longitudinally striate (dry) but terete; clasping roots produced

in groups or series on stems and often ramifying. Leaves opposite, subopposite, or occa-

sionally alternate, petioles 3-5 mm long, ca. 1.3 mm thick, somewhat sulcate above or

slightly winged; laminae 4-8 (12) cm long, 1-4 (7) cm broad, narrowly elliptic-oblong to

broadly elliptic, ovate, or lanceolate, tapering gradually to an acuminate to acute apex

(rarely rounded), obtuse to acute at the base with the margins decurrent on the petiole,

margin thin and becoming slightly revolute, laminae drying stiffly chartaceous and often

brittle, smooth and glabrous, the 3-7 pairs of major secondary veins often difficult to

distinguish from the tertiaries, the teritary veins obscure or slightly raised on the dried

surfaces. Inflorescences solitary or 2-3 in the axils of leaves, racemose panicles 2-7 cm

long, the 8-14 triads evenly spaced in usually opposite or subopposite pairs along the

slender (0.7-1.4 mm) rachis, peduncles of the triads 1-3 (5) mm long with a small (3 mm)

caducous bract at the apex, triads with the central flower sessile and the lateral flowers

sessile or very short- (0.3-1 mm) pedicellate; male flowers 6-8 mm long, ovary and

calyculus 1.5-2 mm long, perianth cream color, 5-6 mm long, becoming reflexed, stamens

with very short filaments, anthers 1-1.6 mm long and almost versatile; female flowers 4-6

mm long, the perianth ca. 3.5-4 mm long, narrow, yellow-green, staminodes with re-

duced anthers ca. 0.6 mm long, style 3.3-4 mm long with a capitate stigma. Fruit ovoid,

becoming 7 mm long and 5 mm in diameter, the central fruit of a triad sessile, the laterals

usually pedicellate.

Epiphyte-like hemiparasites of moist evergreen forest formations between about

1,000 and 3,000 m elevation; flowering collections have been made between Feb-

ruary and July. The species has been collected only from around the eastern edge

of the Meseta Central and the adjacent volcanoes (Irazu and Turrialba) and in the

Chiriqui Highlands (Panama) in our area. The species is also known from Brazil

(but see below).

Struthanthus marginatus is distinguished by its (sub)opposite leaves on smooth

often climbing stems, thin laminae that often have short-acuminate apices, evenly

54 FIELDIANA: BOTANY

spaced short-pedunculate triads with sessile central flowers, and the lack of

apparent bracts beneath the flowers or at the apex of the peduncle in flowering

and fruiting stages. The restriction to the eastern edge of the central volcanic area

is a further unusual characteristic. Specimens of S. quercicola have frequently been

placed under this name, and the two species are similar in appearance. They seem

to share the same habitats, and yet, if present collections are truly representative,

they are not found in the same localities. It may be that S. marginatus is restricted

to the premontane and lower montane wet forests, while S. quercicola is found in

the moister rain forest life zones as defined by Tosi & Holdridge.

There is some question regarding the use of the name Struthanthus marginatus

in our area. A photograph of the type and material from eastern Brazil, where the

type was collected, differ from our specimens in having smaller (0.5 mm) anthers

more versatile on filaments which average about 1.5 mm long (dry). Both male and

female flowers of the Brazilian material are smaller than their Central American

counterparts. However, it seems best to let current usage stand until this very

difficult genus receives monographic treatment.

Struthanthus oerstedii (Oliv.) Standley & Calderon, Lista Prelim. PI. Salvad.

74. 1925. Loranthus oerstedii Oliver, Vidensk. Meddel. Dansk Naturhist. Foren.

Kjoebenhavn 1864:171. 1865. Figure 7.

Shrubs, unisexual, branches not usually over 1 m long, leafy internodes 0.4-5 (8) cm

long, 1.4-4 mm thick, terete, usually becoming rough and reddish brown with many small

(0.1-0.2 mm) lenticels; epicortical roots developed from the stem and numerous. Leaves

opposite, subopposite, or occasionally alternate, articulate at the base, petioles 2-5 mm

long, ca. 1.5 mm broad, slightly sulcate with edges continuous with the lamina margins;

laminae 1.5-8 (10) cm long, 1-3 (5) cm broad, elliptic to slightly ovate or rhombic, tapering

gradually or abruptly to an acute, obtuse or rounded (rarely acuminate) apex, obtuse to

attenuate at the base, margin entire and revolute on drying, decurrent on the petiole,

laminae drying very stiffly chartaceous to subcoriaceous, smooth and glabrous, venation

pinnate with 2-6 pairs of major secondary veins difficult to distinguish from the tertiaries,

the proximal secondaries usually arcuate-ascending, tertiary veins often raised above.

Inflorescences 1 to several in a leaf axil, 1-2 cm long, usually with ca. 4 triads, primary

peduncle 0-5 mm long, peduncles of the triads 1-5 mm long, bracts subtending the

flowers ca. 1 mm long; flowers sessile, unisexual, the male 4-6 mm long, petals slightly

dimorphic and reflexed at anthesis, ca. 4 mm long, anthers subsessile, ca. 0.7 mm long,

yellow; female flowers ca. 4-5 mm long and narrowed to the apex before anthesis, stami-

nodia fused with the petals, stigma undifferentiated. Fruit ovoid, becoming ca. 5 mm long,

green (immature?) truncated at the apex and with a thickened area interior to the calycu-

lus, bracts persisting but small.

Epiphyte-like hemiparasites of deciduous and evergreen forest formations be-

tween sea level and 2,100 m elevation; flowering collections have been made

between June and September and in January and February. The species appears

to be endemic to the area between central Nicaragua and central Costa Rica, but

it has not been found below 500 m elevation on the Caribbean slope.

Struthanthus oerstedii is recognized by the reddish brown lenticellate stems,

small elliptic leaves blunt at the apex, and the very small inflorescences with

sessile flowers. The southernmost collections of the species come from La Suisa

(Cartage) on the Caribbean slope and from Puntarenas on the Pacific slope. An

unusual collection from the Pacific coast near Nicaragua (Howell 20243) was segre-

gated by Kuijt as species number 38 in his study of 1964. Comparison with

material from Nicaragua, where Oersted collected the type, suggests that this

specimen falls within the pattern of variation encompassed by S. oerstedii.

BURGER: FLORA COSTARICENSIS 55

Struthanthus orbicularis(H.B.K.) Bl., in Schult., Syst. Veg. 7, pt. 2:1731. 1830.

Loranthus orbicnlaris H.B.K., Nov. Gen. & Sp. 3:434. 1820. Loranthus chordocladus

Oliver, Vidensk. Meddel. Dansk Naturhist. Foren. Kjoebenhavn 1864:173. 1865.

S. chordocladus (Oliv.) Eichl., Fl. Brasil. 5, pt. 2:70. 1868. S. belizensis Lundell,

Lloydia 2:83. 1939. Figure 7.

Scandent shrubs, often climbing over small trees or with pendent stems to 2 m long,

leafy internodes 3-9 cm long, 1.5-5 mm thick, quadrangular or flattened with 2 longi-

tudinal ridges, smooth and striate (dry); epicortical roots from base and stems. Leaves

opposite or subopposite (occasionally alternate), at first lanceolate and recurved and func-

tioning to climb over adjacent vegetation, the later leaves as described, articulate at the

base, petioles 4-12 mm long, ca. 1.5 mm broad with lateral ridges continuous with the

lamina margins; laminae 2.5-6 (8) cm long, 1.2-3 (6) cm broad, obovate to suborbicular or

elliptic, bluntly obtuse to rounded at the apex with apiculate to emarginate tip, obtuse to

rounded at the base, margin entire, becoming slightly revolute, the edge thin and smooth,

decurrent on the petiole, the laminae drying stiffly chartaceous to subcoriaceous, venation

pinnate with 3-7 pairs of major secondary veins arising throughout the length of the

midvein. Inflorescence solitary in axils of leaves (occasionally fallen leaves), 3-16 cm long,

unbranched spicate or racemose panicles, the triads 3-11 mm distant on the smooth,

striate (dry) and ridged rachis 1-3 mm thick, peduncles of the triads 0-2 mm long,

subtended by caducous bracts 3-6 mm long (rarely seen on dry specimens), pedicels of the

flowers 0-1 mm long at anthesis, not expanded at the apex beneath the ovary (the floral

bracts not apparent); male flowers 6--8 mm long, yellowish, ovary ca. 1.7 mm long with

slightly flared calyculus 0.5 mm long and 1-1.8 mm broad at the apex, perianth 5-6 mm

long, anthers subsessile or on very short filaments at 2 different levels on alternating

petals, anthers 0.8-1.5 mm long, 1 mm broad, the connective apiculate; female flowers

more tubular, 4-6 mm long, staminodes reduced to ridges on the perianth, stigma sub-

capitate. Fruit borne on pedicels becoming 8 mm long and 2 mm thick with conspicuous

(0.7 mm) brownish lenticels, fruit becoming 10-13 mm long, 6-8 mm thick, truncated at

the apex, ellipsoid to cylindrical, purple to blue.

Epiphyte-like hemiparasitic climbers in evergreen or partly deciduous forest

formations from sea level to 1,200 m elevation; collected with flowers from Octo-

ber to June in Costa Rica. The species ranges from southern Mexico to Brazil and

Peru.

Struthanthus orbicularis is distinguished by its usually opposite rounded leaves

on smooth climbing or trailing stems, spicate inflorescences with subsessile triads

of flowers that develop expanded lenticellate pedicels in fruit, and moist lower

elevation habitats. The species is apparently rare on the Caribbean slope of Costa

Rica.

Struthanthus panamensis(Riz.) Barlow & Wiens, Brittonia 25:39. 1973. Phrygi-

lantlms panamensis Rizzini, Ann. Missouri Bot. Card. 47:270. 1961. Figure 6.

Shrubs or clambering branch parasites, apparently bisexual, leafy internodes 1-6 cm

long, 1-4 mm thick, terete or slightly ridged in early stages, smooth and grayish. Leaves

opposite or subopposite, articulate at the base, petioles 7-22 mm long, 1-2 mm thick,

narrowly sulcate above with thin adaxial margins continuous with the lamina margins;

laminae 3.5-10 cm long, 1.5-6 cm broad, ovate to ovate-oblong, acute to acuminate or

caudate-acuminate at the apex, the tip often elongate, truncate to obtuse or rounded at the

base, margins entire and slightly revolute (dry), the edge differentiated and smooth, the

laminae drying stiffly chartaceous to subcoriaceous, venation pinnate, the 3-7 pairs of

major secondary veins usually obscure on both surfaces. Inflorescences 1-3 in the axils of

leaves, at first subtended by 4-6 pairs of stiff decussate imbricate bracts resembling bud

scales, racemose, 1-3 (4) cm long with flowers borne on pedicels bracteate at their apex,

pedicels 1-2 mm long and 0.5-1 mm broad, the pediceled flowers (opposite monads) 2-6

mm distant along the thick (1 mm) strongly 4-angled rachis, the abaxial bract decurrent on

the peduncle with free apex ca. 2 mm long, lateral bracts ca. 1 mm long, persisting; the

56 FIELDIANA: BOTANY

flowers and bracts drying black, flower buds becoming 8-12 mm long, 2 mm thick distally

but narrowed to an acute and often slightly curved apex, yellow, calyculus minute and

entire, petals 6 and narrowly acute at the apex, yellow-green, anthers sessile at 2 different

levels on adjacent petals, ca. 1.2-1.8 mm long, apiculate, style ca. 5 mm long, stigma only

slightly thickened. Fruit apparently globose, 5-8 mm long and equally thick, yellow, apex

with a thick central disklike area within the thin calyculus, bracts and bractlets persisting

in fruit.

Epiphyte-like hemiparasites of evergreen montane (lower montane rain) forest

formations between 1,800 and 2,200 m elevation in western Panama. The species

is known only from Chiriqui, Panama. Flowers were collected in March (Davidson,

431, the type, and 392) and May (Luteyn 3786), and mature fruit, in January

(Wilbur & Teeri 1310a).

Struthanthus panamensis is recognized by its stiff opposite leaves on terete stems,

laminae often ovate and caudate-acuminate at the tip, short racemose in-

florescences, and single flowers on short stalks with one larger (abaxial) bract and

two smaller lateral bracts that persist in fruit. The flowers appear to be bisexual,

but may be functionally unisexual and male in the type, the only flowering

material seen. The very narrow petal tips are unusual among our species of

Struthanthus.

The correct genus for this species is in doubt, and its placement in Struthanthus

should be considered provisional. See the discussion by Kuijt in Ann. Missouri

Bot. Card. 65:761, 1978. The term pedicel is used in this description for stalks

bearing a single flower or monad.

Struthanthus quercicola (S. & C.) Blume, in Schult., Syst. Veg. 7:1731. 1830.

Loranthus quercicola Schlecht. & Cham., Linnaea 5:173. 1830. L. cansjeraefolius

Oliver, Vidensk. Meddel. Dansk Naturhist. Foren. Kjoebenhavn 1864:173. 1865, e

descr. et photo. Struthanthus cansjeraefolius (Oliv .) Eichler, in Martius, Fl. Brasil. 5,

pt. 2:87. 1868. Figure 7.

Shrubs or with climbing or pendent stems, to over 3 m long, leafy internodes 2-12 cm

long, 1-4 mm thick, smooth, strongly 4-angled or with 4 longitudinal ridges but becoming

terete, pale gray; epicortical roots common on stems. Leaves opposite, subopposite, or

occasionally alternate, articulate at the base, petioles 3-8 (11) mm long, 1-2 mm broad,

slightly sulcate or flattened and continuous with the lamina margins; laminae (3.5) 5-10

(13) cm long, (1) 2-5 (7) cm broad, ovate to elliptic-ovate, gradually tapering to the

cuspidate or long-acuminate apex, obtuse at the base with the margin decurrent on the

petiole, margin entire and becoming slightly revolute, the laminae drying very stiffly

chartaceous, smooth and glabrous, venation pinnate with 3-6 pairs of major secondary

veins often difficult to distinguish from the tertiary veins, the tertiaries usually slightly

raised above. Inflorescences usually solitary (3-5) in the axils of leaves, racemose panicles

2-6 cm long with 5-12 triads 3-10 mm distant on the slender (0.7-1.5 mm) rachis, the

rachis usually with 4 longitudinal ridges, peduncles of the triads (1) 2-3 (6) mm long,

bracts beneath the flowers 0.5-2 mm long, deciduous or occasionally persisting; the central

flowers sessile and the laterals subsessile or very short pedicellate, male flowers 4-5 mm

long, ovary and the very short calyculus ca. 1 mm long, perianth 3-4 mm long, becoming

reflexed, stamens with very short filaments, anthers ca. 0.8 mm long; female flowers

3.2-4.5 mm long, perianth 2-3 mm long and not becoming reflexed, staminodes small

with very slender nonfunctional anthers, style 2-3 mm long, stigma capitate and faintly

3-lobed. Fruit becoming ca. 5 mm long and 4-5 mm thick, ellipsoid to ovoid, bright orange.

Epiphyte-like hemiparasites of evergreen montane forest formations of the Cor-

dillera de Tilaran, Central Volcanic Highlands, and the Cordillera de Talamanca

between (600) 1,200 and 2,300 (2,800) m elevation; flowering material has been

collected from December to June. The species, as here interpreted, ranges from

Jalapa, Mexico, to western Panama.

BURGER: FLORA COSTARICENSIS 57

Struthanthus quercicolais distinguished by its opposite and subopposite leaves on

smooth stems, relatively thin laminae with long-acuminate apices, evenly spaced

triads on slender peduncles, inconspicuous floral bracts, and orange fruit. Stru-

thanthus quercicola has been collected along the Caribbean side of the Central

Highlands (Zarcero, Volcan Poas, Volcan Barba, La Palma, to the vicinity of

Cartago) and along the Talamanca mountains to above San Isidro del General.

This distribution differs from that of S. marginatus which is found on the slopes

of Volcan Irazu and Volcan Turrialba. The two distributions overlap in the area

between San Jose and Cartago, and yet there are no collections of the two species

from the same locality, such as Tres Rios or the campus of the university. Material

of this species has usually been identified as S. marginatus; see the discussion

under S. marginatus. Struthanthus papillosus Standl. & Steyerm. of Guatemala may

prove to be this species.

Included here are specimens from between 600 and about 1,100 m altitude on

the Pacific slope of Costa Rica that were segregated as a separate entity (No. 37)

by Kuijt in 1964. These plants tend to have more narrow leaves, shorter in-

florescences, and longer peduncles, and the little bracts subtending the flowers

are less often persisting. While they do appear quite different from most of the S.

quercicola material collected from the Central Highlands, they do not appear as

different when compared with material from Chiriqui, Panama. This group of

specimens ranges from the area of San Ramon (Brenes 1704 and 22889) to the

highlands bounding the Pacific side of the General Valley (Allen 5891, Burger &

Baker WW5 and 10108, Kuijt 2446), all between about 800 and 1,100 m elevation.

A very unusual specimen (Skutch 3954) with small (5 by 1.5 cm) lanceolate laminae

and very short (2 cm) inflorescences from the area of El General at 610 m altitude

is also included here.

Kuijt (1978) has recently identified a number of Panamanian collections as

Struthanthus aff. dichotrianthus Eichler. These have a leaf form and habit very

similar to those of S. quercicola, but they differ in having longer and narrower

leaves and much shorter inflorescences with fewer triads. These characteristics

have not been seen in Costa Rican collections.

Struthanthus rotundatus Rizzini, Rev. Brasil. Biol. 10:401. 1950. Figure 7.

Climbing branch parasitic shrubs, leafy internodes (2) 4-18 cm long, (1.5) 2-6 mm thick,

smooth but with longitudinal ridges and somewhat 4-angled, soon becoming terete;

epicortical roots few from the internodes, more often from the nodes. Leaves opposite or

subopposite, articulate at the base, petioles 5-20 (28) mm long, 1.1-2 mm thick, often

somewhat sulcate above with adaxial ridges continuous with the lamina margins; laminae

2.5-7 cm long, 2-6 cm broad, broadly obovate to suborbicular or broadly oblong, usually

rounded and emarginate at the apex, obtuse to subtruncate at the base, margin entire and

decurrent on the petiole, often undulate and slightly revolute when dry, the laminae

drying subcoriaceous to stiffly chartaceous and brittle, smooth and glabrous, venation

pinnate with 4-8 pairs of major secondary veins arising from throughout the length of the

midvein, ascending at angles of ca. 45°. Inflorescences solitary in the axils of leaves or fallen

leaves, 5-15 cm long, unbranched racemose panicles with opposite or subopposite triads

3-12 mm distant on the rachis, rachis 1.5-2 mm thick (dry) and with longitudinal ridges,

peduncles of the triads 1-3 mm long with the subtending (abaxial) bract adnate and

persisting, these bracts ca. 6 mm long from base of the peduncles to the acute or obtuse

tip, apparently fleshy, the sessile flowers subtended by shorter (2-3 mm) floral bracts

similar to the larger bract in texture; male flowers ca. 8 mm long, ovary and calyculus 2 mm

long and 2-2.7 mm in diameter distally, perianth parts (petals) ca. 6 mm long, stamens

with very short (0.5 mm) filaments, borne at 2 levels on adjacent perianth parts, anthers

1-1.7 mm long, connective apiculate; female flowers similar in size and form to the male,

58 FIELDIANA: BOTANY

calyculus subentire and thin, staminodes with reduced (0.5 mm) anthers on very short

filaments, style ca. 5 mm long with thickened stigma. Fruit ca. 10 mm long and 6 mm thick,

ellipsoid or somewhat truncated at base and apex, becoming red, borne on thick (2 mm)

peduncles to 5 mm long with occasional round brownish lenticels, bracts persisting at the

base of the sessile fruit.

Epiphyte-like hemiparasitic climbing plants of the wet montane forests of west-

ernmost Panama between 1,200 and 2,000 m elevation; probably flowering

throughout the year, but collected with flowers only in January, March, June, July,

and September. The species is known only from western Panama and Brazil (fide

Rizzini).

Struthanthus rotundatus is distinguished by its opposite rounded leaves on

smooth climbing stems often slightly quadrangular in cross section, racemose

inflorescences with sessile flowers in bracteate and pedunculate triads, the bracts

adnate to and persisting on the apex of the peduncles, and the restricted higher

elevation habitat. The anthers and staminodes with very short filaments further

distinguish this species from the very similar S. orbicularis. While not yet recorded

from Costa Rica, this species is to be expected in the montane forests adjacent to

Panama. Struthanthus macrostachyus Lundell from Chiapas, Mexico, appears to be

a close relative of S. rotundatus.

Struthanthus woodsonii Cufodontis, Arch. Bot. Sist. 10:31. 1934. Figure 6.

Twining shrubs with slender stems, leafy internodes (2) 4-12 cm long, 1-4 mm thick,

terete, smooth and usually drying dark; epicortical roots not usually found on flowering

stems. Leaves opposite, subopposite, or alternate, articulate at the base, petioles 2-7 mm

long, 0.6-1 mm thick, sulcate above with the adaxial margins continuous with the lamina

margins; laminae (2.5) 4-10 cm long, (0.6) 1.5-4 cm broad, very narrowly ovate to lanceo-

late or narrowly elliptic-ovate, gradually tapering to the acute to long-acuminate apex,

acute to obtuse at the base, margin entire, the laminae drying stiffly chartaceous, the 2—4

pairs of major secondary veins usually arising from the proximal half of the midvein and

strongly ascending, occasionally with the basal pair of secondaries very prominent and the

leaf appearing trinerved, secondary and tertiary veins usually slightly raised on both

surfaces when dry. Inflorescences 1 or 2 in the axils of leaves or fallen leaves, 2-6 cm long

and becoming 10 cm long in fruit, the male with sessile or subsessile triads and the

inflorescences spicate, the female becoming racemose panicles, the 10-16 triads 3-5 mm

distant on the slender (1 mm) ridged rachis, peduncles of the female triads 0.5-3 mm long,

ca. 0.5 mm thick, bract at the apex of the peduncle persisting or deciduous; male flowers

5-6 mm long, ovary narrowed at the base and expanded at the minutely denticulate or

erase distal margin of the calyculus, anthers subsessile, 1-1.4 mm long and apiculate;

female flowers ca. 4 mm long, ovary only slightly expanded distally, with the calyculus

obscurely denticulate (erose), staminodes minute (0.2 mm), style ca. 3 mm long, stigma

capitate. Fruit ca. 5 mm long and 3 mm thick, cylindrical or ellipsoid, slightly constricted

around the middle, orange, the triads borne on peduncles to 8 mm long, the lateral fruits

of a triad on pedicels to 3 mm long and 1.5 mm thick.

Epiphyte-like hemiparasites of aerial branches in evergreen lowland forest for-

mations between sea level and about 500 m elevation found mostly on the Carib-

bean side of Costa Rica; probably flowering throughout the year, but collected

with flowers only in October, February, March, and May. This species appears to

be endemic to the area of Costa Rica and Bocas del Toro, Panama. A single

collection from the Pacific slope (Brenes 22889, Rio Jesus de San Ramon, Alajuela-

Puntarenas) appears to be this species.

Struthanthus woodsonii is recognized by its slender twining stems, the distant,

usually lanceolate leaves with thin laminae and prominent ascending secondary

veins (when dry), the calyculus with minutely denticulate or erose margin, and

usual restriction to the lowland Caribbean area.

BURGER: FLORA COSTARICENSIS 59

VISCACEAE

Small- to medium-sized shrubs, mostly with some chlorophyll and hemiparasitic, para-

sitic on the aerial stems of dicotyledonous trees and shrubs (in ours) or less often on

gymnosperms, attaching to the host by means of a primary haustorium which often

branches within the host, young stems terete, angular, or compressed, often articulate

above the node, glabrous (in ours) or rarely puberulent, the plants unisexual or bisexual,

with foliage leaves or with all the leaves reduced to minute scale leaves, paired opposing

scale leaves present on the inflorescences (and often called bracts), on lateral branches

above branching nodes, and on percurrent stems in leafy species or defining all the nodes

in leafless (squamate) species; stipules absent, paired lateral prophylls occasionally persist-

ing above the base of lateral branches, epicortical roots absent. Leaves present or all

reduced to scale leaves (squamate), opposite and decussate in ours, the foliage leaves in

leaf-bearing species usually articulate at the base, simple and entire, usually thick and

drying stiff or brittle, venation palmate to pinnate, glabrous in ours; the small pairs of

opposite scale leaves often united at the base across the stem, generally less than 5 mm

long. Inflorescence basically a spike, borne in the axils of leaves or scale leaves, rarely

terminal, not ensheathed by deciduous scale leaves in early stages, bisexual or unisexual,

with the male and female flowers in a variety of arrangements, the spikes composed of

internodes defined by small paired scale leaves (bracts), the lower (proximal) internode(s)

sterile and functioning as a peduncle, the flowers arising individually from depressions in

the rachis of the fertile internodes, arranged in longitudinal ranks or other configurations

on the rachis; the flowers very small and unisexual, the same color as the stems in ours,

perianth of a single whorl, a calyculus absent, the perianth united with the inferior ovary

beneath and usually 3-parted or 3-lobed distally, valvate in bud; male flowers with the

stamens equal in number and opposite the perianth parts, stamens often borne on the

perianth, on short filaments or sessile, anthers 1- or 2-thecous in ours, opening by pores

or irregular slits, pollen mostly spherical; a style-like pistillode present or absent; female

flowers without staminodes, ovary inferior with single locule, style and stigma 1, ovules

not differentiated. Fruit a fleshy 1-seeded berry, seed solitary and surrounded by viscous

tissue, endosperm with chlorophyll.

A family of about 11 genera and several hundred species; most diversified in

tropical areas, but extending into milder parts of the temperate zone. Dendroph-

thora and Phoradendron are the New World counterparts of Viscum which is indig-

enous to Africa and parts of Europe and Asia. The epiphyte-like habit, parasitic

attachment to aerial parts of trees and shrubs, stiff opposite leaves or absence of

foliage leaves, and minute flowers arising from depressions in spikelike in-

florescences distinguish these plants from virtually all other Costa Rican families.

Only Oryctanthus of the Loranthaceae (s.s.) is easily mistaken for a genus of

Viscaceae.

A discussion of inflorescence types (mentioned in some of the descriptions) can

be found in Kuijt's "A study of heterophylly and inflorescence structure in Den-

drophthora and Phoradendron" (Acta Bot. Neerl. 8:506-546, 1959).

KEY TO THE GENERA OF VISCACEAE IN COSTA RICA

la Anthers with a single theca (locule), the flowers in 2, or less often 4-6, longitudinal

ranks on each fertile internode, the fertile rachis often somewhat flattened with the

floral ranks on opposing narrower edges or the 6 ranks somewhat irregularly arranged

on a terete rachis; plants lacking expanded foliage leaves (in 2 species) or with ex-

panded foliage leaves and then lacking paired scale leaves on percurrent stems and

with paired scale leaves ca. 5-10 mm above branching nodes on lateral stems; growing

as high as 3,500 m, but rarely found below 1,800 m elevation Dendrophthora

Ib Anthers with 2 thecae (locules), the flowers usually in 4-6 longitudinal ranks on each

fertile internode or rarely in broad circular areas, the fertile rachis terete or 4-angled

in cross section; plants always with expanded foliage leaves, percurrent stems with

or without paired scale leaves above the leaf-bearing node, lateral stems with paired

scale leaves above the branching node (in ours); plants growing from sea level to ca.

3,000 m elevation . Phoradendron

60 FIELDIANA: BOTANY

DENDROPHTHORA Eichler

REFERENCES: Job Kuijt, A Revision of Dendrophthora. Wentia 6:1-145, 1961. A

Revision of the Loranthaceae of Costa Rica. Bot. Jahrb. Syst. 83:250-326. 1964.

Shrubs, usually small (20-50 cm) but occasionally to 2 m tall (as in D. ambigua), branch

parasites of a great variety of dicotyledonous (rarely gymnospermous) trees and shrubs,

often densely branched and olive green or yellow-green in color, unisexual or bisexual,

stems terete and usually articulate (winged, ridged, or compressed in some species),

paired scale leaves often present on lateral stems above branching nodes, many species

with all the leaves reduced to scale leaves (i.e., squamate); haustorial system apparently

simple (lacking lateral expansion with aerial shoots), epicortical roots unknown. Leaves

present or apparently absent and represented by minute paired scale leaves, opposite and

decussate, foliage leaves with poorly differentiated petioles or subsessile, often articulate

at the base, simple and entire, the scale leaves acute to obtuse, often united at the base

across the stem, occasionally tubular. Inflorescences solitary in the axils of leaves or scale

leaves or in groups forming compound inflorescences, rarely solitary and terminal (as in

D. terminalis), unisexual or bisexual, basically a spike with 1 to several basal sterile inter-

nodes functioning as a peduncle, flowers arising from depressions in the rachis of the

fertile internodes and arranged in 2-6 longitudinal ranks (series) on each internode (1-3

ranks above each bract in a variety of arrangements); flowers unisexual and very small,

perianth of a single 3-parted or 3-lobed whorl (calyx), stamens sessile on the calyx-lobes,

anthers 1-thecous and dehiscing by a transverse slit, female flowers with inferior uni-

locular ovary and single short style, stigma scarcely differentiated, ovules not differ-

entiated. Fruit a berry, a layer of fleshy tissue and viscin surrounding the single seed;

endosperm almost completely surrounding the undifferentiated embryo and bright green.

A genus of about 53 species in tropical America; especially numerous in eastern

Cuba, Hispaniola, and the higher mountains from southern Mexico to the Boliv-

ian Andes. Almost all the continental species and a great majority of the insular

ones are restricted to mountainous areas.

The unilocular anther used to distinguish this genus from the closely allied

Phoradendron is a very difficult character to ascertain in flowers that are so small.

Most workers have, nevertheless, been able to work with these plants without

having to examine the male flowers of each species. The complete or almost

complete lack of foliage leaves immediately distinguish D. squamigera and D.

terminalis from all our species of Phoradendron. Dendrophthora costaricensis looks

very much like some species of Phoradendron, but has the male flowers terminal

and opening later than the female flowers of the same internode, and the arrange-

ment of flowers on the internodes is often quite irregular. Dendrophthora lacks the

paired scale leaves found above percurrent stems (above nonbranching nodes)

found in some species of Phoradendron.

la Foliage leaves absent or a few pairs of leaves present near the base of the plant, almost

all nodes with small scale leaves in opposing pairs; spikes axillary or terminal. ... 2

Ib Foliage leaves present on the stems, small scale leaves present on inflorescences and

above branching nodes; spikes only from the axils of leaves 3

2a Spikes numerous and arising from the axils of scale leaves, in opposing pairs;

scale leaves to 2 mm long, prophylls obscure; common plants of high montane

formations, 2,400-3,400 m D. squamigera

2b Spikes fewer, solitary and terminal on short stems; scale leaves to 5 mm long,

prophylls ca. 2 mm long and persisting above the branching node; rare plants

known only from around 1,700 m D. terminalis

3a Plants to ca. 1 m in diameter, leaves without a stiff leathery margin, drying thin and

yellowish; spikes solitary in leaf axils, flowers in 6 irregular ranks on each fertile

internode, flowering rachis terete; common plants from (1,500) 1,800-3,300 m

elevation D. costaricensis

3b Plants to 2 m in diameter, leaves with a stiff leathery margin, drying thick and dark

in color; spikes 1 or several in the leaf axils, flowers in 2 opposing ranks or up to 6

PHORADENDRON

quadrangulare

FIG. 9. Viscaceae: species of Dendrophthora and two species of Phoradendron.

61

FIG. 10. Viscaceae: species of Phoradendron, second group.

62

Fie. 11. Viscaceae: species of Phoradendron, third group.

63

64 FIELDIANA: BOTANY

ranks on a fertile internode, variable in the number of ranks but often with the rachis

flattened and the flower ranks on the opposing edges of the rachis; rare plants

probably not growing below 2,500 m D. ambigua

Dendrophthora ambigua Kuijt, Wentia 6:29-30. 1961. Figure 9.

Shrubs or subshrubs to 2 m in diameter, bisexual, leafy internodes 5-30 mm long, 2-6

mm thick, becoming striate and dark on drying, terete or slightly flattened beneath the

nodes, branching vegetative stems with a pair of scale leaves ca. 1 cm above the branching

node, the scale leaves (cataphylls) appressed to the stem in life but becoming separate in

herbarium material, the node bearing the scale leaves swollen in living material. Leaves

present throughout the plant, symmetrical, petioles 4-12 mm long but not clearly differ-

entiated from the lamina, lateral ridges of the petiole continuous with the lamina margins;

laminae 2-5 (7) cm long, 1.5-3 (5) cm broad, obovate to spatulate or elliptic-obovate,

rounded at the apex, gradually narrowed at the base, entire and with a brown leathery

margin, the laminae drying very dark and stiffly coriaceous, venation palmate, the 3-5

primary veins usually obscure. Inflorescences solitary or several in the axils of leaves, up

to 9 cm long with 2 or 3 fertile internodes 2-3 cm long, arrangement of sexes and flowers

very variable, both sexes may be present on the same internode or even above the same

bract (in the same rank), female flowers normally in an apical position in bisexual inter-

nodes and most commonly in a single median rank (series), the fertile internodes most

often with 2 opposing longitudinal ranks, but also with 2 or 3 ranks on each side above

the bract (2-6 ranks or series per internode in 2A or IB arrangements); flowers 1-2 mm in

diameter. Fruit a translucent white berry, flattened apically and discoid, ca. 6 mm long and

5 mm thick, flesh not viscid but the seed with thin viscid coating, endosperm dark green.

Branch parasites of evergreen montane forest formations known from only two

collections in Costa Rica: Kuijt 2420 and 2468, both from near Villa Mills between

2,500 and 2,800 m elevation. The species is also known from Colombia and

Ecuador at elevations over 2,500 m.

Dendrophthora ambigua is recognized by its thick rounded leaves, terete stems

with paired scale leaves on lateral branches, and the very unusual spikes. Al-

though these plants are very difficult to distinguish from species of Phoradendron,

there are usually some spikes present with only two flower-series on opposing

sides of the fertile internode. Also, there are spikes in which the two or three

ranks above each bract are separated from those above the opposing bract by a

somewhat flattened rachis. The impressions in the rachis made by fallen flowers

and fruit are often continuous between the points of floral attachment to give a

more or less continuous longitudinal groove.

Dendrophthora costaricensis Urban, Ber. Deutsch. Bot. Ges. 14:285. 1896.

vhoradendronflavescensKuntze, Rev. Gen. 2:587. 1891, non Nuttall 1847. P. crispum

Trel., Gen. Phorad. 77. 1916. P. alleniilrel., in Woodson & Schery, Ann. Missouri

Bot. Card. 27:307. 1940. D. costaricensis subsp. poasensis Kuijt, Wenria 6:51. 1961.

Figure 9.

Small shrubs to 1 m tall, bisexual, leafy internodes 0.3-3.5 (6) cm long, 1-3 mm thick,

glabrous and becoming wrinkled on drying, terete, lateral stems with 1 (2) pair of scale

leaves 5-18 mm above the branching node, percurrent stems without paired scale leaves.

Leaves well developed, petioles 1-6 mm long but not clearly distinguished from the

laminae, 1-2 mm wide, the lateral margins continuous with the lamina margin; laminae

1.5-4.5 (6) cm long, 1-3 (4.5) cm broad, broadly obovate to elliptic-oblong or oblong (rarely

suborbicular), abruptly rounded to bluntly obtuse at the apex, gradually narrowed to the

attenuate base, margin entire and of somewhat thinner differentiated tissue (dry), ve-

nation palmate with the 3 major veins usually obscure on both surfaces. Inflorescences

solitary in the axils of leaves (2 per node), 1-3 (4) cm long, the basal sterile internode 2-8

BURGER: FLORA COSTARICENSIS 65

mm long with distal paired scale leaves fused at the base and flaring distally to become 2-3

mm broad, flowering internodes usually 2 (3, 4), the fertile internodes producing 10-30

flowers in basically 6 vertical (longitudinal) ranks but these often becoming obscured, male

flowers 0.5-1 mm broad and distal on the internodes, as many as 12; female flowers

forming later and proximal on the internodes but sometimes opening earlier, lacking

staminodes. Fruit opaque white, globose to lobed, up to 6 mm in diameter, the calyx lobes

persistent on the apex, seeds ca. 1.5 mm long and 1 mm thick, endosperm dark green.

Branch parasites of moist montane evergreen forest formations between (1,500)

1,800 and 3,300 m elevation; flowering throughout the year, but collected most

often in the dry season: December to March. The species ranges from El Salvador

to western Panama.

Dendrophthora costaricensis is recognized by its small opposite leaves rounded at

the apex, terete stem with united scale leaves on branching stems, small spikes

solitary in leaf axils, distinct basal flowerless internode, and crowded flowers in

(more or less) six longitudinal ranks on each internode. The regularly spaced

nodes and regular branching often produce a symmetrical treelike form. These

plants are difficult to distinguish from species of Phoradendron.

.*

Dendrophthora squamigera(Benth.) Kuntze, Rev. Gen. 585. 1891. Viscumsqua-

migerum Bentham, PI. Hartw. 190. 1845. Phoradendron squamigerum(Benth.) Oliver,

Danske Vidensk. Meddel. 176. 1864. Dendrophthora biserrula Eichler, in Martius,

Fl. Brasil. 5:104. 1868. D. geniculata Rizzini, Rodriguesia 18-19:219-220. 1956.

Figure 9.

Small subshrubs (5) 10-40 cm tall, leafless and much branched with short crowded

internodes, bisexual, the nodes demarked by paired scale leaves, internodes 2-50 mm

long, 1-3 mm thick, glabrous and becoming roughly wrinkled when dry, terete. Leaves

absent or 2 or 3 pairs of leaves at the base of the plant and grading into the scale leaves

above, the scale leaves 1-2 mm long and opposite, partly united and with 2 sinuses or

almost completely united and entire distally, 2-5 mm broad distally or becoming broader

at branching nodes; the early-formed basal leaves up to 12 mm long, lanceolate and

succulent, usually absent on flowering collections. Inflorescences solitary in the axils of

scale leaves, 2 per node, 5-40 mm long, bisexual with the female flowers distal in an

internode, male flowers formed later and proximal on the fertile internode, the spikes with

a basal sterile internode (peduncle) 2-13 mm long and terminated by the cupulate united

scale leaves, fertile internode usually 1 (2 or very rarely to 4), the 2 opposing spikes of a

node often mirror images of each other, the fertile internode often somewhat flattened

with the 2 longitudinal ranks of flowers imbedded in the narrow opposing edges (in a 2A

arrangement), to as many as 40 flowers per internode and the flowering rachis to 4 mm

broad, flowers ca. 1 mm broad. Fruit globose or somewhat oblate (compressed at base and

apex), pearly to translucent white, 4-7 mm in diameter, drying yellowish, leaving shallow

depressions in the rachis ca. 1 mm wide.

Branch parasites of higher montane forest formations between 2,400 and 3,500

m elevation; flowering throughout the year. The species ranges from the northern

Andes of Colombia and Venezuela northward at higher elevations to Honduras

and Guatemala.

Dendrophthora squamigera is recognized by the lack of foliage leaves, short inter-

nodes and symmetrical growth that often give a dense treelike form, flowers

arranged in only two ranks, and the absence of terminal inflorescences. The

restriction to higher altitudes and the usual yellowish green color of the plants

further characterize this species. These plants are often dense and compact in

their form, but it is not unusual to find individuals with longer thin internodes

and a much more open habit. It is the most common parasite of the higher (above

3,000 m) subparamo formations that are dominated by ericaceous shrubs.

66 FIELDIANA: BOTANY

Dendrophthora terminalis Kuijt, Wentia 6:109-110. 1961. Figure 9.

Small subshrubs ca. 20 cm high, leafless and with open regular branching, bisexual but

female flowers rare, the nodes demarked by conspicuous paired scale leaves, internodes

quite uniform in length, ca. 2 cm long and 1-2 mm thick (dry), glabrous and becoming

wrinkled on drying, terete; prophylls prominent (2 mm long) and resembling the scale

leaves, lateral and persistent on the base of branching stems. Leaves absent, the scale

leaves 2-5 mm long, acute, opposing scale leaves united only near the base, the united

basal margins concave and minutely erose, the paired scale leaves widely (4-6 mm) spread

at branching nodes and when dry. Inflorescences terminal in those seen, made up of 3 or

4 fertile internodes, each fertile internode 5-10 mm long, the fertile rachis 1-1.5 mm thick,

somewhat flattened with the 2 longitudinal ranks of flowers imbedded in the opposing

narrow edges with the broader flat surface between the ranks, (2) 3-7 flowers per rank,

flowers 1-1.5 mm broad, yellow.

This species is known only from a single collection by Alexander Skutch (3570)

from Vara Blanca de Sarapiqui between Poas and Barba volcanoes at about 1,680

m elevation. The plant was collected in February and was found as a parasite on

Clusia, which itself may have begun as an epiphyte.

Dendrophthora terminalis is very similar to D. squamigera and differs primarily in

the terminal inflorescences, the prominent prophylls, and the larger, more acute

scale leaves united only near the base. The regular branching and lack of foliage

leaves are characteristic of both species. This species seems to have thinner and

somewhat more openly branched stems than most specimens of D. squamigera, but

some individual specimens of D. squamigera do have thinner stems branching

more distantly. Lateral branches develop in the axils of scale leaves one or two

internodes below the spikes, producing a somewhat dichasial habit.

PHORADENDRON Nuttall

Small- to medium-sized (3 m) shrubs, aerial (branch), parasites and attached to the host

by specialized roots but with chlorophyll in the leaves and stems, bisexual or unisexual,

glabrous in ours, young stems terete or angular, the nodes with a narrow constriction

around the circumference (articulate), often breaking easily, sometimes thickened, the

stems often bearing pairs of small (1-5 mm) basally united scale leaves just above the base

of lateral branches and/or between the leaf-bearing nodes. Leaves opposite and decussate,

articulate at the base, petioles often not clearly differentiated from the laminae, our species

with well-developed coriaceous laminae, margins entire or slightly undulate, venation

palmate to pinnate. Inflorescence axillary or rarely terminal, a thick spike with partly

imbedded flowers, divided by opposing pairs of scale leaves into 1 basal sterile and 1 to

several distal fertile internodes, the internodes with basal intercalary growth, the oldest

flowers distal on the internodes and the youngest emerging from the proximal scale leaves

(bracts), the flowers often in 4 longitudinal ranks with 2 alternate distal flowers (1A type),

less often in 6 ranks per internode (IB type), and rarely in annular or in broad groups

among our species, the spikes unisexual or bisexual; flowers very small and unisexual,

perianth of 1 series of 3 (2-5) lobes, united near the base, valvate in bud, the male flowers

with sessile or subsessile stamens at the base (opposite) the calyx lobes, anthers with 2

thecae; female flowers with inferior 1-locular ovary and 1 basal ovarian papilla, style short,

stigma capitate. Fruit a fleshy, 1-seeded berry with viscid pulp, the endosperm copious

and green; sepal lobes persistent.

A large American genus, mainly tropical, but extending into the temperate

zones in the United States and in Argentina. The genus was reviewed in 1916

when 240 species were recognized and more than half described as new (see

William Trelease, The Genus Phoradendron, A Monographic Revision, Urbana,

Illinois).

The shrubby semiparasitic and epiphyte-like habit is often associated with thick

semisucculent leaves, stems, and inflorescences. The articulate nodes, strictly

BURGER: FLORA COSTARICENSIS 67

opposite leaves, glabrous parts (in ours), and flowers arising from depressions in

the rachis further characterize the genus. The flowers in usually more than four

vertical (longitudinal) ranks and lacking subtending bracteoles distinguish Pho-

radendron from similar plants found in most species of Dendrophthora and Or-

yctanthus. Some leafy species of Dendrophtlwra are very much like Phoradendron,

but the difference in anthers seems to be a consistent distinction between the two

genera.

The following account of Phoradendron species is probably the least adequate of

any genus yet treated for the Flora Costaricensis series. At first it seemed that

inadequacy of collection, the fragile and fragmentary nature of many specimens

and unusual within-species variation were responsible for making species delim-

itation so difficult in Phoradendron. However, occasional collections suggest that

hybridization between species may be another factor. The illustrations and keys

are an attempt to characterize the commonly collected material. Neither keys nor

illustrations are of much help when working with unusual collections that may

represent variants, hybrids, or species not yet recognized in our flora. Never-

theless, we need more collections of these plants so that future workers can solve

some of the problems posed by the material at hand.

la Branching always dichotomous with the inflorescence solitary and terminal, pairs of

small scale leaves present just above the branching node and midway between

leaf-bearing nodes; rare plants of evergreen areas P. dichotomum

Ib Branching various, dichotomous or not, the inflorescences axillary to leaves. . . . 2a

2a Leaf-bearing nodes difficult to see because of the slightly expanded leaf bases,

laminae lanceolate to falcate (curved), narrowed toward the base but widened and

somewhat auriculate at the stem P. dipterum

2b Leaf-bearing nodes not obscured by the leaf bases, petioles narrowed to the

stem 3a

3a All stems between leaf-bearing nodes with small (1-5 mm) pairs of scale leaves,

these sometimes just above the node and obscure or not visible on older nodes, stems

usually terete or only slightly 2- to 4-angled in early stages 4a

3b Percurrent stems between leafy nodes lacking small pairs of scale leaves, scale leaves

present on the lateral stems above branching nodes (note that branching nodes may

occasionally appear to be percurrent when 1 branch fails to develop or is lost), stems

sometimes flattened or angled in age 9a

4a Paired scale leaves tubular or splitting, ca. 4 mm long, borne 2-8 mm above the

branching node; laminae lanceolate to falcate and often acute apically; female

spikes with the proximal (basal) half of the fertile internode lacking flowers;

branching usually dichotomous; rare in collections P. acinacifolium

4b Paired scale leaves short-tubular or a tube not apparent, borne 0-5 mm above

the node; female spike with flowers throughout most of the length of the fertile

internodes 5a

5a Laminae usually somewhat rounded at the apex; branching often dichoto-

mous; stems between leaf-bearing nodes almost always with only 1 pair of scale

leaves 6a

5b Laminae usually acute to acuminate at the apex; branching not usually dichot-

omous; stems between leaf-bearing nodes occasionally with 2 pairs of scale

leaves 7a

6a Laminae usually oblong, mostly rounded at the apex, often emarginate with a

minute succulent tip, surface slightly rough in life, the plants often grayish or

olive green; common in deciduous and semideciduous areas, 0-1,000 m

elevation P. wbustissimum

6b Laminae usually narrowly ovate to lanceolate, often tapering gradually to the

blunt apex and rarely emarginate, the surface smooth and lustrous in life,

entire plant often with a reddish cast; common in eastern areas of the Central

Highlands, evergreen areas from (0) 1,000-2,000 m elevation P. obliquunt

7a Venation pinnate, the major veins flat and inconspicuous when the leaves are

68 FIELDIANA: BOTANY

dried, the laminae tapering gradually to base and apex, 1-4 (6) cm broad;

0-1,500 m elevation but most common in the evergreen lowlands

P. piperoides

To Venation palmate, the major veins usually becoming raised and prominent

when dry, the laminae often rounded at the base 8a

8a Spikes usually borne in the axils of leaves; flowers in 6 longitudinal ranks on

each internode; the fruit obscuring the rachis and nodes of the spike, pale

orange-yellow; laminae 2-6 (8) cm broad; 600-2,000 m elevation . . .P. flavens

8b Spikes not usually borne in the axils of leaves, often in 2-4 groups along the

stem between leaf-bearing nodes; flowers in 4 longitudinal ranks with 2 alter-

nate distal flowers on each internode; fruit not usually obscuring the

rachis, yellowish white; laminae (2) 4-10 cm broad; 0-1,000 (1,500) m elevation

in evergreen formations P. crassifolium

9a Laminae lanceolate to linear-lanceolate, to 15 (20) cm long but ra/ely more than 2.5

cm broad; plants of higher altitudes, (700) 1,000-3,000 m elevation lOa

9b Laminae elliptic to narrowly ovate or narrowly oblong 12a

lOa Male flowers in 2 opposite disklike areas on each fertile internode, female

flowers usually in an annular ring around the fertile internode (a whorl of

usually 6 1-flowered ranks); leaves rarely curved and often drying dark; plants

often found on Quercus, 1,500-3,000 m P. annulatum

lOb Male flowers never found in broad disklike groupings, female flowers with

at least some ranks of 2 or more flowers, never a whorl of 1-flowered

ranks lla

lla Laminae generally curved (falcate), to 20 cm long, but rarely more than 2 cm

broad; male spikes with 6 ranks (of ca. 10 flowers each) per fertile internode;

female spikes with usually 4 ranks (of 2 flowers) and 2 alternate distal flowers;

most commonly found on the Pacific slope, (300) 1,000-2,000 m . .P. tonduzii

lib Laminae only occasionally curved, never so long and narrow 12a

12a Laminae usually elliptic to narrowly ovate or lanceolate, usually tapering to an acute

apex, often more than 8 cm long 13a

12b Laminae obovate, ovate, to narrowly oblong or very narrowly elliptic, usually

rounded at the apex, rarely more than 7 cm long 14a

13a Laminae with pinnate venation, lateral stems with 2 or 3 pairs of scale leaves

between the branching node and the first leaf-bearing node; (900) 1,100-

2,500 m P. undulatum

13b Laminae with palmate venation, lateral stems with a single pair of scale leaves

about halfway between the branching node and the first leaf-bearing node;

2,000-3,000 m P. sp. aff. undulatum

14a Lateral stems with 2 pairs of scale leaves between the branching node and the first

leaf-bearing node, young stems usually 2-angled or somewhat flattened; 1,000-2,000

m elevation in western Panama P. corynarthron

14b Lateral stems with a single pair of scale leaves between the branching node and the

first leaf-bearing node, young stems usually 4-angled in cross section; plants of

lowland (0-1,000 m) deciduous and evergreen formations. (Note that specimens

with rounded stems from above 1,400 m elevation keying to this dichotomy are

probably plants of the genus Dendrophthora.) 15a

15a Laminae usually narrowly oblong or elliptical, bluntly rounded at the apex; fruit with

a smooth or wrinkled surface; common in Costa Rica P. quadrangulare

15b Laminae usually broadly obovate, with an apical notch or bilobed at the apex (rarely

rounded); surface of the fruit with small tubercles or projections; rarely collected in

Central America P. mucronatum

Phoradendron acinacifolium Eichler, in Martius, Fl. Brasil. 5, pt. 2:117.

1868. P. robaloense Woodson ex Rizzini, Ann. Missouri Bot. Card. 47:282. 1962.

Figure 11.

Shrubs, 1-2.5 m tall, basal stems up to 7 cm in diameter, apparently unisexual, leafy

internodes 2-10 cm long, 2-6 mm thick, terete, stems usually branching dichotomously

and percurrent stems absent, 1st lateral internodes with united pairs of scale leaves form-

ing a tube ca. 4 mm long a little (2-8 mm) above the branching node. Leaves usually

BURGER: FLORA COSTARICENSIS 69

asymmetric and slightly curved, petioles 4-15 mm long, lateral ridges continuous with the

lamina margins; laminae 4-20 cm long, 1.4-5 cm broad, lanceolate to falcate or narrowly

ovate, tapering gradually to the usually acute (rarely blunt and rounded) apex, sometimes

with a minute succulent tip, tapering to the attenuate and often unequal base, margin

entire but undulate on drying, venation palmate with 5-7 primary veins, the major veins

often obscure. Inflorescences usually 2 (3) per leaf axil, 4-6 per node, the sterile rachis

(peduncle) usually drying very (1-2 mm) thin; male spikes to 8 cm long, with 3 or 4 fertile

internodes, each to ca. 2 cm long, male flowers present throughout the length of the

internode or absent near the base, in 4 longitudinal ranks of 6-10 flowers each; female

spikes 3-5 cm long, rachis with 3 or 4 fertile internodes bearing flowers only in the distal

half, each fertile internode 2-15 mm long, female flowers in 4-6 longitudinal ranks of only

1-3 flowers each. Fruit a large ovate berry with acute apex ca. 8 mm long and 6 mm thick

but much smaller (5x2 mm) and narrowly ovoid or subcylindrical when dry.

Branch parasites of evergreen montane forests between 700 and 2,000 m ele-

vation in Costa Rica; flowering and fruiting collections have been made between

January and August. The species apparently ranges southward to Paraguay.

Phoradendron acinacifolium is recognized by the usually dichotomous branching,

slightly curved somewhat falcate leaves, and slender spikes with the fertile inter-

nodes (of the female) bearing flowers only in the distal part. Description of the

male flowers and inflorescences is based on only two collections: Stone 3121 from

above Tapanti, Cartago, and Stern et al. 1992 from near Boquete, Chiriqui,

Panama.

Phoradendron angustifolium(H.B.K.) Nuttall, J. Acad. Nat. Sci. Philadelphia,

ser. 2, 1:185. 1847. Loranthus angustifolius H.B.K., Nov. Gen. Sp. PI. 3:442. 1820.

Viscum angustifolium (H.B.K.) DC., Prodr. 4:281. 1830. V. ensifolium Pohl, in DC.,

loc. cit. Phoradendron ensifolium (Pohl) Nuttall, loc. cit.

In the paper on mistletoes of Panama, P. corynarthron and P. tonduzii are placed

in synonymy under P. angustifolium (Kuijt, Ann. Missouri Bot. Card. 65:744,

1978). In the present treatment, both P. corynarthron and P. tonduzii are retained,

because they are usually easy to recognize and distinguish in Costa Rica, and

because they do not conform very well with the type description of P. an-

gustifolium or with a majority of the South American material associated with that

name. If the broader concept of P. angustifolium proves sound, then the rank of

subspecies might be considered as appropriate for the two groups of specimens

here placed under P. corynarthron and P. tonduzii.

Phoradendron annulatum Oliver, Vidensk Meddel. Dansk Naturhist. Foren.

Kjoebenhavn 1864:176. 1865. Figure 10.

Large shrubs 1-2 m tall, probably unisexual, leafy internodes 2-8 cm long, 2-9 mm

thick, slightly flattened at first but soon terete, drying dark, percurrent stems without

paired scale leaves, lateral stems with distinct paired scale leaves 5-15 mm above the

branching node. Leaves symmetrical or somewhat curved and asymmetrical, petioles 5-15

mm long but gradually merging with the lamina and the lateral ridges continuous with the

lamina margins; laminae 8-15 cm long, 1-2.5 cm broad, linear-lanceolate to somewhat

falcate, tapering very gradually to the acute or blunt apex, tapering gradually to the

attenuate base, margin entire, laminae drying stiffly coriaceous and the margins occa-

sionally slightly undulate, venation palmate to subpalmate with 3 major veins, venation

obscure. Inflorescences solitary in the leaf axils (2 per node), very short (1-3 cm) and thick,

drying dark; male spikes with 4 or 5 very short (4-6 mm) fertile internodes, each fertile

internode with 2 opposite groups of 10-15 densely crowded flowers, the flowers forming

a well-defined cushion-like area circular to ovate in outline and without floral ranks or

seriation; female spikes 1-3 cm long, usually of 4 rounded internodes 3-5 mm long and

70 FIELDIANA: BOTANY

each bearing only a single whorl (annulus) of 6 flowers deeply imbedded in the rachis (6

longitudinal ranks of 1 flower each from a modified 1A arrangement). Fruit not seen,

fruiting spikes ca. 3.5 cm long.

Branch parasites of higher montane evergreen forest formations between 1,500

and 3,000 m elevation; fertile collections have been made in July, August, and

December. The species, as presently understood, is endemic to the Cordillera de

Talamanca in Costa Rica and the adjacent highlands of western Panama.

Phoradendron annulatum is distinguished by its thick linear-lanceolate leaves on

thick rounded stems, the scale leaves about 1 cm above the base of lateral

branches, lack of scale leaves on percurrent nodes, the unusual arrangement of

male flowers in opposite circular areas, and the equally unusual "annulus" of

flowers on the female spikes. These plants often have long unbranched distal

stems with short thick internodes (as in Davidson 996 and Evans et al. 202). This

species is very closely related to P. conzattii Trel. of Mexico, but that species has

longer, more falcate leaves and the female flowers more numerous in each fertile

internode. Material ascribed to this species in the Flora of Guatemala was appar-

ently misidentified (Fieldiana, Bot. 24, pt. 4:69. 1946), as neither P. annulatum nor

P. conzattii have been collected from the area between Mexico and Costa Rica.

Phoradendron corynarthron Eichler, in Martius, Fl. Bras. 5, pt. 2:115. 1868, e

descr. P. davidsoniae Standl., Publ. Field Mus. Nat. Hist., Bot. Ser. 22:17. 1940.

Figure 9.

Small shrubs of yellowish brown appearance, leafy internodes 1-5 cm long, 1-3 mm

thick, somewhat flattened or 2-ridged in early stages but soon becoming terete, scale

leaves absent on percurrent stems, the 1st lateral internodes with 2 pairs of scale leaves,

the lowest ca. 1 cm above the base of the lateral branch and the 2nd usually 1-2 cm above

the 1st. Leaves usually symmetrical and straight (rarely curved), petioles ill defined, 1-5

mm long, the lateral winglike margins continuous with the lamina margins; laminae 4-8

cm long, 8-14 mm broad, lanceolate to very narrowly elliptic-oblong, gradually or abruptly

narrowed at the usually blunt apex, very gradually narrowed at the cuneate to attenuate

base, margin entire and drying as a rim of thinner slightly differentiated tissue, laminae

drying very stiffly chartaceous or subcoriaceous but thin, venation palmate, the 3 central

primary veins reaching the distal half of the lamina but obscure beneath. Inflorescences

solitary in the axils of leaves, 2 per node, 1-3 cm long or becoming 4 cm long in fruit,

flowering internodes with a basal sterile rachis 2-8 mm long and ca. 1 mm thick (dry); male

spikes not seen; female spikes with 2-4 fertile internodes, female flowers in 6 vertical ranks

of usually 1 or 2 flowers each (ca. 8-14 flowers per internode). Fruit becoming globose and

white, 3-5 mm in diameter.

Branch parasites of premontane and lower montane wet forest formations be-

tween 1,000 and 2,200 m elevation in western Panama; collected with flower or

fruit between March and July. The species is known only from around Volcan

Baru (Chiriqui) and the Boquete district in the province of Chiriqui, Panama.

Phoradendron corynarthron is recognized by the smaller very narrow symmetrical

leaves, the consistent presence of two pairs of scale leaves on the lateral stems

above the branching node, solitary spikes in the leaf axils, few-flowered inter-

nodes with a basal sterile rachis often equaling in length the fertile portion, and

the restricted mid-elevation habitat. While not known in Costa Rica, it may be

expected in the eastern part of the Talamanca range. This species is related to P.

tonduzii; see the discussion under that species and under P. quadr angular e.

Recently, P. corynarthron has been placed into synonymy under P. angustifolium;

see the discussion under that species.

BURGER: FLORA COSTARICENSIS 71

Phoradendron crassifolium(Pohl ex DC.) Eichler, Fl. Brasil. 5, pt. 2:125. 1868.

Viscum crassifoliumPoh] ex DC., Prodr. 4:280. 1830. P. crassifolium var. pittieriTrel.,

Gen. Phorad. 145, pi. 215. 1916. Figure 11.

Shrubs, apparently bisexual, stems usually percurrent, leafy internodes 3-15 cm long,

(1.5) 2-6 mm thick, terete, percurrent stems with a small pair of scale leaves above the leafy

node, usually with 2-4 pairs of deciduous scale leaves evenly spaced between the leafy

nodes. Leaves opposite, articulate at the stem, petioles ca. 5 mm long, difficult to dis-

tinguish because of the attenuate lamina base, margins of the lamina continuous with the

lateral ridges of the petiole; laminae 6-16 cm long, (2) 4-10 cm broad, ovate to elliptic-

ovate, elliptic-oblong, or oblanceolate, tapering gradually to the acute to acuminate apex,

tapering gradually or abruptly to the cuneate or attenuate base (sometimes rounded

basally in the broader lamina), margin entire but often undulate on drying, the laminae

drying stiffly coriaceous, venation subpalmate with the 5 major veins free or united for 1-2

cm above the base, major veins obscure in fresh material but often raised on both surfaces

and readily visible when dried. Inflorescences usually in the axils of small scale leaves and

occasionally in the axils of larger leaves, 1 or 2 (rarely 3) per axil (2, 4, or rarely 6 per node),

spikes 1-3 cm long, usually with 4-7 small closely spaced fertile internodes, flowerless

rachis ca. 1.5 mm thick (dry), paired fertile scale leaves ca. 1.5 mm long and 2 mm broad

(apex to apex), flowers in 4 vertical ranks of only 1 or 2 (3) flowers each, distal flowers in

the internode may be staminate. Fruit 2-3 mm long, ca. 2.5 mm thick (dry), pale orange-

yellow when ripe, somewhat ellipsoid, 4-8 fruit per internode, fallen fruit leaving a

depression 1-2 mm broad with a minutely erase margin.

Branch parasites of wet evergreen forest formations and known in Costa Rica

only from the southern half of the Pacific slope (western area of the General Valley

to Golfo Dulce) from 30 to 800 m elevation, but to 1,500 m in adjacent Panama;

apparently flowering throughout the year. The species ranges from Guatemala

southward to Peru and Brazil.

Phoradendron crassifolium is recognized by the thick leaves, with subpalmate

venation usually tapering at both apex and base, and the short spikes usually

borne in the axils of two to four pairs of small scale leaves on the leafless stems

between pairs of foliage leaves. The species is poorly represented in collections,

and it may be more widely distributed in Costa Rica than present collections

indicate. This is the only Central American Phoradendron in which the in-

florescences arise from the axils of scale leaves (intercallary cataphylls) as well as

from the axils of leaves or fallen leaves (but not all flowering stems exhibit this

peculiarity).

Phoradendron dichotomum (Bert.) Krug & Urban, Bot. Jahrb. 24:48. 1897.

Viscum dichotomum Bertero, in Sprengler, Syst. 1:488. 1824, non D. Don, Prod. Fl.

Nepal. 147. 1825. Figure 10.

Shrubs, bisexual, with slender dichotomizing growth, leafy internodes 3-12 cm long,

1.5-4.5 mm thick, round or slightly angular in early stages, branching dichotomous

because of terminal inflorescences (percurrent stems absent), a pair of scale leaves present

about midway between the leaf-bearing nodes and a pair of scale leaves also present just

above the leaf-bearing node. Leaves symmetrical or slightly asymmetric, petioles 2-8 mm

long but gradually merging with the lamina base; laminae 5-12 cm long, 1.7-4 cm broad,

lanceolate to narrowly ovate, gradually tapering to the acute apex, acute to attenuate at the

base, margin entire and usually drying undulate, the laminae drying coriaceous but

relatively thin, venation palmate or subpalmate with usually 5 primary veins, the major

veins usually obscure. Inflorescences 1 and terminal or occasionally with 2 additional

lateral spikes on 1 side of the dichotomizing twigs, the spikes 3-6 cm long, with up to 6

(7) fertile internodes, a sterile basal internode often present, the fertile internodes to 16 mm

long and apparently bisexual with the male flowers distal, the flowers in 6 longitudinal

FIELDIANA: BOTANY

ranks or 4 ranks with 2 alternate distal flowers (IB or less often 1A), flowers borne in deep

depressions with entire margins. Fruit and undeveloped flowers often present in the same

areas of the fertile internodes, fruit globose and ca. 3 mm in diameter (dry).

Apparently rare branch parasites of evergreen forest formations and known in

Central America from only two collections: Tonduz 13142, near Tucurrique, Car-

tago, at 635 m altitude, and Williams et al. 29155, from near Sta. Maria de Ostuma,

Cordillera Central de Nicaragua, at 1,400 m altitude. These collections were made

in January and February. The species ranges to northern South America and the

West Indies.

Phoradendron dichotomum is the only species of the genus in our area with truly

terminal spikes; however, these may fall off, and paired lateral spikes may be

found at some nodes. In any event, virtually all the stems exhibit dichotomous

branching. Scale leaves present in the middle of stems between leaf-bearing nodes

as well as at the base of all stems and the bisexual(?) fertile internodes of the spikes

further distinguish this species.

Phoradendron dipterum Eichler, in Martius, Fl. Brasil. 5, pt. 2:109-110. 1868.

Figure 10.

Shrubs, bright green to olive green, bisexual, leafy internodes 2-9 cm long, 2-6 mm

thick, somewhat flattened or 4-angled in cross section (in ours) or prominently 2- to

4-winged, becoming terete, percurrent stems without paired scale leaves, 1st lateral inter-

nodes with a pair of scale leaves at the base just above the branching node. Leaves

symmetrical or asymmetrical, sessile and the petiole absent; laminae 5-13 cm long, 1.5-3

(4) cm broad, narrowly lanceolate or falcate to ovate-lanceolate, or narrowly oblong, taper-

ing gradually or abruptly to the rounded apex, tapering gradually to the obtuse or rounded

and clasping base, margin at the base usually revolute downward and almost auriculate,

5-10 mm broad across the base, the lamina edge orange in life, entire or undulate dried,

venation palmate with 3-5 primary veins slightly raised on the upper surface (dry).

Inflorescences 1 or 2 in the axils of leaves (2-4 per node), unisexual, male and female at

different but sometimes adjacent nodes, flowers often irregularly arranged in the fertile

internodes; male inflorescences to 8 cm long with 4-7 fertile internodes, each fertile

internode ca. 1 cm long, the flowers usually in 6 longitudinal ranks of 6 or 7 flowers each,

united pairs of scale leaves (fertile bracts) ca. 2 mm long and 3-4 mm broad apically; female

inflorescences to 5 cm long with 3-5 fertile internodes, the female flowers usually in 6

longitudinal ranks of ca. (2-) 5 flowers each. Fruit an orange- tipped white spherical berry

ca. 2 mm in diameter, flesh transparent.

Apparently rare branch parasites of partly deciduous and evergreen forest for-

mations between about 1,000 and 2,000 m elevation in Central America. The

species is known in Costa Rica only from a single collection (Kuijt 2587} from El

Alto, Cartago, and flowering in August. The species, as interpreted by Kuijt

(1964), ranges from southern Mexico to northern Argentina.

Phoradendron dipterum is recognized by its narrow leaves clasping the stems at

their base with recurved auriculate-like margins, the palmate venation, the long

internodes with ridges or wings in early stages, the male and female spikes

sometimes found at adjacent nodes, and the fertile internodes with usually more

than 25 flowers. The species is known from only a few collections in Central

America. Material from Oaxaca, Mexico, under the name P. auriculatum Trelease

has the stems prominently winged and quadrangular, and the leaves are shorter

and more ovate-oblong. With so few samples at hand, it is not possible to decide

at this time the proper status of these northern elements. In most of these col-

lections, the flowers are deeply imbedded in the rachis, and the edges of the floral

depressions are minutely erose.

BURGER: FLORA COSTARICENSIS 73

Phoradendron flavens (Sw.) Grisebach, Fl. Brit. W. Ind. 313. 1864. Viscum

flavens Swartz, Prodr. Veg. Ind. 32. 1788. P. quinquenenrium Krause, Notizbl.

Konigl. Bot. Gart. Berlin 5:264. 1912. P. supravenulosum Trel., Gen. Phor. 154.

1916. Figure 11.

Shrubs 1-2 m broad, branch parasites, apparently unisexual, leafy internodes 2-7 cm

long, 1.3-4 (6) mm thick, terete or somewhat 2- or 4-angIed in early stages, percurrent

stems with a pair of keeled scale leaves ca. 1 cm above the basal node, the 1st lateral leafy

internode with a pair of scale leaves near the base and a 2nd pair at ca. Vs the length to

the next leafy node. Leaves opposite, somewhat articulate at the stem, petioles 1-6 mm

long, not clearly distinguished and with lateral ridges continuous with the lamina mar-

gins; laminae 4-11 (14) cm long, 2-6 (8) cm broad, ovate to elliptic-ovate or elliptic,

occasionally lanceolate or asymmetric and curved, tapering gradually or abruptly to the

acute to acuminate apex (occasionally obtuse), tapering gradually or abruptly to the atten-

uate base, margin entire, drying stiffly chartaceous to subcoriaceous, venation palmate or

subpalmate with 3 or 5 primary veins, the 3 central primaries reaching the distal part of

the lamina, the major veins often raised on the surface of the dried leaves, secondaries

obscure. Inflorescences at first solitary but often with 1 or 2 additional developing in leaf

axils (2-6 per node), the spikes 2-6 cm long, secondary spikes subtended by sterile scale

leaves, the paired fertile scale leaves ca. 3 mm broad (tip to tip) and 2-3 mm long and

keeled, sterile portions of the rachis ca. 0.8-2 mm thick (dry), with 6 vertical ranks of 2-7

flowers each, male spikes with usually 6-15 flowers per internode, the female with 30-50

flowers. Fruit becoming contiguous and obscuring the rachis, 2-3 mm in diameter, glo-

bose, yellowish white, pulp viscid and transparent, fertile internodes to 15 mm long.

Branch parasites of wet evergreen and moist mountain forest formations be-

tween 600 and 1,900 m elevation on both the Caribbean and Pacific slopes of Costa

Rica; probably flowering throughout the year, but not collected in July- August or

October and December. The species ranges from Guatemala and Belize to north-

ern South America and the southern Caribbean Islands.

Phoradendron flavens is recognized by the scale leaves present on the often

slender percurrent stems, leafy nodes often with two older and four younger

spikes, laminae often narrowed to both apex and base with the palmate venation

readily apparent when dry, and the flowers usually in six vertical ranks. This

species is closely related to P. piperoides, but is easy to separate by the obviously

palmate venation of the usually broader leaves. Typical material of this species is

distinguished by the ovate to oblong leaves with three primary veins distinctly

raised above when dried and the succulent fruits becoming contiguous so that the

nodes are not visible on the spike.

Phoradendron woodsonii Trelease, discussed by Kuijt (1974) in his recent review

of the Panamanian species, would key out here if found in the Costa Rican flora.

This species has two to four pairs of scale leaves on its internodes, and it differs

from P. flavensin other ways. Phoradendron flavens, P. crassifolium, P. piperoides, and

P. woodsonii appear to form a closely related group, and it is possible that hybrid-

ization occurs among them.

Phoradendron mucronatum (DC.) Krug & Urban, Bot. Jahrb. 24:34. 1897. Vis-

cum mucronatum DC., Prodr. 4:282. 1830.

Small shrubs, leafy internodes to ca. 3 cm long and 3 mm thick, usually with 4 promi-

nent longitudinal ridges and square in cross section, glabrous, percurrent stems without

scale leaves but often with an articulation just above the node, lateral stems with a pair of

scale leaves ca. 3 mm long just above the node. Leaves short and broadly rounded, petioles

1-3 mm long; laminae 1-4 cm long, 1-3 cm broad, broadly obovate to suborbicular,

abruptly rounded distally and emarginate to bilobed at the apex, abruptly narrowed at the

base, margins entire, laminae drying yellowish green and subcoriaceous, venation palmate

74 FIELDIANA: BOTANY

with the 3 or 5 primary veins often visible on the upper surface. Inflorescence 0.5-3 cm

long, usually solitary in the axils of leaves or fallen leaves, each fertile internode 2-5 mm

long, usually with only 1 flower in each of 4 ranks resulting in a whorl of 4 flowers (rarely

6) per internode. Fruit ca. 2.5 mm in diameter, with persisting and erect perianth lobes ca.

0.5 mm high, surface covered with small papillae that give a muricate or tuberculate

appearance.

A species of evergreen or partly deciduous lowlands often found near man-

groves, lakes, and rivers in Central America; from sea level to 800 m elevation.

Rarely collected in Mesoamerica, but ranging from the Yucatan peninsula and the

West Indies to southern Brazil.

Phoradendron mucronatum is a very distinctive member of the genus with small

rounded leaves deeply emarginate or bilobed at the apex, short four-ridged inter-

nodes, short spikes often with only one whorl of flowers per fertile internode and

unusual surface of the fruit. Although known from Panama and Guatemala, this

species has not been collected from Costa Rica; it is likely to occur in the Caribbean

lowlands.

Phoradendron obliquum (Presl) Eichler, Fl. Brasil. 5, pt. 2:134. 1868. Figure 11.

Large shrubs to 2 m in diameter, often pendent, unisexual, the branches often forking

dichotomously, leafy internodes 2-15 cm long, 2-8 mm thick, young stems slightly flat-

tened or round and the older terete, percurrent stems rare and with paired scale leaves just

above the node, lateral stems with paired scale leaves just above the branching node and

somewhat tubular distally (a second pair of scale leaves rarely present ca. 1 cm above the

node). Leaves often asymmetric and somewhat curved, petioles to 15 mm long but difficult

to distinguish from the laminae, the winged margins continuous with the lamina margins;

laminae 6-15 (23) cm long, 2-7 (9) cm broad, narrowly ovate to lanceolate and sometimes

slightly curved, tapering gradually to the acute or bluntly rounded apex, rounded or

gradually to abruptly narrowed at the attenuate base, margin entire but drying undulate,

the laminae drying stiffly coriaceous, venation palmate, subpalmate, or pinnate with the

secondaries arising in the lower 4th of the midvein, the major veins usually 5 and only the

central 3 subparallel, often obscure (dry). Inflorescences 3-5 cm long, the male as many as

12 per node, the female 2-6; male spikes unknown (see discussion); female spikes with 3

or 4 fertile internodes with 4 longitudinal ranks of 2-4 flowers each and 2 distal alternating

flowers. Fruit ellipsoid to somewhat cylindrical, ca. 3-5 mm long and 2-3 mm thick (dry),

depressions in the rachis to 3 mm in diameter and with an entire edge.

Branch parasites of evergreen and partly deciduous forest formations between

sea level and 1,400 m elevation; probably flowering throughout the year. This

species ranges from Costa Rica to Peru; our material has been collected in the

Central Highlands and their slopes and southwestern Costa Rica.

Phoradendron obliquum is recognized by the frequent dichotomous branching,

scale leaves usually present just above the node on all stems, thick larger leaves

usually tapering to a narrowed apex and often slightly curved, and flowers

throughout the length of the fertile internodes. In life, the plants are often char-

acterized by a reddish color and leaves with a shiny-glossy surface. The latter

character differentiates this species from the closely related P. robustissimum which

has rough-surfaced leaves in life. Unfortunately, this character disappears on

drying, and some specimens may be very difficult to place in one or the other

species. The two species are very closely related, and intermediate plants or

populations may exist: see the discussion under P. robustissimum.

A survey (Kuijt, 1964) of almost 30 plants near the continental divide between

Tres Rios and Cartago disclosed no staminate plants or spikes. A more intense

study of plants on the University campus near San Pedro found that male flowers

are extremely rare (1% to 2%), occurring as the lowest (proximal and last devel-

BURGER: FLORA COSTARICENSIS 75

oping) flowers of a floral rank in occasional fertile internodes. The fact that these

populations were producing fruit suggests that these plants may have been re-

producing asexually (Kuijt, 1964).

This species has been the subject of a thesis by R. E. Zurcher: Las Loranthaceae

que parasitan el laurel, Cordia alliodora, en Costa Rica y sus posibilidad de control

con inyecciones de hierbicidas al tronco del huesped, Inst. Interamer. Ciencias

Agric, Turrialba, 1958.

Phoradendron piperoides (H.B.K.) Trelease, Genus Phoradendron 145. 1916.

Loranthus piperoides H.B.K., Nov. Gen. & Sp. 3:443. 1820. P. biolleyi Krause,

Notizbl. Kdnigl. Bot. Gart. Berlin 5:264. 1912. Figure 11.

Small shrubs, epiphyte-like hemiparasites, bisexual, the stems often trailing, leafy inter-

nodes 2-8 (10) cm long, 1.5-4 mm thick, terete, percurrent stems with a pair of reduced

opposite scale leaves just above the node and often difficult to see, the first lateral inter-

node with 1-5 pairs of scale leaves, 1 pair often near the center of the 1st lateral internode.

Leaves opposite, petioles 1-4 (7) mm long, 1-2 mm broad, margins of the lamina con-

tinuous with the wings of the petiole; laminae 3-11 (14) cm long, 1-4.5 (6) cm broad,

elliptic to oblong or narrowly obovate, acuminate to acute at the apex, usually tapering to

the acute or attenuate base, margin entire but undulate on drying, the laminae drying

coriaceous, glabrous and smooth, venation pinnate with 2-3 pairs of major secondary

veins but these usually obscure. Inflorescences 1-3 per leaf axil (2-6 per node), very rarely

terminal, the spikes 3-8 cm long with 1-3 sterile internodes near the base, the paired

fertile scale leaves ca. 2.3 mm broad (apex to apex), sterile portion of the spike ca. 1 mm

thick (dry), fertile internodes 6-11, usually with 4 longitudinal ranks of 2-4 flowers and

2 alternate distal flowers, rarely to 28 flowers per internode. Fruit spherical to ellipsoid, ca.

3 mm in diameter and yellowish to orange when ripe, separate from each other when

dried.

Branch parasites of evergreen and partly deciduous forest formations between

sea level and about 1,500 m elevation on both the Caribbean and Pacific sides of

Costa Rica; apparently flowering throughout the year, but not collected in October

and November and with the majority of collections made in February and August.

The species ranges widely over the American tropics.

Phoradendron piperoides is recognized by the small paired scale leaves on all the

slender and usually terete stems, medium-sized laminae tapering to both apex

and base with usually obscure pinnate venation, and the infloresences usually in

axillary clusters at leafy nodes. The scale leaves just above the nodes of percurrent

stems are often difficult to see or are represented only by opposite scars. This is

one of Costa Rica's most abundant species of Phoradendron, especially common in

the wet lowlands. It is particularly abundant in the Limon region on cacao and

shade trees.

Phoradendron quadrangulare (H.B.K.) Krug & Urban, Bot. Jahrb. 24:35. 1898.

Loranthus quadrangularis H.B.K., Nov. Gen. & Sp. 3:444. 1818. P. ceibanum Trel.,

Genus Phoradendron 110. 1916. P. rensoniiTrel., loc. cit. 105. P. venezuelenseTrel.,

loc. cit. 111. P. corynarthron var. seibertii Trel., Ann. Missouri Bot. Card. 24:187.

1937. P. paquitanum Trel., Publ. Field Mus. Nat. Hist., Bot. Ser. 18:405. 1937. P.

herrerenseTrel., Ann. Missouri Bot. Card. 27:307. 1940. P. sonanumTrel., loc. cit.

308. P. seibertii (Trel.) Rizzini, loc. cit. 47:285. 1960. Figure 9.

Small shrubs, bisexual(?), branches usually erect and often branching dichotomously,

leafy internodes, 1-8 cm long, 1-4 (6) mm thick, with 4 distinct longitudinal ridges but

becoming rounded in age, often slightly thickened beneath the node, percurrent stems

lacking fused pairs of scale leaves, 1st lateral stems with a pair of fused scale leaves just

76 FIELDIANA: BOTANY

above the branching node. Leaves usually symmetrical and straight, subsessile or with

poorly defined petioles 0-4 (6) mm long, with lateral ridges continuous with the lamina

margins; laminae 2.5-6 (7.5) cm long, 1-2.3 cm broad, very narrowly oblong, to narrowly

elliptic or very narrowly obovate, rounded to bluntly obtuse at the apex, cuneate to

attenuate at the base, margin entire, the laminae drying very stiffly chartaceous to subcor-

iaceous and often becoming grayish green, venation palmate with 3 or 5 primary veins, the

3 central veins reaching the distal part of the leaf, the major veins slightly raised or obscure

(dry). Inflorescences 1 or less often 3 in each leaf axil (2-6 per node), spikes 1.5-3 (5) cm

long, peduncle 2-6 mm long, flowering axis with 4-6 fertile internodes, the fertile pairs

of scale leaves often small and inconspicuous, male and female spikes quite similar, the

flowers in 4 vertical ranks of 2-4 flowers (very rarely to 7 in male spikes), and 2 alternating

distal flowers, usually up to 18 flowers per fertile internode. Fruit globose, to 4 mm in

diameter, yellow to yellow-orange.

Branch parasites most often found in evergreen and partly deciduous (tropical

moist and premontane moist) forest formations below 1,000 m in elevation; flow-

ering throughout the year. The species ranges from southern Mexico to northern

South America and the West Indies.

Phoradendron quadrangulare is characterized by the small- to medium-sized

leaves that are very narrow and abruptly rounded at the tip, the four-angled

young stems, short slender spikes with few fertile internodes (or the internodes

difficult to distinguish because of the very small paired scale leaves), and the

apparent preference for semideciduous lowland habitats. The present circum-

scription of this species includes considerable variation in inflorescence

morphology, but the laminae are rather uniform. For example, one of the very few

collections from the wet Caribbean lowlands (Lent 1926 from Tirimbina, Sarapiqui,

Heredia) has very narrow stems, distinctly petiolate leaves, and short spikes with

very few (three to five) flowers per fertile internode. Material from Venezuela

often has short-cylindrical or ellipsoid fruit, whereas ours appears to be uniformly

globose as interpreted from the dry collections.

Several collections from near Tilaran (Standley & Valeria 44983 and 45577, Wil-

liams, Molina, & Williams 26606) key to this species, but are perhaps more closely

related to P. corynarthron and P. tonduzii. The leaves of these specimens are a bit

larger, but of much the same shape, as P. quadrangulare and P. corynarthron.

However, they lack the angled stems of P. quadrangulare and the second set of scale

leaves (on lateral stems) of P. corynarthron. These collections also lack the falcate

leaves and two-ridged stems of P. tonduzii. All three species and the Tilaran

material have very similar inflorescences and appear to be closely related. Perhaps

the collections from Tilaran represent an unusual population of mixed parentage.

The type of P. dodgei (Dodge 6200) from the same area, while having many of the

characteristics of P. tonduzii, does resemble the specimens referred to above and

may be intermediate between them and typical P. tonduzii. All these collections

come from altitudes beneath those at which P. corynarthron and P. tonduzii are

usually found.

Phoradendron robustissimum Eichler, in Martius, Fl. Brasil. 5, pt. 2:122. 1868.

P. falcifolium Trel., Gen. Phorad. 79. 1916. P. pergranulatum Trel., Ann. Missouri

Bot. Card. 27:308. 1940. Figure 11.

Shrubs to 1 m or more, dark green or grayish green, unisexual, usually branching

dichotomously, leafy internodes 2-10 cm long, 2-6 mm thick, usually terete, percurrent

stems infrequent, both percurrent and lateral stems with a single (rarely 2 or 3) pair of

united scale leaves ca. 5 mm above the node, 2 spikes often peripheral to the aborted apex

with vegetative laterals developing axillary to prophylls of the spikes or vice versa. Leaves

BURGER: FLORA COSTARICENSIS 77

symmetric or the larger often asymmetric, petioles 2-15 mm long but not easily dis-

tinguished from the attenuate leaf base, broader than thick and with the lateral margin

continuous with the lamina margin; laminae 3.5-10 (14) cm long, 2-5 (8.5) cm broad,

oblong to ovate-oblong or somewhat spatulate, abruptly rounded at the apex, often emar-

ginate with a minute succulent tip, abruptly narrowed at the attenuate base, margin entire

with a thin brownish edge, the laminae drying stiffly coriaceous, surfaces with a slightly

rough texture in life and, when dry, venation palmate to pinnate with the secondaries

arising from the lower 3rd of the midvein, usually subpalmate, with usually 5 major veins

and these often obscure. Inflorescences dimorphic, the male much thinner than the fe-

male, 1 to several per leaf axil or the males as many as 12 on older nodes; male spikes to

11 cm long, with 4-6 fertile internodes 1-2 cm long, sterile rachis ca. 1-2 mm thick, male

flowers in 4 (6) longitudinal ranks of ca. 10 flowers each and with 2 alternating terminal

flowers (1A arrangement) female spikes to ca. 6 cm long with 3-5 fertile internodes, the

central (longer) internodes 1-2 cm long with 4 ranks of 4 or 5 flowers each and 2 alternate

terminal flowers (1A arrangement). Fruit elliptic-ovate or cylindric (dry) and about 4 mm

long and 2 mm thick, yellowish to dusty yellow-green.

Branch parasites of both evergreen and deciduous lowland forest formations

below about 1,000 m in Costa Rica; apparently flowering throughout the year. The

species ranges from southern Mexico to Venezuela.

Phoradendron robustissimum is recognized by the stiff oblong leaves usually with

bluntly rounded apices, frequent dichotomous forking of the stems, slender male

spikes often more than eight at older nodes, and usual restriction to lower (0 to

500 m) altitudes on the Pacific slope. This species is related to P. oblicjuum and may

be difficult at times to separate from specimens of that species; see discussion

under P. obliquum.

Phoradendron tonduzii Trelease, Gen. Phorad. 67. 1916. P. cooperi Trel. , loc.

tit. P. dodgeilrel, Publ. Field Mus. Nat. Hist., Bot. Ser., 18:405. 1937. P. novae-

helvetiaelrel., Ann. Missouri Bot. Card. 27:307. 1940. Figure 10.

Shrubs, often orange-brown or yellow-brown, to ca. 1 m broad and often pendulous,

unisexual, leafy internodes 1-10 cm long, 1-5 mm broad, with 2 opposing ridges and

somewhat flattened in early stages, scale leaves absent on percurrent stems, the 1st lateral

internode with 1 or rarely 2 pairs of scale leaves near the base. Leaves opposite, articulate

at the stem, usually asymmetric, petioles 5-20 mm long but not really distinguishable from

the lamina base; laminae 6-15 (20) cm long, 0.5-2 (3) cm broad, very narrow and usually

curved (falcate) to narrowly lanceolate, tapering gradually to the acuminate apex, tapering

gradually to the attenuate base, margin entire or undulate, laminae drying very stiffly

chartaceous, smooth and glabrous, venation palmate with 3-6 primary veins the 3 central

primaries subparallel and reaching the distal half of the lamina, slightly raised and usually

visible (dry). Inflorescences 1-3 in the axils of leaves (2-6 per node), spikes 3-7 cm long,

usually without basal sterile scale leaves; the male spikes with ca. 4-8 fertile internodes,

flowerless rachis 0.5-1.5 mm thick (dry), the male flowers usually in 6 ranks, of ca. 20

flowers each (ca. 70 flowers per internode), the paired scale leaves almost completely

united and only slightly cleft, 2-2.5 mm broad at the apex; female spikes with 2-5 fertile

internodes, female flowers usually in 4 ranks of 2-4 flowers with 2 alternate distal flowers,

ca. 10-20 flowers per internode, the united scale leaves becoming slightly cleft. Fruit ovoid

to globose, 3-4 mm long, white but becoming orange in the sun, flesh translucent orange.

Branch parasites of wet evergreen and partially deciduous forest formations

between (300) 1,000 and 2,000 m elevation and only rarely collected on the Carib-

bean slope; apparently flowering throughout the year. As here defined, the

species is endemic to the area between Tilaran, Guanacaste, and western Panama

(see below).

Phoradendron tonduzii can often be recognized almost immediately because of the

long and very narrow curved leaves. The young stems with two opposite ridges,

lack of scale leaves on the percurrent stems, and male spikes more slender and

78 FIELDIANA: BOTANY

with many more flowers than the female further distinguish this species. Among

our species, the relationships of P. tonduzii are with P. corynarthron and P. quad-

rangulare. A number of collections from Tilaran may represent an unusual popu-

lation of P. tonduzii (Kuijt, 1964) or a population of hybrid origin. However, these

plants are here keyed to and discussed under P. quadmngulare. Recently, P. ton-

duzii has been placed into synonymy under P. angustifoium; see the discussion

under that species.

Phoradendron undulatum Eichler, Fl. Brasil. 5, pt. 2:122. 1868. P. gradlispicum

Trel., Gen. Phorad. 130. 1916. Figure 10.

Shrubs, often pendent, bisexual, leafy internodes 1-9 cm long, 2-8 mm thick, with 2

opposite ridges and somewhat flattened in early stages but becoming terete, percurrent

stems without paired scale leaves, first lateral internodes with (1) 2 or 3 pairs of scale leaves

with the distal often in the middle of the internode. Leaves opposite, articulate at the base,

often slightly asymmetric, petioles 2-10 (16) mm long, with lateral ridges continuous with

the lamina margins; laminae 5-17 (23) cm long, 1.5-4.5 (6) cm broad, lanceolate to very

narrowly elliptic-ovate or narrowly elliptic, tapering very gradually to the acute to acu-

minate apex, abruptly to gradually narrowed at the obtuse to acute base, margin entire or

undulate when dry, the lamininae drying stiffly coriaceous, venation pinnate with 2 or 3

pairs of major secondary veins but the secondaries often obscure. Inflorescences 1 or 2 (3

or 4) in leaf axils (2-8 per node), spikes 1.4-5(7) cm long, with 6-12 fertile internodes 4-10

mm long, with 4 (-6) longitudinal ranks of 1-3 (4) flowers each, with usually 10-14 flowers

per internode, terminal internode usually with only 2 flowers; male flowers few and

usually in an apical position, pairs of fertile scale leaves often forming a distal edge

perpendicular to the rachis and very slightly erose. Fruit a white globose berry ca. 4 mm

in diameter, often drying cylindrical, flesh transparent and viscid.

Branch parasites of wet evergreen montane forest formations between (900)

1,100 and 2,500 m elevation in Costa Rica; probably flowering throughout the year,

but collected most often in December and February. The species ranges from the

Cordillera Central de Nicaragua southward to Bolivia and Brazil.

Phoradendron undulatum is recognized by the lack of scale leaves above per-

current internodes, flattened young stems, stiff leaves gradually tapering to the

apex and with pinnate venation, spikes with short and few-flowered internodes,

and the montane habitat. Small plants of P. undulatum are easily confused with

P. piperoides, but that species has paired scale leaves on the percurrent internodes.

Phoradendron sp. aff. undulatum. Figure 10.

Shrubs, green, apparently unisexual, branches to ca. 1 m long, leafy internodes 2-5 (7)

cm long, 1.5-6 mm thick, often with 2 longitudinal ridges and somewhat flattened or even

quadrangular, older nodes conspicuously thickened, percurrent stems lacking scale

leaves, lateral stems with 1 (rarely 2) pair of united scale leaves 8-30 mm above the

branching node. Leaves usually symmetrical, petiole 5-15 mm long but gradually expan-

ding into the leaf with lateral margins continuous with the lamina margins; laminae 5-11

cm long, 2-4 cm broad, lanceolate to narrowly ovate, tapering gradually to the acute or

acuminate apex, gradually or abruptly narrowed to the attenuate base, the larger laminae

more abruptly narrowed and somewhat asymmetric at the base, the laminae drying thin-

to rigid-coriaceous with the margin often becoming undulate, venation palmate with

usually 5 major veins, the venation often obscure. Inflorescences 1-3 from the leaf axils

(2-6 per node), 2-3 cm long; male spikes with 2-5 fertile internodes, each fertile internode

with 6 longitudinal ranks of ca. 4 or 5 flowers each; female spikes with 3 or 4 fertile

internodes, sterile rachis of the fertile internode often as long (2-4 mm) as the fertile

rachis, sterile rachis 0.6-1 mm thick (dry), female flowers in 6 ranks (4 plus 2) of only 1 or

2 flowers each. Fruit not seen.

BURGER: FLORA COSTARICENSIS 79

Branch parasites of wet evergreen montane forest formations between about

2,000 and 3,000 m elevation; probably flowering throughout the year. The species

is known only from the Central Volcanic Highlands and Cordillera de Talamanca

in Costa Rica.

Phoradendron species (aff. undulatum) is recognized by the paired scale leaves

about halfway up the first lateral internodes (to the first leaves), angular stems,

acute to acuminate laminae, short spikes, and few-flowered female internodes.

The description of female spikes is based on Kuijt 2469; no other female material

is known. This species is very similar in overall appearance to P. undulatuni, but

that species has leaves with pinnate venation, more scale leaves on the lateral

stems above a branching node, and undulating leaf margins.

ARISTOLOCHIACEAE

By Kerry Barringer

REFERENCES: F. C. Hoehne, Aristolochiaceae, in Fl. Brasil. 15(2):1-141. 1942.

Howard W. Pfeifer, Revision of the North and Central American hexandrous

species of Aristolochia. Ann. Missouri Bot. Card. 53:115-196. 1966. O. C. Schmidt,

Aristolochiaceae, in Engler & Prantl, Nat. Pflanzenfam. ed. 2. 16b:202-242. 1935.

Lianas, herbs, or shrubs, often aromatic, wood with broad medullary rays; true stipules

absent, clasping pseudostipules sometimes present. Leaves alternate, simple, petiolate;

lamina entire or lobed, usually cordate and reniform, often palmately veined, often with

eglandular pubescence. Flowers solitary or racemose, bisexual; calyx enlarged, petaloid,

usually tubular, radially or bilaterally symmetrical, the lobes valvate; petals usually absent

or reduced to scales; stamens 5, 6, or multiples thereof, free or connate, anthers extrorse,

dehiscing by longitudinal slits; ovary inferior or half inferior, 4- to 6-locular, rarely of free

carpels; ovules many in each locule, axile or parietal, anatropous. Fruit a septicidal capsule

or follicle, usually pendent, rarely indehiscent; seeds with copious endosperm.

A family of seven genera and about 600 species, the Aristolochiaceae previously

were placed in an order with the Rafflesiaceae and Hydnoraceae. They are now

generally classified into an order by themselves and are believed to be derived

from the Magnoliales, with affinities to the Annonaceae. Many authors have

noted the similarities between the lianoid species of this family and the Di-

oscoreaceae. Only Aristolochia has been found in Costa Rica.

ARISTOLOCHIA Linnaeus

Woody or herbaceous lianas, rarely shrubs or trees; true stipules absent but broad,

clasping pseudostipules formed by the first leaf of axillary buds often present. Leaves

alternate, distichous, simple, petiolate, entire or lobed, usually palmately veined with 3-7

primary veins. Flowers solitary and axillary or in racemose clusters which may be axillary

or from older parts of the stem, bisexual, variously colored with yellow, brown, and red,

often fetid smelling; calyx tubular, bilaterally symmetrical due to bending of the tube,

forming an inflated, ovoid utricle at its base, a narrowed or funnelform tube, and an

expanded, 1- to 3-lobed, variously structured limb; stamens in multiples of 3 or 5, connate

into a gynostemium which appears to also function as stigmas; ovary inferior, narrowly

cylindric, 5- or 6-loculate. Fruit a dry, septicidal capsule, pendent; valves and pedicels

usually splitting from the base of the capsule, the valves remaining attached at the apex,

sometimes with persistent, lattice-like septa connecting the valves; seeds compressed,

often with paired, lateral wings.

A genus of 400 to 500 species, chiefly pantropical. The Costa Rican species are

all lianas whose flowers have six stamens and six-loculate ovaries. Little is known

about the pollination of Aristolochia in the wild. Recent work in Costa Rica indi-

A. veraguensis

A. constricta

Fruit

ARISTOLOCHIA

maxima

A. tonduzii

A. Translucida

FIG. 12a. Aristolochiaceae: Aristolochia maxima and its Costa Rican allies; note the un-

usual fruit.

80

A. gigantea

ARISTOLOCHIA

odoratissima

A. grandiflora £l

A pilosa

A. anguicida

A. triloba

FIG. 12b. Aristolochiaceae: six Costa Rican species of Aristolochia, two with large flowers.

81

FIELDIANA: BOTANY

cates that butterfly larvae feed on the leaves of some species, thereby obtaining

substances which make them distasteful to predators. Extracts of the stems and

roots have been used against snakebite and to alleviate pain during childbirth.

Sterile material is sometimes confused with species of Dioscorea, but Aristolochia

can usually be recognized by the distichous leaf arrangement on young shoots.

la Pseudostipules rounded, clasping, at the base of leaves 2

Ib Pseudostipules absent 6

2a Leaves 3-lobed; calyx limb with a long apical appendage A. trilobata

2b Leaves entire; calyx limb without apical appendage 3

3a Leaf blade broadly reniform, glaucous below; flowers bilabiate . . . . A. ringens

3b Leaf blade triangulate, glabrous or slightly pubescent below; flowers uni-

labiate 4

4a Leaf apex rounded; fruit with a prominent beak A. littoralis

4b Leaf apex acute to cuspidate; fruit without beak 5

5a Capsule narrowly cylindric, 7-10 cm long; calyx limb peltate, acuminate; leaves

usually hastate A. odoratissima

5b Capsule ovoid, 2-3 cm long; calyx limb triangulate, folded; leaves triangulate

A. anguicida

6a Plants hirsute with brown septate hairs A. pilosa

6b Plants glabrous, strigose, or tomentose with white hairs 7

7a Leaves broadest at base; fruit valves free 8

7b Leaves broadest at middle; fruit valves usually connected by the persistent, lattice-

like septa 12

8a Petioles 10-15 cm long 9

8b Petioles 2-8 cm long 10

9a Leaves variegated with yellow or white along veins; flowers in racemose clusters on

old wood A. veraguensis

9b Leaves green, not variegated. Flowers solitary on older stems A. grandiflora

lOa Lower leaf surface white tomentose; petioles 5-7 cm long A. gigantea

lOb Lower leaf surface glabrous; petioles 2-4 cm long 11

lla Capsule narrowly cylindric, 7-10 cm long; calyx limb peltate, acuminate; leaves

usually hastate A. odoratissima

lib Capsule ovoid, 2-3 cm long; calyx limb triangulate, folded; leaves triangulate

A. anguicida

12a Leaves ovate, long acuminate, deeply cordate; fruits 5-6 cm long . . . . A. constricta

12b Leaves oblong, short acuminate to cuspidate, truncate to shallowly cordate; fruits

6-10 cm long 13

13a Leaf base truncate; calyx 5-7 cm long A. maxima

13b Leaf base shallowly cordate; calyx 8-13 cm long 14

14a Calyx limb translucent, with green spots A. translucida

14b Calyx limb opaque, with dark elevated maculae A. tonduzii

Aristolochia anguicida Jacq., Enum. Syst. PI. 30. 1760. A. loriflora Mast., Bot.

Jahrb. Syst. 8(1):220. 1887. Figure 12b.

Herbaceous lianas, younger stems glabrous; pseudostipules often present, rounded,

clasping, 1.5-2 cm wide. Leaves with petioles 3-4 cm long; laminae triangulate, 5-6.5 cm

long, 4.5-6 cm wide, acuminate, base cordate with a sinus ca. 1 cm deep, 5 primary veins

diverging from the base, glabrous. Flowers solitary in leaf axils or on short leafy shoots,

ovary and pedicel 2.5-3 cm long; utricle horizontal, ovoid, 8-12 mm long, 5-7 mm wide;

tube erect, funnelform, 15-25 mm long, limb erect, unilabiate, triangulate, folded, 15-25

mm long. Capsule ovoid, 2-3 cm long, valves 6-8 mm wide, horizontally striate, finely

pubescent; seeds triangulate, bilaterally winged, 6 mm long, 4 mm wide.

This species is found along the seasonally dry Pacific slope of Costa Rica and

into the Meseta Central. It occurs in similar habitats from Peru to Guatemala.

Flowering material has been collected from December through March, but it

probably blooms throughout the year. The flowers are similar to those of A.

linearifolia, a native of Haiti, and of A. bilabiata of the Greater Antilles.

BURGER: FLORA COSTARICENSIS 83

Aristolochia constricta Griseb., Abh. Konigl. Ges. Wiss. Gottingen 7:225.

1857. A. securidata Mast., Bot. Gaz. 33:256. 1902. Figure 12a.

Woody lianas, older stems with ridged, corky bark, younger stems glabrescent; pseudo-

stipules absent. Leaves with petioles 3-5 cm long; laminae ovate, 10-20 cm long, 7-13 cm

wide, long acuminate, bases deeply cordate with sinuses 2-4 cm deep, 5 primary veins

diverging from the base, glabrous above, glabrescent below. Flowers in axillary racemes;

utricle horizontal, ovoid, 1 cm long; tube erect, funnelform, 2 cm long; limb erect, uni-

labiate, 2-3 cm long. Capsules broadly ellipsoid, 5-6 cm long, valves 2 cm wide, horizon-

tally striate, glabrous, septa persistent, lattice-like, connecting the valves after dehiscence;

seed cordate, with broad lateral wings, 7 mm long, 12 mm wide.

Native to Costa Rica, northern Panama, and southern Nicaragua, growing in

evergreen forests below 1,000 m. Flowering material collected between October

and May.

This species can be recognized by its ovate, long-acuminate, deeply cordate,

glabrous leaves. The species seems to be closely related to A. maxima. The fruits

have persistent septa which connect the valves of the fruit after dehiscence,

making the fruit look like a hanging basket. Similar fruits are found in A. maxima,

A. chapmaniana, A. tonduzii, and probably A. translucida.

Aristolochia gigantea Mart. & Zucc., Nov. Gen. Sp. PI. 1:75. t. 48. 1824. A.

sylvicola Standley, J. Wash. Acad. Sci. 15:5. 1925. Figure 12b.

Woody lianas, older stems with a deeply ridged, corky bark, younger stems glabrous;

pseudostipules absent. Leaves with petioles 5-7 cm long; laminae ovate to triangular,

15-16 cm long, 11-16 cm wide, apices acuminate, bases truncate to slightly cordate, with

5 major veins diverging from the base, glabrous above, villous with white hairs below.

Flowers borne on older stems; utricle pendent, turbinate, 7 cm long, 3 cm wide; tube

strongly reflexed, funnelform, 3 cm long, limb erect, peltate, 15-17 cm long, 12-13 cm

wide, with purple-brown markings between the veins. Capsule cylindric, 13 cm long,

valves 8-10 mm wide, glabrous, horizontally striate; seeds ovate, without wings, 7 mm

long, 5 mm wide.

Currently known from the wet lowland forests from Panama to Amazonian

Brazil; it is likely that this species will be found in Costa Rica. Flowering material

has been collected from September to March.

This species can be recognized by the leaves, which are villous on the lower

surface, and the broadly peltate calyx limb. The Central American material differs

from the South American material in having smaller flowers, no pseudostipules,

and white hairs on the lower leaf surface. The two groups may be distinct taxa,

but these characters are known to vary within some species. Aristolochia gigantea

is closely related to A. littoralis.

Aristolochia grandiflora Swartz, Prodr. Veg. Ind. Occ. 126. 1788. Figure 12b.

Lianas, older stems with a corky bark, younger stems glabrous; pseudostipules absent.

Leaves with petioles 10-17 cm long; laminae triangulate, 8-15 cm long, 9-17 cm wide,

apices acute, bases deeply cordate with sinuses 3-6 cm deep, glabrous. Flowers solitary on

older stems, fetid; utricle pendent, narrowly ellipsoid, 9-11 cm long, with prominent

veins, tube strongly bent, 2-3 cm long, bullate at base of limb, limb funnelform, pendent,

15-20 cm long, with a long caudate appendage to 1 m long at its apex, veins prominent,

purple-brown on veins within. Capsule cylindric, 10 cm long; seeds triangulate.

Common in wet lowland forests below 1,000 m along both coasts. Plants have

been cultivated in San Jose. The species is found from Mexico to Panama and

throughout the West Indies.

84 FIELDIANA: BOTANY

The leaves of this species resemble those of A. veraguensis, but are not varie-

gated. The flowers are among the largest of any neotropical plant and are ex-

tremely fetid. The flowers are distinctive because of their size and because of the

long apical appendage.

Aristolochia littoralis Parodi, Anales Soc. Ci. Argent. 5:155. 1878. A. elegans

Mast., Card. Chron. n.s. 34:301. t. 61. 1885.

Lianas, older stem with a ridged bark, younger stems glabrous; pseudostipules rounded,

clasping, 1.5-2.5 cm wide. Leaves glabrous, petioles 3-5 cm long; laminae triangulate,

broader than long, 3-5 cm long, 4-6 cm wide, apex rounded, base cordate with a sinus 1

cm deep, 5 primary nerves diverging from the base. Flowers solitary in leaf axils; utricle

horizontal, oblong, 3.5 cm long, tube erect, 3 cm long, limb erect, peltate, 10 cm long.

Capsule ellipsoid, 4-5 cm long, with a 6 mm long beak at its apex, valves 4-5 mm wide,

horizontally striate, glabrous; seeds triangulate, slightly winged, 5-6 mm long, 4-5 mm

wide.

Native to South America, widely cultivated and escaped in Central America.

The species flowers through most of the year and appears to set viable seed

without insect pollinators.

This species can be recognized by the triangulate leaves with pseudostipules

and the peltate calyx limb. It is similar to A. gigantea in the structure of the

flowers, but the flowers of A. gigantea are much larger. The fruit is distinctive due

to the short beak at its apex.

Aristolochia maxima Jacq., Enum. Syst. PI. 30. 1760. Howardia hoffmanni

Klotzsch, Aristoloch. Berl. Herb. 621. 1859. A. maxima var. angustifolia Duchartre,

in DC., Prodr. 15(1):457. 1864. Figure 12a.

Woody lianas, older stems with a deeply ridged, corky bark, younger stems glabrescent

with strigose hairs; pseudostipules absent. Leaves with petioles 1-2 cm long; laminae

oblong to obovate, 7-15 cm long, 3-7 cm wide, apices acuminate, base truncate to slightly

cordate, glabrous above, glabrescent beneath. Flowers in loosely branched rhipidia from

the axils of leaves or from the base of the stem; ovary and pedicel 3 cm long, utricle

horizontal, ovoid, 2-3 cm long, 1 cm wide, tube bent, funnelform, 2 cm long, limb erect,

unilabiate, ovate, slightly hooded, 3-5 cm long, purple-brown. Capsule ovoid, 10-15 cm

long, valves 2 cm wide, glabrous, held together after dehiscence by persistent, lattice-like

septa; seed triangulate, winged, 1 cm long, 1.5 cm wide.

In thickets and forests along both coasts below 1,000 m and rarely found in the

Meseta Central, where it may have escaped from cultivation. The species is found

throughout Central America and into Colombia and Venezuela.

The species can be recognized by the oblong leaves which are glabrous when

mature and which usually have truncate bases. The inflorescences are unusual in

the genus, not only because of their branching pattern, but also because they may

appear from the base of the plant. The species is closely related to A. bicolor of

Colombia. Pittier reported that the young fruits are edible.

Aristolochia odoratissima L., Sp. PI. 1362. 1763. A. pandurata Jacq., PI. Hort.

Schoenbr. 4:49. t. 497. 1804. Figure 12b.

Lianas, glabrous; pseudostipules rarely present, 1-2 cm wide, clasping. Leaves with

petioles 4 cm long; laminae hastate-triangulate, 10-15 cm long, 8-10 cm wide, apices

acute, bases cordate with a sinus 5-15 mm deep, with 3 major veins diverging from the

base, glabrous. Flowers solitary in leaf axils; ovary and pedicel 6-7 cm long, utricle

pendent, ovoid, 2-3 cm long, 1 cm wide, tube erect, 1-2 cm long, expanding slightly

BURGER: FLORA COSTARICENSIS 85

above, limb erect, unilabiate, peltate, 5-10 cm long, 3-5 cm wide, acuminate, dark purple-

brown. Capsule narrowly cylindric, curved, valves 5 mm wide, glabrous; seeds rhombic

to triangulate, without wings, 3 mm long, 2 mm wide.

Widely distributed in wet forests throughout the neotropics. This species shows

considerable geographic variation throughout its range and has been divided into

many species in South America. In Costa Rica, plants have been collected near

Pto. Limon where they seem to bloom throughout the year.

Aristolochia odoratissima can generally be recognized by the hastate leaves, but in

cases where the leaf shape is variable, the peltate and acuminate calyx limb is

distinctive. It is related to the other Aristolochia species with peltate calyx limbs,

including A. gigantea and A. littoralis.

Aristolochia pilosa H.B.K., Nov. Gen. & Sp. 2:146. t. 113. 1817. Hawardia

costaricensis Klotzsch, Aristoloch. Berl. Herb. 614. 1859. A. costaricensis (FQotzsch)

Duchartre, in DC., Prodr. 15(1):450. 1864. A. costaricensis var. zamorensis Heiron.,

Bot. Jahrb. Syst. 20(Beibl. 49):5. 1895. Figure 12b.

Lianas, all parts hirsute with brown, septate hairs, older stems with a deeply ridged,

corky bark; pseudostipules absent. Leaves with petioles 3-7 cm long; laminae deltoid to

ovate, 10-14 cm long, 8-10 cm wide, apices acute to rounded, margins ciliate, bases deeply

cordate with sinuses 1-3 cm deep, glabrous above, hirsute on veins and white pubescent

on areolae below. Flowers solitary in leaf axils; pedicel and ovary to 6 cm long, utricle

horizontal, ovoid, 15-20 mm long, 10-12 mm wide, tube erect, 15-25 mm long, flaring

slightly at mouth, limb erect, unilabiate, ligulate, 30-40 mm long, 15-25 mm wide, fim-

briate to efimbriate, maculate. Capsules broadly ellipsoid, 5-7 cm long, valves 5 mm wide;

seeds obovate, acute, without wings, 4 mm long, 3 mm wide.

Plants of the wet lowlands on the Atlantic coast and the Osa Peninsula, from

sea level to 850 m elevation. Probably blooming throughout the year. This species

is distributed in wet forests from southern Mexico to the Amazon.

Aristolochia pilosa is easily recognized by the conspicuous brown pubescence on

most of the plant; it is variable in leaf shape and in the pigmentation and vestiture

of the limb. Closer study of the species throughout its range may show it to be a

group of closely related species.

Aristolochia ringens Vahl, Symb. Bot. 3:93. 1794.

Woody lianas, older stems with a ridged, corky bark, younger stems glaucous; pseudo-

stipules rounded, clasping, 2-3 cm wide. Leaves with petioles 10-11 cm long, glaucous;

laminae broadly reniform, 9-10 cm long, 13-15 cm wide, apex rounded, base deeply

cordate with a sinus 3 cm deep, glaucous, with 7 primary veins diverging from the base.

Flowers solitary in leaf axils; utricle pendent, turbinate, 5-6 cm long, tube bent, fun-

nelform, 2-2.5 cm long, limb horizontal, bilabiate, the upper lobe spatulate, 8 cm long, the

lower lobe lanceolate, 8-9 cm long, 2-3 cm wide. Capsules cylindric, 8 cm long, glaucous;

seeds cordate, bilaterally winged, 12 mm long, 7 mm wide.

Native to South America and widely cultivated in tropical and subtropical areas.

In Costa Rica, plants have been grown in San Jose. It is the only bilabiate species

known from Costa Rica and can be recognized by the broad reniform leaves and

pseudostipules. The bilabiate aristolochias are a distinctive group within the ge-

nus; their taxonomy has yet to be satisfactorily worked out.

Aristolochia tonduzii Schmidt, Repert. Sp. Nov. 23:284. 1927. A. chapmaniana

Standley, Contr. Arnold Arbor. 5:60. 1933. Figure 12a.

86 FIELDIANA: BOTANY

Woody lianas, older stems with a ridged, corky bark, younger stems glabrescent, stri-

gose; pseudostipules absent. Leaves with petioles 1.5-3 cm long, strigose hairy; laminae

ovate to oblong, 10-16 cm long, 4-8 cm wide, short acuminate, bases cordate with a sinus

5-20 mm deep, glabrous above, densely strigose below. Flowers racemose on short axillary

branches; ovary and pedicel 2-3 cm long, strigose, utricle horizontal, ellipsoid, 3-4 cm

long, 1.5 cm wide, strigose, tube slightly bent, 1.5-3.3 cm long, strigose, flaring at mouth,

limb erect, unilabiate, hooded or folded back, 4-6 cm long, 1-2 cm wide, maculate on inner

surface, purple. Capsule ovoid, 10 cm long, valves 2 cm wide, held together after de-

hiscence by persistent, lattice-like septa; seeds deltoid, cordate, 9 mm long, 15 mm wide,

winged.

This species is found in the wet lowland forests of the Atlantic coast. Flowering

specimens have been collected from April through July, and fruiting specimens

have been collected throughout the year.

The cordate leaf base and strigose lower leaf surface distinguish A. tonduzii from

A. maxima. The flowers of A. tonduzii are also larger, and the inflorescence is not

as strongly branched. Aristolochia tonduzii is very closely related to A. schmidtiana,

native to Amazonian Peru.

Aristolochia translucida H. W. Pfeifer, Brittonia 28(3):349. 1976. Figure 12a.

Woody lianas, older stems with a ridged, corky bark, young stems strigose; pseudo-

stipules absent. Leaves with petioles 2 cm long; laminae oblong-ovate, 10 cm long, 5 cm

wide, apices slightly acuminate, bases cordate with sinuses to 1 cm deep, with 3 major

veins diverging from the base, glabrous above, strigose below. Flowers on short axillary

branches; pedicel and ovary to 4 cm long, utricle horizontal, ovoid, 4 cm long, tube erect,

funnelform, 2 cm long, limb unilabiate, hooded, translucent, 8-9 cm long, with elevated

green maculae within. Fruit and seed unknown.

Known only from the type collection, found growing at "La Selva" in the

Caribbean lowlands near Puerto Viejo de Sarapiqui. Living material was collected

without flowers and bloomed later in a northern greenhouse. Aristolochia trans-

lucida is very similar to A. tonduzii, differing chiefly in the pigmentation of the

flowers. More collections are needed to determine the variability and range of this

species.

Aristolochia trilobataL., Sp. PI. 960. 1753. Figure 12b.

Woody lianas, young stems glabrous; pseudostipules rounded, clasping, 1-2.5 cm wide.

Leaves with petioles 3 cm long; laminae 3-lobed, 7-9 cm long, 7-11 cm wide overall, the

middle lobe 2.5-3.5 cm wide, apex acute to rounded, base truncate to slightly cordate,

glabrous. Flowers solitary in leaf axils; ovary and pedicel to 6 cm long, utricle horizontal,

ovoid, 3 cm long, with a 6-lobed, starlike hypanthium at its base, tube bent, erect, 2 cm

long, limb unilabiate, triangulate, 1.5 cm wide, with a long, pendent, caudate apex, over

15 cm long. Capsule ovoid, 5 cm long, valves 1.5-2 cm wide, glabrous, horizontally striate;

seeds triangulate, bilaterally winged, 8 mm wide, 8 mm long.

A circumcaribbean species found in lowland woods and thickets on the Atlantic

coast. In Costa Rica, the species has been found near Limon. This is the only Costa

Rican species which has lobed leaves. Its relationships are obscure, but it is most

likely not closely related to the other Caribbean species with lobed leaves.

Aristolochia veraguensis Duchartre, in DC., Prodr. 15(1):458. 1864. Figure 12b.

Woody lianas, older stems with ridged, corky bark, to 10 cm diameter, younger stems

glabrous; pseudostipules absent. Leaves with petioles 10-12 cm long; laminae variegated

with white or yellow along veins, triangulate, 13-17 cm long, 13-15 cm wide, acuminate,

deeply cordate, with a sinus 3-4 cm deep, glabrous. Flowers in dense, racemose clusters

BURGER: FLORA COSTARICENSIS 87

on older stems; ovary and pedicel 5-6 cm long; utricle pendent, ovoid, 1.5-2 cm long, 1

cm wide, tube strongly bent, 1-1.5 cm long, limb funnelform, 2.5-3.5 cm long, acute,

yellow with purple marking around the edge. Capsule cylindric, 14-20 cm long, valves 5

mm wide, glabrous; seeds ovate, without wings, 3 mm long, 1 mm wide.

Plants of the seasonally dry, lowland forest in Guanacaste, Puntarenas, and San

Jose. Generally found below 1,000 m, but cultivated at higher elevations. Also

known from Panama. The plants bloom from December to May. This species is

very distinctive due to its variegated leaves. It is probably related to A. schippii of

southern Mexico and Belize.

HYDNORACEAE

By Luis D. Gomez

Parasitic plants on roots of the host, lacking chlorophyll, stems all subterranean, lacking

leaves and roots, underground stem (rhizome) producing terete or 3- to 6-angled rhizoids

with numerous, elongated and wartlike protuberances. Flowers solitary or groups borne

on the rhizome and usually with only the distal part of the flower above the gound, large

and bisexual, thick-fleshy, perianth of 1 whorl of 3 or 4 valvate sepals (tepals), the fleshy

perianth united below to form a thick tube and opening distally, the perianth lobes either

opening and diverging or remaining united at their tips and opening below to form a

lantern-like structure; stamens borne on the perianth tube, anthers very large and numer-

ous, united into a ringlike mass on the perianth tube or forming synandria of tightly

united undulating thecae, the synandria as many as the perianth lobes in Hydnora; ovary

inferior, of 3 fused carpels with a single locule containing many reduced ovules on placen-

tas which are leaflike or lamellate, the placentation parietal or pendulous from the apex of

the large locule, stigma broad and rounded or button-like, sessile. Fruit usually formed

underground, a large thick-walled fleshy berry; seeds minute, with copious endosperm

and a verrucose or pitted testa.

An unusual family of parasites with very large fleshy flowers that remain par-

tially underground and are pollinated by beetles. The Hydnoraceae are probably

most closely related to the Rafflesiaceae and Balanophoraceae from which they

differ in the bisexual flowers. Hydnora, with some six to 10 species in Africa and

Madagascar, and Prosopanche, with two species in southern South America and a

newly discovered species in Central America, are the only genera in the family.

PROSOPANCHE DeBary

REFERENCE: A. E. Cocucci, Estudios en el genero Prosopanche 1. Revision tax-

ondmica. Kurtziana 2:53-74. 1965.

Herbs, completely parasitic, vegetative stem hypogean, a thick mass of tissue from

which arise the flowers which are its only aerial parts and long underground 4- to 5- (6-)

angled rhizoids bearing numerous wartlike haustorial rudiments on the aristae. Leaves

absent. Flowers bisexual, perianth fused at the base to form a tube of variable length, the

upper edge of which subtends the stamens, tepals (perianth lobes) 3, thick and valvate;

stamens 3, opposite the tepals, filaments much reduced, more or less deltoid and fused to

the perianth tube on one side and with an anther at the distal end, anthers with 15-30

thecae concrescent to form a conical or ovoid synandrium, dehiscing longitudinally, ex-

trorse; an inner whorl of 3 bilobed interstaminal appendages alternating with the stamens;

pistil 1 and united with the perianth tube, ovary inferior, 3 carpellate, the solitary locule

completely filled by numerous lamellar parietal placentae the upper ends of which are

united to form a button-like stigmatic surface which plugs the upper end of the locule, the

ovules very reduced, embedded in the placental walls, absent near the stigmatic surface.

Fruit a fleshy berry; seeds numerous, small, with pitted or verrucose episperm, dark

brown or black at maturity, very hard.

FIG. 13. Hydnoraceae. Prosopanche costaricensis; vertical scale at the right is 5 cm high.

BURGER: FLORA COSTARICENSIS 89

The genus is represented by P. americana (R. Br.) Baillon, which is parasitic on

Leguminosae, mainly Prosopis, and P. bonacinae Spegazzini, which parasitizes

several dicotyledonous families. Both species are found only in the drier parts

of southern South America. Prosopanche costaricensis is the only species known

from Central America and northern South America.

Prosopanche costaricensis Gomez & Gomez-L., Phytologia 49:53. 1981. Fig-

ure 13.

Thick-fleshy herbs, found in clumps forming extensive colonies, parasitizing the roots

of leguminous trees, rhizoids 4- or 5-angular, up to 15 mm thick, with numerous fragile

haustorial rudiments. Flowers pedicellate, pedicels up to 150 mm long and 15 mm thick,

the flowers averaging ca. 6 cm long and 2 cm thick when in bud, sepals 3, up to 90 mm

long and 35-42 mm broad, externally flocose or with lenticel-like warts, inside smooth and

white, valvate and remaining joined at the tip with the perianth segments forming a

narrow lantern-like structure when open, perianth tube short (ca. 7 mm), the exterior

diameter 25-32 mm; synandrium conical, 19-23 (29) mm long, 11-22 mm broad at the

widest point, anthers white when young, dark brown or black when mature, thecae ca.

25, dehiscing longitudinally; ovary ca. 50-60 mm long and 40-53 mm in external diameter

when mature, also flocose-warty on the exterior, stigmatic surface 3-radiate with fewer

than 12 pairs of lamellae lobes in frontal view. Seeds elliptic, 1-1.3 mm long, 0.6-0.7 mm

wide, buff when young, black in maturity, testa rugulose-verrucose.

This species is known only from cacao groves near Siquirres at about 50 m

elevation at the edge of the Caribbean coastal plain in evergreen wet forest for-

mations. The plants are parasitic on Inga orstediana and possibly Gliricidia saepium.

Unlike its South American congeners, this plant inhabits a tropical rain forest

region. Its closest affinity is with P. americana from which it differs in various

aspects, particularly size of structures, seed ornamentation, and folds or lobes of

the stigmatic surface.

Editor's note: The collection of Prosopancheby Luis D. Gomez and Jorge Gomez -

Laurito in June 1981 was an exciting find. This is not only a major geographical

disjunction for the genus, but a new ecological niche. The brown warty flowers

are easily mistaken for stems and roots on the forest floor. Surprisingly, an earlier

collection from Costa Rica in Field Museum's fruit collection had been overlooked.

This specimen was collected by W. R. Hatch on July 22, 1937, identified as Corynea

crassa, and annotated as Prosopanche by Bertel Hansen in 1976. The only informa-

tion this collection gives us is the word Reventazon, a river that descends the

highlands and flows close by Siquirres.

RAFFLESIACEAE

By Luis D. Gomez

REFERENCES: J. Kuijt, The Biology of Parasitic Flowering Plants, Univ. of Califor-

nia Press, 1969; I. Vattimo, Contribuicao ao conhecimento da tribo Apodanthea R.

Br. Parte 1, Conspecto das Especies. Rodriguezia 26, no. 38:37-62, 1971.

Herbaceous parasites, without chlorophyll and without roots, stems short and fleshy or

absent and flowers emerging directly from the tissues of the host, borne on the aerial

woody stems or on the subterranean roots of the host. Leaves absent or represented by

bracts or scales. Inflorescences racemose or the flowers borne directly from the stems or the

roots of the host, neotropical genera with small (less than 1 cm) to medium size (up to 3.5

cm in diameter) flowers (Rafflesia with the world's largest flower, ca. 1 m in diameter).

Flowers radially symmetric, usually unisexual by abortion, rarely bisexual, perianth of

4-10 imbricate (rarely valvate) or spreading segments or calyx lobes, united at the base to

90 FIELDIANA: BOTANY

form a short tube; stamens 5 to many, filaments united into a tube, column or synandrium,

anthers sessile, arranged in 1-3 whorls around the apex of the column or staminal tube,

1- or 2-thecous, opening by a longitudinal slit, pollen decussate, acolpate or 3- or

4-colpate, without sculpturing, sometimes shed in tetrads; pistil superior to inferior,

carpels 4, 6, or 8, ovary always unilocular, placentae 4, 6-20, or more, placentation parietal

over the inner wall or toward the distal (apical) portion of the locule (in transection the

locule appears circular, cruciform, or lamellate, with many parietal projections from the

wall), ovules very numerous and sessile, stigma solitary, terminal on a slender style or

stout column, capitate to discoidal, undivided or obscurely lobate. Fruit a fleshy berry with

many seeds, opening irregularly, often concrescent in Bdallophyton; seeds very small, with

a hard testa, often with few (8) cells rich in fatty substances.

This unusual family of parasites, containing about nine genera and 50 species,

is well represented in the Old World, with the best known examples being Cytinus

and Rafflesia. In the neotropics, the family is represented by the genera Apodanthes

and Pilostyles of the Apodantheae, and Bdallophyton and Mitrostemon of the Cy-

tineae. Bdallophyton is the only genus endemic to the American tropics. The

biology of the family, which is little known, has been summarized by Kuijt (1969).

Plants of Apodanthes and Pilostyles are easily passed by since they appear only as

small flowers on the thicker stems of the host, often only 5 mm high.

la Parasites on aerial branches of Flacourtiaceae or Leguminosae; flowers solitary and

terminal, subtended by a short peduncle covered by closely appressed bracts, borne

directly on the stem of the host, often with scars of previous flowers evident on the

bark of the host in the immediate vicinity; Apodantheae 2a

Ib Parasites on the horizontal roots of Burseraceae or Fagaceae; flowers several to many

in racemose inflorescences; stems evident and single or branched above the tissue of

the host, stems with leathery foliose or triangular bracts; Cytineae 3a

2a Locule of ovary cruciform in transection, placentation restricted to the swollen

longitudinal ribs; flowers subtended by 4 alternate bracts, exserted, 4-segmented,

subepigynous, not persisting; parasites on Flacourtiaceae Apodanthes

2b Locule of ovary evenly circular in transection with placentation on the entire inner

surface; flowers subtended by a wide base covered with imbricate bracts and

concealed, parasites on Leguminosae Pilostyles

3a Plants large, up to 25 cm tall, stems well developed and with many foliose bracts,

these spirally arranged but often with the internodes so reduced as to produce bracts

in pseudowhorls; flowers axillary on a racemose axis; ovary inferior; parasites on the

roots of Burseraceae, Ficus and Cochlospermum Bdallophyton

3b Plants smaller, usually 4-8 cm tall; stems not well developed and often concealed by

many bracts; ovary superior or subinferior; parasitic on roots of Quercus and perhaps

Carpinus (Betulaceae) Mitrastemon

APODANTHES Poiteau

Very small plants parasitic on the aerial stems of the host (almost always species of

Flacourtiaceae, especially Casearia), aerial stems very short (less than 5 mm), leaves re-

duced to small scales, only the cupular stem apex with whorled scale leaves and flower or

fruit visible on the stem surface of the host. Flowers solitary from a woody cupule borne

on the bark of the host (but flowers usually occurring in groups on the surface of the host),

the cuplike stem apex with 3 whorls of imbricate scales, the 1st whorl with 2 free scale

leaves, the 2nd with 4 scales united at the base, and the 3rd whorl with thinner rounded

petal-like parts that are deciduous. Flowers unisexual, small, radially symmetrical, the

male flowers with a central "stylar " column broadened and stigma-like at the top beneath

which 2 series of many (ca. 20) subsessile stamens are attached; female flowers with a

partly inferior ovary, 1-locular with many ovules, stylar column cylindrical and narrow

above, stigma annular. Fruit baccate, subtended by the woody and scaly cup; seeds with

hard testa.

Seven species have been reported for the genus, ranging from Mexico to Gua-

temala and Honduras, Panama, and northern South America. The Mexican plants

BURGER: FLORA COSTARICENSIS 91

may all be forms of A. caseariae Poit., while in South America all described

material may be referable to either A. flacourtiae Karst. or A. tribracteata Rusby.

Only one species is known from Costa Rica.

Apodanthes caseariae Poit., Ann. Sci. Nat. (Paris) 3:422. 1824. A. panamensis

Vatt., nom. nud. Figure 8.

Small, leafless, parasitic herbs, emerging from bark of trunk and branches of Flacourtia-

ceae. Flowers solitary. Pistillate flowers white, subtended by a receptacle, 1-2 mm high

and 2-3 mm wide, formed in the outer bark of the host plant; basal bracts 2, suborbicular

cucullate, leathery, 2-3 mm long; the upper bracts 4, apically rounded, almost twice as

long as the basal ones, fused to the lower l/2 or Vs of the ovary and pressed against the

ovary; calyx segments 4, obovate, imbricate, slightly inequilateral, borne in a shallow and

small depression near the apex of the ovary, petals lacking; ovary ovoid, creamy white to

dirty yellow to orange, 3-4 mm long; ovules numerous, parietal on folds or invaginations

within the unilocular ovary, style thick, 2 mm long, dilated toward the apex, stigma finely

papillate. Staminate flowers with a short stylar column which subtends 2 series of many

stamens each, stamens almost sessile. Fruit a berry.

Only known from the General Valley where it grows abundantly on Caesearia

nitida. The most evident stage is the fruit, which becomes yellow or orangish in

maturity and is eaten by Hnaupis birds. The flowers are visited by Trigona bees.

These parasites have been reported as growing on Burseraceae and Meliaceae, in

addition to Flacourtiaceae, in northern Central America. On Barro Colorado Is-

land, Panama, these parasites have been seen only in August and September.

BDALLOPHYTON Eichler

This is the only genus of the Rafflesiaceae endemic to the neotropics. The plants

are parasitic on the roots of Burseraceae, fleshy, with well-developed stems bear-

ing lanceolate, acute, foliose bracts. Racemose inflorescence often producing con-

crescent fruit. Two species have been described: B. americanum (A. Br.) Harms and

B. oxylepis (B. L. Robinson) Harms, the latter of which differs from the former only

in size of plants, which are slightly smaller, and in the reduced internodes of the

infructescence which give it the appearance of a small pineapple. I consider the

genus as monotypic.

Bdallophyton americanum (A. Br.) Harms, in Engler & Prantl, Nat. Pflanzen-

fam. ed. 2, 17b:281. 1935, based on Cytinus americanum R. Br., Trans. Linn. Soc.

(London) 19:246. 1845. B. andreuxiiEichl, Bot. Zeitung. (Berlin) 39:715, t. 8B. 1872.

B. cerantantherum Eichl. loc. cit. t. 8A, figs. 1-5. B. bambusarum (Liebm.) Harms,

loc. cit. B. oxylepis (B. L. Robinson) Harms, loc. cit. Figure 14.

With characters of the genus and those given in the generic key.

Plants when alive a drab purplish olive green, the pigment dissolving in alcohol

leaving all parts an opaque olive brown. Anthers sometimes relatively short and

swollen or elongate and rather narrow (4-5 x as long as wide), horny projections

straight in the immature flowers, curving outward just before dehiscence of the

pollen. Pollination carried out by flies, apparently attracted only by the morbid

color since no odor was perceived at the time of observation. The seed are carried

around by rodents, the ripe fruit often showing the signs of sharp incisors.

In Costa Rica, the plants are known only from one locality, the riparian forest

along the lower course of Rio Calera, Parque Nacional Sta. Rosa, Guanacaste

Province. Blooms in the months of October to early December.

Bdallophyton

americanum (A.Br.) Harms

FIG. 14. Rafflesiaceae. A— H, Bdallophyton americanum: A, habit of female plant showing

concrescent fruits; B, polar view of 4-colpate pollen grain; C, discoid and lobate stigma; D,

E, types of anthers and anther appendages; F, young shoot covered by imbricate bracts; G,

female flower; H, diagrammatic view of transection of ovary.

92

BURGER: FLORA COSTARICENSIS 93

PILOSTYLES Guillemin

No representatives are so far recorded from Costa Rica, where they may occur

on the Leguminosae of the tropical dry forest and at middle elevations of the

northern mountain ranges of the country. Pilostyles thurberi A. Gray and P. globosa

S. Watts are known from northern Central America. Pilostyles covillei Rose is

described from Colombia. Like Apodanthes, these plants appear as small (5 mm)

projections or pustules on the branches of the host. These projections give rise to

the flowers which are often found in considerable number and give the parasitized

branch a peculiar warty appearance.

MITRASTEMON Makino

Originally described under the name Mitrastemma, Makino later not only

changed the generic name to its present form, but also transferred the genus from

Rafflesiaceae to Mitrostemonaceae (in Bot. Mag. [Tokyo] 25:253, 1911), proposing

also a separate order, Mitrostemonales. Unlike all other Rafflesiaceae, Mitrostemon

has a subinferior to superior ovary. In other characters, it conforms with the

family concept. One species, M. matudai Yamamoto, in Matuda (Bull. Torrey Bot.

Club 74:133, 1947), is described and illustrated from Mexico and Guatemala,

where it is a parasite on the roots of Quercus (Fagaceae). It might possibly occur

in Costa Rica in the same habitats as Conopholis (Orobanchaceae), also parasitic on

Quercus and ranging from the central portions of Mexico southward into northern

South America. Mitrastemon matudai is the only species in the New World; the

others are from Japan, Formosa, and southeastern Asia to Borneo.

BALANOPHORACEAE

By Luis D. Gomez

REFERENCE: B. Hansen, Balanophoraceae, Flora Neotropica, Monograph

23:1-80, 1980.

Fleshy parasitic herbs devoid of chlorophyll, consisting of a cylindrical tuberous rhi-

zome, long-creeping and then dichotomously branching or globose and massive, epigeous

or hypogeus, the surfaces tomentose, squamose or naked, erect stems represented by the

inflorescence peduncles or absent. Leaves reduced to scales present only on the peduncles

of the spadices in imbricate spirals, transformed into hexagonal peltate scales which form

a crustose covering of the spadix and fall before anthesis, or reduced to an annulus of

leathery laciniate tissue. Inflorescences spadix-like, short to long-pedunculate, flowering

portion globose to cylindrical or clavate. Flowers small and densely crowded on the surface

of the spadix (capitulum) or intermixed with a dense mat of filiform paraphyses (hairs),

unisexual or bisexual. Flowers small and radially symmetrical, unisexual, male flowers

naked or with a 3- to 8-lobed valvate perianth, stamens generally equal in number to the

perianth lobes and opposite them, filaments free or connate to form a tubular column,

anthers free or united, usually 2-thecous, attached by their basal or dorsal surface, de-

hiscing by a lateral longitudinal slit or by an anterior-apical fissurate pore. Female flowers

lacking a perianth or the perianth reduced and adnate to the ovary, the free limb small,

truncate, 2-lobed or tublar, ovary globose-ellipsoid to prismatic-obovoid, 1- to 3-locular,

styles terminal, 1 or 2, filiform and elongate or short and subclavate, the stigmas simple

and discoid to capitellate. Fruit small 1-seeded achenes or slightly fleshy and drupelike;

seeds globose or compressed, testa thin or lacking, endosperm rich in fatty substances.

A family of about 18 genera and twice as many species in urgent need of more

intensive study. The relationship of the Balanophoraceae to other families of the

Santalalean alliance is at best obscure and likely to prove unnatural. The alliance

FIG. 15. Balanophoraceae. 1—4, Langsdorffia hypogaea: 1 & 2, inflorescences; 3, female

flower; 4, male flower. 5- 9, Helosis cayennensis: 5, inflorescence; 6, peltate scale; 7, female

flower, lateral view; 8, female flower, frontal view; 9, male flower. 10— 14, Corynaea crassa:

10, inflorescence; 11, male flower; 12, peltate scale; 13, paraphysate trichomes, uniseriate

and crotaliform apices; 14, female flowers. 15 & 16, Corynaea sphaerica: 15, inflorescence; 16,

peltate scale. Refer to Figure 16 also.

94

Fie. 16. Balanophoraceae: see Figure 15 for explanation.

95

96 FIELDIANA: BOTANY

of this family may be with the Gunneraceae. These unusual parasitic plants are

represented by three genera and perhaps four species in Costa Rica.

la Peduncle of the inflorescence ensheathed by many stiff imbricate, lanceolate scales to

3 cm long; rhizome elongate and branched, tomentose or velutinous; flowers not

imbedded in a mat of filiform hairs; style 1 Langsdorffia

Ib Peduncle of the inflorescence not covered by large imbricate scales, peduncle of

variable length or the inflorescence almost sessile, always covered by a crustose layer

of small peltate scales caducous at anthesis; flowers imbedded in a dense mat of

filiform hairs (paraphyses); styles 2 2a

2a Perianth of male flowers with 3-6 tepals, connate in the lower half to form a tube and

with the lobes always reflexed; stamens free at the apex but connate in the lower half

(the apex often held together as a unit by an adhesive mucus of the anthers), anthers

with 3 pollen sacs, 1 anterior and 2 posterior Helosis

2b Perianth of male flowers completely connate or the parts free in the upper half, the

perianth lobes never reflexed but the margins always somewhat crenate; stamens

connate in their entire length to form a tube, anthers with 2 lateral pollen sacs

Corynaea

CORYNAEA Hooker f.

Herbaceous root parasites, rhizomes nodular, globose or very rarely elongate, varying

in size from a few centimeters to 35-40 cm in diameter. Inflorescence usually monoecious

and proterogynous, rarely dioecious, peduncle emerging from an annular basal volva

which disintegrates quickly, glabrous except for occasional annular appendages, capitu-

lum or spadix globose to cylindrical or claviform, before anthesis covered by a compact

layer of 6-sided, peltate and umbonate scales under which a substrate of dense filiform

paraphyses surround the flowers, distal peltate portion of the scales caducous at anthesis,

the pedicels of the scales forming areolae with male flowers along edges and corners of the

areolae and female flowers within the areolae around the pedicles of the hexagonal (but

deciduous) scales. Male flowers with a tubular or subcampanulate perianth, entirely con-

nate or free in the upper half, perianth lobes 3, always crenate and never reflexed; stamens

3, exserted, filaments united into a column, anthers each with 2 lateral pollen sacs and

forming a synandrium of 6 thecae; pollen 3- (5-) colpate, globose; some male flowers have

a much-reduced pistillode. Female flowers without an apparent perianth or the perianth

adnate to the pistil and reduced to a low epigynous collar with 2 inconspicuous segments;

pistil 2-carpellate, ovary compressed, ovules 2, of which only 1 matures, styles 2, filiform

and divergent, deciduous, stigmas slightly capitate. Fruit a 1-seeded achene with a hard

epicarp and endocarp.

A genus of yellowish brown to deep purple root parasites that have not been

found below 1,000 m elevation. Hansen (1980) considers the genus to be com-

posed of a single species, Corynea crassa, but Costa Rican material does appear to

fall into two species.

la Inflorescences pinkish in color, capitulum globose to ellipsoid or fusiform in

shape C. crassa

Ib Inflorescences golden yellowish brown in color, capitulum spherical to globose-

elongate in shape C. sphaerica

Corynaea crassa Hooker f., Trans. Linn. Soc. London 22:31. t. 13. 1856. Fig-

ure 16.

Inflorescence pink by the concentration of pigments in the glandular apices of the

paraphysate trichomes which are 1.5-2.4 mm long, uni- or pluriseriate in which case the

apex is crotaliform. Capitulum globose to elliptical or fusiform, up to 12 cm long and 4 cm

in diameter, when immature covered by scales 3-4 mm in diameter, peltate on a pedicel

3-4 mm long. Staminate flowers up to 5 mm long with a 3-lobed perianth, the lobes 3-3.5

mm long and 1-1.4 mm wide, free in the upper % below which they form a tight tube of

wide base; stamens 3, connate into a thick and long staminal column, anthers clavate and

massive; pistillodes absent; pistillate flowers lacking a perianth, ovary obovoid, styles 2,

BURGER: FLORA COSTARICENSIS 97

divergent, filiform and tapering at the apex, stigmas subdiscoid-capitate. Fruit obovoid,

large (up to 1.5 mm long).

Parasitic on Quercus, Chusquea, and Vitis in the mixed forest at high elevations.

The inflorescences are visited by many flies and occasionally by Bombus bees; the

fruit are scattered by staphylinid beetles which forage on the paraphysate tri-

chomes and also rear their larvae in the outer layers of the rhizome which is of a

bright cadmium yellow when freshly cut and rapidly oxidizes into a rusty brown.

Although the rhizomes are often exposed and seem fleshy enough to attract

herbivores, they contain a bitter principle that probably detracts predation.

Corynaea sphaerica Hooker f., loc. cit. t. 14. Figure 16.

Rhizome tuberous, irregularly lobate, often velutinous and not more than 10 cm in

diameter. Inflorescences bisexual, solitary or rarely 2 per rhizome, peduncles very short

(less than 1 cm) or almost lacking, emerging from a persistent and thick annular volva,

capitulum spherical or globose-elongate, up to 6 cm in diameter, when immature covered

by spirally arranged rows of shortly peltate, wide scales, paraphysate trichomes abundant

and dense, golden brown, up to 1.5 mm long, mostly uniseriate. Staminate flowers sub-

immersed, with an obscure perianth of 3 connate lobes forming a campanulate in-

fundibulum, stamens 3, fused into a tube 2.5 x as long as the width of the perigonial

funnel, anthers and pollen like those of C. crassa; laterally compressed pistillodes some-

times present. Pistillate flowers lacking a perianth, styles 2, rather short and parallel,

stigma discoid, ovules 1. Fruit not seen.

Parasitic on Chusquea and Euterpe (?) in the high elevations of Talamanca. It

differs little from the preceding species, the most important character being the

mixed distribution of male and female flowers and the golden brown color of the

paraphysate trichomes.

HELOSIS Richard

Herbaceous root parasites, rhizomes fleshy, tuberous, emitting elongate hypogean

branches from which emerge the erect peduncles (flowering stems or spadices). In-

florescences borne on short or long peduncles that are annulate at the base or higher,

capitulum broadly ovoid or globose, at first covered with hexagonal peltate valvate scales

but these deciduous before anthesis, the inflorescences bisexual, the mixed male and

female flowers in mammillate areoles and subimmersed in a dense mat of slender clavate

paraphysate trichomes. Male flowers with a cylindrical perianth united near the base and

with 3 ovate, concave, valvate and strongly reflexed distal lobes; stamens 3, filaments

connate to form a tube up to 7 mm long, often with the filaments free in the upper third

and directed outward or the anthers remain joined and the filaments become arcuate

beneath them, anthers 3, basifixed and united into a 9-thecous synangium (each anther

3-thecous and cordate); pollen 3-colpate, ca. 30 /im in diameter. Female flowers with

superior perianth united to form a tube near the base and 2-labiate distally with 2 deltoid

obtuse lobes; ovary ellipsoid, 1-locular, styles 2, long and filiform, stigmas capitate. Fruit

a small 1-seeded achene with hard pericarp and oily endosperm.

Helosis includes only a single species, ranging from central Mexico through

Central America and northern South America to Brazil and Bolivia. The plants

range from sea level to 2,000 m elevation.

Helosis cayennensis(Sw.) Sprengel, Syst. Veg. 3:765. 1826. Cynomorium cayen-

nense Swartz, Nov. Gen. & Sp. 12. 1788. H. guyanensis Richard, Mem. Mus. Hist.

Nat. 8:416, t. 20. 1822, nom. illeg. superfl. Caldasia cayennensis (Sw.) Mutis ex

Steudel, Nom. ed. 2. 1:255. 1840. H. mexicana Liebmann, Forh. Skand. Naturf. 4,

Mote 1844:181. 1847. H. cayennensis var. mexicana (Liebm.) B. Hansen, Svensk Bot.

Tidskr. 72:188. 1978. Figures 15 and 16.

98 FIELDIANA: BOTANY

Plants whitish, light brown or ferrugineous, glabrous, arising from a coralline tuberous

rhizome. Peduncles solitary or clustered, erect, up to 10 cm long and 1 cm thick, with or

without an annular process near the base or higher, capitulum ovoid or ellipsoidal-globose

up to 6 cm long and 3 cm in diameter, before anthesis covered by peltate (stalk 2 mm)

umbonate scales 3 mm in diameter which correspond to mammilate areoles densely cov-

ered by paraphysate claviform trichomes 1-2.5 mm long and densely pigmented. Perianth

of staminate flowers a tube 2-3 mm long, 3-lobed, lobes ovate, 1-3 mm long, 0.5-1 mm

wide; stamens 3, filaments connate, free at the apex when mature, anthers forming a

synandrium 0.5-1 mm long, 0.5-1 mm in diameter. Pistillate flowers with a 2-Iabiate,

fimbriate perianth, styles 2, 1.5 mm long, caducous, stigmas capitate. Fruit up to 2.25 mm

long, ca. 0.5 mm broad.

Plants parasitic on the roots of a number of woody plants, found from the

lowlands to elevations up to 1,600 m, in Costa Rica. The plant resembles a fungus,

and like the species of Corynaea, it is called "hongo" by the local residents. Two

other species have been described under Helosis, but I cannot find consistent

differences to warrant their segregation. In Standley's Flora of Costa Rica (1937)

and in Woodson & Schery's Flora of Panama (1960), this plant is cited or described

as H. mexicana Liebmann, the apparent confusion stemming from the peculiar

behavior of the staminal tube which may be of connate filaments throughout its

entire length, have the apices free or the anthers held together, while the imme-

diate portion of the filaments are free and curved outward like a lantern (cf.

Howard, Rhodora 61:79-81, 1959; Hansen, 1980).

Helosis cayennensis has been divided into two varieties by Bertel Hansen (1980);

both of these varieties have been found in Costa Rica. The following key dis-

tinguishes them:

la Peduncle with an involucre of 2-6 minute triangular scales near the base, the length

of the peduncle divided by the length of the involucre (or annulus) to the base giving

a value greater than 2.8; plants usually found below 1,000 m var. cayennensis

Ib Peduncle with the involucre represented by a more or less oblique, slightly swollen,

ringlike mark (annulus) near the middle, the length of the peduncle giving a value of

less than 2.8 when divided by the length of the ringlike involucre to the base; plants

usually found above 1,000 m var. mexicana

LANGSDORFFIA Martius, in Eschwege

Herbaceous fleshy root parasites with creeping or upright rhizome-like tubers dichoto-

mously branched and tomentose-velutinous or glabrescent, stems (peduncles) bearing the

inflorescences unbranched, surrounded at the base by a short toothed sheath (volva),

covered by many imbricate stiff scales (scaly leaves). Leaves represented by the imbricate

scales, ovate near the base and becoming progressively more lanceolate toward the apex

surrounding the inflorescence. Inflorescences unisexual (rarely bisexual and monoecious)

the flowering capitulum globose, to ellipsoid, capitate-clavate or ovoid, the male in-

florescence taller than the female, the female often discoid or hemispherical. Male flowers

pedicellate, with a perianth of 3 (rarely 2) concave, valvate tepals; stamens as many as there

are tepals and opposite them, exserted, with short filaments connate into a column;

anthers united into a synandrium with 6 pollen sacs, 4 at the base and 2 near the apex.

Female flowers tightly congested, linear-prismatic, the perianth reduced to an obscurely

2- to 4-lobed collar, pistil solitary and the locule not apparent, ovule 1 and fused to the

ovary wall, style 1 and caducous, stigma papillate. Fruit a small 1-seeded achene or

drupaceous with a fleshy epicarp and hard endocarp.

A genus of yellowish or reddish root parasites resembling Thonningia, with one

neotropical species ranging from southern Mexico to Brazil and Bolivia and a

second, recently described, species from New Guinea. The stiff scaly covering of

the flowering stems (inflorescences) readily distinguishes this genus.

BURGER: FLORA COSTARICENSIS 99

Langsdorffia hypogaea Martius, in Eschwege, J. Brasil. 2:179, t. 5. 1818. Fig-

ure 15.

Herbaceous and thick-stemmed root parasites with the characters of the genus. Male

inflorescences with peduncles (stems) up to 7 cm long, entirely covered by scales, the larger

scales lanceolate, 4 cm long and 1 cm broad, the basal ones smaller and sparsely ciliate and

stiff, capitulum ellipsoidal, 7-8 cm long and up to 4 cm in diameter; staminate flowers on

long (10 mm) pedicels, petals 3, connate at the base to form a tube, free above with reflexed

lobes, glabrous, petals 3-4 mm long and 1.5-2 mm wide; stamens 3 with filaments entirely

connate, anthers subglobose-cordate dehiscent by slits, pollen 4-colpate, globose, smooth.

Pistillate inflorescence of smaller dimensions, the capitulum or head discoidal to sub-

globose; female flowers with a short 2-lobed perianth 1.0-1.5 mm long and up to 0.5 mm

in diameter, closely appressed to the style, the style up to 1.5 mm long, spirally twisted

and laterally compressed, stigma subcapitate and minutely papillate.

Parasitic on the roots of Ficus, Sapium, and Palmae in elevations above 1,500 m

in the Cordillera Volcanica Central. The pistillate flowers form a mat of coalescent

structures due to a resinous or waxy secretion which burns easily.

POLYGONACEAE

Herbs, shrubs, trees, or climbers, terrestrial or semiaquatic, bisexual or unisexual, gla-

brous or puberulent, stems often with thickened nodes, internodes solid or hollow; stip-

ules forming a sheathing tube (the ocrea or ochrea) that surrounds the stem, usually

deciduous and leaving a scar encircling the stem (absent in Eriogonum of N. America).

Leaves alternate and simple, in a spiral or distichous (absent in Muehlenbeckia platyclada),

petiolate, laminae entire or pinnately lobed, usually pinnately veined. Inflorescences usu-

ally an unbranched axis with alternating fascicles of flowers, less often with the flowers

solitary, axillary, or cymose, the axes racemose or spikelike but occasionally borne on a

leafless terminal twig and forming a compound paniculate arrangement, the flower fas-

cicles usually subtended by a sheathing (ocreate) bract and the flowers subtended by thin

translucent (ocreolate) bracteoles. Flowers usually radially symmetrical, bisexual or func-

tionally unisexual, usually small, often borne on articulated pedicels, the perianth 3- to

6-parted, in 1 or 2 whorls of 2, 3, or 5 parts, the inner and outer whorls often not clearly

differentiated (tepals), the outer parts often persistent or enlarging in fruit; stamens (4)

6-9, often in 2 whorls, filaments free or united at the base, borne on the base of the

perianth or on the edge of a short hypanthium, anthers 2- or 4-thecous, often versatile and

introrse; gynoecium of a solitary pistil with 1 locule and 1 basal orthotropous ovule, ovary

superior and often 3-angled or lenticular, stigmas and carpels usually 3 (2-4). Fruit an

achene with usually smooth hard surface, often tightly enclosed within the persistent and

expanded outer perianth, the perianth sometimes forming wings, hooks, or tubercles as

aids to dispersal; seed with a straight or curved embryo and mealy endosperm.

A cosmopolitan family of about 40 genera and 800 species, well represented in

the north temperate zone as well as the tropics and subtropics. The unusual

sh'pular tube leaving an encircling scar around the often thickened node is a

distinctive vegetative feature. Some Commelinaceae have similar leaf bases, but

the leaves are never pinnately veined. The small (less than 1 cm) flowers, often

borne on spikelike axes and subtended by thin scarious bracteoles, are also dis-

tinctive.

KEY TO THE GENERA OF POLYGONACEAE IN COSTA RICA

la Climbing plants with tendrils borne at the ends of inflorescences or leafless shrubs

with flat ribbonlike stems; usually found in gardens but occasionally escaped; the

stipules reduced to a thickened line around the stem at the nodes 2a

Ib Plants with leaves and without tendrils; the stipules usually forming a tube (ocrea)

around the shoot apex, the ocrea persisting or deciduous and leaving a line around

the stem. . 3a

100 FIELDIANA: BOTANY

2a Climbing plants with tendrils terminating the inflorescence; laminae cordate to

triangular; plants often growing along fences in gardens, with bright pink (or

white) sepals Antigonon

2b Shrubs or very short many-branched trees, the distal green stems 1-3 mm thick

and 1-2 cm broad with transverse lines Muehlenbeckia platyclada

3a Plants herbaceous, stems not woody except sometimes near the base; flowers bisex-

ual or less often unisexual 4a

3b Plants with woody stems, trees, shrubs, or climbers; flowers unisexual or bi-

sexual 6a

4a Leaves with palmate venation; plants grown in gardens for food Rheum

4b Venation pinnate; plants not cultivated for food 5a

5a Perianth of 2 whorls of 3 parts, the inner 3 expanding in fruit and often with

grainlike tubercles, the flowers often in verticels, filaments shorter than

the anthers, stigmas fimbriate; leaves often dimorphic with broader basal

leaves Rumex

5b Perianth of 3-5 subequal parts, not expanding in fruit; filaments longer than the

anthers, stigmas capitate; leaves rarely dimorphic (in ours); plants often found

in wet sites Polygonum

6a Achenes enclosed but not greatly exceeded by a dry or succulent perianth, the

fruiting perianth without long distal wings, perianth usually 5-parted; trees, shrubs

or climbers, without spines 7a

6b Achenes enclosed and exceeded by a thin, dry, expanded perianth, the fruiting

perianth with 3 conspicuous long thin distal wings; perianth 6- (rarely 3-) parted;

trees or spine-bearing shrubs 9a

7a Trees or large erect shrubs, often with thick stems, not growing wild above

1,000 m in our area; inflorescences with thick (more than 3 mm) axes, flowers

functionally unisexual and with a cuplike hypanthium Coccoloba

7b Climbers and scrambling shrubs, growing wild or cultivated above 1,000 m

elevation in our area; inflorescences with slender axes 8a

8a Native plants found only above 1,400 m elevation; laminae cordulate and with

more than 6 pairs of secondary veins; flowers unisexual, pedicels articulated

beneath the flowers and lacking wings Muehlenbeckia

8b Cultivated or escaped plants only known from around 1,100 m elevation;

laminae cordate and with 3-5 pairs of secondary veins; flowers bisexual, pedi-

cels with wings and articulated at the base Bilderdykia

9a Fruiting perianth with the wings expanded from the back of the midrib and de-

current on the pedicel; stems with solid internodes and short terminal spines and

with small (3 mm) rounded short shoots from which leaves and flowers arise in

fascicles; laminae less than 8 cm long and with fewer than 7 pairs of secondary veins

Podopterus

9b Fruiting perianth with the wings expanded distally and long-spatulate or

oblanceolate; stems lacking spines, flowers in racemose or paniculate arrange-

ments lOa

lOa Internodes hollow and often with biting ants; leaves with more than 12 pairs of major

secondary veins, 15-35 cm long; achene with 3 sharp ridges and 3 flat or concave

faces, tightly enclosed within the fruiting perianth Triplaris

lOb Internodes solid; leaves with fewer than 12 pairs of major secondary veins, 3-18 cm

long; schene with 3 deep longitudinal sulci separating 3 rounded sides, loosely

enclosed within the fruiting perianth Ruprechtia

ANTIGONON Endlicher

Climbing plants with tendrils borne on the ends of the inflorescences, bisexual, glabrous

or more often puberulent, stems thin and herbaceous or slightly woody; stipules reduced

and leaving a small (often obscure) thickened line around the stem. Leaves alternate in a

spiral, petiolate, laminae cordate to deltoid or sagitate, acute to acuminate at the apex and

often with the midvein extended to form a small (1-3 mm) mucronate tip. Inflorescences

basically a single racemose axis with alternating clusters of flowers, solitary and axillary

but occasionally terminal and forming paniculate arrangements by the failure of sub-

tending leaves to develop, the flower-fascicles subtended by small floral bracts. Flowers

bisexual, borne on articulated pedicels, perianth of 5 parts with the outer 3 usually broader

BURGER: FLORA COSTARICENSIS 101

than the inner 2, persisting and enlarging in fruit, stamens 8, free from the perianth, the

filaments united to form a short tube, anthers 4-Iocular, ovary trigonous, styles 3 with 3

peltate stigmas. Fruit enclosed within the persisting and enlarged perianth parts, achene

strongly 3-ridged in the upper (distal) part, surface smooth.

A genus of several poorly defined species, apparently native to Mexico and

northern Central America. The plants are now widely cultivated because of the

attractive inflorescences with long-persisting rose red or white perianth parts. The

climbing habit, tendril-bearing inflorescences, and colorful stiff dry persisting

perianth parts immediately distinguish this genus. These plants resemble the

genus Bougainvillea in a superficial way. Collections of Antigonon from Costa Rica

are few; the species have become naturalized only rarely.

la Laminae not decurrent on the petiole and not forming conspicuous lateral wings on

the upper part of the petiole; plants recorded from Costa Rica 2a

Ib Laminae decurrent on the petiole and forming conspicuous lateral wings (or margins)

on the petiole; floral bracts small (less than 4 mm) and inconspicuous; plants not

included in the descriptions 3a

2a Petioles short (less than 10 mm) and often obscured by the broad basal lobes

of the lamina; floral bracts ca. 5 mm long and ovate; outer perianth parts be-

coming 3.5 cm long in fruit, with rounded apex and overlapping cordate basal

lobes A. guatemalense

2b Petioles to 30 mm long and usually easily seen within the broad sinus of the

lamina lobes (in pressed material); floral bracts 1-3 mm long, lanceolate; outer

perianth parts rarely more than 1.5 cm long in fruit, acute at the apex and

subcordate at the base A. leptopus

3a Perianth reddish in color, outer perianth parts becoming 2 cm long in fruit, broadly

rounded in outline; staminal tube very short (?M- cinarascens Mart. & Gal.

3b Perianth whitish or yellowish, outer perianth parts becoming 1.3 cm long in fruit,

acute at the apex; staminal tube half the length of the pistil . . (?)A. flavescens Wats.

Antigonon guatemalense Meisn., in DC., Prodr. 14:184. 1856. Polygonum gran-

diflorum Bertol., Fl. Guat. 412. 1840, not P. grandiflorum Willd., 1799. A. grandi-

florum (Bertol.) Robinson, Proc. Amer. Acad. 44:513. 1909. A. macrocarpum Britton

& Small, Sci. Surv. Porto Rico and Virgin Isl. 5:266. 1924. Figure 19.

Climbers, usually in thickets and shrubs to 2 m high, leafy internodes 1.5-15 cm long,

0.5-4 mm thick, densely short (0.3 mm) brownish puberulent. Leaves with petioles 2-10

mm long, densely brownish tomentulose, often obscured by the lamina base; laminae 6-15

cm long, 3.5-10 cm broad, broadly ovate to ovate-triangular, tapering gradually to an acute

apex, cordate to cordulate at the base with the basal sinus usually narrow and obscuring

petiole and node, margin entire, laminae drying thin chartaceous, densely soft puberulent

below, venation pinnate with 3-6 pairs of major secondary veins. Inflorescences solitary

and axillary with a single axis but occasionally terminal with several axes in a paniculate

arrangement, to ca. 25 cm long, the alternating flower fascicles 5-25 mm distant on the

rachis, subtended by conspicuous broadly ovate floral bracts, ca. 5 mm long and with a

cuspidate apex, puberulent, pedicels 8-25 mm long, densely and minutely (0.2 mm)

puberulent; flowers with the perianth parts ca. 4 mm long before anthesis, staminal tube

relatively short and thick. Fruit enclosed within the persisting perianth, the outer perianth

parts becoming 3.5 cm long, cordate at the base with the basal lobes usually overlapping,

rounded to emarginate at the apex, venation prominent on the surface of the perianth, fruit

narrowly ovoid, with 3 sharply defined longitudinal ridges on the upper half, ca. 10 x 5

mm, lustrous and pale brown or greenish.

Plants cultivated in gardens or found as escapes in secondary growth between

(0) 200 and 1,500 m altitude. The species appears to be native in Guatemala and

is now widely distributed in the tropics and subtropics.

Antigonon guatemalense is recognized by the usually cordate leaf bases obscuring

the short petioles and nodes, the generally dense brownish puberulence, the

meisnenanum

hydropiperoids

10 cm hispidum

acuminatum

FIG. 17. Polygonaceae: species of Polygonum in Costa Rica.

102

RUM EX

obtusifolius

5 mm

'nepalensis

FIG. 18. Polygonaceae: species of Rumex in Costa Rica.

103

MUEHILENBECKIA

tamnifolia

ANTIGONON

leptopus

BILDERDYKIA

aubertii

FIG. 19. Polygonaceae: wild and ornamental climbing plants in the genera Antigonon,

Bilderdykia, and Muehlenbeckia.

104

PODOPTERUS

mexicanus

RUPRECHTIA

costata

1 cm

FIG. 20. Polygonaceae: trees and shrubs with winged fruits; species of Podopterus, Ru-

prechtia, and Triplaris.

105

106 FIELDIANA: BOTANY

broad little floral bracts at the base of the flower clusters, and the large pink

perianth parts enclosing the fruit. The common names Bellisima, Colacion, Confite,

Confitillo, and San Andres have been used for these plants in Central America.

Antigonon leptopus Hook. & Am., Bot. Beechey Voy. 308, pi. 69. 1839-1840.

A. cordatum Mart. & Gal., Bull. Acad. Roy. Sci. Bruxelles 10, pt. 1:14. 1843. Fig-

ure 19.

Climbers to over 10 m high but more often only 1 or 2 m high, leafy internodes 1-10 cm

long, 1-4 mm thick, sparsely to densely puberulent. Leaves with petioles 4-30 mm long,

0.5-1.5 mm thick; laminae 3-15 cm long, 2-10 cm broad, broadly ovate to sagitate, acu-

minate to acute at the apex, subcordate to cordate at the base with a usually broad open

sinus, margin entire or undulate, laminae drying thin chartaceous, sparsely to densely

puberulent beneath with small (0.1-0.3 mm) brownish hairs, venation pinnate with 5-9

pairs of major secondary veins. Inflorescences solitary and axillary with a single axis but

also terminal and forming a compound panicle, to 25 cm long, flower clusters 5-20 mm

distant on the rachis, floral bracts 1-3 mm long, lanceolate, sparsely puberulent, the

pedicels 2-10 mm long; flowers ca. 4 mm long when buds open for anthesis, staminal tube

ca. half the length of the pistil. Fruit enclosed within the persistent and enlarged perianth

parts, the perianth parts to 15 mm long and 12 mm broad (rarely? to 25 x 12 mm), acute

at the apex, subcordate at the base, very sparsely or very minutely (0.5 mm) puberulent,

the venation raised on the abaxial surfaces when dry, body of the fruit ovoid, 3-angled

with 3 longitudinal ridges near the apex, ca. 9 x 5 mm, lustrous and pale brown.

Plants cultivated in gardens or occasionally escaped and climbing over low

shrubs (especially along river banks), rarely found above 1,000 m altitude in

Central America. The species is probably native to Mexico, but is now widely

planted in warmer parts of the world.

Antigonon leptopus is recognized by the broadly cordate or subcordate leaf bases,

the inconspicuous floral bracts, the persisting perianth parts which are pink to

rose or deep rose red, and the usual association with gardens. This species is

called Bellisima, Confite, Flor de San Miguel, and San Miguel in Central America.

BILDERDYKIA Dumort

Woody or herbaceous, climbing or decumbent, slender stemmed; ocreate stipules

present and persisting. Leaves alternate in a spiral, petiolate, laminae deltate to cordate or

sagittate, glabrous or very minutely puberulent. Inflorescences solitary, axillary or termi-

nal, flowers usually in small fascicles subtended by ocreate bracts, on spikelike or raceme-

like axes that may be part of a paniculate arrangement (as in ours); flowers bisexual and

regular, borne on articulate pedicels, perianth composed of 5 parts united near the base,

the 3 outer perianth parts with longitudinal keels or wings on the midrib abaxially,

stamens 6-9 and borne on the edge of a shallow cup, filaments of 2 lengths, ovary superior

and free of the cup, styles absent, stigma capitate. Fruit a 3-angled nut, usually included

in and not exceeding the persisting perianth, perianth becoming pink and the nut black.

A small genus once included in Polygonum and native to Europe and Asia.

Bilderdykia aubertii (Henry) Moldenke, Revista Sudamer. Bot. 6:29. 1939.

Polygonum aubertii Louis Henry, Rev. Hort. 1907:82-83. 1907. P. baldschuanicum

auct., non Regel. Figure 19.

Woody climbers to 5 m high, the distal stems thin and pendent, bisexual, leafy inter-

nodes 1-10 cm long, ca. 2 mm thick, glabrous, drying pale in color and with longitudinal

ridges; ocreate stipules 2-3 mm long, thick, brown, the basal parts persisting. Leaves

alternate, petioles 1-4 cm long, slender and glabrous, with 2 adaxial ridges forming a

narrow sulcus; laminae 2-9 cm long, 1.5-5 cm broad, ovate to ovate-triangular in outline,

tapering to an acuminate apex, cordate to subcordate at the base and decurrent on the

BURGER: FLORA COSTARICENSIS 107

petiole, margin entire, drying chartaceous, smooth and essentially glabrous on both sur-

faces, venation pinnate or subpalmate with 3-5 pairs of major secondary veins. In-

florescences solitary and axillary or terminal, to 15 cm long, paniculate with alternate

racemose branches, flower-bearing rachis slender and with minutely papillate-puberulent

longitudinal ridges, pedicels ca. 2 mm long, articulate near the base and with 3 wings

enlarged distally; flowers bisexual, ca. 1.5 mm long (open) and 3 mm broad, the 3 outer

perianth parts with conspicuous wings that are continuous with the wings of the pedicel,

inner perianth parts smaller, stamens mostly 8 or 9, ovary strongly 3-angled and with large

papillate stigmas. Fruit not seen.

A trailing climber in shrubs about 2.5 m high and apparently growing wild in

the city of San Jose at 1,100 m elevation; flowering in September. We have no

other record of this genus and species in Central America.

Bilderdykia aubertii is recognized by the glabrous cordate leaves on thin hanging

stems, the small white flowers in open paniculate inflorescences, and the thin

wings on the back of the outer perianth parts decurrent on the pedicel. This

species is planted in Europe as an ornamental climber; it is native to western China

and Tibet. This species also is referred to as Fallopia aubertii (Louis Henry) J. Holub

or as Polygonum aubertii Louis Henry. English names are "Silver Lace Vine" and

"China Fleece Vine."

COCCOLOBA Patrick Browne ex Linnaeus

REFERENCES: R. A. Howard, Studies in the genus Coccoloba, VII. A synopsis and

key to the species in Mexico and Central America. J. Arnold Arbor. 40:176-220.

1959. R. A. Howard, in R. E. Woodson, Jr., and R. W. Schery, Coccoloba in the

Flora of Panama. Ann. Missouri Bot. Card. 47:340-353. 1960.

Shrubs or trees, rarely scramblers or lianas (none in Central America), usually unisexual,

leafy stems usually thick and terete, glabrous to puberulent or with glandular excretions

resembling small trichomes, often longitudinally striate, the pith solid or the internodes

sometimes hollow, the nodes usually thicker than the internodes; stipules united to form

a tube (the ocrea) around the stem, the ocrea initially enclosing and protecting the shoot

apex and then splitting as the apex develops, the distal sheathing part of the stipule

usually thin and deciduous, the basal part usually thicker and often persisting, occa-

sionally extending below the petiole base, glabrous to puberulent, leaving a scar around

the stem. Leaves alternate and simple, persistent or deciduous, often varying considerably

in size on the same plant, the leaves of juveniles and adventitious shoots often much larger

than leaves on distal branches; petiole borne at the base or above the base of the differ-

entiated stipule enclosing the node, usually canaliculate or sulcate (grooved) above (ad-

axially); laminae entire or undulate, venation usually pinnate, often quite stiff, glabrous to

puberulent or with glandular excretions. Inflorescences terminal or terminal on short

lateral shoots, unisexual, basically a single unbranched axis but occasionally branched near

the base or the axes in a paniculate arrangement on a short leafless terminal shoot, the

peduncle (flowerless base) usually very short and not differentiated from the rachis, the

rachis usually minutely puberulent and with longitudinal ridges, flowers borne singly or

in fascicles on thickened areas of the rachis, subtended usually by a stiff bract and 1 or

more thin ocreolate bracteoles, pedicels shorter or longer than the subtending bracteoles,

the female flowers usually solitary, the male flowers usually several together in a fascicle.

Flowers functionally unisexual but with organs of both sexes, perianth and staminal bases

forming a small or distinct cuplike hypanthium, perianth 5- (6-, 7-) parted, imbricate in

bud, becoming reflexed in male flowers but usually remaining appressed to the ovary and

succulent in fruit in the female, stamens usually 8, functional stamens of the male flowers

exserted, filaments free above, staminodes of the female flower included, and pistillode of

the male flowers remaining included, the functional pistil of female flowers with 3 exserted

styles and expanded stigmatic surfaces, ovary trigonous. Fruit included within the ex-

panded and usually fleshy or succulent hypanthium and/or perianth lobes (essentially a

drupe with loosely enclosed achene); the achene usually trigonous, surface smooth and

brownish in color, outer wall hard, seed with ruminate endosperm.

COCCOLOBA

caracasana

FIG. 21. Polygonaceae: two species of Coccoloba with larger rounded leaves.

108

COCCOLOBA A acapulcensis

COCCOLpBA

tuerckheimii

FIG. 22. Polygonaceae: species of Coccoloba with petioles arising above the base of the

associated tubular stipule.

109

COCCOLOBA

acuminata

FIG. 23. Polygonaceae: common species of Coccoloba with petioles arising near the base

of the associated stipules.

110

COCCOLOBA

belizensis

FIG. 24. Polygonaceae: rarely collected species of Coccoloba with petioles arising at or

above the base of the associated stipules.

Ill

112 FIELDIANA: BOTANY

A genus of the American tropics with more than 100 species, best represented

in the West Indies and South America. Our members of the genus can be recog-

nized by the shrub or tree habit, ocreate stipules at first enclosing the shoot apex

and leaving a scar around the stems, the usually thick twigs often with thickened

nodes, usually stiff entire leaves with pinnate venation, small flowers subtended

by thin tubular bracteoles and borne on a relatively thick and often unbranched

axis, and a trigonous achene tightly enclosed within a thin succulent perianth.

Papaturrois a commonly used name for members of the genus in Central America.

None of our species grow above 1,000 m elevation in Costa Rica.

la Petiole arising well above the base of the tubular stipule (ocrea), the diameter of the

petiole scar smaller than the distance from the base of the petiole scar to the differ-

entiated stipular base 2a

Ib Petiole arising at about the same level as the clearly differentiated base of the sheath-

ing stipule 6a

2a Inflorescence paniculate; distal internodes covered by the stiff persisting

ocreae, leafy stems 6-15 mm thick, laminae usually over 18 cm long and 10 cm

broad; wet evergreen forests below 1,000 m elevation C. tuerckheimii

2b Inflorescences usually solitary spikes or racemes; distal internodes usually

visible between the ocreae, leafy stems to 7 mm thick, laminae generally not

over 18 cm long; found mostly on the Pacific slope in Costa Rica 3a

3a Flowering and fruiting pedicels 2-15 mm long, the flowers clearly separate

in early stages on a rachis only 1-2 mm thick; achenes not coronate at the

apex 4a

3b Flowering and fruiting pedicels 1-4 mm long, the flowers densely congested

in early stages, flowering rachis 2-3 mm thick; achenes coronate at the

apex 5a

4a Laminae ovate to suborbicular, to 10 cm long, cordate to subcordate or

peltate near the base, petioles 12-45 mm long, 0.6-1.5 mm thick; rare

plants of the Pacific slope between ca. 500 and 1,000 m elevation

C. acapulcensis

4b Laminae oblong to obovate, to 20 cm long, sometimes rounded at the base

but never cordate to peltate, petioles to ca. 25 mm long and 1-3 mm thick

(dry), plants of dry deciduous lowland areas in Guanacaste

C. guanacastensis

5a Petioles 8-25 (60) mm long; fruit sessile or short (1-3 mm) pedicellate; laminae

elliptic to obovate, abruptly narrowed at both apex and base (but never rhom-

bic); evergreen or partly deciduous formations of the Pacific slope. C. obavata

5b Petioles 8-11 mm long; fruit clearly pedicellate; laminae usually conspicuously

and gradually narrowed to both apex and base, often rhombic in outline; low

elevations around the Caribbean but not recorded from between Honduras and

Central Panama and not included in the descriptions C. coronata

6a Inflorescence paniculate; terminal shoot at first enclosed in a spatulate or obovoid

and bluntly rounded densely tomentulose ocreate stipule 1-4 cm long; laminae

becoming 50 cm long and 25 cm broad on relatively short petioles; Caribbean

lowlands C. belizensis

6b Inflorescence a spike or raceme and these not in a paniculate arrangement; shoot

apex of terminal shoots not enclosed in a large spatulate covering 7a

7a Laminae almost as broad as long to broader than long, stiffly subcoriaceous to

coriaceous, rounded to emarginate at the apex 8a

7b Laminae usually distinctly narrower than long, usually acute to acuminate at the

apex (or rounded in C. venosa) 9a

8a Plants usually found near the sea shore but also planted for ornament; laminae

usually broader than long and with 3-6 pairs of major secondary veins,

coriaceous C. uvifera

8b Plants of seasonally very dry areas along the Pacific slope below 500 m; laminae

usually narrower than long and with 6-12 pairs of major secondary veins

C. caracasana

9a Bracts and bracteoles subtending the flowers very small (less than 1 mm) and incon-

BURGER: FLORA COSTARICENSIS 113

spicuous, the flowers usually separate along the rachis and clearly pedicellate with

the inflorescence racemose, both male and female flowers often solitary; plants

glabrous and rare in collections lOa

9b Bracts and bracteoles subtending the flowers usually more than 1 mm long, con-

spicuous and often obscuring the pedicels and the inflorescence appearing spicate

(at least in early stages), puberulent, male flowers usually borne in groups of

2-5 lla

lOa Flowers 2-4 mm long; laminae with the tertiary venation obscure above

when dry; trees of evergreen forests of the Pacific slope between sea level and

1,000 m C. padiformis

lOb Flowers 4-6 mm long; laminae with the tertiary venation forming a fine (xiO)

reticulum on the upper surface (dry); plants of the deciduous lowlands of

Guanacaste C. guanacastensis

lla Laminae usually broadest above the middle, drying very stiffly chartaceous or sub-

coriaceous; inflorescences not usually exceeding 14 cm in length, the flowers usually

closely crowded and the rachis not clearly visible; plants of the deciduous and partly

deciduous formations of the Pacific slope below 300 m elevation C. venosa

lib Laminae usually broadest at the middle (occasionally obovate in C. lehmanii), drying

chartaceous to stiffly chartaceous; inflorescences usually exceeding 14 cm with the

flowers or flower fascicles separate along the rachis 12a

12a Inflorescence racemose, pedicels usually clearly visible, bracts usually perpendicular

to the rachis; laminae broadly elliptic to elliptic-obovate, acute to short acuminate;

wet evergreen formations, 0-1,000 m C. lehmannii

I2b Inflorescence spicate, pedicels usually hidden within the bracts, the bracts at an acute

angle to the rachis; laminae narrowly elliptic and acuminate; evergreen or partly

deciduous areas, below 300 m elevation C. acuminata

Coccoloba acapulcensis Standley, Proc. Biol. Soc. Wash. 33:66-67. 1920. C.

browniana Standl., Trop. Woods 10:4. 1927. C. wercklei Standl., Publ. Field Co-

lumbian Mus., Bot. Ser. 4:304. 1929. C. cardiophylla Standl., Publ. Field Columbian

Mus., Bot. Ser. 8:8. 1930. Figure 22.

Shrubs or small trees 2-6 (8) m tall, glabrous throughout, leafy internodes 0.5-8 cm

long, 1-4 mm thick, smooth and glabrous, becoming striate, becoming dark brown and

lenticellate; stipules 1-2 cm long, originating 7-12 mm below the base of the petiole,

glabrous or very minutely puberulent, drying reddish brown and usually persisting, the

apex of the ocrea often resembling a ligulate development on the leaf base. Leaves alternate

in a spiral or distichous, often peltate in younger plants, petioles 10-45 mm long, 0.6-1.5

mm thick, smooth and drying brown, becoming articulate near the middle of the ocrea;

laminae 4-14 cm long, 2.5-11 cm broad, ovate to suborbicular, rounded distally and very

short-acuminate at the apex, rounded at the subcordate or peltate or truncate base, occa-

sionally peltate with the petiole as much as 15 mm distant from the base of the laminae,

margin entire or somewhat crenate-undulate, the edge becoming revolute, the laminae

drying chartaceous pale brown or grayish, smooth and glabrous above and below, with

4-8 pairs of major secondary veins. Inflorescences terminal on axillary short shoots, 3-8

cm long, solitary and racemose, rachis 0.5-2 mm thick, glabrous, both male and female

flowers solitary, bracts and bracteoles ca. 1 mm long; male flowers with filaments 2-3 mm

long, anthers ca. 0.5 mm long; female flowers not seen. Fruit retained within the succulent

perianth, about 2 cm long and 1.2 cm in diameter, globose or obovoid, perianth lobes only

ca. 3 mm long at the apex of the fruit, fruiting pedicels 5-10 (15) mm long; achene difficult

to remove from the perianth when dry, ca. 16 x 10 mm, rounded-trigonous.

Plants of the seasonally dry Pacific slope and known in Costa Rica from only

two collections: Werckles.n. from near El Coyolar, and Brenes 21872 from San Pedro

de San Ramon, both from the province of Alajuela between 100 and 1,000 m on

the Pacific slope. The species ranges from Guerrero and Yucatan in Mexico along

the Pacific slope of Central America to central Costa Rica.

Coccoloba acapulcensis is recognized by its relatively small rounded (sometimes

peltate) leaves on slender petioles articulated well above the stipular base, the lack

114 FIELDIANA: BOTANY

of pubescence on nearly all parts, small inflorescences with conspicuous pedicels,

and relatively large fruit. The species is very poorly known in Costa Rica, and it

may be restricted to the partly deciduous (tropical moist and premontane moist)

forest formations between 100 and 1,000 m on the Pacific slope.

Coccoloba acuminataH.B.K., Nov. Gen. 2:176. 1817. Figure 23.

Shrubs or small slender trees 1.5-5 (8) m tall, leafy internodes (0.2) 1-6 cm long, 1-3 (6)

mm thick, glabrous or brownish puberulent, usually drying dark and smooth, sparsely

lenticellate; stipules 3-15 mm long, the free distal portion of the tube truncate at the apex

and caducous, densely brownish hirsutulous or (less often) very sparsely hirsutulous or

with reddish brown glandular excretions. Leaves alternate and distichous, petioles 6-15

mm long, 0.8-1.5 mm thick, slightly sulcate above, hirsutulous or glabrous, arising from

the base of the ocrea; laminae 7—16 (24) cm long, 2-6 (8) cm broad, narrowly elliptic to

elliptic-oblong or lanceolate, gradually tapering to the long-acuminate apex, acute to

obtuse at the base, margin entire, the laminae drying thin-chartaceous to stiff-chartaceous

and often dark in color above, smooth and glabrous above or minutely papillate-

puberulent on the midvein, glabrous below except for tufts of hairs in the axils of the major

veins, often with small punctate glandular excrescences sparsely distributed over the

lower surface, with 7-15 pairs of major secondary veins, the tertiary veins very thin and

inconspicuous, the midvein raised above. Inflorescences terminal and solitary, 15-30 (45)

cm long, the flower fascicles clearly separate along the slender (0.7-1.5 mm) rachis,

peduncle 2-5 cm long, the rachis minutely (0.2-1 mm) puberulent to short-hirsute, thick-

ened below each flower cluster, pedicels shorter than the subtending bracteoles; male

flowers usually solitary, ca. 3 mm long, fertile pistil ca. 1 mm long with 2 or 3 styles. Fruit

included within the succulent perianth, becoming globose and 6-8 mm in diameter (dry),

the perianth parts united only near the base (at first) but the basal areas expanding and the

lobes finally covering Vz-Va of the fruit, translucent white at maturity and drying black,

pedicels 1-2 mm long and not exceeding the bracteoles; achene ca. 5-6 mm long and 4 mm

broad, trigonous with rounded edges, broadest near the base, dark brown to tan, smooth

and lustrous.

Plants of the lowland wet evergreen and partly deciduous forest formations

below 300 m elevation; flowering and fruiting throughout the year, but collected

most often in March and April. The species ranges from the Caribbean side of

Guatemala, Honduras, and Nicaragua through Costa Rica and Panama to Peru

and Brazil.

Coccoloba acuminata is recognized by the thin narrow acuminate leaves, long-

pendulous spicate inflorescences with the flower groups clearly separate along the

slender rachis, short pedicels hidden within the bracteoles (ocreolae) and the wet

lowland habitat. The leaves resemble those of many tropical wet forest species, but

differ strikingly from other members of the genus in our area. The species is

known only from three collections in Costa Rica: Allen 5226 from near the Rio

Terraba, Puntarenas, Holm & Iltis 710 from the Rio Frio, and Standley 40189 from

near the Rio Grande de Tarcoles, both in the province of Alajuela. Recently, new

records for Costa Rica have come from Santa Cruz, Guanacaste, near the Nic-

araguan border. Despite the paucity of Costa Rican collections, the species ap-

pears to be common in southeastern Nicaragua and central Panama.

Coccoloba belizensis Standley, Trop. Woods 16:38. 1928. C. hirsuta Standley,

Publ. Field Columbian Mus., Bot. Ser. 4:303. 1929. Figure 24.

Small to large trees, reaching 30 m in height, leafy internodes 1-8 (12) cm long, 4-8 (14)

mm thick, at first densely short (0.3 mm) hirsute but quickly becoming glabrous, (densely

puberulent to tomentulose on adventitous and young shoots), the hairs yellowish brown

to dark brown; stipules 1-4 cm long, forming an unusual spatulate cap over the shoot

BURGER: FLORA COSTA RICENSIS 115

apex, rounded at the apex and arising above the petiole base, densely brownish hir-

sutulous, splitting irregularly as the shoot expands. Leaves alternate, variable in size and

often quite large, petioles 1.5-5 cm long, 3-8 mm thick, densely brownish hirsutulous

with hairs 0.1-1 mm long, longitudinally striate and somewhat canaliculate above; lami-

nae 16-30 (50) cm long, 8-14 (30) cm broad, oblong to broadly ovate, tapering to the

usually bluntly obtuse apex, obtuse to rounded at the base and often cordulate at the

petiole in larger leaves, margin entire and revolute on drying, subcoriaceous, quickly

becoming glabrous and grayish-lustrous above, remaining minutely puberulent on the

veins beneath (but remaining densely brownish hirsutulous in some), with 6-10 pairs of

major secondary veins, the midvein raised within a depression above, tertiary veins

obscure above but prominent and often subparallel beneath. Inflorescences to 20 cm long,

paniculate terminal arrangements of nearly equal racemose or spicate branches, flower

fascicles separate on the minutely puberulent and deeply ridged rachis, pedicels not

exceeding the bracteoles; male flowers several per fascicle, filaments ca. 1 mm long,

anthers ca. 0.5 mm long; female flowers solitary. Fruit enclosed within the succulent

f>erianth ca. 7 mm long and 6 mm in diameter, the perianth lobes free for about half the

length of the fruit; achene ca. 5-6 mm long and 5 mm thick, ovoid to subglobose, weakly

trigonous with rounded edges, surfaces lustrous brown.

Trees of the Caribbean lowlands below 800 m elevation; flowering and fruiting

from May to October. The species, as presently known, ranges from Belize to

eastern Nicaragua.

Coccoloba belizensis is easily recognized because of the unusual size and form of

the ocreate stipules covering the shoot apices, the large leaves with relatively few

secondary veins, and the paniculate inflorescences. The leaves often dry with a

very smooth, but not lustrous, upper surface that is pale gray or pale yellowish

brown. While not yet recorded from Costa Rica or Panama, the species has been

collected in eastern Nicaragua (Shank 93) and is said to be common along the Rio

Grande in the Department of Zelaya (Molina 2509).

Coccoloba standleyana Allen, based on sterile material and described in The Rain

Forests of Golfo Dulce (1956, pp. 177 & 409), probably belongs under C. belizensis,

according to Howard (1959, p. 215).

Coccoloba caracasana Meisner, in DC., Prodr. 14:157. 1856. Figure 21.

Small- to medium-sized trees 2-12 (20) m tall, often with multiple trunks and rounded

crown, leafy internodes 0.5-6 (9) cm long, 2-6 (10) mm thick, minutely puberulent to

hispidulous with brownish hairs 0.1-0.8 mm long, becoming glabrous and pale gray;

stipules 10-20 (40) mm long, the distal tube membranaceous and deciduous, minutely

(0.1-0.3 mm) puberulent or with resinous excretions. Leaves alternate and usually di-

stichous, petioles 8-25 (40) mm long, 1.5-4 mm thick, inserted just above the thickened

node on the base of the ocrea, densely reddish brown puberulent or glabrescent; laminae

(5) 8-20 (37) cm long, (4) 6-15 (28) cm broad, broadly ovate to broadly oblong or

suborbicular, rounded and often emarginate at the apex, rounded and truncate to subcor-

date at the base, margin entire and slightly undulate, the edge flat, the laminae drying

stiffly chartaceous to subcoriaceous and dark brown or pale gray, smooth and glabrous

above, minutely (0.1-0.5 mm) puberulent on the veins beneath or glabrous but with hairs

in the axils of the major veins, with 6-10 (12) pairs of major secondary veins, the major and

minor veins slightly raised on the upper surface when dry. Inflorescences terminal on

short axillary shoots, 15-25 cm long, the rachis densely and minutely puberulent, 1.5-3

mm thick; male flowers 3-4 per fascicle and the fascicles often crowded on the rachis (but

the rachis usually visible), filaments ca. 2 mm long, anthers ca. 0.4 mm long; female flowers

solitary in the fascicles and the fascicles separate along the rachis, perianth lobes 1-1.5 mm

long on a hypanthium 0.5 mm long. Fruit enclosed in the white semisucculent perianth,

4-6 mm long and ca. 5 mm in diameter when dry, the perianth lobes free almost to the

base (covering % of the achene), the fruit borne on short (2-3 mm) pedicels; achenes 3-5

mm long, strongly 3-angled with 3 distinct longitudinal ridges, broadest near the base,

becoming dark brown, smooth and lustrous.

116 FIELDIANA: BOTANY

Trees of the seasonally very dry and deciduous forest formations along the

Pacific slope of Costa Rica below 500 m elevation; flowering and fruiting from

January to June. The species ranges from Chiapas, Mexico, and the Pacific slope

of Central America to Panama, Colombia, and Venezuela.

Coccoloba caracasana is recognized by the usually broad stiff leaves rounded at

both apex and base, the short pedicels that hardly exceed the bracteoles and make

the inflorescences appear to be spikes, the enlarged perianth lobes covering half

the fruit, and the seasonally dry forest habitat. There is no evidence of an associ-

ation with biting or stinging ants in our area. The inflorescences become pen-

dulous with the fleshy edible white drupes, and it is from these that the common

name Papaturro Blanco is derived.

Coccoloba guanacastensis W. Burger, Phytologia 49:387. 1981. Figure 24.

Trees to over 10 m tall, leafy inter nodes 0-3 cm long, 1.5-4 mm thick, glabrous, be-

coming pale grayish and longitudinally striate; stipules (ocreae) 4-8 mm long, the distal

tube 2-3 mm broad, thin and caducous, the basal portion persisting with the leaves and

extending below the petiole base, glabrous. Leaves often borne close together near the

ends of branches, deciduous, petioles 1.5-3 cm long, 1-2.5 mm thick, terete except near

the lamina base, glabrous, becoming articulated 2-6 mm above the differentiated base of

the ocrea; laminae (7) 9-23 cm long, 3-11 cm broad, broadly oblong to elliptic-oblong,

obtuse to rounded at the apex and often very short acuminate, obtuse to rounded at the

base, margin entire or drying undulate, the laminae drying very stiffly chartaceous and

often pale grayish green in color (the newly expanded leaves drying thin and brown),

smooth and glabrous on both surfaces, with 6-10 pairs of major secondary veins, the

tertiary venation forming a very fine (0.2-0.6 mm) reticulum that is slightly raised on both

surfaces and paler in color than the enclosed leaf areas on the lower surface (dry). In-

florescences terminal or axillary, 1 or 2 from the same node, 10-15 cm long at anthesis,

clearly racemose with the flowers usually solitary (rarely 2 together) and separate along the

thin (0.5-1 mm) glabrous rachis, the rachis longitudinally ridged but not thickened at the

pedicel bases, the subtending bracts and bracteoles less than 0.5 mm long, deciduous,

pedicels 2-4 mm long; the flowers appearing to be bisexual, 4-6 mm long and 7 mm broad,

the hypanthium narrowed for 0.5-2 mm above the articulation with the pedicel, hypan-

thium 2-3 mm long with perianth lobes 2-3 mm long, filaments 2-3 mm long, anthers ca.

0.8 mm long, stigmas 3, exserted and thickened. Fruit (immature?) partly enclosed in the

perianth base (hypanthium), perianth lobes about half the length of the developing fruit,

free and separate from the upper half of the fruit; achene apparently somewhat turbinate

with an expanded and rounded apical half.

Trees of the riparian forest in the seasonally very dry deciduous forest for-

mations below 200 m elevation on the Pacific slope of northern Costa Rica; flow-

ering from February to April. This species is known only from the province of

Guana caste in Costa Rica.

Coccoloba guanacastensis is recognized by the lack of pubescence, petioles articu-

late well above the base of the stipular sheath, minute and deciduous floral bracts,

and relatively large and long-pedicellate flowers. This species appears to be closely

related to C. padiformis and is known only from deciduous forest formations near

Canas, Bagaces, and Santa Rosa National Park.

Coccoloba lehmannii Lindau, Bot. Jahrb. Syst. 49:6-7. 1895. C. changuinolana

Standl., Publ. Field Columbian Mus., Bot. Ser. 8, pt. 1:9. 1930. C. williamsii

Standl., Field Mus. Nat. Hist., Bot. Ser. 11, pt. 5:148. 1936. C. allenii Lundell,

Contr. Univ. Michigan Herb. 6:8. 1941. Figure 23.

Small trees to 10 (rarely? 20) m tall, leafy internodes 2-7 cm long, 1.8-4.5 mm thick,

glabrous or with resin-like excretions resembling short hairs, becoming longitudinally

BURGER: FLORA COSTARICENSIS 117

striate (dry) and grayish or pale brown; stipules 5-15 (20) mm long, 1-3 mm thick, with

or without glandular excretions, arising above the petiole base, caducous. Leaves alternate

and distichous, petioles 8-15 (25) mm long, 1-3 mm thick, narrowly sulcate above (cana-

liculate) above near the lamina base, minutely puberulent, resinous punctate or glabrous;

laminae (6) 9-18 (26) cm long, (3) 4.5-9 (16) cm broad, broadly elliptic to elliptic-obovate,

abruptly narrowed to the acute or short acuminate apex (occasionally rounded distally),

obtuse or slightly rounded at the often unequal base, margin entire or slightly undulate,

drying stiffly chartaceous, smooth and glabrous above and below or minutely puberulent

on the midvein, with longer crooked hairs in the axils of major veins, with 5-10 pairs of

major secondary veins, the tertiary venation often subparallel and slightly raised on both

surfaces when dry. Inflorescences terminal and solitary, often on short axillary shoots,

8-22 (35) cm long, peduncle 1-3 cm long, flower fascicles separate along the rachis or

closely approximate but not congested, rachis 0.5-2 (3) mm thick, prominently ridged,

very minutely papillate puberulent and brownish, pedicels equaling the bracteoles in

length; male flowers in groups of 2-4, filaments to 2 mm long, pistil 1.5 mm long with 3

styles. Fruit enclosed in the succulent perianth, 5-7 mm long and 4-5 mm in diameter, the

perianth lobes free almost to the base of the fruit, borne on pedicels to 6 mm long that

clearly exceed the bracteoles; achenes 5-7 mm long, ca. 5 mm in diameter, broadest near

the base and ovate in outline, strongly trigonous, pale brown to light tan and lustrous.

Trees of the wet evergreen forests of both the Caribbean and Pacific slopes

between sea level and 1,000 m elevation; flowering collections have been made

from March to June, and fruiting collections have been made between March and

August in our area. The species ranges from central Costa Rica through Panama

and Colombia to Venezuela and Peru.

Coccoloba lehmannii is recognized by its usually broad thin leaves, the stipule

usually completely caducous to leave only a small scar on the stem, the relatively

long racemes of close or separate (but not congested) flowers, and the evergreen

lowland habitat. The species is known in Costa Rica from only three collections:

Dunlap 467 from along the Rio Sixaola valley, Lent 2464 along the Rio Parismini,

both in the province of Limon, and A. Jimenez 2444 from near Golfito in southern

Puntarenas province. An interesting collection with more narrowly obovate leaves

has recently been made by Luis Poveda (1105) near Bijagualito-Tarcoles on the

Pacific slope. This collection lacks flowers and fruit, so it is not possible to say

whether it is an unusual representative of this species or perhaps a new and

closely related species.

Coccoloba obovata H.B.K., Nov. Gen. 2:176. 1817. C. riparia Lundell, Contr.

Univ. Michigan Herb. 6:11. 1951. Figure 22.

Shrubs or trees 3-8 (15) m tall, leafy internodes 0.5-5 cm long, 2-4 (7) mm thick,

glabrous and terete, becoming grayish or reddish brown and often splitting longitudinally

when dry; stipules 5-10 (20) mm long, 3-6 mm broad, appressed and with the thin distal

tube deciduous or persisting, longitudinally striate, originating 2-8 mm below the petiole

and the lower portion usually persisting. Leaves alternate and distichous, petiole 0.8-4 (6)

cm long, 1-3 mm thick, minutely puberulent or glabrous, striate when dry, becoming

articulate 2-8 mm above the differentiated (stipular) ocrea; laminae (5) 12-22 (24) cm long,

(3) 5-14 cm broad, elliptic to elliptic-obovate or occasionally ovate, tapering or occasionally

rounded to the short acuminate apex, narrowed or rounded to the obtuse, truncate, or

cordulate base, often unequal at the petiole with 1 side or lobe larger than the other, entire,

drying chartaceous to stiffly chartaceous and usually dark above, smooth and glabrous

above and below but sometimes minutely puberulent in the axils of the major veins, with

4-9 pairs of major secondary veins, tertiary veins slightly raised on both surfaces (dry),

often with minute (0.1 mm) circular peltate scales on the lower surface. Inflorescences

solitary and terminal, on tips of branches or short axillary shoots, 8-25 cm long, spicate

or racemose and the flower fascicles densely crowded, subtended by bracteoles ca. 1.5 mm

long, rachis ca. 2 mm thick, and minutely papillate-puberulent to short pilose; male

118 FIELDIANA: BOTANY

flowers borne on pedicels not exceeding the bracteoles, filaments ca. 1 mm long, anthers

0.3-0.4 mm long; female flowers on inflorescences to 17 cm long, pedicels becoming

exserted only in fruit, ovary with 3 styles. Fruit included within the appressed succulent

perianth, globose, or somewhat turbinate, 8-10 mm in diameter, developing thick longi-

tudinal ribs, the free perianth lobes covering only the distal 2-3 mm of the fruit, borne on

pedicels to 3 mm long and 0.5 mm thick (dry); achene difficult to separate from the

enclosing perianth when dry, trigonous, tan to dark brown.

Plants of the seasonally dry partly deciduous or evergreen forest formations

between sea level and 1,000 m elevation; flowering and fruiting throughout the

year, but collected most often between June and September. The species ranges

from the Pacific side of central Costa Rica to Panama and Colombia.

Coccoloba obovata is recognized by the petiole arising well above the base of the

stipular tissue surrounding the node, the thin often obovate and glabrous leaves,

densely flowered spikes becoming racemose in fruit, and seasonally dry but

mostly evergreen habitats of the Pacific slope between the Bay of Nicoya and

Chiriqui, Panama.

Coccoloba padiformis Meisner, in DC., Prodr. 14:166. 1856. Figure 24.

Trees to 30 m tall, leafy internodes 2-6 cm long, 2-4 (6) mm thick, terete and glabrous,

becoming pale gray; stipules 6-18 mm long, 2-5 mm broad, acute at the apex, glabrous,

persisting or deciduous, becoming stiff basally, arising at about the level of the petiole

attachment. Leaves alternate and usually distichous, apparently deciduous and produced

in flushes, petioles 8-18 (26) mm long, 1-3 mm thick, glabrous, with a shallow adaxial

groove above (canaliculate), becoming articulate at or 1-2 mm above the base of the ocreate

stipule; laminae 6-18 (26) cm long, 2-8 (12) cm broad, elliptic-ovate to elliptic-oblong,

tapering or rounded to the short-acuminate apex (sometimes acute or obtuse), rounded to

obtuse at the base, the margin entire or slightly undulated, the edge becoming slightly

revolute, the laminae drying stiffly chartaceous to subcoriaceous and often grayish or

pinkish in color, smooth and glabrous on both surfaces, with 4-9 (11) pairs of major

secondary veins, arcuate ascending near the margin, tertiary venation usually flat on both

surfaces and obscure above. Inflorescences axillary (or terminal on very short axillary

shoots?), 1-3 together, 3-8 cm long, racemose, both the male and female flowers sub-

tended by very short (1 mm) bracts and bracteoles (ocreolae), rachis ca. 1 mm thick with

minute (0.2 mm) reddish hairs; male flowers 1 or 2 per fascicle, pedicels 2-3 mm long, the

hypanthium narrowed at the base (above the pedicel articulation), filaments ca. 2 mm

long, anthers ca. 0.5 mm long; female flowers solitary, pistil ca. 2 mm long. Fruit included

within the succulent perianth, ca. 9 mm long and 7 mm thick, free perianth lobes covering

only 2-3 mm at the top of the fruit, fruiting pedicels 3-5 mm long; achene ca. 8 mm long

and 6 mm in diameter, ovoid to subglobose, weakly trigonous, lustrous pale brown.

A poorly known species of evergreen or partly deciduous forests of the Pacific

below 1,000 m elevation in southern Costa Rica; collected with flowers and fruit

between February and April. The species ranges from the Golfo Dulce area of

Costa Rica to Colombia and Venezuela.

Coccoloba padiformis is recognized by the glabrous vegetative parts, stiff leaves

drying pale tan and with obscure tertiary venation (above), short racemose in-

florescences that are usually axillary, and distinctive pedicels subtended by very

small bracts and bracteoles. Included here are two collections of Paul Allen (5944

and 5964) that are said to form small stands at the heads of valleys on the summits

of high (300-800 m) hills above Palmar Norte. These trees were listed under the

name C. roseiflora Standl. & L. Wms. (never validly published) in Allen's book,

The Rain Forests of Golfo Dulce, 1956, p. 177.

Coccoloba tuerckheimii Donnell Smith, Bot. Gaz. 37:213. 1904. Figure 22.

BURGER: FLORA COSTARICENSIS 119

Trees 7-15 (20) m tall, leafy internodes 0.5-4 on long, 6-15 mm thick, often obscured

by the persisting stipules, becoming striate and glabrous; stipules 2-5 cm long, over-

lapping near the apex of the stem, to 25 mm broad, stiffly chartaceous and striate, thicker

near the base and arising below the petiole attachment, persisting with the leaves. Leaves

usually clustered near the ends of branchlets, petioles 5-45 mm long, 1.5-8 mm thick,

sulcate or canaliculate above, very minutely brownish puberulent or glabrescent, be-

coming striate or deeply ridged, borne and becoming articulate on the ocrea 6-15 (25) mm

above the stipular base; laminae (13) 20-45 (70) cm long, (7) 10-25 (40) cm broad, obovate

to broadly elliptic-oblong, abruptly narrowed at the short-acuminate apex, gradually

narrowed at the acute or rounded base, decurrent or subtruncate at the petiole, margin

entire or slightly undulate, the laminae drying stiffly chartaceous, and often dark brown,

smooth and glabrous on both surfaces or minutely puberulent beneath, the axils of the

major veins often with tufts of hairs, with 6-10 pairs of major secondary veins, tertiary

venation slightly raised beneath and not usually subparallel (between the secondaries).

Inflorescence a paniculate arrangement of 10-20 racemes of nearly equal length, terminal

on axillary short shoots, 20-40 cm long, the common peduncle and main axis less than 5

cm long, peduncles of the racemes less than 1 cm long, rachis very minutely (0.1 mm)

puberulent and with longitudinal ridges, 1-2 mm thick and drying dark brown; male

flowers in fascicles of 4-8, bracts and bracteoles 0.5 mm long or less, pedicels 2-3 mm long,

filaments 1.5-3 mm long, anthers ca. 0.5 mm long; female flowers solitary, functional pistil

3 mm long. Fruit enclosed within the succulent perianth, 11-14 mm long and 6-9 mm in

diameter, ovoid, the free perianth lobes covering only the apex of the fruit, the base

constricted into a stalk 1-2 mm long above the pedicel articulation, fruiting pedicels 1-3

mm long; achene obtusely trigonous, dark brown.

Plants of the wet evergreen forest formations below 1,000 m elevation; fertile

collections have been made in July and August. The species ranges along the

Caribbean side of Central America from Guatemala to central Panama and is

found in the evergreen forests of the Pacific slope in southern Puntarenas Prov-

ince.

Coccoloba tuerckheimii is very distinctive because of the unusually large leaves

borne on large striate ocreae enclosing the nodes on thick brownish stems and the

paniculate arrangement of long racemose inflorescence branches. Very few flow-

ering or fruiting collections have been made of this species.

Coccoloba uvifera(L.) L., Syst. Nat. ed. 10:1007. 1759. Polygonum uviferaL., Sp.

PI. 365. 1753. Figure 21.

Shrubs or short widely branching trees 2-8 (15) m tall, the trunk not usually straight,

leafy internodes 0.3-4 (10) cm long, 2.5-10 mm thick, very minutely puberulent or

sparsely glandular but quickly becoming glabrous, longitudinally striate, pale brown to

gray; stipules 3-12 mm long, beginning just beneath the petiole base, very minutely

puberulent to pilose, splitting as the shoot expands and becoming reddish brown. Leaves

usually distichous and broader than long, petioles 5-15 mm long, 3-7 mm thick, arising

just above the stipular base and becoming articulate on the ocrea, minutely papillose in

ours; laminae 6-15 (18) cm long, 8-20 (27) cm broad, reniform to broadly transversely

elliptic or suborbicular, often with the lateral halves unequal in area, rounded to emar-

ginate at the apex, rounded to the cordulate or subcordate base, 1 basal lobe often extend-

ing around the petiole, margin entire, often slightly undulate with the edge revolute, the

laminae thick, fleshy, and pale green when alive but drying pinkish gray and coriaceous,

smooth and essentially glabrous above and below but with minute (0.05 mm) hairs on the

major veins beneath and with larger hairs in the axils of the major veins, with 3-5 pairs

of major secondary veins, the tertiary venation minutely reticulate or obscure, minutely

punctate on both surfaces. Inflorescences usually solitary and terminal (rarely branched at

the base), 15-30 cm long, racemose, the rachis ca. 3 mm thick and minutely puberulent,

flower fascicles subtended by small (1-1.5 mm) bracts and bracteoles, the flower fascicles

separate and not closely crowded along the rachis; male flowers in groups of (1) 2-7,

pedicels 1-2 mm long, filaments 2-4 mm long, anthers 0.5-0.8 mm long; female flowers

solitary, perianth 3-4 mm long. Fruit included within the succulent perianth, ca. 2 cm long

120 FIELDIANA: BOTANY

and 1.5 cm in diameter, narrowed at the base above the pedicel articulation, obpyriform

to obovoid, the perianth lobes remaining free only at the top 2-4 mm of the fruit; achene

difficult to separate from the perianth when dry, black at maturity.

Plants of the sandy seashore vegetation just above and behind the high water-

line on both the Pacific and Caribbean coasts in Costa Rica; probably flowering and

fruiting throughout the year, but with very few collections made in January or in

May and June. The species ranges from Mexico and southernmost Florida

(U.S.A.) through the West Indies and Central America to northern South Amer-

ica. This species is occasionally grown in gardens as an ornamental tree.

Coccoloba uvifera is distinguished by its thick stiff leaves, usually broader than

long on very short petioles, thick stems with ocreae covering the nodes, long

racemose inflorescences, and sandy coastal (or garden) habitats. Because the spe-

cies is so characteristic of seaside vegetation and so easily recognized, it is only

occasionally collected and hence poorly represented in herbaria. The plants are

known as Uva, Uva de Playa, Papaturro, and Sea Grape in Central America. The

juicy fruit is edible with a slightly acid flavor and has been used to make an

alcoholic beverage in the West Indies. The hard heavy wood has been used in

making furniture and for charcoal.

Coccoloba venosa L., Syst. Nat. ed. 10:1007. 1759. Campderia floribunda Ben-

tham, Bot. Voy. Sulphur 159, tab. 52. 1846. Campderia mexicana Meisner, in DC.,

Prodr. 14:171. 1856. Coccoloba floribunda (Benth.) Lindau, Bot. Jahrb. 13:217. 1890.

Coccoloba mohnaeStandl. & L. Wms., Ceiba 3:198. 1952. Figure 23.

Shrubs or small trees 4-7 (10) m tall, often with a twisted trunk and thick spreading

crown of many branches, leafy internodes 0-3 cm long, 2-6 mm thick, at first very

minutely papillate-puberulent but quickly glabrescent, becoming pale grayish and longi-

tudinally striate, sometimes resinous punctate, becoming lenticellate, the nodes con-

spicuously thicker than the internodes; stipules 5-15 mm long, originating at or just below

the petiole attachment, glabrous or very minutely puberulent, deciduous or persisting

with the leaves. Leaves alternate, often borne on short shoots, distichous on the long

shoots, petioles 3-10 (15) mm long, 1-2.3 mm thick, glabrous or minutely puberulent,

often reddish punctate or reddish brown throughout, often borne above the thickened

base of the node or on the differentiated base of the ocreate stipule; laminae 5-14 (20) cm

long, 2.5-7 (10) cm broad, obovate or obovate-oblong, acute to obtuse or rounded at the

apex, obtuse to rounded at the base, often with small (3 mm) and unequal cordulate lobes

at the petiole, entire, drying undulate and very stiffly chartaceous to subcoriaceous,

smooth and glabrous above and below (in ours) except for the presence of crooked hairs

in the axils of the major veins, with 5-8 pairs of major secondary veins, tertiary venation

often slightly raised above and below when dry. Inflorescences usually solitary, terminal

or (apparently axillary) on short axillary shoots, (4) 6-12 (16) cm long, the flowerless

peduncle less than 5 mm long, flower fascicles densely crowded and the rachis often

difficult to see, the rachis 1-2.5 mm thick, very minutely puberulent or glabrous, usually

reddish brown and deeply ridged, basal bracts of each fascicle 1-2.5 mm long, dark and

stiff, the thin bracteoles 2-3 mm long, pedicels ca. 2 mm long; male flowers several per

fascicle, filaments 1-2 mm long, anthers 0.3-4 mm long; female flowers solitary, flower

buds ca. 2 mm long. Fruit enclosed within the persistent slightly succulent perianth, to 6

mm long, perianth lobes separate almost to the base, narrowed at the base above the

articulation with the petiole; achenes 4-5 mm long to 4 mm broad, strongly trigonous,

broadest near the base and ovate or trangular in outline, dark brown and lustrous.

Plants of the seasonally very dry deciduous (tropical dry and tropical moist

transition) forest formations below 200 m elevation in Costa Rica; probably flow-

ering sporadically throughout the year, but fertile collections have been made

primarily in January and February and in July and August. The species ranges

from central Mexico to the northwestern Pacific lowlands of Costa Rica, and from

BURGER: FLORA COSTARICENSIS 121

Hispaniola and Puerto Rico southward through the Lesser Antilles to Trinidad

and Venezuela.

Coccoloba venosa is recognized by the short-petiolate obovate-oblong leaves

borne on thick grayish stems with thickened nodes, compact spikes with densely

crowded flowers, and deciduous lowland habitat. The species is occasionally

found near the seashore. Common names that have been used for the species in

our area include Papaturro, Papaturro de Playa, Papaturro Negro Macho, Papaturro

Rastero, and Gateador.

MUEHLENBECKIA Meisner

Nomen Conservandum

Small shrubs, slender climbers, or lianas, unisexual or bisexual, stems mostly glabrous

and longitudinally striate; stipules united to form a thin tube (ocrea) surrounding the

stem, deciduous or persisting. Leaves alternate and simple, sessile or petiolate; laminae

entire, glabrous. Inflorescence basically fasciculate and axillary, often racemose or spike-

like with the fascicles alternating along leafless stems or the stems branched and pan-

iculate, the flowers occasionally borne directly from the axils of persisting leaves; fascicles

subtended by thin stipule-like (ocreolate) bracts. Flowers unisexual (in ours) or bisexual,

borne on short pedicels articulate at the base of the perianth, perianth usually deeply

5-lobed (rarely 4- or 6-lobed), segments subequal, united near the base to form a shallow

cup, stamens 8 (9), inserted on the perianth cup, filaments free, anthers introrse, stami-

nodes present or absent in the female flowers, a pistillode present or absent in the male

flowers; ovary 3-angled, style 3, short, stigmas fimbrillate to capitate. Fruit remaining

enclosed in the persistent perianth, the perianth becoming somewhat larger and slightly

succulent in many species; achenes trigonous, the edges rounded or acute, the surface

often dark and lustrous.

A genus of about 20 species best represented in montane and southern South

America, New Zealand, and Australia. The genus is characterized by woody

climbing or clambering stems (or subshrubs of high mountains), very small flow-

ers in small fascicles, and five tepals that enlarge just enough to enclose the fruit

and become slightly fleshy.

la Distal stems flat and ribbon-like; leaves usually absent; flower fascicles borne alter-

nately along the edges of the phyllodes just above transverse lines; cultivated orna-

mentals or occasional escapes M. platyclada

Ib Stems terete or angular; leaves always present 2a

2a Large shrublike plants with clambering or climbing stems to over 3 m long; leaves

often cordulate at the base, to 12 cm long; flower fascicles in spikelike or paniculate

arrangements; 1,500-3,000 m M. tamnifolia

2b Small subshrubs with stems less than 80 cm long; leaves elliptic, to 10 mm long;

flowers solitary or fasciculate in the leaf axils; plants of high montane paramo vegeta-

tion above 3,000 m M. volcanica

Muehlenbeckia platyclada (F. v. Muell.) Meisner, Bot. Zeitung 23:313. 1865, as

platydados. Polygonum platycladum ¥. v. Muell., Trans. & Proc. Philos. Inst. Victoria

2:73. 1858. Homalocladum platycladum (F. v. Muell.) Bailey, GentesHerb. 2:58. 1929.

Shrubs, 0.5-3 m tall, with many closely crowded branches, glabrous, unisexual, basal

and primary stems terete, the distal stems flat and straplike, less than 1 mm thick (dry) and

5-15 mm broad, with distinct transverse lines ca. 1-2 cm distant and demarking the

nodes, the surface with closely parallel ribs 0.2-0.5 mm apart; stipules (ocreae) repre-

sented by a transverse line across the flattened stem. Leaves usually absent, sessile;

laminae 2-5 cm long and 0.5-1.5 cm broad, cuneate to hastate at the base, narrowly

lanceolate distally, acute at the apex, glabrous, quickly caducous. Inflorescences small

flower fascicles borne alternately along the edges of the stems just above the transverse

122 FIELDIANA: BOTANY

lines, pedicels ca. 1 mm long; flowers small and unisexual, perianth ca. 2 mm long. Fruit

included within the sweet red succulent perianth, ca. 5 mm long; achene ca. 3 mm long.

Unusual plants native to the southwestern Pacific and adjacent areas. These

plants are often grown in gardens and in pots for their dense branching and

unique stems. They escape and grow in the wild on occasion, but do not appear

to persist in our area. The flat stems with transverse line and small flower fascicles

alternating along the edges of the distal phyllodes distinguish this species from

virtually all other terrestrial plants. The species is often called Solitaria or Tenia in

Spanish and "Ribbon Bush" or "Tapeworm Plant" in English.

Muehlenbeckia tamnifolia (H.B.K.) Meisner, Gen. PI. 2:227. 1840. Polygonum

tamnifolium H.B.K., Nov. Gen. & Sp. 2:180. 1817. Figure 19.

Woody vines or lianas, scrambling or climbing to 15 m high, often forming dense

masses, unisexual or bisexual, glabrous, leafy internodes 0.5-3 (8) cm long, 0.8-4 mm

thick, often with 2 or 4 longitudinal ridges, drying reddish brown and striate; stipules

tubular (ocrea), thin and translucent, 5-15 mm long but often deciduous, entire distally,

leaving a distinct scar around the stem. Leaves alternate and usually distichous, slightly

succulent when alive, glabrous, petioles 4-16 (20) mm long, with 2 adaxial ridges con-

tinuous with the lamina margins; laminae 3-9 (12) cm long, 1-4 (6) cm broad, elliptic-

oblong to elliptic-ovate or elliptic, tapering gradually or abruptly to the acute or acuminate

apex, usually rounded near the base and auriculate or cordulate with a relatively wide

sinus or the laminae sometimes gradually tapering to an obtuse or acute base (rarely

auriculate and acute on the same plant), margin entire, usually drying thin chartaceous,

venation pinnate with 7-14 pairs of major secondary veins, the basal secondaries often

dichotomizing halfway to the margin. Inflorescences 1-3 from the axils of leaves, simple

and racemose or spikelike or branched and paniculate (sometimes with reduced leaves),

unisexual or bisexual, very variable (2-15 cm) in length, flowers in alternating fascicles

subtended by a number of ocreolate bracts 1-3 (5) mm long, rachis 0.3-0.6 mm thick,

pedicels 0.5-3 mm long and articulate beneath the perianth. Flowers unisexual, small, pale

yellowish to greenish white, the perianth usually 5-parted, united near the base, ovate;

male flowers ca. 2 mm long, stamens 8 (10), borne on the lower half of the tepals, ca. 1 mm

long, anthers ca. 0.5 mm long, a pistillode present or absent; female flowers 1.5-3 mm

long, ovary 3-angled, styles 3 and spreading, stigmas fimbrillate, staminodia usually

present. Fruit loosely enclosed within the persistent perianth, perianth enlarging slightly

to cover the fruit; achenes bluntly 3-angled, 2.5-3.5 mm long, 2-3 mm broad, dark and

lustrous.

Vinelike plants of evergreen moist and wet montane forest formations, from

(1,400) 1,800 to 2,900 (3,100) m elevation in Costa Rica; probably flowering

throughout the year, but collected most often between September and March.

This species is known from southern Mexico and Guatemala, central Costa Rica

to western Panama, and western South America.

Muehlenbeckia tamnifolia is recognized by the clambering stems with annular

scars left by the ocreate stipules, thickened nodes with reddish brown surfaces,

glabrous leaves often cordulate at the base, and the small fasciculate flowers in

spikelike or panicle-like inflorescences. Plants placed under this name exhibit an

interesting polymorphism as regard the shape of the leaf base. In Costa Rica, about

one-third of the collections have an acute or obtuse leaf base, while the majority

have a slightly auriculate or cordulate leaf base. The latter often have a con-

spicuous (2-5 mm) basal bract with dark midvein subtending the flower fascicles,

whereas plants with acute leaf bases seem to lose these bracts early or to have

them less well developed. The plants with acute leaf bases also seem to have their

inflorescences more often branched. A closer examination of these differences in

the field might prove rewarding.

BURGER: FLORA COSTARICENSIS 123

Muehlenbeckia volcanica(Benth.) Endlicher, Gen. PI. Suppl. 4:51. 1847. Po/y-

gonum volcanicum Bentham, PI. Hartweg. 81. 1841.

Small shrubs or dwarf shrubs, 5-50 cm tall, stems to 75 cm long, usually prostrate or

pendent, leafy internodes with conspicuous ridges, very minutely puberulent; stipules

with a tube 1-2 mm long. Leaves alternate, closely spaced, petioles 0-3 mm long; laminae

3-10 mm long, 2-5 mm broad, elliptic to rhombic-elliptic, tapering gradually to both acute

apex and attenuate base, drying coriaceous. Flowers axillary, solitary or in fascicles,

pedicels very short, flowers 2-3 mm long, greenish white. Fruit fleshy and black, loosely

enclosed in a calyx 3-4 mm long; seed an ovoid-trigonous achene.

Plants found only on Costa Rica's highest mountains above 3,100 m elevation,

but ranging from 2,400 to 4,000 m in Guatemala. The species ranges from southern

Mexico to Guatemala, Costa Rica and Colombia to Bolivia.

Muehlenbeckia I'olcanica is recognized by its very small stature, tubular ochreate

stipules, very small leaves, and high altitude habitat. Arthur Weston's careful

collecting on Chirripd in 1976 first disclosed the presence of this species in Costa

Rica.

PODOPTERUS Humboldt & Bonpland

Shrubs or small trees, bisexual, branches usually terminating in stiff spines, leaves and

inflorescences usually borne on small rounded short shoots (spur shoots or brachyblasts);

ochreate stipules small, caducous. Leaves alternate in a spiral or in fasciculate clusters on

small axillary short shoots, deciduous and the plants leafless for much of the year, lamina

thin and entire, venation pinnate. Inflorescences fascicles of flowers borne on small axil-

lary short shoots, pedicels articulate. Flowers bisexual and regular, small, perianth of 2

whorls of 3 parts, united near the base, the outer 3 perianth parts larger and each with a

longitudinal keel along the midrib that quickly expands to form a thin wing decurrent on

the pedicel, the inner smaller perianth whorl with flat erect parts enlarging slightly in fruit;

stamens 6, free, filaments filiform, anthers ovate; ovary trigonous, styles 3, stigmas cap-

itate, ovule basal and subsessile. Fruit a trigonous achene, included within the persisting

and enlarged perianth, each of the 3 outer perianth parts with a thin expanded longi-

tudinal wing arising from the back (abaxial) side of midrib, the wing rounded near the

apex of the perianth part and long-decurrent on the pedicel.

A genus of three species previously known only from Mexico, Belize, and

Honduras. The genus has a number of unusual characters, including spiny

branches with small rounded short shoots, deciduous leaves usually borne in

fascicles, flowers and fruits also in fascicles borne on the rounded short shoots,

and fruit with persisting perianth and unusual wings that are decurrent on the

pedicels. The genus is only found in deciduous formations with a severe dry

season.

Podopterus mexicanus Humboldt & Bonpland, PI. Aequin. 2:89. 1812. Fig-

ure 20.

Shrubs or small trees, 2-5 (8) m tall, usually branched near the base of or with a short

trunk, stems straight or angled and with short rounded spur shoots and terminal spines,

leafy internodes 0.3-2 cm long, 1.5-5 mm thick, smooth and usually dark gray in color,

glabrous; ochreate stipules not usually apparent, stipule scars encircling the young stems

often difficult to see, older stems with the stipule scars obscure except near the leaf base.

Leaves usually borne in fascicles of 2-5 on short shoots ca. 3 mm long, deciduous, petioles

(3) 5-20 mm long, ca. 0.6 mm thick, minutely puberulent with very short (0.1-0.2 mm)

straight hairs; laminae 2-7 cm long, 1.5-4.5 cm broad, broadly elliptic to obovate, rounded

to bluntly obtuse at the apex, gradually narrowed to the cuneate base, margin entire,

laminae drying stiffly chartaceous, smooth on both surfaces, glabrous or very minutely

puberulent along the midvein beneath, the 3-6 pairs of major secondary veins arising at

124 FIELDIANA: BOTANY

an angle of ca. 50° from the midvein. Inflorescences fascicles of flowers borne on short (3-4

mm) thick short shoots (brachyblasts) alternate along the stems, with 5-20 flowers per

fascicle, pedicels articulate near the base; flowers small and said to be green but becoming

white and with the 3 outer perianth parts soon developing longitudinal wings along their

midribs. Fruit borne within the thin whitish to pale brown or yellowish pink perianth

parts, 2-3 cm long including the pedicel, wings of the outer 3 perianth parts 2-3 mm broad

and 10-15 mm long, decurrent on the pedicels for 5-10 mm, the wings thin and trans-

lucent, achene ca. 5 mm long and 3 mm broad, with 3 narrow longitudinal ribs, the surface

pale brown and lustrous.

Trees and shrubs of the dryest seasonally deciduous woodlands and scrub

formations along the Pacific side of Central America between sea level and 700 m

elevation; fruiting collections have been made from late December to March. The

species ranges from central Mexico through western Guatemala and Honduras

and has recently been collected in northwestern Costa Rica.

Podopterus mexicanus is recognized by the spine-tipped branches with alternat-

ing rounded short shoots (spur shoots), small fasciculate leaves rounded at the

apex and cuneate at the base, the small fasciculate white flowers, and the fruit

with outer perianth parts bearing broad thin wings that are long-decurrent on the

pedicel. The restriction to seasonally very dry areas and the (often obscure) stipule

scars further distinguish these plants. They are called Escambron and Escambron

negroin Honduras. This species was not known from Costa Rica until early Decem-

ber 1977, when Daniel Janzen collected it growing near the swamp at Palo Verde,

Guanacaste. A collection by William Haber (74) from the same locality was made

on March 17, 1977.

POLYGONUM Linnaeus

Herbs, climbers or rarely shrubs, annual or perennial, bisexual, often growing in or near

water, stems often with thickened nodes and longitudinal ridges, glabrous to strigose and

occasionally with gland-tipped hairs; stipules and leaf base developed into a tubular leaf

sheath that surrounds the stem and is called an ocrea, the ocrea often with cilia or stiff

awns on its distal margin. Leaves alternate and simple, some species articulate above the

leaf sheath, subsessile to short-petiolate; laminae linear to ovate and occasionally cordate

to sagittate or hastate, margins entire, glabrous or puberulent, usually pinnately veined,

often glandular or pellucid punctate. Inflorescences basically fasciculate, the flowers in the

axils of leaves or leaflike bracts but more often subtended by ocrea-like bracts that are borne

close together on a raceme-like or spikelike axis, the flowering axes solitary or several in

a panicle-like arrangement, terminal or axillary near terminal leaves, flowers borne on

pedicels articulated near their apex. Flowers bisexual or occasionally functionally uni-

sexual, radially symmetrical, small, perianth of 4-6 subequal tepals in 1 or 2 whorls, the

inner often smaller and of thinner texture, perianth united near the base, green to white

or pink and red in color, imbricate in bud and persisting in fruit; stamens (3) 5-8 (9), often

unequal in length, included or protruding on filaments free or adnate to the perianth near

the base, anthers small, introrse, usually isodiametric and versatile (appearing peltate);

ovary superior, lenticular or trigonous, styles 2 or 3 and united near the base, stigma small

and capitate, locule 1 with an erect basal ovule. Fruit an achene, lenticular or trigonous,

ovate to orbicular in outline, usually included in the dry persisting perianth, the apex often

beaked with a persistent style base, surface of the fruit often dark and lustrous.

A genus of about 150 species often growing in open wet habitats and with the

great majority of species in the North Temperate Zone. With the exception of the

common Polygonum punctatum, the Costa Rican species of the genus are poorly

represented in collections. This is probably due to the fact that most live in wet

habitats that collectors have difficulty working in. Additionally, populations are

quite variable, and delimitation of species is often difficult. The genus is seriously

BURGER: FLORA COSTARICENSIS 125

in need of monographic revision, and one suspects that a great many species will

have to be redefined as much broader concepts than those currently in use.

Most of our species of Polygonum are easily recognized by their herbaceous

habit, narrow alternate pinnately veined leaves, flowers in spikelike or raceme-like

inflorescences, and the very unusual stipular structure (ocrea) enclosing the node.

Some Commelinaceae have similar stipular developments, but their leaves are not

pinnately veined.

la Flower groups borne on long (1.5-15 cm) spikelike or raceme- like axes; common

plants 2a

Ib Flowers in small groups in the axils of leaves or on short (0.5-1.5 cm) raceme-like or

spikelike groups at the ends of much longer slender flowerless peduncles; plants

rarely collected in Costa Rica 6a

2a Outer perianth parts and/or the ocreate bracts subtending the flowers with dis-

tinctive minute (0.1 mm) disklike gland dots; flower fascicles often somewhat

separate on the flowering rachis; leaves gradually narrowed to apex and base; a

common species P. punctatum

2b Outer perianth parts and ocreate floral bracts lacking disklike gland dots; flower

fascicles often crowded on the floral rachis; less common species 3a

3a Distal margins of the ocreae (stipular tube above the leaf base) with very short

(1-2 mm) setae or ciliate awns or these lacking; plants rarely with conspicuous

hairs, pubescence often confined to leaf margins and midveins, stems often slen-

der; leaves usually narrow and tapering gradually to apex and base 4a

3b Distal margins of the ocreae with long (2-20 mm) stiff setae, plants often

conspicuously pubescent, lower stems often more than 1 cm thick; fruit

lenticular 5a

4a Distal branches of the inflorescence lacking minute gland-tipped hairs,

flower fascicles often somewhat separate on the flowering rachis; fruit lentic-

ular or trigonous; mostly from the Caribbean lowlands (in ours)

P. hydropiperoides

4b Distal branches of the inflorescence with minute gland-tipped hairs, flower

fascicles usually crowded; fruit lenticular with 2 flat faces P. segetum

5a Stems and inflorescence branches with longer strigose hairs and minute gland-

tipped hairs; green leaflike flanges often present on the distal margin of the

ocreate stipules; laminae somewhat rounded near the base P. hispidum

5b Stems and inflorescence branches with appressed ascending strigose hairs,

gland-tipped hairs absent; stipules with setae to 25 mm long but lacking flanges;

laminae tapering gradually to the base P. acuminatum

6a Laminae broadly ovate to triangular and deeply cordate, petioles usually more

than 1 cm long; introduced vining weeds of open sites, naturalized in a few areas

in Panama but not recorded from Costa Rica and not included in the species

descriptions P. convolvulus

6b Laminae never broadly ovate or deeply cordate, (rarely subcordate to hastate),

petioles less than 5 mm long; plants erect or clambering but not vines 7a

7a Laminae 5-15 cm long; flowers clustered in fascicles at the ends of long slender

inflorescence branches; native plants of swamps and marshes, 0-1,800 m

P. meisnerianum

7b Laminae 0.5-4 cm long; flowers in fascicles in the axils of leaves; introduced

weeds of short (less than 0.5 m) stature, 1,500 m and above P. aviculare

Polygonum acuminatum H.B.K., Nov. Gen. & Sp. 2:178. 1817. P. guatemalense

Gandoger, Bull. Bot. Soc. France 66:225. 1919. Figure 17.

Herbs to 1.5 (rarely 2) m tall, usually erect, leafy internodes 2-4 (12) cm long, 4-8 (15)

mm thick, with appressed ascending strigose hairs but soon becoming glabrous; stipules

1—4 cm long with a usually dense covering of slender ascending hairs 1-4 mm long, distal

margin with 10 or more stiff setae (awns) 10-25 mm long; leaves alternate in a spiral,

subsessile or with petioles 1-15 mm long, lateral margins of the petioles continuous with

126 FIELDIANA: BOTANY

the lamina margins, appressed strigillose; laminae 8-23 (30) cm long, 1.5-3.5 (5) cm broad,

narrowly lanceolate to linear-lanceolate, occasionally curved and falcate, tapering gradu-

ally to the acuminate or acute apex, gradually tapering to the base and abruptly decurrent

on the petiole, margin entire and densely strigillose, laminae drying chartaceous and

smooth to the touch, the surfaces usually covered with appressed hairs distally oriented

and 0.1-1 mm long, the hairs often more dense along the midvein and margins, venation

pinnate with 13-25 pairs of major secondary veins. Inflorescences terminal or axillary from

near-terminal leaves, to 30 cm long, raceme-like or spikelike with a single axis or with 1-3

branches and paniculate, primary peduncle (2) 4-7 cm long, appressed strigose, branches

of the inflorescence 6-16 cm long with the flowering portion (1) 3-11 (14) cm long, ca. 1

cm in diameter at full anthesis, fascicles of flowers closely crowded and partly overlapping,

bracts of the fascicles 2-3 mm long and with distal cilia 0.5-2 mm long, pedicels exceeding

the bracts by 1-2 mm; flowers ca. 3-4 mm long, whitish or pink, perianth united only near

the base, usually 4-parted, stamens usually 6 and becoming exserted beyond the perianth,

anthers ca. 0.5 mm long, versatile, styles 2, 2-3 mm long. Fruit a thick lenticular achene

2-2.5 mm long, ovate in outline and with convex faces, dark brown or black, lustrous.

Plants of swamps, streamsides, wet depressions, and dense aquatic (rooted or

floating) vegetation from sea level to about 1,500 m elevation; probably flowering

throughout the year in Central America. The species ranges from southern Mexico

and the West Indies to southern South America.

Polygonum acuminatum is characterized by the often robust stems, general pres-

ence of thin ascending appressed (strigillose) hairs, long-setose stipules, ciliate

floral bracts, crowded flower fascicles, and thick lenticular fruit. The species is

known from only two collections in Costa Rica: Brenes 14353 from near San

Ramon, Alajuela, and Standley 48465 from near Cairo, Limon province.

Polygonum aviculareL., Sp. PL 362. 1753.

Annual herbs, prostrate or decumbent-ascending, simple or much branched, often

densely leafy with short internodes, leafy internodes 0-10 (30) mm long, 0.5-1.5 (3) mm

thick, smooth and glabrous, often ridged or angled and becoming longitudinally striate

when dry; stipules forming short (2-5 mm) tubular ocreae, the ocreae very thin and

translucent, smooth and lustrous, becoming lacerate distally. Leaves alternate in a spiral,

often bluish green in color, glabrous, clearly articulate just above the sheathing ocreate

base, subsessile or very short (0-3 mm) petiolate; laminae 0.5-3 (4) cm long, 1.5-5 (8) mm

broad, very narrowly elliptic to narrowly elliptic-oblong, bluntly acute at the apex, acute

and decurrent on the base, margin entire, laminae drying stiffly chartaceous, venation

pinnate with 3-5 pairs of major secondary veins. Inflorescences of fasciculate flowers in the

axils of leaves or leaflike bracts, 1-5 flowers per fascicle, pedicels to 2 mm long; flowers 2-3

mm long, perianth greenish white to pinkish or white, tepals rounded at the apex, fruit

a 3-angled achene, ovate in outline, dark brown, dull or slightly lustrous.

A weed of roadsides and open ground originally from north temperate areas but

now naturalized at higher altitudes in the tropics. The species is known in Costa

Rica from only three collections, all from the southern slope of Volcan Irazu near

Tierra Blanca, Cartago.

Polygonum aviculare is recognized by its small narrow leaves, many-branched

and often procumbent stems, translucent ocreae on the leaf bases, articulate small

narrow leaves, flowers in axillary fascicles and usually open weedy habitat. This

species has become naturalized in Guatemala, where it probably does not persist

below 1,500 m elevation; it is not likely to be found in our area below 1,500 m.

Polygonum hispidumH.B.K., Nov. Gen. & Sp. 2:178. 1817. Figure 17.

Herbs, 0.5-1.5 m tall, perennial and often with robust viscid stems to 2 cm thick, leafy

internodes 1-8 cm long, 3-10 mm thick, usually densely strigose to hispid with stiff

pale-colored hairs 2-6 mm long and minute (0.2-0.5 mm) reddish gland-tipped hairs;

BURGER: FLORA COSTARICENSIS 127

stipules forming a narrow tube 2-20 mm long with setae on the distal margin 2-5 mm long

and often with distal rounded reflexed or spreading flanges, long ascending and minute

gland-tipped hairs covering the surface. Leaves alternate in a spiral, petioles 1-6 cm long

with lateral ridges continuous with the lamina margins, strigose-hispid and with gland-

tipped viscid hairs; laminae 7-30 cm long, 3-12 cm broad, ovate- triangular to narrowly

ovate-elliptic or broadly lanceolate, tapering gradually to the acute or acuminate apex,

abruptly rounded near the base but gradually decurrent on the petiole, margin entire and

strigose, the laminae drying thin chartaceous, smooth to the touch but strigose-hispid

along the veins above and below, usually glabrous between the veins on older leaves,

venation pinnate with 10-25 pairs of major secondary veins, the lower surface punctate.

Inflorescences to 30 cm long, usually paniculate with 3 flowering branches, the primary

peduncle 5-10 (15) cm long and ca. 3.5 mm thick, branches 4-16 cm long and bearing

flowers over more than half their length, flowerless axes of the inflorescence with strigose

hairs 1-2 mm long and minute gland-tipped hairs, flower fascicles congested or closely

approximate, spikelike and raceme-like, 6-18 mm thick, fascicles subtended by ocreate

bracts 3-4 mm long, bracts usually with few hairs on the back and a ciliate margin of stiff

hairs 0.5-1 mm long, pedicels becoming slightly longer than the bracts; flowers 3-5 mm

long, deep pink to dark red (rarely greenish), perianth lobes 5, stamens 5, filaments 1.5-2

mm long, styles 2 and ca. 2 mm long, united for half their length. Fruit a lenticular achene,

3-4 mm long, ovate to orbicular in outline, with 2 flat or slightly concave faces, becoming

black or very dark brown, lustrous, style persisting and ca. 1 mm long.

Plants growing in and along streams and in seasonally wet swampy sites from

sea level to 1,500 m elevation; flowering throughout the year in Central America,

but collected most often in August. The species ranges from Guatemala and the

West Indies to southern South America.

Polygonum hispidum is recognized by the combination of larger strigose or hispid

and minute gland-tipped hairs, the unusual herbaceous flanges on the stipule-

margins (not always developed), the larger laminae rounded above an attenuate

base, the colorful flowers, and the lenticular fruit. In living material, the stems and

leaves are usually viscid. Virtually all the Central American collections come from

the Pacific slope, but the species has not yet been collected in Costa Rica. The

plants are called Tobacon and Tobaquillo in Honduras.

Polygonum hydropiperoides Michaux, Fl. Borealis 1:239. 1803. Figure 17.

Herbs, to ca. 1 m tall, lower stems usually slender and often rooting at the nodes, leafy

internodes 0.5-6 cm long, 1-4 mm thick, glabrous or with a few minute strigillose hairs,

becoming longitudinally striate when dry; stipules forming a narrow tube 8-20 mm long,

the distal margin entire or with small (1-2 mm) thin setae (ciliate awns). Leaves alternate

in a spiral, petioles 2-5 mm long and not clearly differentiated from the lamina; laminae

5-12 (15) cm long, 0.4-1 (1.5) cm broad, very narrowly elliptic or elliptic-lanceolate to

linear-lanceolate, tapering very gradually to the acute apex, tapering very gradually to the

attenuate base, margin entire and minutely strigillose, laminae drying chartaceous and

smooth to the touch, minutely (0.1-0.5 mm) strigillose on the midveins and margins and

often glabrous between, venation pinnate with 10-18 pairs of major secondary veins,

pellucid-punctate in transmitted light. Inflorescences to ca. 15 cm long, paniculate or

unbranched, the spikelike or raceme-like flowering portions 1-5 cm long and 0.5-1 cm in

diameter, ocreate bracts subtending the fascicles 2-3 mm long; flowers 2-3 mm long,

pedicels becoming ca. 1 mm longer than the subtending bract; flowers 2-3 mm long,

usually 5-parted, the outer perianth lobes 1.5-2 mm long, white or pink, stamens usually

8, anthers less than 0.5 mm long, ovary trigonous or lenticular, styles 2 or 3. Fruit a

3-angled or lenticular achene (mostly lenticular in Costa Rica), 2-3 mm long and ca. 1.5

mm broad, ovate in outline, brown to black, becoming lustrous.

Plants of river edges, wet depressions, and sandy or gravelly soils near water;

flowering throughout the year, but seen most often during the wet season. Al-

though these plants grow from sea level to 1,500 m elevation on the Pacific side

of northern Central America (El Salvador, Honduras, and Guatemala), they have

128 FIELDIANA: BOTANY

only been collected from below 200 m on the Caribbean slope in Costa Rica and

Panama. The species ranges from the southern United States to South America,

where it appears to merge with P. persicarioides.

Polygonum hydropiperoides is recognized by the almost glabrous aspect of the

plants, the slender stems, the narrow leaves tapering gradually to both apex and

base, slender inflorescences with flower fascicles not closely crowded, and fruit

that may be either lenticular or trigonous. As used here, P. hydropiperoides includes

Central American material previously identified as P. persicarioides H.B.K. Speci-

mens placed here may include individuals of P. punctatum lacking the character-

istic gland dots on perianth and floral bracts. Likewise, specimens of P. segetum

lacking gland-tipped hairs may find themselves placed under this name. The use

of P. hydropiperoides as something of a catchall is, I believe, justified by the very

poor state of our knowledge of these semiaquatic plants in Central America. James

S. Wilson has annotated specimens of this species as P. persicaria L. in the herbar-

ium of the University of Wisconsin; a Linnean epithet may be the ultimate

repository for this material.

Polygonum meisnerianum Cham. & Schlecht., Linnaea 3:40. 1828. Figure 17.

Herbs, to ca. 1m tall, decumbent or somewhat scandent, leafy internodes 1-5 (8) cm

long, 1.2-3 mm thick, glabrous or with stiff retrorse barbs or prickel-like hairs (these often

at the base of the node), occasionally with gland-tipped hairs; stipules narrow and tubular,

8-30 mm long, distally with short (1-2 mm) setae or entire, glabrous or with strigose or

glandular hairs. Leaves alternate in a spiral, sessile or with petioles 1-4 mm long, retrorse

hairs covering the petiole or only along the adaxial ridges continuous with the lamina

margins; laminae 5-15 cm long, 0.6-1.6 (2) cm wide, linear lanceolate, tapering very

gradually to the acute apex, abruptly narrowed at the base (very rarely subcordate or

hastate at the base in Central America), margin entire and with minute (0.1-0.2 mm)

ascending and/or glandular hairs along the edge, the laminae drying thin-chartaceous,

with stiff curved retrorse or ascending hairs to 1 mm long on the midvein beneath or

occasionally scattered over the leaf surface, venation pinnate with 15-30 pairs of major

secondary veins. Inflorescences terminal, panicles 5-20 cm long, primary peduncles 2-8

cm long, glabrous or with minute gland-tipped hairs, usually branching dichotomously,

usually with 2-5 thin branches 1-8 cm long and bearing flowers only in the distal 10-16

mm, branches of the inflorescence subtended by minute small ocreate bracts and with few

to many minute gland-tipped hairs; flowers ca. 3-4 mm long, borne on pedicels not

exceeding the floral bracts, perianth lobes less than 2 mm long, usually white. Fruit ca. 3

mm long, 3-angled, ovate in outline, lustrous pale brown, with a short (0.2 mm) beak.

Plants of lake edges, swamps, and marshes from 500 to 1,800 m elevation in

Central America and apparently flowering throughout the year. The species

ranges discontinuously from central Mexico to Brazil and the West Indies.

Polygonum meisnerianum is distinctive because of its open few-branched panicles

with relatively few (1-5) flower-fascicles at the ends of long thin secondary pe-

duncles. The glandular hairs usually found on the secondary peduncles and the

stiff retrorse hairs on leaves and nodes further distinguish this species. Only two

collections have been seen from Costa Rica; one from Lago Arena!, Guanacaste (A.

Jiminez 2699), and the other from Canas Gordas, Puntarenas (Pittier 11089). The

aquatic habitat of this species probably accounts for the fact that so few collections

have been made in our area.

Polygonum punctatum Elliott, Sketches Hot. S. Carol. & Georgia 1:445. 1817.

P. acreH.B.K., Nov. Gen. & Sp. 2:179. 1817. Figure 17.

BURGER: FLORA COSTARICENSIS 129

Herbs to ca. 1 m tall, annual or perennial, often with creeping stems, leafy internodes

0.5-2 (4) cm long, 1.3-10 mm thick, glabrous or rarely with ascending appressed strigose

hairs, longitudinally ridged when dry, partly included in the tubular stipule of the pre-

vious node; stipules tubular and narrow, (5) 12-25 mm long, glabrous or rarely with

appressed hairs, the distal margin setose ciliate with hard yellowish setae (bristles or

awns) 2-8 mm long (rarely smaller). Leaves alternate in a spiral, petioles 1-10 (15) mm

long, gradually merging with the sheathing ocreate (stipular) base and enclosing the stem;

laminae (2) 4-10 (17) cm long, (0.5) 1-2 (4) cm broad, very narrowly elliptic to lanceolate

or linear-lanceolate, tapering gradually to the acute or acuminate apex, decurrent on the

petiole, margin entire and minutely (0.3 mm) strigose, the laminae drying chartaceous and

slightly rough to the touch, usually glabrous above except for the margins and midvein,

glabrous below or with appressed ascending (strigose) hairs on the major veins, hairs ca.

0.2-0.5 mm long, venation pinnate with 5-15 (25) pairs of major secondary veins, the

lamina pellucid punctate when viewed by transmitted light. Inflorescences solitary or less

often 2-3 from the axils of near-terminal leaves, spikelike or raceme-like with a single

major axis or occasionally with 1 or 2 branches and paniculate, 5-20 cm long, flowering

portion 2-10 (15) cm long and ca. 0.5 cm thick, flower fascicles separate along the axis and

not closely crowded, fascicles subtended by tubular ocreate bracts ca. 2-3 mm long, entire

or awned apically, usually glandular punctate; flowers exserted from the bracts on slender

(0.2 mm) pedicels, ca. 4 mm long and articulate at the apex, flowers ca. 3—4 mm long,

gradually narrowed to the base, perianth lobes ca. 2 mm long, glabrous but with raised

pellucid dots, white to pale greenish in color, the 3 larger outer parts imbricate and

including the 2 inner perianth parts; stamens usually 8, filaments 1-1.5 (2) mm long,

anthers less than 0.5 mm in diameter, subglobose, ovary trigonous, style ca. 1 mm long,

connate basally. Fruit a 3-angled achene (in ours), ovate in outline, 2.5-4 mm long, 1.5-2

mm broad, dark brown to black and lustrous, beak inconspicuous.

Herbs of wet depressions in open areas, moist forest edges, and stream sides

from sea level to 2,000 m altitude; flowering throughout the year in Costa Rica, but

with most of the collections made between December and April and in July and

August. This species ranges from southern Canada to Argentina and is found in

Asia.

Polygonum punctatum is readily recognized among our species by the pellucid

punctate perianth and floral bracts. The species is further distinguished by the

narrow leaves tapering gradually to both apex and base and the inflorescences

with flower fascicles never closely crowded. This is the most commonly encoun-

tered species of Polygonum in Central America and often referred to as Chile de

perro in Costa Rica. This species is very similar to P. hydropiperoides among our

species; P. hydropiper L. of north temperate areas is a closely related species.

Polygonum punctatum is variable over its wide range, and a number of varieties

have been proposed (see Norman Fassett, The variations of Polygonum punctatum,

Brittonia 6:363-393, 1949).

Polygonum segetumH.B.K., Nov. Gen. & Sp. 2:177. 1817. P. mexicanum Small,

Bull. Torrey Bot. Club 19:256. 1892. Figure 17.

Herbs to ca. 1 m tall, mostly annuals, sometimes decumbent, leafy internodes (0.5) 2-9

cm long, 1.3-6 (10) mm thick, smooth and glabrous or rarely with gland-tipped hairs,

somewhat longitudinally striate when dry; stipules tubular and narrowly enclosing the

stem, 5-35 mm long, glabrous, entire or with a few short (1 mm) cilia distally. Leaves

alternate in a spiral, petioles 2-15 mm long, with lateral margins continuous with the

lamina margins, sparsely short-strigose; laminae 5-16 cm long, (0.5) 1-2.5 cm broad, linear

to lanceolate, occasionally curved and falcate, tapering very gradually to the acute or

acuminate apex, tapering gradually to the base and decurrent on the petiole, margin entire

but with appressed ascending minute stiff hairs along the edge, laminae drying char-

taceous, usually with minute (0.1-0.4 mm) appressed ascending stiff hairs on the major

veins, margins and near the tip, glabrous and minutely punctate between the veins,

130 FIELDIANA: BOTANY

venation pinnate with 10-30 pairs of major secondary veins. Inflorescences solitary or less

often 2 or 3 from the axils of near-terminal leaves, spikelike or raceme-like with a single

major axis or occasionally with 1 or 2 branches and paniculate, to 20 cm long, flower-

bearing rachis 1-6 cm long, 8-18 mm thick, primary peduncle and inflorescence branches

usually with minute (0.1-0.3 mm) gland-tipped hairs, flowers in crowded fascicles sub-

tended by ocreate bracts 2-4 mm long, bracts entire distally or with very short (1 mm) thin

cilia, glabrous or minutely gland punctate, pedicels equaling or slightly exceeding the

bracts; flowers 3-4 mm long, to 5 mm in fruit, pale pink or white, outer perianth lobes

2-2.5 mm long, ovary lenticular, styles 2. Fruit (2) 3-4 mm long, 2-3 mm broad, orbicular

in outline, lenticular with 2 flat faces, pale brown becoming very dark brown or black,

lustrous.

Aquatic or semiaquatic plants of lake edges, wet depressions, seasonally

flooded land or occasionally forming floating mats on open water; between sea

level and 1,500 m elevation and probably flowering throughout the year, but

collected most often in November and from January to March. The species ranges

from Mexico to Colombia and the Greater Antilles.

Polygonum segetum is distinguished by the glandular hairs on peduncles, flowers

often closely grouped on the floral axes, lenticular fruit, narrow leaves that taper

gradually to apex and base, and preference for aquatic and temporarily flooded

habitats. Specimens lacking the distinguishing gland-tipped hairs will key out to

P. hydropiperoides, and that species may prove to be a catchall for a variety of

atypical material. Vegetatively, the plants are also similar to P. punctatum, with

appressed ascending hairs usually confined to the major veins and margins of the

leaves.

Polygonum mexicanum is placed under P. segetum following C. D. Adams in the

Flowering Plants of Jamaica (1972). The plants placed here are also closely related

to P. pennsylvanicum L. of the eastern United States.

RHEUM Linnaeus

Herbs, with thick woody rhizomes, aerial stems short (1 m) or lacking; stipules thin and

loose, without cilia. Leaves often large, petiolate, the laminae palmately veined, often

sinuately dentate or palmately lobed. Inflorescences panicles with small clusters of flowers;

the flowers bisexual or functionally male, perianth 6-parted, subequal or the outer whorl

smaller, not enlarging in fruit; stamens usually 9, anthers ovate; ovary trigonous, styles 3,

short and recurved. Fruit an achene, narrowly or broadly 3-winged, embryo straight.

A genus of about 30 species best represented in eastern Asia and originally

ranging to eastern Europe. A few are cultivated for their large showy leaves and

one (Rheum rhabarbarum) is widely grown as a vegetable for its succulent petioles

which are used with sugar as a vegetable or a filling for pies.

Rheum rhabarbarum L., Sp. PI. 372, 1753.

A large perennial herb, most of the leaves borne from the base of the rhizome and lacking

a prominent stem but producing inflorescences to 2 m tall. Leaves to 1 m long, the petiole

thick and succulent, often reddish in color; laminae broadly ovate to suborbicular, 50-80

cm broad, deeply cordate at the base, the margin undulate, glabrous, venation palmate

with usually 5 primary veins. Inflorescences 1-2 m tall, a narrowly pyramidal leafy pan-

icle, flowers small and pale green to white, on slender articulated pedicels, similar to those

of Rutnex.

Rhubarb (Ruibarbo) is cultivated at middle elevations in Costa Rica; the reddish

succulent petioles are often seen in vegetable markets of the Meseta Central. The

name Rheum rhaponticum L. has been used for this species, but refers to a species

of Bulgaria that is little cultivated.

BURGER: FLORA COSTARICENSIS 131

RUMEX Linnaeus

REFERENCE: K. H. Rechinger, Die siid- und zentralamerikanischen Arten der

Gattung Rumex. Ark. Bot. 26, no. 3:1-58. 1934.

Perennial herbs, rarely annual or woody shrubs, unisexual or bisexual, glabrous or

sparsely puberulent, stems often longitudinally striate and hollow; stipules united to form

a sheathing tube (ocrea) around the stem, usually very thin and translucent, persistent but

lacerate and withering. Leaves alternate, simple, often dimorphic with basal or lower

cauline leaves differing from those near the inflorescences, early leaves often forming basal

clusters, petioles sheathing the stem at their base and united with the ocreae; laminae

entire or minutely dentate, often slightly succulent. Inflorescences racemose or paniculate

but basically composed of flower fascicles borne along an unbranched axis (and these axes

usually in panicle-like arrangements), the fascicles at first enclosed in stipule-like (ocre-

olate) bracts, the flowers sometimes numerous within the fascicle and appearing to be

verticillate or whorled, the flowers borne on simple or articulate (ebracteolate) pedicels.

Flowers bisexual or unisexual, mostly wind-pollinated, radially symmetrical, the perianth

of 6 parts in 2 whorls of 3 each, the outer whorl usually smaller than the inner whorl and

reflexed, remaining small, the inner 3 perianth parts usually larger and often enlarging in

fruit, both whorls persisting in fruit, stamens 6, free, filaments very short, anthers

2-thecous, narrowly oblong, usually longer than the filaments; ovary 3-angled (trigonous),

styles 3 and spreading or reflexed, stigmas fimbriate or penicillate, locule and ovule 1. Fruit

usually enclosed within the stiff dry persisting perianth parts of the inner whorl, in some

species at least 1 of the enlarged perianth parts develops a grainlike tubercle on the back

(abaxially) of the midrib; fruit a 3-angled achene, the 3 longitudinal ribs usually acute, the

surface smooth and lustrous or rough and dull.

A genus of perhaps 200 species, most of which grow in the Northern Hemi-

sphere. The genus is recognized by the very thin stipules encircling the stem, the

leaves often varying in form on the same stem, the small flowers often in verticel-

like groups along the inflorescence branches, perianth parts in two whorls of three

with the parts persistent and closely enclosing the fruit. In some species, at least

1 of the inner perianth parts enclosing the achene develops a grainlike tubercle on

the back of the midrib. This remarkable and conspicuous (pale colored) structure

may serve as a seed-mimic and reduce predation on the achenes. Of the four

species represented in Costa Rica, three are widely naturalized weeds recognized

by Linnaeus over 200 years ago, while the fourth species is a unique high-

mountain endemic that may include the tallest plants known in the genus.

la Flowers borne on pedicels becoming 3 cm long; plants regularly becoming more than

2 m tall, stems to 10 cm thick at the base; usually found only in moist protected sites

above 2,800 m elevation R. costaricensis

Ib Flowers borne on pedicels usually less than 1 cm long; plants rarely becoming more

than 1.5 m tall 2a

2a Small plants only occasionally becoming 50 cm tall; leaves 1-5 cm long, basal leaves

often with divergent lobes (hastate); flowers unisexual, 2 mm long or less; fruiting

perianth without grainlike tubercles on the back of the midrib; 1,500-3,500 m elevation

R. acetosella

2b Larger plants regularly becoming 50 cm or more tall, leaves 2-40 cm long, basal leaves

never hastate; flowers bisexual, usually more than 2 mm long; fruiting perianth often

with a grainlike tubercle developed on the back of the midrib 3a

3a Enlarged sepals (valves) with entire margins in fruit, without sharp projections or

spines, flowers and fruit often crowded on the inflorescence axis; largest basal leaves

usually less than 4 cm broad, rarely truncate or subcordate; common weeds, 1,000-

2,000 m R. crispus

3b Enlarged sepals with spine-like projections along the edge in fruiting stages; flowers

and fruits often in separate verticels along the flowering rachis; larger leaves com-

monly truncate to cordate; uncommon weeds, 1,000-2,800 m elevation 4a

4a Enlarged sepals with a few straight spines along the margin; larger leaves usually more

132 FIELDIANA: BOTANY

than 5 cm broad, broadest below the middle, laminae margins arising at about the

same level on the petiole R. obtusifolius

4b Enlarged sepals with distinctly hooked (uncinate) spines along the margins; larger

leaves 2-10 cm broad and often broadest at the very base of the laminae, margins

usually arising 1-5 mm distant along the apex of the petiole R. nepalensis

Rumex acetosella L., Sp. PI. 338. 1753. Figure 18.

Perennial herbs 10-40 (60) cm tall, usually branched only at the inflorescence and near

the base, spreading by slender rootstocks, usually unisexual, glabrous, leafy internodes

0-3 cm long, with 6-9 distinct longitudinal ribs; stipules 0.5-1 cm long, thin and trans-

lucent, lacerate. Leaves often dimorphic with basal and distal leaves of different form,

petioles weakly differentiated from the laminae, 4-40 mm long, with lateral margins

continuous with the laminae margins; upper (distal) laminae 1-3 cm long, 1-4 mm broad,

very narrowly elliptic and tapering gradually to both apex and base, lower (basal) laminae

1-5 cm long, often with divergent basal lobes (hastate), lanceolate or narrowly elliptic to

obovate above the basal lobes and 2-12 mm broad, acute to obtuse at the apex, usually

attenuate on the petiole below the divergent lobes, margins entire, the laminae drying

chartaceous, (often slightly succulent and with a pleasant sour taste in life), venation

subpalmate with usually 3 major veins. Inflorescence usually a panicle of spicate branches

(a simple spikelike or raceme-like axis on small plants), usually unisexual, the flowers

borne in fascicles resembling verticels on the inflorescence branches, each fascicle sub-

tended by an ocreate (stipule-like) bract and with 1-6 (9) flowers, the flowers borne on

slender pedicels 0.5-3 mm long. Male flowers 1-2 mm long, perianth of 3 narrow outer

parts and 3 inner slightly broader parts, a pistillode absent. Female flowers 0.5-1.5 mm

long, outer perianth parts small and narrow, the inner perianth parts broader and expand-

ing in fruit, ovary strongly 3-angled, the fimbriate stigmas ca. 0.5 mm long. Fruit included

within the persistent perianth, larger perianth parts ca. 1.5 mm long and closely appressed

to the fruit, entire; achene 1-1.5 mm long, 3-angled, surfaces flat or convex, ovate to

orbicular in outline, smooth, dull or slightly lustrous, brown.

Introduced weeds of open and disturbed sites at higher elevations. In Costa

Rica, the plants are commonly found between 2,200 and 3,500 m elevation and

occasionally down to 1,500 m; flowering and fruiting throughout the year. The

species, a native of Europe and Asia, is now widely naturalized.

Rumex acetosella is recognized by its often hastate basal leaves that are usually

slightly succulent and have a pleasant sour taste. The small stature, very small

reddish flowers and fruit, and open weedy high-altitude habitats further dis-

tinguish this species. The plants are often called Ruibarbilloin Central Ameria. This

species and its close relatives have sometimes been placed in a separate genus,

Acetosella.

Rumex costaricensis Rechinger, Repert. Sp. Nov. 40:300. 1936. Figure 18.

Large single-stalked herbs 2-5 (7) m tall, bisexual, glabrous, the main stems usually

unbranched below the flowering nodes, leafy internodes to 15 cm long, 5-10 cm thick,

hollow, longitudinally striate, glabrous and smooth; stipules (ocreae) to 18 cm long and

surrounding the stem (at upper nodes), thin and translucent, pale reddish brown, with

many subparallel veins. Leaves alternate in a spiral, somewhat polymorphic with the

upper leaves on the plant attenuate to the stem and the lower leaves petiolate and rounded

near the base, petioles 0-3 cm long near the inflorescence and 2-12 cm long lower on the

stem, 3-10 mm broad, with lateral margins continuous with the laminae margins; laminae

20-60 cm long, 5-15 cm broad, narrowly elliptic, tapering gradually to the acute or

acuminate apex, gradually tapering to the attenuate base (distal laminae near the in-

florescence) or abruptly rounded at the base (more basal cauline leaves), margin entire

(sometimes very slightly undulate), the laminae drying thin chartaceous, smooth and

glabrous above and below, venation pinnate with 20-40 pairs of major secondary veins.

Inflorescence paniculate, terminal or 1-3 from the axils of leaves, bisexual, flowers borne

in fascicles of 10 or more, separate and distant along the flowering axes, inflorescences

BURGER: FLORA COSTARICENSIS 133

usually with many male and few female flowers; the flowers unisexual and glabrous, borne

on long (to 3 cm) slender pedicels. Male flowers 4-7 mm long, the outer tepals 1.5-2 mm

long, inner tepals 4-6 mm long, stamens with very short filaments, anthers 2.5-3.3 mm

long, a small pistillode present. Female flowers ca. 2-3 mm long, inner perianth whorl ca.

2 mm long, ovary ca. 1 mm long with fimbriate styles 0.5-1 mm long. Fruit loosely

enclosed within the persisting perianth, larger perianth parts becoming 4-6 mm long, and

2-5 mm wide, emarginate; achene ca. 3 mm long (immature?), strongly 3-angled, with

convex faces ca. 2 mm broad and broadly ellipsoid in outline, becoming dark greenish

brown.

Unusual plants apparently restricted to partly open sites on moist slopes and

stream edges in areas protected from strong winds and found only from (2,200?)

2,800 to 3,400 m elevation; probably flowering throughout the year, but collected

with flowers and fruit only from December to March and in June and July. The

species is endemic to Costa Rica and known only from the Cerro de las Vueltas

(Heredia) and on the Cerro de la Muerte (along the Interamerican Highway

bordering the provinces of San Jose and Cartago).

Rumex costaricensis is recognized by its tall single-stalked habit, hollow thin-

walled stems, large leaves with many veins, large tissue-thin stipules, and pan-

iculate inflorescences to 50 cm long. This is one of Costa Rica's most unusual

high-mountain endemics and seems to be restricted to very moist areas subject to

wet Caribbean winds. This species apparently attains the largest size known for

the genus and is closely related to R. peruanusRech. and R. tolimensis Wedd. of the

high Andes of northern South America.

Rumex crispusL., Sp. PL 335. 1753. Figure 18.

Erect perennial herbs 0.3-1.5 m tall, with a thick taproot, the stems usually unbranched

below the inflorescence, bisexual, leafy internodes 1-5 (10) cm long, 3-10 mm thick,

glabrous, longitudinally striate, hollow; stipules 0.5-5 cm long, thin and translucent.

Leaves quite variable on the individual plant, often forming dense clusters at the base of

the plant, petioles 1-5 cm long (to 10 cm on basal leaves); clasping the stem at the base;

laminae 3-25 cm long, 0.5-4 cm broad, linear-lanceolate to narrowly oblong or linear-

oblong, acute at the apex, attenuate at the base (rounded or cordate on basal leaves),

margin undulate and crisped, minutely crenate-dentate, the laminae drying chartaceous,

smooth and glabrous above and below, venation pinnate with 10-20 pairs of major second-

ary veins. Inflorescences terminal and paniculate with racemose branches or axillary and

racemose, the flowers in verticil-like fascicles on the flowering axes, the verticils (whorls)

closely approximate, and subtended by ocrea-like bracts, flowers borne on slender pedicels

1-5 (12) mm long; flowers bisexual, outer tepals 1-2 mm long and not expanding in fruit,

oblong, inner tepals ovate to cordate and becoming 3-5 mm long in fruit, stamens 6 with

filaments less than 0.5 mm long, anthers 1-3 mm long, ovary 3-angled, stigmas fimbriate.

Fruit tightly enclosed within the persisting perianth, the perianth parts 3-5 mm long and

ca. 3 mm broad in fruit, entire, often with an expanded ellipsoid tubercle ("grain") on the

base of the midrib abaxially becoming 3 mm long and 1.5 mm broad; achene 1.5-3 mm

long, ca. 1.3 mm broad, 3-angled with 3 flat or slightly convex faces, ovate to orbicular in

outline, lustrous dark brown.

Introduced weeds from Europe and naturalized in areas of extensive agricultural

and similar activity. The species has only been collected in the Central Highlands

(around the Meseta Central) between 1,000 and 2,000 m in Costa Rica; flowering

and fruiting material has been collected from December to May and in August.

The species is now naturalized in many parts of the world.

Rumex crispus is recognized by the densely flowered inflorescences, undulate

and obscurely dentate leaves, and the entire perianth parts which (in the inner

whorl) enlarge in fruit and often have a grainlike development on their back

(abaxially). The young leaves are eaten after cooking in the United States, but this

134 FIELDIANA: BOTANY

practice is not recorded from Central America. The plants are called Lengua de vaca,

Lengua de caballo, and Lechugon in Guatemala.

Rumex nepalensis Sprengel, Syst. Veg. ed. 16, 2:159. 1825. Figure 18.

Herbs, 0.3-1 m tall (in ours), leafy internodes 0-10 cm long, 1.5-6 cm thick, glabrous,

longitudinally ribbed; stipules thin and soon tearing off. Leaves basal or cauline but not

differing greatly in form, petioles 1.5-10 cm long; laminae 5-15 (21) cm long, 1.5-5 (10) cm

broad, ovate-oblong to narrowly triangular and broadest at the base, acute to bluntly

obtuse at the apex, truncate to cordate or somewhat hastate at the base, margins entire and

usually arising 1-5 mm distant on the petiole, laminae drying thin-chartaceous with 9-15

pairs of major secondary veins. Inflorescences terminal or axillary, 10-50 cm long, the

flower fascicles 5-12 mm distant on the deeply ridged glabrous rachis, the fascicles form-

ing verticil-like (whorled) clusters with short (1-4 mm) pedicels articulated near the base;

flowers ca. 2 mm long at anthesis. Fruit enclosed in valves (enlarged sepals) 3-5 (6) mm

long and with numerous hooked (uncinate) spines 1-3 mm long on the margins, the

tubercles (grains) on the abaxial surface poorly developed or absent but the venation

prominent.

An apparently recently introduced weed collected only in San Jose (Godfrey

67336) and on the grounds of the University (Croat 593 and 959), both in 1965.

These specimens are smaller in almost every respect than those seen from other

parts of the world, but the valves with their hooked spines are diagnostic.

Rumex obtusifoliusL., Sp. PI. 335. 1753. Figure 18.

Erect perennial herbs, 0.5-1.5 m tall, often forming dense colonies, bisexual, leafy

internodes 0-8 cm long, 2-8 mm thick, longitudinally striate, glabrous; stipules (ocreae)

to 5 cm long, very thin, pale yellowish and translucent. Leaves often polymorphic on the

same plant, the basal leaves often in dense clusters and long-petiolate, petioles 1.5-15 cm

long, 0.5-3 mm broad, clasping the stem at their base; laminae 4-25 (40) cm long, 1-10 cm

broad, tapering gradually to the acute apex, the upper (distal) laminae elliptic to lanceolate

or ovate-elliptic and tapering to the obtuse base, the lower (basal) laminae much larger and

oblong to ovate-oblong, truncate to cordate at the base, margin entire or crisp-undulate

and minutely crenate-dentate in larger leaves, the laminae drying thin-chartaceous,

smooth and glabrous above, minutely (0.2 mm) puberulent on the veins beneath (the

trichomes rather thick and irregular), venation pinnate with 4-14 pairs of major secondary

veins. Inflorescences racemose or paniculate, the flowers borne in somewhat separate

verticil-like fascicles, the fascicles subtended by ocreate bracts, the flowers borne on

articulate pedicels to 8 mm long. Flowers bisexual, 2-3 mm long, the 3 outer perianth parts

ca. 0.5 mm broad, the inner parts broader (1-1.5 mm) and longer (2-3 mm), enlarging and

developing unusual lobes in fruit, anthers ca. 1.5 mm long, stigmas fimbriate. Fruit tightly

enclosed in the persisting and enlarged perianth, inner perianth parts (valves) 3-7 mm

long, with narrow or teethlike lobes along the margin, an ellipsoid tubercle (grain) 2-3 mm

long often developing on the back of the midrib; achene 2-3 mm long, strongly 2-angled

with 3 convex faces, ovate or elliptic in outline, lustrous brown, the perianth becoming

reddish.

Introduced weeds of open sites in montane habitats between 1,200 and 2,800 m

elevation; probably flowering primarily in the wet season and early part of the dry

season (June to February). Native of Europe and Asia and now widely naturalized.

Rumex obtusifolius is recognized by the raceme-like inflorescences, with separate

whorls (actually fascicles) of flowers, articulate pedicels, unusual teeth and thick-

ened "grains" on some of the fruiting perianth parts, and the long-petiolate basal

leaves. The stems are said to be eaten for stomach ailments in Costa Rica.

RUPRECHTIA Meyer

REFERENCE: A. E. Cocucci, Revision del Genero Ruprechtia, Kurtziana 1:217-269.

1961.

BURGER: FLORA COSTA RICENSIS 135

Trees or shrubs, unisexual (dioecious), stems solid; stipules forming a tube around the

stem (ocrea), small and thin, caducous or evanescent. Leaves alternate and simple, decid-

uous, petiolate, the laminae pinnately veined, glabrous or puberulent. Inflorescence race-

mose or of racemose axes in a paniculate arrangement on a leafless terminal twig, the

flowers borne in fascicles subtended by several thin ocreolate bracts, the flowers pedi-

cellate (in ours), flowers very small at anthesis. Male flowers with 3 sepals, free or united

at the base, usually obtuse and membranaceous, the 3 inner petals (tepals) similar to the

sepals and alternate with them; stamens 9, the outer whorl with 6 stamens in pairs

alternating with the petals and the inner whorl of 3 stamens opposite the petals, a disk

usually present, a pistillode absent. Female flowers with 3 sepals, free or united and with

or without a conspicuous tube, greatly enlarging in fruit, 3 petals alternate with and much

smaller than the sepals, glabrous or pilose, 9 very small staminodes usually present; ovary

with 3 short styles and 3 globular or linear stigmas. Fruit retained within the persisting

perianth, the sepals always enlarging in fruit, often with 3 long narrowly spatulate wings,

stiffly chartaceous in texture; achene with 3 deep longitudinal sulci separating the rounded

faces of the fruit, narrowly ovoid to ellipsoid and narrowed to the persisting styles and

stigmas, seed deeply 3-sulcate.

A New World genus of 17 recognized species, with several species in Mexico

and a majority in southern South America. These separate centers of diversity

probably reflect the fact that most of the species are restricted to seasonally dry

and deciduous habitats. These plants are recognized by the small flowers in

fasciculate groups subtended by ocreolate bracts, the unusual three-winged fruit

very similar to those in Triplaris, and the restriction to seasonally very dry areas

of northwestern Costa Rica. In addition to the differences listed in the key,

Ruprechtia differs from Triplaris in chromosome number (n = 14 in Ruprechtia and

11 in Triplaris) and the much smaller stipules that are often difficult to see.

Ruprechtia costata Meisner, in DC., Prodr. 14:180. 1857, sensu lato. R. deamii

Robinson, Proc. Amer. Acad. Sti. 43:51. 1907. R. kellermanii Donn. Smith, Bot.

Gaz. 47:260. 1909. Figure 20.

Medium-sized to tall trees, 2-10 (25) m tall, trunk to ca. 35 (60) cm in diameter, uni-

sexual, leafy internodes 0.3-1.5 (5) cm long, 0.7-2 (5) mm thick, at first with thin ascending

hairs ca. 0.5 mm long, soon becoming glabrous and gray, lenticellate; stipules forming a

short (1 mm) thin tube around the stem and leaving a very obscure ring around the stem.

Leaves usually distichous, articulate at the base (below the ocrea), petioles 2-6 mm long,

with lateral margins continuous with the lamina margins; laminae 3-11 (18) cm long, 1.5-5

(8) cm long, elliptic or slightly elliptic-obovate to elliptic ovate, tapering to a short-

acuminate apex (in ours), acute to obtuse at the base, margin entire or slightly undulate-

crenate, the laminae drying chartaceous (in ours) or subcoriaceous, glabrous above and

with few thin ascending hairs on the midvein beneath, the midvein prominent above and

below, venation pinnate with 5-7 (11) pairs of major secondary veins. Inflorescences

racemose, 1-3 from the axils of leaves or fallen leaves, occasionally in paniculate arrange-

ments on leafless stems, flower fascicles subtended by small (1 mm) thin-translucent

ocreolate bracts, flowering axes to ca. 3 cm long in ours (to 10 cm in others). Male flowers

on puberulent pedicels 1-2 mm long, perianth ca. 2 mm long, filaments ca. 3 mm long,

anthers ca. 0.6 mm long; female flowers not seen. Fruit enclosed within the enlarged dry

perianth and borne on pedicels 2-4 mm long, pedicel and perianth with thin erect hairs

0.2-0.5 mm long, fruiting perianth 2-4 cm long, (only 2-2.5 cm long in ours with wings

ca. 15 mm long and 3—4.5 mm wide), pale rose brown when dry; achene becoming 1 cm

long, 3-4 mm thick with 3 deep longitudinal sulci separating the 3 rounded sides of the

fruit, narrowly ovoid to ellipsoid and usually puberulent on the upper half.

Trees of the seasonally very dry deciduous (tropical dry) forest formations of

Guanacaste province below 300 m elevation; flowers and fruit have only been

collected in January, February, and March. The species ranges from Guatemala to

the Pacific lowlands of northern Costa Rica and central Panama.

136 FIELDIANA: BOTANY

Ruprechtia costatais recognized by its distinctive 3- winged fruiting perianth with

the texture of stiff paper and basal tube loosely enclosing the 3-furrowed achene.

The ocreate stipules are very small and difficult to see, while the male in-

florescences are densely (though often minutely) puberulent, with the flower

fascicles subtended by a cluster of small thin ocreolate bracts. The fruiting peri-

anth resembles that of Triplaris, but does not enclose the fruit so tightly.

The current usage of the name Ruprechtia costata, following Cocucci, includes

specimens of diverse aspect. Specimens from Guatemala with subcoriaceous

leaves to 18 cm long, with long male inflorescences and fruiting perianth to 4 cm

long are quite different from the Costa Rican material with thin leaves, male

inflorescences usually only about 2 cm long, and fruit becoming only 2.5 cm long.

In addition, our material has foliage with fewer secondary veins. There are Gua-

temalan collections that appear to be intermediate between these extremes; more

collections are needed to ascertain the importance of these variations. Our mate-

rial from Guanacaste is closely matched by specimens from Mexico identified as

R. pringlei Greenman, a species not treated in the recent monograph. (Earlier, our

material had been placed under the name of a South American species, R. cum-

ingii Meissn.)

TRIPLARIS Loefling

Trees, unisexual (dioecious), distal stems and branches hollow and often populated by

fiercely stinging or biting ants, young stems densely strigose to glabrescent; stipules united

and forming a cap over the shoot apex (as in Ficus), leaving a raised annular scar around

the stem at each node, the annular ridges often present on older stems (5-10 cm in

diameter), the stipular structure (ocrea) united to the petiole only at the base, deciduous.

Leaves alternate and usually distichous, folded (plicate) in bud, short-petiolate, the lami-

nae generally elliptic to ovate or oblong, often unequal at the base, entire, strigose or

sericeous to glabrescent, venation prominent beneath, the younger leaves often with

longitudinal striae or folds reflecting the development within the bud. Inflorescence basic-

ally an unbranched axis (spicate or racemose), the spikes or racemes in groups of 1-8 in

the axil of a leaf or undeveloped leaf, often in alternate groups on a terminal twig on which

the leaves fail to develop or are soon deciduous and thus forming a panicle-like compound

inflorescence, the flowers usually in fascicles (male) or solitary (female), at first enclosed

in ocrea-like bracts (ocreolae), the bracts opening along the adaxial side and soon decid-

uous, the flowers unisexual, subsessile to short-pedicellate, often densely sericeous or

strigose on the outside, regular. Male (staminate) flowers 1-5 per fascicle, tepals 6, linear

to ovate, uniseriate and equal or subequal, stamens 9, exceeding the tepals on slender

filaments, anthers versatile, introrse, 4-thecous, a pistillode absent. Female (pistillate)

flowers pedicellate and mostly solitary, perianth of 2 very different series, an outer sepa-

loid series of 3 broad thin parts that are united to form a distinct tube in the lower Vz-Vi,

the inner series of 3 very narrow stiff perianth parts free or partly adnate to the tube and

equal to or only slightly exceeding the length of the tube, the perianth strongly accrescent

in fruit with the 3 outer lobes becoming spatulate to oblanceolate paperlike wings, stami-

nodes absent or forming an annular disk (rarely with functional stamens), ovary strongly

3-angled with acute edges, styles 3 with stigmatic areas on their inner faces, locule 1, ovule

1. Fruit an achene included within the persistent and enlarged 3- winged perianth tube, the

achene usually 3-angled and ovate in outline, the surfaces lustrous or dull, occasionally

verrucose or pustulate.

A neotropical genus with probably fewer than 20 species, most of which are

confined to South America. These trees are quite distinctive, with large colorful

compound inflorescences, fruit enclosed in a three- winged papery perianth, stip-

ules capping the shoot apex and leaving raised scars around the stems, and the

frequent presence of aggressive ants. Several species of the genus are planted as

ornamental trees.

BURGER: FLORA COSTARICENSIS 137

Variability of the populations, apparent intergradation of what have been called

species, and lack of monographic study in the last 100 years have made this a

particularly difficult genus. I have simplified the Costa Rican situation by adapt-

ing Duke's excellent treatment for the Flora of Panama and assuming that T.

cumingiana finds its northern limit in westernmost Panama. Since no specimen

from Costa Rica can be unambiguously keyed to T. cumingiana (following Duke),

the decision to exclude that species from our account is, I believe, justified. Our

material will probably prove to be the northern element of a more broadly circum-

scribed species, such as T. cumingiana.

Triplaris melaenodendron (Bertol.) Standley & Steyermark, Publ. Field Mus.

Nat. Hist., Bot. Ser. 23:5. 1943, Velasquezia melaenodendron Bertoloni, Fl. Guatem.

40. 1840. T. macombii Donn. Sm., Bot. Gaz. 19:257. 1894. Figure 20.

Trees, 5-15 (25) m tall, unisexual, trunk with circular annular rings in early stages, the

bark flaking off in broad flat patches, biting or stinging ants often present in the hollow

distal stems, leafy internodes 1-4 (8) cm long, 3-10 mm thick, with thin appressed and

ascending strigose hairs but soon becoming glabrous, often reddish brown in color, longi-

tudinally striate; stipules 1.5-3.5 (5) cm long, densely strigose on the outer surfaces,

leaving an annular scar around the stem, caducous. Leaves usually distichous, articulate

at the base, longitudinally folded in bud, petioles 4-20 mm long, 2-5 mm thick, strigose

abaxially and along the 2 lateral margins, glabrous in the upper (adaxial) shallow sulcus;

laminae 15-35 cm long, 6-18 cm broad, ovate-elliptic to ovate-oblong, acute or very short

acuminate at the apex, rounded to the obtuse and often unequal base, entire, the laminae

drying chartaceous to stiffly chartaceous, smooth or very slightly scabrous above, glabrous

or sparsely strigose above, strigose beneath with thin stiff hairs 1-2 mm long on the major

and minor veins, venation pinnate with (12) 14-20 pairs of major secondary veins. Male

inflorescences spikelike, 1-8 from the axil of a leaf or fallen leaf, often panicle-like when

leaves have fallen or are undeveloped on terminal twigs bearing spikes, the spikes 10-25

cm long, flowers borne in fascicles at first enclosed in narrowly ellipsoid or narrowly ovoid

bracts borne spirally on the rachis of the spike, rachis and bracts usually densely yellowish

brown strigose, the primary floral bracts deciduous and each fascicle producing several

flowers; male flowers ca. 5 mm long, perianth of 3 linear and 3 narrowly deltoid tepals

united near the base, filaments to 7 mm long, anther 0.5-1 mm long. Female inflorescence

basically a compact raceme, solitary or paired, usually borne in leafless axils on a terminal

twig and paniculate in form, the spikelike racemes 5-20 cm long, the compound terminal

inflorescence to 40 cm long, the flowers solitary and at first enclosed in narrowly ovoid

bracts (resembling the stipules), bracts and rachis densely strigose with yellowish brown

hairs 1-2 mm long, bracts ca. 1 cm long and narrowed at the base, deciduous, flowers

solitary from within the bract on short pedicels; female flowers 10-15 mm long at early

anthesis, pedicels and outer perianth surface densely strigose or sericeous, ovary ca. 3 mm

long, strongly 3-angled. Fruit an achene tightly enclosed and united near the base to the

dry persisting long-winged perianth, fruiting perianth 4-6 cm long, perianth tube ca. 14

mm long and 8 mm thick, the wings 3-5 cm long, oblanceolate, 6-10 mm broad, thin

chartaceous, venation pinnate and raised on both surfaces, the 3 wings usually parallel

and not spreading, the inner perianth parts adnate to the tube for 4-7 mm from the base;

achene ca. 11 mm long and 7 mm broad, ovate in outline and strongly 3-angled, the faces

slightly convex (deeply concave when undeveloped) and lustrous dark brown, tightly

enclosed by the perianth tube (especially near the base) and difficult to free from the tube,

style short, style branches usually breaking off.

Trees of both the seasonally very dry deciduous forest formations and the wet

evergreen formations of the Pacific slope from near sea level to 300 (850) m

elevation in Costa Rica; usually flowering in January and February and with fruit

in February and March. The species, as presently defined, ranges from central

Mexico to southernmost Costa Rica; however, these plants will probably prove to

be the northern representatives of a more broadly defined South American species

(see below).

138 FIELDIANA: BOTANY

Triplaris melaenodendron is readily recognized by its large panicle-like compound

inflorescences, bright pinkish red or reddish brown fruit with three long papery

wings, the hollow distal stems usually inhabited by aggressive little ants, the

shoot apex at first enclosed in a stipule that leaves a prominent scar around the

stem, the rather large leaves with numerous secondary veins, and the bark on

older trees breaking off in flakes or strips. This species appears to intergrade with

T. cumingiana Fischer & Meyer, and some of our collections may be intermediates

between the two (A. Jiminez 3799, Molina et al. 18245, Skutch 4242; all from the

General Valley). Triplaris cumingiana differs in having somewhat narrower leaves

with a greater number of secondary veins and with the inner perianth parts

adnate for only 2-5 mm on the pistillate perianth tube. All these plants will

probably prove to be subspecific elements of more broadly defined species. Con-

sidering the present state of knowledge, I believe it best to follow Duke's treat-

ment in the Flora of Panama (Ann. Missouri Bot. Card. 47:353-359, 1960); see also

the remarks by Standley & Steyermark in the Flora of Guatemala (Fieldiana, Bot.

24, pt. 4:136-137. 1946).

Common names used in Central America for these trees are Canilla de Mula,

Callito, Hormigo, Mulato, Tabaco, Tabaco de Monte, Tabacon. The name Mulato refers

to the patchy, variously colored bark. The Brunka name is Turi-Svan-Kra. Many

of our collections come from along streams and in flood plains, but it may be that

these individuals have more accessible lower branches than those growing in

forests. Included among the collections placed here is a form (Allen 5923 and 6022)

with larger fruiting perianth and whitish trunk; see the remarks in Paul Allen's

book, The Rain Forests of Golfo Dulce (1950, p. 348).

CHENOPODIACEAE

Herbs (in ours), shrubs, or rarely small trees, bisexual or unisexual, glabrous or puber-

ulent, sometimes with inflated or flattened hairs, the stems frequently semisucculent,

often with anomalous anatomy; stipules absent. Leaves alternate, less often opposite or

whorled, simple, often becoming conspicuously smaller and narrower on distal stems,

often slightly succulent (reduced to scales in some genera), sessile or petiolate, margin

entire to lobed or with conspicuous teeth. Inflorescences often of condensed cymes of few

closely approximate or appressed flowers (glomerules), the glomerules often borne on

distal axillary or terminal unbranched (spikelike) or branched (panicle-like or thyrselike)

stems, glomerules often subtended by reduced leaves, the flowers often subtended by a

bract and bracteoles; flowers bisexual or unisexual, usually small and radially symmetrical,

perianth of 1 whorl and calyx-like, thin or semisucculent, of 2-5 parts (rarely absent or 1),

free or united near the base, persisting in fruit, stamens as many as the perianth parts or

fewer, opposite the perianth parts, free or united at the base to the perianth or to a disk,

filaments flattened to filiform, bent inward in bud, anthers dorsifixed and introrse, 2- or

4-thecous, pollen polyporate; ovary superior or united at the base to the perianth or disk,

1-locular, with 1 basal ovule borne on an erect funicle. Fruit a utricle, often tightly enclosed

within the persisting perianth or bracts, indehiscent (rarely circumscissle); seed variously

oriented within the fruit, and with much endosperm, embryo strongly curved, annular to

hippocrepiform (circular to horseshoe-shaped), rarely spiral or conduplicate.

A large family well represented in deserts, semideserts, and saline environ-

ments of the Northern Hemisphere, especially common in eastern Europe and

western Asia. The family is poorly represented in the tropics, and none of its

species seem to be members of Costa Rica's indigenous flora. The slightly succu-

lent or edible alternate leaves, small greenish flowers with stamens borne at the

base of the sepals, lack of petals, sessile stigmas, solitary locule with ovule borne

on a basal funicle, and smooth lenticular seeds help distinguish our species. The

BURGER: FLORA COSTARICENSIS 139

strongly curved embryo surrounding much endosperm is a characteristic of the

order Centrospermae, also called the Chenopodiales or Caryophy Hales. Our spe-

cies are easily confused with members of the Aizoaceae and some Ama-

ranthaceae.

la Plants growing wild, herbaceous weeds of open sites; basal and lower leaves rarely

more than 10 cm long; flowers not subtended by bracts or bracteoles; fruit loosely

enclosed in the dry slightly enlarged and separate perianth parts Chenopodium

Ib Plants cultivated for food and ornament; basal leaves often exceeding 15 cm in length;

flowers subtended by small bracts; fruits tightly enclosed within the united perianth

parts or bracts 2a

2a Flowers bisexual; plants cultivated for the edible root, edible leaves or colorful leaves;

fruit enclosed by the hard and woody perianth, bracts and bracteoles remaining free

(Acelga, Remolacha) Beta

2b Flowers unisexual, plants cultivated for the edible leaves; fruits enclosed in a nut-

like structure formed by the 2 subtending bracts and often with spines

(Espinaca) Spinacia

BETA Linnaeus

Herbs, annual, biennial or perennial, roots often thick and fleshy. Leaves whorled or in

a rosette at the base, alternate on the stems, laminae entire or sinuate, venation pinnate.

Inflorescences small clusters of flowers, solitary in leaf axils or terminal, often on distal

spikelike or paniculate branches, a bract and 2 bracteoles subtending each flower; flowers

bisexual, small, greenish or reddish, perianth urceolate and 5-lobed, united at the base to

the ovary and to other flowers, becoming stiff in fruit, stamens 5, borne on the side of the

ovary, anthers oblong, ovary united to perianth and stamens near the base and appearing

to be sunken into a disk, with 3 (2-5) short stigmas. Fruit enclosed within the hardened

and nearly closed perianth, often the product of several flowers united together, pericarp

free of the seed and united to the perianth at the base; seed horizontal (with the long axis

at right angles to the floral axis), orbicular to reniform, embryo annular or subannular.

A genus of about 12 species native to northern Africa, Europe, and Asia. This

genus includes the many cultivated varieties of beets grown for their nutritious

roots, the leaves used for green vegetables, and, in some varieties, the colored

foliage for ornamental plantings.

Beta vulgarisL., Sp. PI. 222. 1753.

Herbs, annuals, stems to 1.3 m tall and usually produced in the 2nd growing season,

sparsely puberulent with thin whitish hairs. Leaves dimorphic, the basal leaves to 50 cm

long, with long petioles and broad laminae abruptly narrowed at the base, leaves of the

stem becoming progressively smaller upward on the stem. Inflorescences (glomerules)

subtended by lanceolate leaves on distal stems; perianth parts narrowly oblong to spat-

ulate, often keeled on the back.

An important root and vegetable crop, grown at higher elevations in the tropics.

Generally referred to as Acelga and Remolacha, the species can be divided into two

general groups of cultivars:

Cicla group: Includes the leafy vegetable and ornamental varieties, such as the leaf

beet, spinach beet, chard, and Swiss chard. The root is not greatly thickened,

the leaves become as much as 25 cm broad and often have a thick midrib. The

ornamentals often have reddish leaves.

Crassa group: Includes those grown for the root, such as the garden beet, red beet,

yellow beet, and sugar beet. The root may be short or long, but is thick, from

white or yellow to dark wine red.

140 FIELDIANA: BOTANY

CHENOPODIUM Linnaeus

Herbs or subshrubs, annual or perennial, erect or procumbent, often much branched,

often with minute scurfy or inflated farinose hairs. Leaves alternate and simple, often

larger near the base and becoming progressively smaller on the distal branches, the larger

laminae with petioles, laminae entire to pinnatifid, usually gradually narrowed at the base

and decurrent on the petiole, gland dots or pale whitish farinose scales often present.

Inflorescences basically of small groups of flowers (glomerules) solitary or congested

together, axillary or terminal, often subtended by reduced leaves, the axillary or distal

flowering stems often spicate or paniculate in form, the flowers lacking both bracts and

bracteoles; flowers bisexual or unisexual, radially symmetrical, perianth of (3) 5 parts, free

or united near the base, broadly imbricate, white or greenish, drying thin; stamens 3-5,

often variable in number on the same plant, filaments short and flat, free or united at the

base, anthers introrse, ovary superior and subglobose, style short or none, stigmas 2-5,

fimbriate to subulate. Fruit an indehiscent utricle, subglobose to ovoid, the pericarp mem-

branaceous or fleshy, free or adherent to the seed, perianth persisting and loosely en-

closing the fruit; seed usually cochleate-lenticular, held vertically (with long axis parallel

to the floral axis) or horizontally (with long axis at right angle to the floral axis), embryo

coiled into a ring (annular) around the central endosperm.

A chiefly temperate genus with perhaps 100 species, best represented in north-

ern North America. Most species are weedy, and a few are now distributed

around the world. Our representatives seem not to be native, and the key includes

a species not yet recorded but likely to be introduced. Plants of the genus are

sometimes used as green vegetables; the seeds and inflorescences of C. quinoa

Willd. provide a grain for the Andean people of Ecuador and Peru. Some are used

for condiments, and one of our species is used as a vermifuge; see the treatment

of this genus in the Flora of Guatemala (Fieldiana, Bot. 24, part 4:139-143, 1946).

la Plants strongly aromatic, with minute yellowish glands on the undersides of leaves,

not farinose; seeds black and lustrous, ca. 0.7 mm broad, embryo not completely

encircling the endosperm, seeds both vertical and horizontal within a glomerule of

flowers; inflorescences paniculate or spikelike, flowers and glomerules mostly sessile;

occasional in Costa Rica, 1,000-2,600 m elevation C. ambrosioides

Ib Plants not strongly aromatic, yellowish gland dots absent, with gray or whitish in-

flated hairs and appearing farinose (as if dusted with flour); seeds more than 1 mm

broad, dull or lustrous, horizontal, embryo completely encircling the endosperm; rare

or unrecorded in Costa Rica 2a

2a Seed with a blunt edge, 1-2 mm in diameter, testa with radial furrows, the pericarp

easily removed from the seed; inflorescences to 10 cm long and leafless distally; stems

often reddish; a recent introduction C. album

2b Seed with a sharp edge, 1.2-1.5 mm in diameter, testa surface with rounded pits,

pericarp difficult to remove from the seed; inflorescences to 5 cm long, often branched,

leafless except at the base; unrecorded from Costa Rica and not included in the

descriptions: C. murale L.

Chenopodium album L., Sp. PI. 219. 1753.

Herbs to 1.5 m tall, usually much branched, stems often with longitudinal stripes of pale

yellow and green (dry), leafy internodes 0.5-10 cm long, minutely puberulent with

crooked whitish hairs but becoming glabrous. Leaves very variable on the same plant with

the distal leaves becoming linear-oblong, petioles with thin lateral margins; larger laminae

2-8 cm long, 1-5.5 cm broad, broadly rhombic-ovate to narrowly oblong, margins entire

or with up to 10 shallow teeth on each side, upper surface smooth and glabrous, lower

surface with minute (0.1 mm) translucent or whitish oblong scales (inflated hairs) and

farinose in appearance, venation pinnate or subpalmate with a prominent pair of basal

secondaries. Inflorescences of axillary stems with alternate sessile flower clusters and

spikelike or with some flower clusters (glomerules) stalked and panicle-like, distal flower

clusters without subtending reduced leaves; flowers 1-1.5 mm broad, perianth with fari-

nose scales, ovary with 2 stigmas. Fruit with pericarp easily removed from the seed, seed

BURGER: FLORA COSTARICENSIS 141

1.2-1.6 mm in diameter, circular or slightly ovate in outline, thick-lenticular and with a

rounded edge, dull or shiny black.

A weedy species of worldwide distribution, but only recently collected in Costa

Rica by J. M. Orozco near Cartago and by J. Gdmez-Laurito at Faldas del Irazu;

both collections were made in June. The species is recognized by the grayish

coloring and scales. In the area near Faldas del Irazu, it is found as a weed in

potato fields and is called Mejicano.

Chenopodium ambrosioidesL., Sp. PI. 219. 1753. C. anthelminticum L., Sp. PI.

220. 1753. Figure 38.

Herbs, upright or leaning over other plants, 0.1-1.5 (2) m tall, annual or perennial,

usually much branched, stems often with longitudinal pale-colored ridges, leafy inter-

nodes 0.5-30 mm long, 0.5-4 mm thick, glabrous or very minutely and sparsely puber-

ulent, strongly aromatic with a sharp unpleasant odor. Leaves very variable in size and

shape on the same plant, the distal stems with smaller (6-15 mm x 1-4 mm) narrowly

oblanceolate laminae, petioles 1-10 mm long, with narrow lateral margins continuous

with the laminae margins; larger laminae 2-7 (10) cm long, 1-2.5 (4.5) cm broad, elliptic

to obovate in general outline, acute at the apex, gradually narrowed to the base and

decurrent on the petiole, margins entire in small laminae but with 2-6 short teeth or lobes

on each side of the larger leaves, sometimes pinnatifid, laminae drying thin and char-

taceous, smooth and glabrous above, usually glabrous and glandular punctate beneath but

the small gland dots often difficult to see on dried material, venation pinnate with 3-6

pairs of secondary veins. Inflorescences spikelike or paniculate, solitary, axillary or termi-

nal, usually made up of an unbranched rachis with clusters (glomerules) of sessile or

subsessile flowers often subtended by reduced linear leaves, flowering rachis minutely and

sparsely puberulent with thin whitish hairs less than 0.4 mm long; flower buds small and

globose or somewhat flattened distally, 0.6-1.5 mm in diameter, bisexual and male flowers

found on the same plant, perianth 3- to 5-parted, greenish, rounded; anthers ca. 0.3 mm

long, stigmas 3 or 4. Fruit with pericarp easily removed from the seed, seed cochleate-

orbicular in outline, lenticular with rounded edges, smooth and lustrous, black.

A weedy species of open early secondary vegetation. Collected only in the

highlands (1,000 to 2,600 m) in Costa Rica and probably flowering throughout the

year, but with fertile collections made only in January, March, August, and

October. This species ranges widely as a weed in both the tropics and subtropics,

but may be native to North America.

Chenopodium ambrosioides is recognized by its unpleasant odor, alternate leaves

that vary from toothed to entire and become smaller and narrower as they

progress up the stem, small flowers in small clusters of three to eight often on

spikelike stems, and restriction to open weedy habitats. Despite its strong odor,

this species is used for flavoring food in Guatemala. The species has been used

effectively as a medicinal plant for the removal of intestinal parasites.

SPINACIA Linnaeus

Herbs, annual, erect, unisexual (dioecious). Leaves alternate, petiolate, laminae usually

broadest at the base, often with lobes or teeth on the base and lower half, triangular to

hastate, becoming entire and lanceolate distally. Flowers small, rarely bisexual; male flow-

ers in terminal spikes or panicles, perianth 4- or 5-lobed, stamens 4 or 5; female flowers

mostly axillary, subtended by 2 bracts which become thick and united in fruit, ovary with

4 or 5 exserted stigmas. Fruit enclosed within the thick united bracts; seed vertical.

A genus of five species, originally from western Asia and growing best in cooler

climates. This is the species referred to as Espinaca or Spinach in Europe, but these

names are also used for Tetragonia tetragonioides of the Aizoaceae (q,v.) which

grows better in warm climates.

142 FIELDIANA: BOTANY

Spinacea oleraceaL., Sp. PI. 1027. 1753.

Herbs to 0.8 m tall, glabrous. Leaves dimorphic, basal and in a rosette or cauline and

alternate, laminae broadly ovate and with basal lobes to oblong or narrowly obovate,

becoming smaller and narrower up the stem, lanceolate in the distal flowering branches

(inflorescences), glabrous, venation pinnate. Fruit enclosed in the thickened carpel-like

bracts, occasionally with spines, capsule-like.

AMARANTHACEAE

Herbs, shrubs or clambering plants, rarely small trees, bisexual or rarely unisexual,

puberulent or glabrous, stems often marked with reddish coloring; stipules absent. Leaves

alternate or opposite, petiolate, simple and entire (rarely somewhat crenate or serrate), the

lamina often decurrent on the petiole, midvein often extending beyond the lamina as a

small mucronate tip, venation usually pinnate. Inflorescences axillary or terminal, solitary

or several, the flowers usually borne in congested fascicles or small condensed cymose

glomerules and these often arranged in heads or spikes, the flowers usually subtended by

a basal bract and 2 lateral bracteoles, these usually stiff and dry and sometimes colorful,

glomerules occasionally with sterile flowers modified to aid protection or dispersal. Flow-

ers bisexual or unisexual, generally small and radially symmetrical, sessile or subsessile,

perianth (1-3) 4 - or 5-parted in 1 or 2 whorls, usually stiff, dry and imbricate, usually

narrow, greenish to whitish, pink or reddish; stamens usually as many as the perianth

parts and opposite them (rarely 1-3), straight in bud, filaments free or united to form a

short or long tube, the tube often with appendages (pseudostaminodia) between the

anther-bearing filaments, anthers 2- or 4-thecous, medially attached and introrse; ovary

superior and 1-locular, 2- or 3-carpellate, with 1 (more rarely several to many) ovules

pendulous from long erect funicles from a basal placenta, styles 2 or 3 (1-8), stigmas

capitate or filiform. Fruit a 1-seeded utricle or nut, less often a several-seeded capsule or

berry, often with circumscissle dehiscence, sometimes indehiscent, the fruit usually in-

cluded within the persisting bracts, bracteoles, and perianth parts; seed lenticular or

reniform, often cochleate-orbicular in outline, often lustrous and smooth, embryo annular

or U-shaped, with perisperm.

The Amaranthaceae probably exceed 60 genera and 900 species in number,

ranging throughout the tropics and warm temperate areas. They are especially

well represented in Africa and the .Americas. The family is closely related to the

Chenopodiaceae, but that family is often succulent, with the perianth more sepal-

oid and the filaments bent inward in bud. The Amaranthaceae are usually easy to

recognize because of their simple entire estipulate leaves, small flowers, stiff dry

grasslike floral bracts and perianth parts, stamens opposite the perianth parts,

and basal ovule(s) often producing shiny lenticular seeds with curved embryo.

Most of our species are weeds of recently cleared soil or early secondary

vegetation. Some species of Amaranthus, Celosia, Gomphrena, and Iresine are used

in ornamental plantings and in dried flower arrangements because of their col-

orful, often long-persisting, inflorescences. Several species are used as potherbs,

and the grain amaranths (Amaranthus spp.) have been an important source of food

in some cultures. For a recent review of their use as vegetables, see: Vegetables for

the Hot Humid Tropics, Part 6, Amaranthus and Celosia (1979), available from the

Mayaguez Institute of Tropical Agriculture, P.O. Box 70, Mayaguez, Puerto Rico,

00708.

KEY TO THE GENERA OF AMARANTHACEAE IN COSTA RICA

la Leaves alternate along the stem; stigmas 2-8; fruit a 1-seeded utricle or capsule or a

many-seeded capsule or berry, the fruit loosely enclosed within persisting perianth

and bracts 2a

Ib Leaves opposite along the stem (but sometimes alternate within the inflorescences);

stigma 1-3; fruit 1-seeded, usually a thin- walled utricle or indehiscent 5a

BURGER: FLORA COSTARICENSIS 143

2a Fruit a many-seeded capsule (or occasionally berry-like); ovary with several to

many ovules, stigmas 2-8 and usually exerted beyond the perianth parts; fil-

aments united to form a tube or a shallow cup 3a

2b Fruit a 1-seeded utricle; ovary with a single basal ovule, stigmas 2 or 3 and

exerted or included within the perianth; filaments free or united for ca. Va their

length to form a cup 4a

3a Capsule or utricle larger than the perianth parts, the perianth parts often

reflexed; seeds usually more than 10, stigmas 2-8 and equal to or longer

than the style; inflorescences paniculate arrangements of racemes; leaves

usually ovate; shrubs of evergreen forest formations Pleuropetalum

3b Capsules or utricles not exceeding the erect persisting perianth parts; seeds

usually fewer than 9, stigmas 2 or 3 and shorter than the style; inflorescences

spicate (or terminal and broadly flattened or folded and brightly colored in

some horticultural forms); leaves usually lanceolate; herbs of open second-

ary growth or grown in gardens Celosia

4a Erect or prostrate herbs of open secondary vegetation or sometimes cultivated

for the colorful inflorescences (densely flowered spikes in terminal paniculate

arrangements) or for the small edible grains; bracts equal to or larger than the

perianth; flowers mostly unisexual, filaments free; seeds 0.5-1.5 mm broad

Amaranthus

4b Vining or clambering shrubs of primary and secondary vegetation; bracts much

shorter than the perianth parts; flowers bisexual, filaments united at the base,

seeds 1.5-2.5 mm broad Chamissoa

5a Inflorescences elongate (5-20 cm) spikes with the fruiting flowers or flower clusters

(glomerules) becoming reflexed (pointing backward toward the base) on the spike;

fruit tightly enclosed within perianth and bracts, bracts and bracteoles usually with

spine tips; anthers 2- or 4-thecous, style 1 with a capitate stigma 6a

5b Inflorescences of short congested spikes or heads or racemes, often in paniculate

arrangements, flowers or glomerules not strongly reflexed on the inflorescence axis

or branches; fruit loosely or tightly enclosed within the bracts and perianth, bracts

and bracteoles rarely with spine tips (in Alternanthera spp.); anthers 2-thecous, styles

1-3, stigmas filiform, cylindrical, or capitate 7a

6a Spine tips of perianth and bracts straight, not becoming hooked in fruit, flowers

solitary and not borne in glomerules or fascicles together with sterile (modified)

flowers; perianth parts without prominent ribs; seeds 0.8-1.2 mm

broad Achyranthes

6b Spines of some perianth parts and bracts becoming hooked at the apex in late

stages, glomerules composed of fertile and sterile flowers; perianth parts with

prominent longitudinal ribs abaxially; seeds 1-2.5 mm broad Cyathula

7a Leaves mostly linear and succulent, clasping the stem at their base (amplexicaul) and

lacking a well-defined petiole; usually procumbent succulent plants growing at the

seashore; filaments united at the base to form a short tube, style with 2 stigmatic

branches Philoxerus

7b Leaves neither linear nor succulent, with well-defined petioles and not clasping the

stem; plants mostly erect 8a

8a Inflorescences of spikes in paniculate arrangements; flowers unisexual or bisexual,

with a conspicuous tuft of whitish hairs developing from under the fruit or from

under the fruiting perianth; stigmas 1-3 and filiform 9a

8b Inflorescences spikes or gloermules, the spikes and glomerules solitary or in few-

branched cymes or corymbs; flowers bisexual, a tuft of hairs not produced under

flower or fruit in late stages; stigmas 1 or 2 and filiform to capitate lOa

9a Flowers bisexual, stigma solitary and 2-lobed; outer 3 perianth parts much

broader than the inner, basal hairs arising from within (adaxially to) the peri-

anth; clambering shrubs Pfafft°

9b Flowers unisexual or bisexual, stigmas 2 or 3 and filiform to deltoid; perianth

parts subequal, basal hairs arising from the outside (abaxial) base of the peri-

anth; erect herbs or clambering shrubs, often common in secondary growth and

with conspicuous whitish inflorescences Iresine

lOa Stigma 1, capitate or short-cylindric, anthers borne on slender filaments; bracts and

bracteoles much shorter than the perianth parts, bracteoles lacking dorsal (abaxial)

crests or ridges Alternanthera

144 FIELDIANA: BOTANY

lOb Stigmas 2 and filiform, anthers sessile on the staminal tube; bracts and bracteoles

equaling or exceeding the perianth parts, usually with crenate or serrate crests on the

back (abaxially) of the distal half of the bracteoles (in our species) Gomphrena

ACHYRANTHES Linnaeus

Herbs or subshrubs, annuals or perennials, usually becoming woody at the base, erect

or decumbent, puberulent or glabrous; stipules absent. Leaves opposite and simple, pet-

iolate, laminae entire and pinnately veined, usually puberulent. Inflorescence solitary,

terminal or axillary, a spike or spikelike raceme, the flowers on very short pedicels and

becoming reflexed on the inflorescence, flowers subtended by a bract and 2 bracteoles, the

bracts and bracteoles of similar size, often with 1 or 2 stiff straight sharp spines, the

bracteoles with the spine far exceeding the thin translucent lobes (in ours), flowers bisex-

ual, essentially radially symmetrical, sepals 4 or 5, free, subequal, narrow and concave,

longitudinally ribbed, glabrous or puberulent, becoming stiff and dry, persisting but not

enlarging in fruit, stamens usually as many as the perianth parts and opposite them (rarely

2 or 3), united near the base to form a short tube, the tube with pseudostaminodial

appendages, anthers 2- or 4-thecous, opening introrse, ovary ovoid and often narrowed

at the base, 1-locular, the solitary ovule borne from the basal placenta on a long funicle,

style and stigma 1. Fruit an indehiscent utricle (or like an achene when the fruit wall is

stiff), tightly enclosed by bracts and perianth, usually reflexed and appressed against the

axis of the inflorescence; seeds somewhat flattened, the embryo curved.

A weedy genus of probably five to 10 species with two species now widely

naturalized in the neotropics. The genus is thought to have been confined to the

Old World until recently. These plants are characterized by the opposite leaves,

long spikelike inflorescences with flowers and fruit becoming reflexed and ap-

pressed on the rachis, the bracts with stiff straight spines, and the utricles rightly

enclosed within perianth and bracts.

la Larger laminae ovate to elliptic and with acuminate apices; sepals 5-7 mm long;

0-1,200 m A. aspera

Ib Larger laminae broadly obovate to obovate-orbicular, abruptly rounded or truncated

at the apex; sepals ca. 4 mm long; 0-400 m A. indica

Achyranthes aspera L., Sp. PI. 204. 1753. Centrostachys aspera (L.) Standl., J.

Wash. Acad. Sci. 5:75. 1915. Figure 25.

Herbs or subshrubs, 0.4-1.5 (2) m tall, woody in the lower parts and usually erect, leafy

internodes (1) 2-15 cm long, 2-5 mm thick, puberulent with whitish ascending sericeous

hairs ca. 1 mm long, longitudinally striate and brown when dry, usually terete. Leaves

opposite and decussate, often quite variable on the same plant, petioles 3-30 (40) mm long,

densely sericeous, with lateral margins forming an adaxial sulcus near the base and

continuous with the lamina margins distally; laminae 2-18 (25) cm long, 1-8 (10) cm broad,

ovate to elliptic or slightly obovate, apically obtuse on the smaller leaves but acuminate on

the larger leaves, obtuse to attenuate at the base, margin entire or obscurely crenate (dried)

and decurrent on the petiole, the laminae drying thin- to stiff-chartaceous, smooth on

both surfaces, sparsely to densely sericeous to strigillose with the lower surface more

densely puberulent than the upper, with 3-7 pairs of major secondary veins arising at

angles of 40°-60°. Inflorescence usually solitary and terminal, spikes 10-40 cm long, 6-14

mm broad, the flowers at first closely crowded and divergent but soon becoming separate

and reflexed, bracts and bracteoles with 1 or 2 sharp stiff spines, 2-4 mm long, pedicels

0.1-0.5 mm long; flowers essentially subsessile and becoming reflexed on the spike, sepals

5-7 mm long, lanceolate, very stiff. Fruit enclosed by the tightly appressed bracts and

perianth, reflexed and appressed against the axis of the spike, utricle ca. 3 mm long and

1 mm thick, cylindrical with a truncated apex and rounded base, difficult to separate from

the enclosing perianth.

Plants of open weedy sites and early successional stages from sea level to about

1,200 m elevation in both evergreen and seasonally very dry deciduous areas;

BURGER: FLORA COSTARICENSIS 145

flowering and fruiting collections have been made from November to April in our

area. The species appears to be an introduction from the Old World.

Achyranthes aspera is recognized by the long slender spikes with retrorse flowers

and fruits enclosed in spine-bearing bracts and stiff perianth parts, the thin

acuminate leaves, and open-branched growth habit. The larger leaves with acute

or acuminate apices and the perianth parts always 5 mm long or more are consis-

tent traits that distinguish this species from A. indica in our area. The spines on

the bracts and bracteoles are an effective aid in dispersal. The plants are called

Rabo de Chanco, Mosotillo, and Mozote.

Achyranthes indica (L.) Miller, Card. Diet. ed. 8, no. 2. 1768. A. aspera ft indica

L., Sp. PI. 204. 1753. Centrostachys indica (L.) Standl., J. Wash. Acad. Sci. 5:75.

1915. Figure 25.

Herbs or subshrubs, to 1 m tall (rarely taller), woody at the base, leafy internodes (1)

3-15 cm long, 1-4 mm thick, puberulent with thin whitish ascending hairs 0.2-1 mm long,

grayish and longitudinally striate when dry, terete or angular. Leaves opposite and decus-

sate, petioles 1-8 (15) mm long, poorly differentiated from the lamina, with lateral margins

continuous with the lamina margins, densely sericeous; laminae 1.5-6 (8) cm long, 1-4 cm

broad, broadly obovate to obovate-orbicular or rhombic-orbicular (broadly elliptic in small

leaves), rounded or abruptly obtuse at the apex with a small acute tip, usually cuneate at

the base, margin entire and decurrent on the petiole, the laminae drying thin- to

stiff-chartaceous, smooth to the touch and puberulent on both surfaces, densely sericeous

beneath with thin ascending whitish hairs ca. 0.3 mm long, with 5-8 pairs of major

secondary veins. Inflorescences terminal (often terminal on short axillary shoots), 10-30

cm long, 5-10 mm broad, axis of the spike with spreading thin whitish hairs, flowers at

first crowded and divergent but soon becoming separate and reflexed, bracts and bracte-

oles with stiff spines 2-3 mm long; flowers subsessile, perianth parts ca. 4 mm long,

lanceolate, pale green, tightly enclosing the fruit. Fruit tightly enclosed within the spinose

bracts and the stiff dry perianth parts, reflexed on the flowering axis, utricle some-

what cylindrical, ca. 2 mm long and 1 mm thick, difficult to separate from the enclosing

perianth.

Plants of open bare soils and early successional vegetation below 400 m el

evation in both evergreen and seasonally very dry deciduous formations in Cen-

tral America; probably flowering throughout the year, but with collections in our

area only from December through March, July, and August. The species is proba-

bly an introduction from the Old World.

Achyranthes indica is recognized by its long slender spikes with retrorse flowers

and fruits tightly enclosed in spine-bearing bracts and stiff dry perianth parts,

puberulent leaves with blunt rounded apices, and short stature in open or recently

disturbed sites. The bluntly rounded leaves and shorter perianth parts consis-

tently distinguish these plants from A. aspera in our area. The two taxa were

treated as varieties in the Flora of Panama (Ann. Missouri Bot. Card. 48:24, 1961),

but they seem to behave like two different species in Central America.

ALTERNANTHERA Forsskal

Herbs or small subshrubs, annuals or perennials, prostrate, erect or clambering, bisex-

ual, usually puberulent in early stages. Leaves opposite, petioles present, poorly differ-

entiated or absent; laminae simple and essentially entire, puberulent or glabrous, venation

pinnate. Inflorescences axillary or terminal, 1-3 at a node, sessile or pedunculate, capitate

or spicate, each flower usually with a subtending bract and 2 lateral bracteoles, the bracts

and bracteoles similar or differing in size and shape. Flowers bisexual, radially sym-

metrical, sessile or subsessile, perianth 5-parted, the perianth parts free with the inner and

outer whorls similar or differing in size and form, usually concave, dry and stiff, ovate to

ACHYRANTHES

indica

A aspera

FIG. 25. Amaranthaceae: Costa Rican representatives of Achyranthes and Cyathula. Note

the hooked hairs of Cyathula.

146

BURGER: FLORA COSTARICENSIS 147

lanceolate and usually acute at the apex, longitudinal ribs often thickened on the abaxial

side, whitish to purple or pale yellowish, persisting but not enlarging in fruit; stamens 3-5,

united in the lower part to form a tube, usually with staminode-like projections on the

tube between the anther-bearing filaments, anthers oblong and 2-thecous, introrse; ovary

superior and globose to ovoid or obovoid, usually laterally flattened, 1-locular with 1 ovule

borne on an elongate funicle from a basal placenta, style short or long, stigma solitary.

Fruit a thin-walled utricle, indehiscent, tightly or loosely enclosed within the persisting

perianth; seeds cochleate-reniform to orbicular or ellipsoid in outline, usually lenticular in

cross section, smooth and lustrous, embryo annular.

A genus of perhaps 100 species best represented in the Neotropics, but with

many species now widely ranging as weeds in tropical and subtropical to warm-

temperate areas. Our species are distinguished by their small whitish heads

(except in A. costaricensis) , densely crowded little flowers subtended by usually

similar bracts and bracteoles, longitudinal ribs on the back of the outer perianth

parts often thickened, both bracts and perianth parts often terminating in sharp

spinelike tips, and preference for open early secondary growth habitats (except in

A. costaricensis). This genus is very similar to Gomphrena (q.v.), and the nomen-

clature of the two genera has been badly confused (see J. A. Mears in Proc. Acad.

Nat. Sci. Philadelphia 129:1-21, 1977, and in Taxon 29:85-95, 1980). This treat-

ment is based on the annotations and identifications of our material by James

Mears.

la Inflorescences pedunculate, not subtended by leaves; internodes often more than 5 cm

long, leaves usually 4-14 cm long, plants usually erect; seeds lustrous black 2a

Ib Inflorescences sessile in the axils of leaves, less than 12 mm long (except in A. cos-

taricensis); plants erect, clambering or prostrate; seeds lustrous brown or reddish

brown 5a

2a Perianth parts sparsely puberulent or glabrous, inflorescences rarely more than 1

cm long, floral bracts 1-1.5 mm long and often with a sharp tip; 1,000-2,200 m

altitude A. lanceolata

2b Perianth parts densely puberulent or ciliate, inflorescences often more than 1 cm

long, floral bracts more than 2 mm long 3a

3a Floral bracts 4-6 mm long with an erose or ciliate margin and often with very

short (0.5 mm) stiff white hairs; seed ellipsoid, ca. 1.8 mm long; 0-1,000 m but

rarely collected in southern Central America A. brasiliana

3b Floral bracts 2-4 mm long, with an entire margin; 0-1,500 m 4a

4a Perianth with long (4 mm) thin whitish hairs; seeds ca. 0.7 mm long; leaves

lanceolate to very narrowly elliptic; common in Costa Rica in wet evergreen

formations A. laguroides

4b Perianth with short (0.5 mm) hairs; seeds ca 1.5 mm long; leaves elliptic to

elliptic-ovate; rare in Costa Rica and usually found in seasonally deciduous

areas A. pubiflora

5a Plants usually prostrate, laminae rarely exceeding 2.5 cm in length; plants rarely

collected in Costa Rica; fruit truncated at the apex and with the style usually persis-

ting 6a

5b Plants erect,clambering, or occasionally forming low mats; laminae often exceeding 2.5

cm in length; fruit rounded or emarginate (notched) at the apex 7a

6a Perianth with thin hairs minutely hooked (*25) at the ends; fruit tightly enclosed

within the persisting perianth; floral bracts ca. 3 mm long A. caracasana

6b Perianth with thin white hairs lacking very minute hooks at the tip; fruit easily

removed from the persisting perianth; floral bracts 1-2 mm long

A. paronychioides

7a Plants erect or clambering on others and forming mats; often found in open sunny

early secondary sites, common wild plants in evergreen and partly deciduous

areas . . . . 8a

148 FIELDIANA: BOTANY

7b Plants erect; found in gardens or in deep forest shade 9a

8a Laminae to 9 cm long and 4 cm broad; floral bracts 2 mm long, perianth ca. 4 mm

long; fruit tightly enclosed within the persisting perianth; suborbicular

A. tenella

8b Laminae to 6.5 cm long and 2 cm broad; floral bracts 0.5-1 mm long, perianth

ca. 2 mm long; fruit loosely enclosed by the perianth, lenticular, notched at the

apex A. sessilis

9a Laminae 7-16 cm long; inflorescences 1-4 cm long and drying brownish, perianth

parts to 8 mm long; rarely collected plants of wet forest interiors below 1,000 m

elevation A. costaricensis

9b Laminae to 6 cm long, often purplish or marked with varied colors; inflorescences less

than 1 cm long, perianth parts ca. 3 mm long; common ornamental plants of parks and

gardens A. bettzichiana

Alternanthera bettzichiana (Regel) Voss, in Steb. & Voss, Vilmorin's Blu-

mengartnerei 869. 1896. Telanthera bettzichiana Regel, Gartenfl. 11:178. 1862.

Achyranthes bettzickiana (Regel) Standl., N. Amer. Fl. 21:138. 1918. Alternanthera

bettzickiana (Regel) Standl., Publ. Field Columbian Mus., Bot. Ser. 3:254. 1930.

Figure 26.

Cultivated herbs or subshrubs of small (20-80 cm) stature, with many lateral branches

and bushlike in form, leafy internodes 0.5-4 cm long, 0.7-3 mm thick (dry), becoming

sparsely puberulent, ridged or longitudinally sulcate on drying but succulent in life, with

tufts of thin whitish hairs in the leaf axils. Leaves opposite, often with reddish or purple

coloring, petioles poorly differentiated from the lamina, 6-25 mm long, ca. 0.5 mm thick

when dry, with lateral margins continuous with the lamina margins; laminae 0.5-4 cm

long, 0.5-2 cm broad, obovate to oblanceolate, obtuse at the apex, gradually narrowed at

the base and decurrent on the petiole, margins entire, the laminae drying stiffly char-

taceous, sparsely puberulent or glabrous, with 2-4 pairs of major secondary veins. In-

florescences solitary and axillary, capitate, 4—8 mm long, floral bracts 1-2 mm long, stiff

and with a thickened midrib extending to form a sharp tip; flowers with the outer perianth

parts ca. 3 mm long, with 3 longitudinal ribs thickened abaxially, sharply acute at the apex,

with thin whitish hairs on the back. Fruit suborbicular, 1-1.5 mm long; seeds 0.8-1.2 mm

broad, cochleate-orbicular, lenticular in cross section, shiny reddish brown.

Widely cultivated and perhaps occasionally escaping plants of open sunny sites;

to be expected between sea level and about 2,000 m altitude. This taxon may be

unknown as a truly wild plant; it is commonly cultivated from the southern

United States to Brazil and Peru in the New World and in many areas of the Old

World.

Alternanthera bettzichiana is distinguished by its variably colored foliage (from

reddish and purplish to green marked with yellow or white), short succulent bush

habit, small sessile axillary whitish heads, and general restriction to gardens and

cultivated plantings. The dense branching and variously colored foliage make

these plants especially suitable for planting in rows or dense groups to achieve

masses of color irf formal gardens and parks. These plants have been called

Colchdn de nino in Honduras, Coqueta in Belize, Monte negro in Nicaragua, and

Perico in El Salvador. This species is closely related to A. tenella Colla.

Alternanthera brasiliana (L.) O. Kuntze, Rev. Gen. 1:537, 1891. Gomphrena

brasilianaL., Cent. PI. 2:13. 1756. Achyranthes brasiliana (L.) Standl., J. Wash. Acad.

Sci. 5:74. 1915.

Herbs, annual or perennial, erect or prostrate, leafy internodes 3-10 (18) cm long, 0.7-3

mm thick, strigose with thin white ascending hairs 0.5-2 mm long. Leaves opposite,

petioles 3-10 mm long, with narrow lateral margins continuous with the lamina margins;

GOMPHRENA

serrata

ALTERNANTHERA

bettzichiana

ALTERNANTHERA

paronychioides

A caracasana

FIG. 26. Amaranthaceae: species of Alternanthera with sessile inflorescences and a com-

mon species of Gomphrena. The arrow denotes the dorsal crest on the floral bract of

Gomphrena serrata.

149

150 FIELDIANA: BOTANY

laminae 2-10 cm long, 1-6 cm broad, elliptic to ovate-elliptic or lanceolate, tapering

gradually or abruptly to the acute or short-acuminate apex, rounded to obtuse at the base

and decurrent on the petiole, margins entire, the laminae drying thin-chartaceous, puber-

ulent on both surfaces with whitish appressed-ascending hairs 0.2-1.5 mm long, venation

pinnate with 3-6 pairs of major secondary veins. Inflorescences solitary, usually terminal,

compact spikes or capitula 10-20 mm long and 8-14 mm thick, white or yellowish, borne

on slender peduncles 2-16 cm long, with slender ascending stiff hairs, floral bracts 4-6 mm

long, the distal margin erase or cilliate, midrib obscure, glabrous or minutely puberulent;

flowers sessile or subsessile, perianth parts 3-6 mm long and often obscured by the bracts

and bracteoles, ovate-lanceolate, with 3 longitudinal ribs prominent on the basal half,

stamens and pistil hidden within the persisting perianth and bracts. Fruit included within

the persisting perianth and bracts; seed ellipsoid, ca. 1.8 mm long, lustrous black.

Herbs of open sunny sites often in sandy soils below 1,000 m elevation and

flowering throughout the year. Known from a single collection in Nicaragua and

to be expected in Costa Rica; the species is native to Venezuela, the Guianas,

Brazil, and eastern lowland Peru.

Alternanthera brasiliana is recognized by the opposite leaves with conspicuous

hairs, long internodes, terminal and solitary white or yellowish heads on long

peduncles, and large floral bracts. The species has recently been collected near

Esteli in Nicaragua (Molina 23058), but it has not been recorded from Costa Rica

or Panama.

Alternanthera caracasanaH.B.K., Nov. Gen. & Sp. 2:206. 1818. Figure 26.

Densely branched usually prostrate herbs, 5-25 cm tall, leafy internodes 0-4 cm long,

0.7-2 mm thick (dry), densely sericeous or villous with thin whitish hairs 1-2 mm long.

Leaves opposite and small, often clustered at the nodes, petioles 1-5 mm long, with lateral

margins continuous with the lamina margins; laminae 0.5-2 cm long, 3-12 mm broad,

elliptic to obovate, abruptly obtuse or rounded at the apex, gradually narrowed to the

attenuate base, margins entire, the laminae drying chartaceous, sparsely puberulent,

venation often obscure, with 1-3 pairs of secondary veins. Inflorescences solitary and

sessile in the axils of leaves, short spikes 5-10 mm long, floral bracts ca. 3 mm long, the

prominent midvein short-cuspidate at the apex, broadly ovate and thin-translucent,

mostly glabrous; flowers tightly clustered, outer perianth parts ca. 3 mm long, often

obscured by the bracts, the midrib prominent and often extended to form a short (1 mm)

apical spine, 2 lateral longitudinal ribs often thickened near the base, puberulent on the

back (abaxially) with thin whitish hairs that are minutely hooked at the tip. Fruit tightly

enclosed within the persisting perianth; 1.5-2 mm long, truncate at the apex and with a

persisting style; seeds ca. 1.5 mm long, cochleate-ovoid, lenticular in cross section, lus-

trous brown.

Prostrate plants of open early secondary habitats, known only as weeds near

large urban centers in Costa Rica between 1,000 and 1,500 m elevation. The species

flowers throughout the year in northern Central America and ranges from about

500 to 2,500 m elevation (rarely collected below 500 m in Guatemala). The species

ranges from Mexico (in both low and higher altitudes) through Central America

and the West Indies to northern and western South America as far as Bolivia.

Alternanthera caracasana is recognized by its prostrate habit, small leaves and

small sessile inflorescences, and by the unusual "grappling hook" hairs on the

back of the outer perianth parts. These small hairs have very minute terminal

hooks that are quite difficult to see with a 10 x hand lens and are sometimes

interspersed with hairs lacking the hooks. Most of our Central American material

of this species was previously placed under the name A. repens (L.) Kuntze. The

species appears to be quite rare in Costa Rica.

BURGER: FLORA COSTARICENSIS 151

Alternanthera costaricensis O. Kuntze, Rev. Gen. 2:538. 1891. Achyranthes cos-

taricensis (Ktze.) Standl., J. Wash. Acad. Sci. 5:74. 1915. Achyranthes megaphylla

Standl., North Amer. Fl. 21:141. 1917, ex char. Figure 27.

Herbs or subshrubs 20-80 cm tall, apparently perennial, erect and usually unbranched,

leafy internodes 1-9 cm long, 1-3.5 mm thick, with thin usually straight appressed-

ascending yellowish hairs 0.5-1 mm long, stems becoming dark brown and glabrous.

Leaves opposite, petioles 5-10 mm long, puberulent with appressed-ascending hairs,

usually becoming glabrous; laminae 7-16 (17.5) cm long, 2-5 (7.5) cm broad, narrowly

elliptic to narrowly ovate-elliptic or ovate-oblong, tapering gradually to a sharply acu-

minate or acute apex, acute to obtuse at the base, margins entire, the laminae drying

thin-chartaceous, smooth above and below, with scattered thin straight ascending hairs

above and (more densely) below, the 7-10 pairs of major secondary veins arising at angles

of 50°-70°. Inflorescences sessile or subsessile in the axils of terminal or distal leaves,

solitary and spicate, 1-4 cm long, ca. 1 cm thick, greenish in life but drying yellowish

brown or brown, floral bracts 3-4 mm long, ovate but narrowed to a sharp tip ca. 1 mm

long, with a single midrib, rachis visible between the flowers and bracts, densely pilose;

flowers with the outer perianth parts 6-8 mm long, with 3 or 5 prominent longitudinal

ribs, tapering gradually to the sharply acute apex, puberulent at the base. Fruit and seed

not seen.

Rare plants in the understory of wet evergreen forests and known only from

near sea level to 1,100 m altitude on the Caribbean slopes and lowlands; flowering

collections have been made between late March and November. The species has

not been found outside Costa Rica and is presently known from as far north as

Bijagua (Alajuela) and as far south as Siquirres (Limon).

Alternanthera costaricensis stands apart from all our other species of Alternanthera

because of the elongate inflorescences with very stiff, relatively large, dull brown-

ish perianth parts, deep forest habitat, and relatively large thin leaves. The in-

florescences give the impression that this species may be more naturally situated

in another genus. The original description of Kuntze states that the spikes are

borne on peduncles and, indeed, Kuntze's collection from eastern Costa Rica (NY)

gives this impression. However, on closer inspection, the "peduncles" appear to

be terminal internodes, with the distal leaves having fallen off. Note added in

proof: Specimens of this species returned to Field Museum in September 1982 have

been annotated Jamesbondia costaricensis (Ktze.) Mears by Mears.

Alternanthera laguroides (Standl.) Standley, in Standl. & Calderon, Lista PI.

Salvador. 74. 1925. Achyranthes laguroides Standl., Contr. U.S. Natl. Herb 18:90.

1916. Figure 27.

Herbs or subshrubs, erect or clambering, to over 2 m high or 3 m long, older nodes

slightly thickened, leafy internodes 5-15 (20) cm long, 0.5-4 mm thick, at first densely

sericeous with thin ascending whitish hairs ca. 1 mm long but soon becoming glabrescent,

older stems longitudinally striate. Leaves opposite, petioles 1-4 (6) mm long, slightly

expanded at the base and with tufts of hairs in the axils; laminae (2) 4-11 (14) cm long, (0.4)

1-2.5 (3.5) cm broad, lanceolate to narrowly elliptic or very narrowly elliptic-oblong, acute

at the apex and with a slender sharp tip 1-2 mm long, acute at the base and decurrent on

the petiole, margins entire, smooth and sericeous on both surfaces with thin appressed-

ascending whitish hairs ca. 1 mm long, the 3-6 pairs of major secondary veins arising at

angles of 30°- 50°. Inflorescences compact heads or short spikes 4-14 mm (20) mm long,

borne on simple (rarely branched) peduncles 1-4 cm long, solitary or 3 together, axillary

or terminal, leaves subtending the inflorescences sometimes reduced and the terminal

stems and spikes producing panicle-like arrangement, floral bracts 2-4 mm long, with a

single midrib and acute tip; flowers sessile, perianth 4-5 mm long, with long (4 mm) thin

whitish hairs from the base of the perianth, each perianth part with only a single promi-

VCW?', PFAFFIA grandiflora

A. pubiflora

A laguroides

ALTERNANTHERAT costaricensis

FIG. 27. Amaranthaceae: three species of Alternanthera and a species of Pfaffia.

152

BURGER: FLORA COSTARICENSIS 153

nent longitudinal midrib, stamens and pistil hidden within the perianth. Fruit hidden

within the persisting perianth and bracts; seeds ca. 0.7 mm broad.

Plants of moist evergreen forest formations on both the Caribbean and Pacific

sides of Costa Rica from sea level to 1,500 m elevation; flowering collections have

been made from January to June. The species, as presently known, ranges from

central Nicaragua to central Panama.

Alternanthera laguroides is distinguished by its clambering habit, long inter-

nodes, pedunculate heads or short spikes, and the relatively long straight whitish

hairs borne from the base of the outer perianth parts (abaxially). The plants have

been called Botoncillo in our area. The species is well represented by collections

from Costa Rica, but only a very few A. laguroides are from Nicaragua and Panama.

Alternanthera lanceolata (Benth.) Schinz, in Engler & Prantl., Naturl. Pflanz-

enfam. ed. 2, 16O75. 1934. Brandesia lanceolata Bentham, PI. Hartweg. 247. 1846.

Alternanthera lehmannii Hieron. , Bot. Jahrb. Syst. 20, beih. 49:8. 1895. Achyranthes

lehmannii '(Hieron.) Standl., J. Wash. Acad. Sci. 5:74. 1915. Achyranthes panamensis

Standl., Contr. U.S. Natl. Herb. 18:89. 1916. Achyranthes stenophylla Standl., loc.

cit. 18:90. 1916. Alternanthera panamensis (Standl.) Standl., Publ. Field Columbian

Mus., Bot. Ser. 8:9. 1930. Alternanthera stenophylla (Standl.) Standl., loc. cit.

Herbs, 0.3-1 m tall, erect or decumbent, leafy internodes (0.5) 3-15 cm long, 0.7-3 mm

thick, at first densely puberulent with thin ascending hairs 0.5-1 mm long but soon

becoming glabrescent. Leaves opposite, petioles 2-10 mm long, slender and with very

narrow lateral ridges continuous with the laminae margins; laminae 2-8 (15) cm long,

0.5-3 (5) cm broad, elliptic to elliptic-ovate, gradually narrowed to the acuminate apex,

acute to obtuse and somewhat decurrent at the base, margin entire, the laminae drying

thin chartaceous, sparsely puberulent with thin hairs 0.5-1.8 mm long on both surfaces,

with 4-6 pairs of major secondary veins. Inflorescences capitate or very short spikes, 3-10

mm long, solitary or rarely 2-3 on slender (0.3 mm) peduncles (0.5) 2-10 cm long, terminal

or axillary, sparsely puberulent, floral bracts 0.5-2 mm long, cuspidate, with broad thin

translucent margins; flowers 3-4 mm long, subsessile or sessile, perianth parts ca. 3 mm

long, glabrous or very sparsely puberulent, with 3 weakly defined longitudinal ribs,

stamens and pistil included within the persisting perianth. Fruit tightly enclosed within

the persisting perianth, the seeds 1.2-1.5 mm broad, reniform-rounded, lustrous black,

smooth or wrinkled when dry.

Weedy herbs of open and shaded sites in moist and wet evergreen forest for-

mations from 1,000 m to about 2,200 m elevation in Costa Rica; collected with

inflorescences from August through January. The species occasionally forms a

ground cover along stream and brook beds and appears to be restricted to areas

subjected to the very wet Caribbean winds between San Ramon and the Rio

Grande de Orosf drainage in Costa Rica; the species ranges from Guatemala to

Colombia.

Alternanthera lanceolata is recognized by the long slender internodes, usually

solitary capitate inflorescences on long peduncles, long hairs on the upper leaf

surfaces, perianth parts with weakly developed ribs, and the restriction to cloud

forest habitats between 1,000 and 2,200 m (in Costa Rica).

Alternanthera paronychioides St. Hil., Voy. Brasil, 2 (2):43. 1833. Gomphrena

ficoidea L., Sp. PI. 1:225. 1753, non A. ficoides Beauv. 1818. Figure 26.

Herbs, usually prostrate, stems much branched, up to 1 m long and often rooting from

the nodes, leafy internodes 1-5 cm long, 0.5-2 mm thick (dry), at first densely sericeous-

tomentulose but soon becoming glabrous. Leaves opposite, occasionally somewhat dimor-

phic in size, petioles 1-15 mm long, with lateral margins continuous with the lamina

154 FIELDIANA: BOTANY

margins; laminae 0.6-3 cm long, 0.5-1.5 cm broad, obovate to elliptic, abruptly obtuse to

rounded at the apex, gradually narrowed to the attenuate base and decurrent on the

petiole margins, entire, the laminae drying thin-chartaceous, very sparsely puberulent,

with 2 or 3 pairs of major secondary veins. Inflorescences solitary and sessile in the axils

of leaves, capitate or short spikes 4-10 mm long, white, floral bracts 1-2 mm long, thin,

obtuse to acute at the apex, with a single thickened midrib; flowers with the outer perianth

parts 3-4 mm long, 3 longitudinal ribs prominent on the lower half of the abaxial side,

acute and entire, with thin white hairs, anthers 5. Fruit loosely enclosed in the persistent

perianth, obovoid and usually truncate at the apex with the base of the style persisting,

1.2-1.8 mm long and equally broad or broader; seeds 0.8-1.2 mm broad, cochleate-

orbicular, lenticular in cross section, dark brown and lustrous.

Prostrate weeds of open, usually damp, sites from near sea level to about 1,500

m elevation; probably flowering throughout the year, but most often collected in

the wet season. The species ranges from northern Mexico to southern South

America. These plants are often found near the seashore and brackish (salty)

water.

Alternanthera paronychioides is distinguished by its prostrate habit, profuse

branching, small sparsely puberulent leaves, small sessile inflorescences, and fruit

easily removed from the persisting perianth and bracts. This species appears to be

quite rare in southern Central America; I have seen only a single specimen from

Costa Rica: Maxon & Harvey 8750 from near Puntarenas, close to sea level. While

very similar to Alternanthera caracasana, this species lacks the unusual grappling-

hook hairs (20 x) on the back of the outer perianth parts, and the fruit is not rightly

enclosed within the fruiting inflorescence. Standley reported Golondrina as a com-

mon name for this species in Costa Rica, but this species is quite rare and the name

may be misapplied here. Central American specimens of this species have often

been placed under the name A. polygonoides (L.) R. Br. (as in the Flora of Gua-

temala, Flora of Panama, and Flora of Costa Rica), but Mears considers the Lin-

naean basonym a nomen obscurum.

Alternanthera pubiflora (Benth.) O. Kuntze, Rev. Gen. 2:538. 1891. Brandesia

pubiflora Bentham, Bot. Voy. Sulph. 157. 1846. Achyranthes williamsii Standley,

Contr. U.S. Natl. Herb. 18:89. 1916. Alernanthera williamsii (Standl.) Standley, J.

Wash. Acad. Sci. 15:458. 1925. Figure 27.

Herbs, erect, procumbent, or clambering, 0.5-1.5 (2) m tall, leafy internodes 2-11 cm

long, 0.8-5 mm thick, densely strigilose with small (0.2-0.8 mm) whitish ascending or

appressed hairs, becoming glabrescent with age. Leaves opposite, petioles 2-10 (14) mm

long, the lateral edges continuous with the lamina margins; laminae 4—12 cm long, 1.5-4

(5) cm broad, elliptic to elliptic-ovate, acute to short acuminate at the apex, gradually

narrowed to the obtuse or acute base and decurrent on the petiole, margins entire, the

laminae drying thin to stiffly chartaceous, smooth and with thin whitish hairs ca. 0.5 mm

long on both surfaces, venation pinnate with 3-6 pairs of major secondary veins. In-

florescences short spikes or capitate, 6-15 mm long, 1-3 borne in the leaf axils or terminal,

peduncles slender and sericeous, (0.5) 1-8 cm long, floral bracts 2-3 mm long, ovate-

lanceolate, acute at the apex and with a single midrib, bracteoles often with the midrib

prolonged and caudate; translucent white, flowers sessile, perianth parts ca. 6 mm long,

with 3 prominent longitudinal ribs, and stiff little white hairs on the back (abaxially),

stamens and pistil included within the perianth. Fruit enclosed within the persisting

bracts, bracteoles and perianth; seed ca. 1.5 mm long, reniform-oblong, lustrous black.

Weedy plants of open and partly shaded sites in the seasonally dry and decid-

uous or partly deciduous formations between sea level and 800 (1,400) m elevation

along the Pacific slope in southern Central America and collected in flower and

BURGER: FLORA COSTARICENSIS 155

fruit from December to June. The species ranges from Nicaragua and primarily

along the Pacific slope in South America to Chile, Bolivia, and Argentina.

Alternanthera pubiflara is recognized by its larger elliptic leaves, sericeous pubes-

cence on many parts, 3-ribbed perianth parts, smaller bracts and bracteoles, and

long-peduncled inflorescences that arise either from the leaf axils or are terminal.

The terminal inflorescences are sometimes followed by paired axillary shoots that

produce dichotomous branching. While collected quite often in Nicaragua, this

species is apparently rare in Costa Rica.

Alternanthera sessilis(L.) R. Br., Prodr. 417. 1810. Gomphrena sessile L., Sp.

PI. ed. 2, 300. 1762. Figure 26.

Small herbs, erect or decumbent, 10-50 cm tall and to 1 m long, stems usually with only

a few lateral branches, leafy internodes (0.5) 1-8 cm long, 0.5-3 mm thick, very sparsely

puberulent with minute crooked hairs usually confined to 2 narrow longitudinal depres-

sions on opposite sides of the stem, leaf axils often with longer (0.5 mm) whitish hairs, an

interpetiolar ridge usually joining the leaf bases. Leaves opposite, petioles poorly differ-

entiated from the laminae, 0-5 mm long, with lateral margins continuous with the lamina

margins; laminae (1) 1.4-6.5 cm long, 4-14 (20) mm broad, very narrowly elliptic to

oblanceolate, bluntly acute (obtuse) at the apex, tapering gradually to the base and de-

current on the petiole, margins entire, the laminae drying thin-chartaceous and usually

greenish, smooth and glabrous above and below or with a few minute hairs along the

midvein, with 3-6 pairs of major secondary veins. Inflorescences solitary and sessile or

subsessile in the axils of leaves, capitate, 3-5 mm long, whitish or slightly pink, floral

bracts less than 1 mm long, broadly triangular, thin translucent; flowers sessile, outer

perianth parts ca. 2 mm long, thin translucent and with only the midrib thickened. Fruit

a utricle 1.5-2 mm long and not tightly enclosed by the persisting perianth, pale yellowish

brown, emarginate at the apex and broadly obovate in outline with the style sessile in the

apical notch, lenticular in cross section; seeds ca. 1 mm long, pale brown.

Pioneer weeds in open sites, often in wet depressions and river margins, occa-

sionally growing partly submerged in water, from near sea level to 1,200 m

elevation in our area; flowering throughout the year, but collected most often from

February to August in Costa Rica. The species appears to be uncommon in the

seasonally very dry areas of Guanacaste. A pantropical species, ranging from

Guatemala and Belize through Central America and the West Indies to Peru and

Brazil in the New World.

Alternanthera sessilis is distinguished by its small white sessile inflorescences,

unusual fruit that is usually visible within the persisting perianth, the laminae

tapering gradually to the petiole, and the usually wet habitats. The unusual fruit

and the ease with which it can be removed from the dry flowers set this species

well apart from its congeners in Central America.

Alternanthera tenella Colla, Mem. Reale Accad. Sri. Torino 33:131. 1829. Fig-

ure 26.

Herbs, erect or clambering over other small vegetation, stems to 1 m long, leafy inter-

nodes 1-10 cm long, 0.8-3 mm thick, densely puberulent at first with thin or basally

thickened whitish hairs ca. 0.5 mm long, soon becoming glabrous, often with tufts of hairs

in the leaf axils, the hairs often barbed near the base. Leaves opposite, petioles 1-8 mm

long, with lateral margins continuous with the laminae margins; laminae (1.5) 2-9 cm

long, (0.5) 1-4 cm broad, elliptic to ovate-elliptic, obtuse to bluntly acute at the apex,

tapering gradually to the obtuse or acute base and decurrent on the petiole, margin entire,

laminae drying thin-chartaceous, smooth, becoming glabrous or with a few small (0.1-0.5

mm) hairs along the midvein, with 3-6 pairs of major secondary veins. Inflorescences

156 FIELDIANA: BOTANY

solitary and sessile in the axils of leaves, capitate or very short spikes 4-10 mm long, 3-6

mm in diameter, whitish, floral bracts ca. 2 mm long, with a sharp aristate tip and

translucent margins; flowers sessile, outer perianth parts ca. 4 mm long, stiff and with a

sharp tip, with 3 (5) strongly raised longitudinal ridges on the back and short (0. 1-0. 5 mm)

stiff hairs near the base, the hairs occasionally minutely hooked. Fruit a suborbicular

utricle tightly enclosed within the persisting perianth, seed cochleate-oblongoid, 1 mm

long, lustrous dark reddish brown.

Plants of open sunny early secondary growth, from near sea level to about 500

m elevation in both moist evergreen and in seasonally deciduous areas (but appar-

ently very rare in seasonally very dry Guanacaste); flowering throughout the year

in our area, but collected most often between November and February. The

species ranges from southern Mexico and the Yucatan peninsula through Central

America and the West Indies to Bolivia and southern Brazil.

Alternanthera tenella is recognized by the small sessile heads or spikes, the stiff

sharp-tipped perianth parts with prominent longitudinal ribs and thin straight

hairs, lack of hairs on older parts, and low-altitude habitats. Most of our material

had been identified as A. ficoidea (L.) R. Br. until Mears recently determined that

the Linnaean basonym, Gomphrena ficoidea, belongs to the species now known as

Alternanthera paronychioides.

AMARANTHUS Linnaeus

Herbs, rarely perennial, erect or less often prostrate, bisexual or rarely unisexual

(dioecious), glabrous or minutely puberulent, often marked with red or purplish coloring,

unarmed or with spines in A. spinosus; stipules absent. Leaves alternate and simple,

petioles often long and variable in length on the same plant, laminae entire, usually

thin-textured, venation pinnate with the midvein usually extending slightly beyond the

lamina to form a slender mucronate tip. Inflorescences basically of small flower clusters

(glomerules of condensed cymes) in the axil of a bract, glomerules solitary or more often

congested on an unbranched rachis and spicate, the spikes solitary in distal leaf axils or

terminal and near-terminal to form a compound paniculate arrangement of spikes, the

terminal spikes often twice as long as lateral spikes, each flower subtended by a bract and

2 lateral bracteoles, bracts and bracteoles usually similar in form and size. Flowers small,

mostly unisexual, the male and female often borne on the same spike with the male

flowers borne distally and the female near the base, sessile or subsessile, perianth (1- to

3-) 5-parted, free, equal or subequal, thin and stiff, stamens usually the same number as

the perianth parts, filaments slender and free, anthers said to be 2-thecous but dehiscing

by 4 slits, ovary 1-locular, the solitary ovule from a basal placenta, style very short, stigmas

2 or 3, slender. Fruit loosely enclosed within the persisting bracts and perianth parts,

usually a thin-walled utricle or capsule with circumscissle dehiscence or indehiscent; seed

solitary, usually cochleate-orbicular and lenticular in cross section, often dark colored and

very lustrous, smooth, embryo annular.

A genus of about 50 species in both tropical and temperate areas of the world.

Most of the species are pioneers on bare soil or in early successional vegetation and

many have become widespread weeds. The young leaves of a number of species

are used as pot herbs, and the grain amaranths (see under A. hybridus) have been

an important source of food in some cultures (cf. J. D. Sauer in Ann. Missouri Bot.

Card. 37:561-632, 1950). Forms with larger deeply red or purple colored in-

florescences have been used for ornamental planting and in floral arrangements.

Members of the genus are usually recognized by the short-lived weedy growth

habit, unusual paniculate grouping of spikes at the apex of the plant (in most

species), dense clusters of flowers and bracts, fruit loosely enclosed in the persist-

ing perianth and bracts and usually opening by having the distal half fall away,

and the small shiny lenticular seeds. The name Bledo is often used in Central

America to refer to species of Amaranthus.

BURGER: FLORA COSTARICENSIS 157

la Plants bearing paired straight spines (5- 15 mm long) from some of the leaf axils; seeds

0.7-1 mm broad; widespread plants from sea level to 1,600 m elevation

A. spinosus

Ib Plants lacking spines in the leaf axils 2a

2a Fruiting perianth broadest above the middle and becoming partly reflexed with the

distal half rotate; seeds ca. 0.9 mm broad; seasonally dry areas of the Pacific slope

below 900 m elevation A. scariosus

2b Fruiting perianth not broadest above the middle and not becoming reflexed in

fruit 3a

3a Fruit indehiscent and with a rugose yellowish brown surface; perianth 3-parted,

stamens 3, floral bracts 0.5-1 mm long; rachis often visible on the spikes. .A. viridis

3b Fruit usually circumscissely dehiscent and with a smooth surface; perianth 5-parted,

stamens usually 5, floral bracts 1.5-4 mm long; rachis usually obscure in the spicate

inflorescences 4a

4a Floral bracts up to twice as long as the perianth parts and usually with conspicuous

spine tips; seeds more than 1 mm broad; plants of waste places near habitations or

cultivated for the seed or colorful inflorescences, common on the Meseta Central,

1,000-2,000 m A. hybridus

4b Floral bracts equaling the perianth in length; seeds ca. 1 mm broad; weedy plants of

the Caribbean lowlands A. dubius

Amaranthus dubius Martius ex Thellung, Mem. Soc. Sci. Nat. Cherbourg

38:203. 1912. A. tristis Willd., Hist. Amaranth 21. 1790, not A. tristis L., 1753.

Figure 28.

Herbs, 0.2-1 (1.5) m tall, unisexual, branches ascending, leafy internodes 0.5-6 cm long,

1-6 mm thick, at first minutely puberulent with thin curled or scurfy hairs ca. 0. 2 mm long,

becoming glabrous and longitudinally ridged or sulcate. Leaves alternate, petioles 1-10 cm

long, sulcate above with 2adaxial ridges continuous with the lamina margins; laminae 3-8

(12) cm long, 2-6 (8) cm broad, ovate-rhombic to ovate-oblong or ovate elliptic, the smaller

leaves elliptic, bluntly obtuse at the apex and often with a thin 1-2 mm mucronate

extension of the midvein, obtuse to acute at the base and attenuate on the petiole, margin

entire or slightly crenulate, laminae drying thin-chartaceous, sparsely and very minutely

(0.1 mm) puberulent on the veins beneath or glabrescent, midvein impressed above, the

4-12 pairs of major secondary veins prominent beneath. Inflorescences solitary in the leaf

axils or in terminal paniculate arrangements, flower clusters densely aggregated along the

flowering rachis forming spikes 4-9 (12) mm in diameter, terminal spikes to 15 (25) cm

long, lateral spikes 0.5-6 cm long, floral bracts 1.5-2.5 mm long and usually equalling the

flower in length, ovate to lanceolate and ending in a sharp tip; flowers unisexual with the

male less common than the female, perianth 1.5-2 mm long, stamens 5, stigma 3. Fruit

loosely enclosed within the subequal and loosely persisting bracts, bracteoles, and peri-

anth, utricle 1.5-2.5 mm long with circumscissle dehiscence; seeds cochleate-orbicular

and lenticular in cross section, 0.8-1.1 mm broad, dark brown or black and very lustrous.

Plants of open early successional sites at altitudes below 1,000 m and flowering

throughout the year. The species is a common weed in the Caribbean area and

ranges from Mexico to southern South America. The species has become estab-

lished in Europe and Africa.

Amaranthus dubius is recognized by its open weedy lowland habitat, floral bracts

equalling the flowers in length, and the almost circular, lenticular seeds about 1

mm broad. This species has only been collected from the Caribbean Coastal Plain

in Costa Rica and may not occur above 200 m elevation in our area.

Amaranthus hybridus L., Sp. PI. 990. 1753. A. hypochondriacusL., loc. cit. 991.

1753. Figure 28.

Herbs, 0.3-1.5 (2) m tall, erect and often much branched, bisexual, leafy internodes

0.5-3 cm long, 1-5 mm thick, sparsely puberulent with minute (ca. 0.2 mm) thin crooked

hairs in early stages, becoming glabrescent and longitudinally ridged or sulcate when dry.

3 mm

A. hybrid us

hybrid us

scanosus

dubius

spinosus

cruentus

viridis

AMARANTH US

FIG. 28. Amaranthaceae: species of Amaranthus, after Sauer.

158

BURGER: FLORA COSTARICENSIS 159

Leaves alternate, petioles 1-8 (12) cm long, sulcate above with 2 adaxial ridges continuous

with the lamina margins, sparsely and minutely puberulent; laminae (2) 4-10 (15) cm long,

1-7 (10) cm broad, ovate to rhombic or narrowly ovate-elliptic in smaller leaves, bluntly

obtuse in larger leaves or acute in smaller leaves, with a thin 2-mm extension of the

midvein beyond the lamina, margins entire or slightly crenulate, lamina drying char-

taceous, glabrous or very sparsely puberulent with thin crooked hairs only 0.1-0.2 mm

long, midvein impressed above, the 6-10 pairs of major secondary veins pale in color and

prominent beneath. Inflorescences terminal and axillary, spicate or the spikes in terminal

paniculate arrangements, 1-18 cm long, the individual spikes 5-15 mm thick, flower

clusters closely congested and the rachis usually obscure, bracts subtending the flower

clusters 2-4 mm long with almost half the length of the bract made up of a distal stiff spine,

bracts mostly twice as long as the perianth parts; flowers unisexual or bisexual, perianth

5-parted, 1.3-2.3 mm long, thin, stamens 5, stigmas 3. Fruit loosely enclosed in the small

persisting perianth and larger stiff bracts, a thin-walled utricle with circumscissle de-

hiscence near the middle; seeds 1-1.4 mm broad, cochleate-orbicular in outline and thick-

lenticular in cross section, edge with a weakly defined ridge, dark reddish brown to black,

lustrous.

Plants of open fields, roadsides, and early successional habitats and now natu-

ralized throughout the world in both tropical and temperate areas. In Costa Rica,

the species has only been collected between 1,000 and 1,700 m elevation near the

towns and cities of the eastern part of the Meseta Central (San Pedro de Puriscal

to San Jose and Cartago); probably capable of flowering at any time of the year in

our area.

Amaranthus hybridus is recognized by the spine-tipped bracts often twice as long

as the perianth parts, the minute puberulence of younger parts and leaves, long

petioles, and the dark shiny seed usually more than 1 mm broad. The plants are

often marked with pink, red, or purple and seem to be collected only near the

more densely populated or well-traveled areas of Central America.

Amaranthus hybridus subsp. cruentus (L.) Thellung (in Asch. & Graebn., Synop-

sis 5:243, 1914), more often referred to as Amaranthus cruentus L. and also placed

under A. caudatus L., is the commonly cultivated grain amaranth of northern

Central America. These plants are grown for ornament, are occasionally used as

pot herbs, and are an important grain in some areas. They differ in having more

reddish coloring in the floral bracts, the larger utricle exceeding the perianth parts,

the shorter floral bracts, and the lateral spikes of the terminal panicle being more

divergent than those in plants of typical A. hybridus. The two entities, whether

species or subspecies, are very closely related and, though extensively cultivated

in Guatemala, we have seen no authentic material of subspecies cruentus from

Costa Rica.

Amaranthus scariosus Benth., Bot. Voy. Sulph. 158, pi. 51, 1844. Figure 28.

Herbs, 0.5-1.5 (2) m tall, erect and usually much branched, unisexual, leafy internodes

0.5-4 cm long, 1.2-6 mm thick, glabrous or the new growth with minute puberulence,

stems often pink in life, terete. Leaves alternate, petioles 2-10 cm long, sulcate above with

adaxial ridges continuous with the lamina margins; laminae 2-9 (12) cm long, 1-5 cm

broad, ovate to rhombic or elliptic-ovate (ovate-lanceolate in smaller leaves), tapering to an

obtuse apex, the midvein extended less than 1 mm beyond the lamina, obtuse to cuneate

at the base and slightly decurrent on the petiole, margins entire, the laminae drying

membranaceous to very thin-chartaceous, smooth and essentially glabrous above and

below, midvein impressed above, the 7-12 pairs of major secondary veins pale in color and

prominent beneath. Inflorescences terminal or axillary, small spikes to paniculate arrange-

ments of spikes, 0.5-25 cm long, often densely clustered on the distal part of the stem,

floral bracts 2-4 mm long, subulate-lanceolate and sharp-tipped, green along the midrib,

slightly exceeding the flowers; flowers unisexual, female perianth parts 5, ca. 3 mm long

160 FIELDIANA: BOTANY

and 1.5 mm broad, obovate and broadest above the middle, with a green midrib and broad

translucent margins, rounded apically, stamens 5, stigmas 3. Fruit loosely enclosed within

bracts and perianth, the perianth parts becoming partly reflexed and rotate in form, utricle

shorter than the tubelike base of the perianth; seed ca. 0.9 mm broad, orbicular or

cochleate-orbicular, thick-lenticular in cross section, smooth and very lustrous, dark

brownish black.

Weedy plants of open fields and forest edges on the seasonally very dry Pacific

slope of northern Costa Rica below 900 m elevation; flowering and fruiting mate-

rial has been collected from late December through February. The species ranges

from southwestern Mexico along the Pacific lowlands to Guanacaste Province in

Costa Rica.

Amaranthus scariosus is recognized by its unusual female perianth parts which

are broadest above the middle and become reflexed above the middle to form a

tubelike lower basal part and a rotate distal part. The general lack of pubescence,

long-petioled leaves, and restricted habitat in seasonally deciduous areas further

distinguish this species. The species probably occurs in northernmost Puntarenas,

but has not been collected outside of Guanacaste in Costa Rica. It is interesting

that a plant of weedy early successional habitats has such a restricted geographical

range.

Amaranthus spinosusL., Sp. PL, 991. 1753. Figure 28.

Herbs, 0.5-1.5 (2) m tall, erect and with spreading branches, bisexual, leafy internodes

0.3-5 (8) cm long, 0.7-4 mm thick (dry), smooth and glabrous or sparsely and very

minutely puberulent in early stages, often reddish or purplish in life and becoming irreg-

ularly longitudinally striate when dry, paired sharp spines 8-15 mm long and ca. 1 mm

thick at the base often arising from the leaf axils and oriented ca. 90° from the stem, sulcate

adaxially. Leaves alternate, often variable in size on the same stem, petioles 0.5-6 (8) cm

long, less than 1 mm broad (dry), slightly sulcate above with 2 adaxial ridges continuous

with the laminae margins; laminae 1-10 (12) cm long, 0.6-3.5 (5) cm broad, narrowly

elliptic to elliptic-lanceolate or ovate-elliptic, bluntly to sharply acute and rounded at the

tip, often with a slender terminal spine 1-3 mm long, acute or obtuse at the base and

decurrent on the petiole, margin entire, the laminae drying thin-chartaceous or mem-

branaceous, smooth and glabrous, midvein impressed above with 4-7 pairs of major

secondary veins drying pale in color and prominent beneath. Inflorescences terminal or

from the axils of distal leaves, varying (often on the same plant) from small axillary

fascicles or glomerules ca. 10 mm in diameter to densely flowered spikes 1-8 (14) cm long

and 5-10 mm thick, the spikes often subtended by paired spines 3-6 mm long, bracts

variable and often resembling the spines, 2-5 mm long, ovate to subulate; flowers uni-

sexual with the male flowers at the tips of spikes and the female in the lower (proximal)

glomerules, perianth 5-parted, 2-2.5 mm long in the male and ovate-oblong, 1.5-2.5 mm

long and oblong-spatulate in the female, scarious, the midvein green and margins trans-

lucent, stigmas slender, papillate-puberulent, to 1.5 mm long. Fruit loosely enclosed in the

persisting perianth, utricle ca. 1.5 mm long, pale brown with circumscissle dehiscence or

indehiscent; seed cochleate-orbicular, lenticular in cross section, 0.7-1 mm broad, dark

brown and very shiny lustrous.

Plants of open sites and early secondary growth from sea level to 1,600 m

elevation on both the seasonally dry Pacific slope and in the evergreen Caribbean

formations; probably flowering throughout the year in Costa Rica, but collected

primarily in June and July; collections from the deciduous lowlands of Guanacaste

have been made primarily between June and January. This species is now wide-

spread in the warmer parts of the world.

Amaranthus spinosus is recognized by the paired spines arising from the leaf

axils, generally glabrous parts, and small seeds. An unusual collection by John

Taylor (4432 NY) with the floral bracts considerably more spiny than usual and

BURGER. FLORA COSTARICENSIS 161

collected on Volcan Irazu at 3,200 m elevation is included here. However, it is

unlikely that populations of this species persist above 2,000 m elevation in Costa

Rica.

Amaranthus viridisL., Sp. PI., ed. 2, 1405. 1763. A. gracilis Desf., Tabl. Bot. 43.

1804. Figure 28.

Herbs to ca. 1 m tall, erect or decumbent, bisexual, leafy internodes 0.2-3 (5) cm long,

0.8—4 mm thick, glabrous or very sparsely and minutely puberulent in early stages. Leaves

alternate, petioles very slender (dry), 0.3-10 cm long, often very variable in length on the

same plant, slightly sulcate above with adaxial margins continuous with laminae margins;

laminae 0.5-5 (7) cm long, 0.3-3 (4) cm broad, ovate to bluntly triangular or rhombic,

elliptic or elliptic-ovate, rounded at the bluntly obtuse apex and emarginate at the tip with

the midvein extended less than 1 mm beyond the lamina, abruptly narrowed or rounded

at the obtuse to subtruncate base, decurrent on the petiole, margin entire or slightly

crenulate when dry, the laminae drying very thin-chartaceous, smooth and glabrous, with

4-8 pairs of major secondary veins prominent below. Inflorescences usually solitary,

terminal or in the axils of distal leaves, spikelike with clusters of flowers borne on an

unbranched axis, spikes 1-8 (12) cm long, 3-7 mm thick, flower clusters often separate and

the rachis visible, floral bracts and bracteoles subequal, 0.5-1 mm long, thin and scarious;

flowers unisexual, longer than the bracts, perianth 3-parted, 1-1.5 mm long, stamens 3,

stigmas 3. Fruit a rugose utricle clearly exceeding the persisting perianth and bracts, ca.

2 mm long, indehiscent, pale yellowish brown; seed 0.9-1 mm broad, cochleate-orbicular

in outline, thick-lenticular, very dark brown or black, moderately lustrous, with a weakly

defined ridge around the margin.

Small weedy plants of open sandy and gravelly soils and often seen as pioneers

on bare soil from near sea level to 1,500 m elevation; probably capable of flowering

throughout the year in our area. This species is a pantropical weed.

Amaranthus viridis is recognized by its very small stature, glabrous parts, small

bracts and bracteoles, 3-parted perianth, and indehiscent fruit with rugose sur-

face. These plants are often found as pioneers on open soil, but they appear to be

quite rare in our area, with only a few collections from Guanacaste and the eastern

side of the Meseta Central.

CELOSIA Linnaeus

Annual or perennial herbs or subshrubs, erect (in ours) to sarmentose or climbing,

bisexual, glabrous to sparsely puberulent. Leaves alternate, often decurrent on the stem,

petioles often with lateral margins gradually expanding into the lamina margins; laminae

entire and pinnately veined. Inflorescences usually dense spikes (heads), axillary or termi-

nal, solitary, or variously disposed in paniculate arrangements, the spikes usually sub-

sessile, the panicles sessile or pedunculate, the axes bearing the spikes usually un-

branched in our species, each flower subtended by a bract and 2 lateral bracteoles. Flowers

bisexual, sessile or subsessile, perianth of 5 free and equal or subequal parts, scarious,

longitudinally ribbed and slightly convex, persisting but not enlarging in fruit, stamens 5,

the filaments united at the base to form a cup shorter than the ovary, anthers 4-celled, and

introrse, ovary globose to ellipsoid, 1-locular, with several to many ovules on a basal

placenta, style short to long, with 2 or 3 stigmas. Fruit a capsule, usually with circumscissle

dehiscence, loosely included within the persisting bracts and perianth parts; seeds (1) 2 to

many, usually cochleate-lenticular to reniform, smooth and usually lustrous, brown to

black, the embryo annular.

A genus of about 40 species, mostly in Asia and Africa with only the following

two species to be expected in southern Central America. Young leaves and shoots

of C. argentea are sometimes used as vegetables, though the plants are mostly

cultivated for ornament. The native species of the genus are apparently quite local

or rare in Central America.

162 FIELDIANA: BOTANY

la Plants grown for ornament, rarely escaping; inflorescences colorful white to rose red

or magenta spikes, 5-25 cm long and 1.5-3 cm thick, flowers ca. 8 mm long

C. argentea

Ib Plants known in Costa Rica only from the deciduous forest formations of northeastern

Guanacaste Province; inflorescences of small greenish spikes, 0.5-3 cm long, often

borne in paniculate arrangements, flowers ca. 5 mm long C. virgata

Celosia argentea L., Sp. PI. 205. 1753. C. cristate L., loc. cit. Figure 29.

Annual herbs, erect, 0.2-1 (rarely 2) m tall, branched or unbranched, leafy internodes

0.5-9 cm long, 1-7 (12) mm thick, glabrous, becoming longitudinally ridged or grooved.

Leaves alternate in a spiral, petioles 4-30 mm long, with lateral margins that expand

gradually to the lamina margins, decurrent on the stem; laminae 3-15 cm long, 0.5-6 cm

broad, ovate to ovate-elliptic or narrowly lanceolate, acute to acuminate at the apex,

tapering gradually or abruptly to the base and decurrent on the petiole, margins entire, the

laminae drying stiffly chartaceous, glabrous, the 5-8 pairs of major secondary veins

strongly ascending. Inflorescence usually terminal and solitary or a few clustered together,

usually a densely flowered thick (1-2 cm) spike, 1.5-15 (25) cm long, occasionally with

abnormal (fasciated) growth patterns and broadly expanded or divided at the apex, pe-

duncles glabrous, 0.3-12 cm long, floral bracts ca. 6 mm long, lanceolate, lustrous whitish

to pinkish, decurrent on the rachis, carinate and mucronate; flowers 8-10 mm long,

subsessile to short (2 mm) pedicellate, perianth thin and dry (scarious), lustrous, silvery

white to deep rose red, 6-9 mm long, persisting in fruit, style 3-6 mm long, usually

exceeding the perianth. Fruit a circumscissile capsule, 3-4 mm long, hidden within the

persisting perianth and bracts, with (1) 3-9 seeds per fruit; seeds 1.5-2 mm broad,

cochleate-lenticular, very lustrous, black or dark reddish brown.

Plants grown in gardens for their colorful inflorescences and only rarely escap-

ing (especially along streams); flowering throughout the year and rarely encoun-

tered above 1,400 m elevation in Central America. The species is of uncertain

origin and is now widely cultivated throughout the tropics.

Celosia argentea is recognized by its alternate leaves with laminae decurrent on

the petioles, glabrous parts, and thick colorful spikes with stiff dry persisting

bracts and perianth parts. The long-lasting lustrous white to deep rose red in-

florescences make these plants very useful for floral arrangements. In some plants,

the inflorescences develop a broad flattened form that may be digitate or ruffled.

This form is often called variety cristate (L.) Voss, the "Garden cockscomb."

Among the names associated with these plants in Central America are Abanico,

Amaranto, Amor seco, Boria, Cresta de gallo, Flor de mano, Mano de leon, Mono, Mono

de reina, Rabo de gato, and Terciopelo.

Celosia virgata Jacq., Coll. Bot. 2:279. 1788, or a closely related species. Fig-

ure 29.

Erect herbs, 0.5-1.5 m tall, leafy internodes 5-80 mm long 1-4 mm thick, glabrous or

minutely (0.5-2 mm) puberulent with thin crooked hairs, quickly becoming glabrous.

Leaves alternate in a spiral, decurrent on the stems, petioles 1-8 cm long, with lateral

margins gradually expanding to the lamina base; laminae 3-15 cm long, 1-6 cm broad,

ovate to ovate-ellitpic or the smaller leaves often lanceolate, acute to short-acuminate at

the apex, often with a sharp tip 1-3 mm long, acute to obtuse at the base and decurrent

on the petiole, margin entire, drying thin-chartaceous, laminae essentially glabrous, with

5-7 pairs of major secondary veins. Inflorescences basically of short (8-20 mm) spikes,

solitary and axillary to terminal and variously clustered, the paniculate clusters usually

with a single primary axis 5-20 cm long and with separate or congested spikes alternating

along the axis (our material differs from all other collections of the species in having

slender peduncles or with a 2-5 cm long spike-free basal portion on the inflorescence axis,

similar peduncles in other collections rarely exceed 2 cm), bracts and lateral bractets

subequal, 2-3 mm long, cuspidate with a single midvein and thin hyaline margins, nar-

CELOSIA

argentea

CHAMISSOA

altissima

5 mm

PLEUROPETALUM

sprucei

FIG. 29. Amaranthaceae: Costa Rican representatives of the genera Celosia, Chamissoa,

and Pleuropetalum.

163

164 FIELDIANA: BOTANY

rowly triangular to lanceolate; flowers subsessile, ca. 5 mm long, perianth parts with 3-7

abaxially prominent longitudinal ribs in the basal half, oblong-lanceolate. Fruit with (1)

3-8 seeds, utricle shorter than the perianth parts; seeds 0.7-1 mm broad, cochleate-

lenticular, black, lustrous.

Uncommon herbs found in both moist evergreen and in seasonally very dry

deciduous formations, from near sea level to 700 m elevation (rarely to 1,200 m);

flowering and fruiting collections have been made from September to March in

Central America. The species ranges from central Mexico through Guatemala, El

Salvador, Honduras, and has recently been collected near Santa Rosa in north-

western Guanacaste Province, Costa Rica.

Celosia virgata (in a wide sense) is characterized by its short stature, alternate

leaves with laminae decurrent on the petiole, general lack of pubescence, in-

florescences composed of short spikes often aggregated on unbranched axes,

single-veined translucent bracts, stiff multiple-ribbed perianth, and small dis-

tinctive shiny seeds. A single recent Costa Rican collection by Ronald Liesner

(4303) is tentatively placed here. It differs from most specimens of the species in

a number of ways: The leaf axils do not have the little axillary leaves usually found

in the species, the inflorescences of the Costa Rican collection have clearly defined

peduncles, and there are apparently fewer but slightly larger seeds in each fruit.

Nevertheless, stature, leaves, bracts, and flowers are virtually identical to those of

typical Mexican and Guatemalan collections. It may be that the plants from near

Santa Rosa are worthy of subspecific recognition.

CHAMISSOA Humboldt, Bonpland & Kunth

REFERENCE: S. H. Sohmer, A revision of Chamissoa. Bull. Torrey Hot. Club.

104:111-126, 1977.

Shrubs, subshrubs, or lianas, usually climbing or clambering over adjacent vegetation,

stems glabrous or puberulent, terete; stipules absent. Leaves alternate and simple, pet-

iolate, entire or slightly undulate, pinnately veined, glabrous or sparsely puberulent.

Inflorescences terminal or axillary, solitary, leaves of the upper (distal) nodes often un-

developed and the inflorescence a compound panicle, the flowers borne in small cymules

of 2-20 flowers, the cymules usually on unbranched axes and in arrangements of spicate

branches, the flowers subsessile or very short-pedicellate and subtended by a bract and 2

lateral bracteoles, bract and bracteoles subequal, persistent and becoming stiff and dry;

flower small and radially symmetrical, bisexual or functionally unisexual, perianth of 5

ovate-lanceolate imbricate parts, stiff and persisting but not enlarging in fruit, stamens 5,

united near the base to form a short tube, pseudostaminodial appendages absent, pistil

with 1 locule and 1 ovule from a basal placenta, style 1 with 2 slender stigmas. Fruit a

thin-walled capsule opening by circumscissle dehiscence near the top (a pixidium);. seeds

lenticular, black or marked, embryo small with much endosperm.

A genus of two species ranging from Mexico through the West Indies and

Central America to nearly all of South America excepting Chile and southern

Argentina. The alternate leaves, unusual fruit, and dense inflorescences made up

of racemose or spicate arrangements of cymules help distinguish the genus.

la Plants clambering or climbing, often becoming lianas over 2 m high; seeds enclosed by

2-valved arils; flowers bisexual or female, ovary over 2 mm long, tubular or oblong in

form; common lowland plants C. altissima

Ib Plants erect or clambering over others, up to 2 m tall but never becoming lianas;

seeds with minute and inconspicuous arils; flowers always bisexual, ovary usually less

than 2 mm long, ovate to globose in form; plants only rarely collected in Central

America . . . .C. acuminata

BURGER: FLORA COSTARICENSIS 165

Chamissoa acuminata Martius, Nov. Actorum Acad. Caes. Leop. -Carol. Nat.

Cur. 13:286. 1826.

Herbs or subshrubs, erect or clambering, 1-2 m tall, leafy internodes ca. 3-6 cm long,

slender and glabrescent in ours. Leaves alternate, petioles 1-3 (7) cm long, slender; lami-

nae 3-11 (14.5) cm long, 1-4 cm broad, narrowly ovate to lanceolate, tapering gradually

to the acute or acuminate apex, rounded to obtuse at the base, margins entire, glabrous or

puberulent, with 3-7 pairs of major secondary veins. Inflorescences terminal or axillary,

1-6 (30) cm long, usually simple and spikelike, sometimes compound and paniculate,

glomerules with 3-7 flowers, bracts and bracteoles 1.5-2.5 mm long, perianth and bracts

greenish white or yellowish, perianth 5-parted, 2.4-4 mm long, ca. 1 mm broad; stamens

5; ovary ovate to globose, 1.5-2.3 mm long, style 0.5-1.5 mm long, dilated at the base,

stigmas shorter than the style. Fruit a capsule, usually opening by a circular lid; seed

1.5-2.5 mm long, black and lustrous.

Though ranging from southern Mexico to Bolivia and Paraguay, this species has

not been collected in Costa Rica. The type collection of C. acuminata var. swansonii

Sohmer comes from Bocas del Toro, Panama (Wedel 1695), and our plants should

be identical.

Chamissoa acuminata is very similar to C. altissima, but does not become a liana

and differs in floral structure. In addition, the flowers seem to be less robust, and

the stems usually are more slender.

Chamissoa altissima (Jacq.) H.B.K., Nov. Gen & Sp. 2:197, pi. 125. 1817. Celosia

paniculata L., Sp. PI. ed. 2, 298, not L., 1753. Achyranthes altissima Jacquin, Enum.

PI. Carib. 17. 1762. Chamissoa macrocarpa H.B.K., Nov. Gen. & Sp. 2:197. 1817.

Figure 29.

Clambering shrubs to 2 m tall or woody vines to over 10 m long, leafy internodes 2-10

cm long, 1-4 mm thick, very sparsely puberulent with minute (0.1-0.5 mm) hairs, be-

coming longitudinally striate when dry. Leaves alternate and usually in a spiral, petioles

0.5-3 (7) cm long, 0.5-1.5 mm broad, very sparsely and minutely puberulent, sulcate

above with adaxial ridges continuous with the laminae margins; laminae 3-15 (19) cm

long, 1.5-7 (9) cm broad, narrowly ovate to broadly ovate or lanceolate, tapering gradually

to the acute or acuminate apex, acute to obtuse or truncate at the base, often rounded and

truncate in larger laminae, margins entire or slightly undulate, somewhat decurrent on the

petiole, the laminae drying thin to stiff chartaceous, smooth on both surfaces, very

sparsely puberulent with minute whitish hairs with 3-6 pairs of major secondary veins.

Inflorescences basically small cymules densely clustered on a spikelike rachis 1-15 cm

long, the spikelike axes solitary and axillary or in complex terminal paniculate arrange-

ments formed by the failure of distal leaves to develop, the cymules short (1-3 mm)

pedunculate and bearing 2-8 (20) flowers, flowers very short pedicellate and subtended by

a bract and 2 lateral bracteoles; flowers yellowish to white or greenish white to rose,

perianth parts 2-3 (5) mm long, stamens with filaments 2-3 mm long, anthers very small

(0.2 mm) in functionally female flowers, to 1 mm long in functionally male flowers, pistil

2-3 mm long in functionally female flowers, style 0.4-1 mm long, the 2 linear stigmas

minutely papillose. Fruit 3-4 mm long, 1.5-2 mm broad, somewhat ellipsoid with trun-

cated apex and base, opening with a circular lid near the top; seeds lenticular, orbicular

in outline, ca. 2 mm broad, black and lustrous.

Climbing or clambering plants of both the evergreen wet and seasonally very

dry deciduous formations from sea level to 1,500 m elevation, but most common

below 300 m; while occasional flowering plants have been found from May to

November, the majority of flowering and fruiting collections have been made

from December to April. The species ranges from Mexico through Central Amer-

ica and the West Indies to northern Argentina.

Chamissoa altissima is recognized by the vining or clambering sparsely puber-

ulent stems with alternate thin-petioled leaves, spicate or paniculate arrange-

166 FIELDIANA: BOTANY

ments of densely or loosely aggregated flower clusters, flowers with stiff persisting

pale-colored perianth and bracts, and the thin- walled fruit that opens by a circular

lid near the top bearing a short style and 2 stigmas. Plants with flowers pinkish

red to dark purple or with reddish ovaries often having three stigmas have been

designated as variety rubella by Sohmer.

CYATHULA Loureiro

Herbs or subshrubs, annual or perennials, erect, prostrate, or decumbent, usually devel-

oping a woody base and often with long internodes, bisexual; stipules absent. Leaves

opposite and simple, petiolate, laminae entire and pinnately veined. Inflorescences termi-

nal or axillary, solitary or 3 at terminal nodes, spicate with a single unbranched axis

bearing very short-pedunculate flower fascicles (glomerules), the fascicles with 1 to several

fertile bisexual flowers and perianth parts and bracts of highly modified sterile flowers, the

fascicles spicate or capitate and usually subtended by a bract and several bracteoles of

similar size, bracts and bracteoles usually aristate, parts of the sterile flowers developing

elongate spines or bristles ending in sharp hooks as the fruit develop; functional flowers

bisexual, small and radially symmetrical, perianth of 5 free subequal, imbricate parts, stiff

and persisting but not enlarging in fruit, with 1-5 longitudinal ribs, stamens 5, united at

the base to form a tube with small pseudostaminodial appendages, anthers 4-thecous,

dehiscing introrse, ovary ovoid to obovoid, 1-locular, the solitary ovule pendulous from

a long funicle arising from a basal placenta, style 1 with capitate stigma. Fruit an inde-

hiscent utricle, tightly enclosed by the stiff persisting perianth; seeds reniform-oblong to

cochleate-orbicular, testa lustrous brown.

A genus of about 20 species probably native to the tropics of Asia and Africa,

with two species introduced into the neotropics. Our members of the genus are

recognized by their small weedy habit, opposite sparsely sericeous leaves and long

spikelike inflorescences with unusual flower fascicles that become reflexed on the

spike and develop elongate spines with hooked tips in later stages. These plants

resemble and are closely related to Achyranthes.

la Fruiting fascicles developing 5-15 hooked spines to 4 mm long, perianth parts ca.

4 mm long; seeds to 2 mm long; terminal spikes becoming 1-2 (4) x as long as the

subtending leaves; larger laminae usually over 6 cm long, narrowly elliptic and long

acuminate C. achyranthoides

Ib Fruiting fascicles developing over 20 hooked spines to 2 mm long; perianth parts 2-3

mm long; seeds 1-1.5 mm long; terminal spikes becoming 2-10 x as long as the

subtending leaves, larger laminae usually less than 7 cm long, usually rhombic and

obtuse to acute at the apex C. prostrata

Cyathula achyranthoides (H.B.K.) Moq., in DC., Prodr. 13, pt. 2:327. 1849.

Desmochaeta achyranthoides H.B.K., Nov. Gen. & Sp. 2:210. 1817. Figure 25.

Herbs or subshrubs to 1 (2?) m tall, erect or decumbent and rooting at the lower nodes,

leafy internodes 2-18 cm long, 1-4 mm thick, sparsely puberulent with thin ascending

sericeous hairs, more densely puberulent at the nodes. Leaves opposite and decussate,

petioles 2-6 (10) mm long, appressed puberulent, with lateral margins continuous with the

lamina margins; laminae 3-15 (20) cm long, 1.5-5 (7) cm broad, the larger laminae elliptic

to narrowly elliptic and tapering gradually to both apex and base, the smaller laminae

more abruptly tapering and often rhombic-ovate, acuminate to acute at the apex (in larger

laminae) acute or obtuse to attenuate at the base, margin entire or slightly undulate and

decurrent on the petiole, laminae drying thin- to stiff-chartaceous, smooth to the touch

above and below, sericeous with thin whitish appressed-ascending hairs 0.5-2 mm long

on both surfaces but usually more dense beneath, with 4-7 pairs of major secondary veins.

Inflorescences terminal or axillary, spicate, 5-20 cm long, 5-10 mm broad, peduncle 0.5-8

cm long with thin whitish hairs ca. 0.5 mm long, bracts subtending the flower fascicles

(glomerules) 1.5-3 mm long, thin-translucent and broadly lanceolate, mucronate or aris-

BURGER: FLORA COSTA RICENSIS 167

late, each fascicle borne on a very short (0.5 mm) peduncle and becoming reflexed, the

flower fascicles tending to remain closely adjacent even in fruit, the fascicle or glomerule

usually with 1 fertile flower and 1 sterile flower, the hooked (uncinate) spines 1-4 mm long

and often exceeding the perianth; flower with the perianth parts ca. 4 mm long and 1.5 mm

broad, strongly ribbed and sparsely puberulent with thin whitish hairs. Fruit very difficult

to separate from the enclosing perianth, utricle ovoid and operculate, 1.5-2.5 mm long;

seeds somewhat oblongoid, reniform to cochleate-orbicular, 1.5-2 mm long, ca. 1 mm

thick, lustrous brown.

Plants of open or partly shaded sites in both moist evergreen and seasonally

very dry formations, from sea level to 1,000 m elevation but rarely collected above

700 m; flowering and fruiting throughout the year in Costa Rica, but collected

most often from November to May. In seasonally very dry Guanacaste, the col-

lections are primarily from December and January. The species ranges from Mex-

ico and the West Indies to Brazil and throughout much of central Africa.

Cyathula achyranthoides is recognized by the spikelike inflorescences with

densely crowded flower fascicles (glomerules) that become reflexed on the spikes

and develop five to 15 stiff hooked spines. The plants are usually found in recently

disturbed areas or early secondary growth in both open and shaded sites. The

plants are most common in the Caribbean lowlands (0 to 200 m).

Cyathula prostrata (L.) Blume, Bijdr. Ned. Ind. 549. 1826. Achyranthes prostrata

L., Sp. PI. ed. 2, 296. 1762. Figure 25.

Herbs or subshrubs, 0.2-0.8 (1) m tall, erect or procumbent, leafy internodes 2-15 cm

long, 0.6-3 mm thick, sparsely puberulent with thin spreading or ascending whitish hairs

0.3-1.5 mm long, stems 4-angled or with 4 longitudinal ridges. Leaves opposite and

decussate, petioles 2-12 mm long, with narrow lateral margins continuous with the lamina

margins, sparsely puberulent; laminae 1-6 (8) cm long, 0.8-4 cm broad, broadly elliptic to

rhombic, obtuse to acute at the apex and rounded or obtuse at the base, margin entire and

decurrent on the petiole, the laminae drying thin to stiff chartaceous, smooth on both

surfaces, sparsely strigillose above and below with thin usually straight ascending hairs

0.3-1 mm long, with 3-5 pairs of major secondary veins. Inflorescences terminal or

axillary, solitary or 3 at a terminal node, spicate, 5-20 (30) cm long, 4-8 mm broad,

elongating in fruit, peduncles 2-5 cm long, the flower fascicles (glomerules) at first tightly

congested and spreading but becoming separate and reflexed on the narrow (1 mm)

sparsely puberulent rachis, the flower clusters borne on short (0.2-1 mm) peduncles and

subtended by short (1 mm) lanceolate hyaline bracts, usually with 3 bisexual flowers and

2-4 mm long, hooked (uncinate) awns developing as the fruit matures, 1-2 mm long and

not usually exceeding the fascicle in length, often more than 20 per fascicle; flowers with

perianth parts 2-3 mm long, composed of 5 scarious subequal parts. Fruit tightly enclosed

within the stiff persisting perianth parts, utricle ovoid, 1-2 mm long, inconspicuously

operculate; seeds slightly flattened and reniform to cochleate-orbicular, ca. 1 mm long.

Weedy plants of open or shaded sites in disturbed areas, stream beds, and early

successional vegetation in wet and moist evergreen formations between sea level

and 1,000 m elevation; probably flowering throughout the year, but with no

collections having been made from July to October. The species (probably intro-

duced) has a discontinuous range from Mexico and the West Indies to Brazil; the

species ranges widely across central Africa, tropical Asia, and Oceania.

Cyathula prostrata is recognized by the long spicate inflorescences, with fascicles

of flowers that become separate and reflexed on the rachis, and the later devel-

opment of many short hooked spines in each flower fascicle. The small stature of

the plants, their angled stems, small opposite leaves, and the short stiff perianth

that protects the fruit are also characteristic. The plants are often found along

stream edges in evergreen formations in our area.

168 FIELDIANA: BOTANY

GOMPHRENA Linnaeus

REFERENCE: J. A. Mears, The Linnaean species of Gomphrena L. Taxon 29:85-95,

1980.

Annual or perennial herbs, prostrate or erect, stems sometimes slightly thickened at the

nodes, usually puberulent with thin unbranched multicellular hairs. Leaves opposite,

sessile or petiolate, simple and entire, often with a small spine at the lamina apex, usually

puberulent. Inflorescences terminal or axillary, solitary or less often clustered, sessile,

heads or short thick spikes, the bracts, bracteoles, and flowers densely crowded together,

each flower subtended by a single abaxial bract and 2 lateral bracteoles, the bracteoles

usually much larger than the bracts, bracteoles concave and often with a wing or crest

developed from the dorsal (abaxial) side of the midrib, flowers bisexual, sessile, perianth

of 5 equal or subequal perianth parts, wooly (lanate) hairs usually borne on the base of the

perianth and enlarging in fruit, stamens 5, filaments completely united to form a tube with

the anthers sessile near the apex of the tube, 2-lobed pseudostaminodia usually present

at the apex of the tube between the anthers, ovary superior, locule 1, ovule solitary, style

1 with 2 slender stigmas. Fruit an indehiscent utricle, usually tightly included within the

persisting bracteoles and perianth; seeds cochleate-orbicular, smooth, embryo annular.

A pantropical genus of nearly 100 described species. Our species of Gomphrena

are easily recognized by their dense heads and spikes subtended by two small

leaves and by the unusual dorsal crests on the upper half of each of the paired

lateral bracteoles. These dorsal (abaxial) crests are serrate or crenate along their

margins. A similar species, G. filaginoides Mart. & Gal., with small (1 to 3 cm)

leaves and white heads almost 2 cm broad, has recently been collected in Hon-

duras and Nicaragua. Gomphrena elegansMart. and G. t uerckheimii (Vatke) Uline &

Bray have been reported from Guatemala and Honduras, but not Nicaragua.

These last two species are quite similar, with terminal heads that are not sub-

tended by small leaves and bracteoles that do not obscure the flowers or bear

conspicuous dorsal crests. These last two species look very much like some of our

species of Alternanthera (q.v.).

la Flower heads not subtended by a pair of small opposite leaves, the paired lateral

bracteoles lacking conspicuous dorsal crests, tips of the flowers usually visible within

the bracteoles. Weedy plants of northern Central America but not yet collected

from Costa Rica. See the discussion under the genus and compare with species of

Alternanthera.

Ib Flower heads of spikes subtended by a small pair of leaves, the paired lateral bracteoles

with conspicuous serrate to crenate dorsal crests, flowers hidden within the bracteoles

both in flower and fruit 2a

2a Plants commonly grown in gardens and used in floral arrangements; heads of spikes

ca. 2 cm thick, often bright pink or magenta, flowers and bracteoles long-persisting

G. globosa

2b Plants native weeds of open sunny sites below 1,300 m elevation; floral heads or spikes

ca. 1 cm thick, usually white, older flowers and bracteoles deciduous and only the

smaller (3 mm ) floral bracts persisting on the lower portion of older inflorescences

G. serrata

Gomphrena globosa L., Sp. PI. 1:224. 1753.

Erect or ascending annual herbs to 1 m tall. Leaves opposite, laminae 2-10 cm long,

0.5-4 cm broad, narrowly elliptic to elliptic-oblong, acute at the apex with a small spine

tip, with thin ascending appressed hairs on both upper and lower surfaces. Inflorescences

solitary and terminal or rarely paired and axillary, globose heads or thick (2 cm) short (2-4

cm) spikes, the heads or spikes subtended by small (0.5-2 cm) ovate leaves, the primary

floral bracts small (3 mm) and hidden by the much larger (7-12 mm) lateral bracteoles, the

paired lateral bracteoles with broad dorsal crests arising from the upper half of the midrib

(abaxially), the crests irregularly crenate-serrate. Flowers hidden within the bracteoles,

BURGER: FLORA COSTARICENSIS 169

perianth parts 5-8 mm long, stamens forming a tube 4-8 mm long. Fruit an indehiscent

utricle tightly enclosed within the persisting perianth and bracteoles; the seed cochleate-

orbicular, flattened on the sides, 1.5-2 mm broad, reddish brown.

Plants commonly grown as ornamentals in Central America and flowering

throughout the year. The species is not known to have become naturalized in

Costa Rica. The flower heads vary from pink and white to magenta or pink-purple

in color, and because they persist long after flowering, retaining both their color

and form, they are often used in floral arrangements. Common names are Amor

seco, Baton, Inmortal, San Diego, and Siemprevivain Central America. "Batchelor's

button" and "Globe amaranth" are common English names. This species is widely

cultivated in the tropics and subtropics.

Gomphrena serrataL., Sp. PI. 224. 1753. G. decumbens]acq., PI. Hort. Schoenbr.

4:41, t. 482. 1804. G. dispersa Standl., Contr. U.S. Natl. Herb. 18:91. 1916. Fig-

ure 26.

Small herbs 10-50 (80) cm tall, prostrate or erect, annual or perennial, leafy internodes

1-15 cm long, 0.6-2 (3) mm thick, sparsely to densely sericeous with thin ascending

multicellular whitish hairs 1-3 mm long. Leaves opposite, often subsessile, somewhat

articulate at the base, petioles 2-6 mm long, ca. 1 mm broad, little differentiated from the

lamina, with 2 lateral margins continuous with the lamina margins; laminae 1.5-5 (7) cm

long, 0.5-2 cm broad, narrowly elliptic to elliptic or ovate-elliptic, obtuse to acute at the

apex, often with a short (0.5-1 mm) sharp stiff tip, the lamina decurrent on the petiole,

entire, drying chartaceous, sparsely to densely sericeous above and below with thin

ascending hairs 1-3 (4) mm long, venation pinnate with 2 or 3 pairs of major secondary

veins. Inflorescences terminal and solitary (rarely 2), usually subtended by a pair of small

(5-15 mm) leaves, internode below the inflorescence and its subtending leaves (peduncle

if the leaves are considered bracts) 1-10 cm long, heads or becoming spikes as much as 3

cm long with the earlier basal flowers and their lateral bracteoles deciduous but the smaller

(2-3 mm) bracts persisting, the smaller bracts thin whitish translucent with thickened

midrib forming a sharp mucronate tip, bracteoles 4—6 mm long, stiff whitish (rarely

pinkish) with an apically broadened serrate crest on the back (abaxially) of the midrib;

flowers equaling or somewhat smaller than the bracteoles and included within them,

perianth parts ca. 4 mm long and very narrow above the base, with a dense mass of

villous-lanate hairs on the lower third, ovary tightly enclosed within the perianth. Fruit

enclosed within the persisting bractlets and tightly enclosed within the thin stiff perianth

parts, the lanate hairs becoming 3 mm long; seed lenticular with flattened sides, cochleate-

orbicular in outline, ca. 1.3 mm long, lustrous brown, with an apical notch.

Common weedy plants of open secondary sites in both wet evergreen areas and

in seasonally dry deciduous formations from sea level to 1,200 m elevation in

Costa Rica; flowering throughout the year, but collected infrequently in the dry

season (February to May). The species ranges from the southern United States

through Central America and the West Indies to Bolivia and Argentina.

Gomphrena serratais recognized by its short (sometimes prostrate) habit, slender

thin ascending hairs on almost all parts, small opposite leaves with sharp tips,

dense whitish heads and spikes with characteristic bracts and larger lateral brac-

teoles with dorsal serrate crests, and the flowers with a dense woolly covering of

cotton-like hairs. These are common weedy plants of open sunny sites not col-

lected from above 1,200 m elevation in our area. The distinctions used to separate

G. serrata and G. dispersa do not appear to be consistent or important in collections

seen from southern Central America. These plants resemble some of our species

of Alternanthera, but the large paired lateral bracteoles with dorsal crests immedi-

ately distinguish this species.

170 FIELDIANA: BOTANY

IRESINE P. Browne

Herbs, shrubs, lianas, or rarely trees, erect or often clambering over other vegetation,

unisexual or bisexual, internodes terete or strongly ridged, usually puberulent in early

stages, an interpetiolar line or ridge often present and the nodes usually thickened in age.

Leaves opposite or subopposite but often becoming alternate in the inflorescence, petioles

with adaxial ridges continuous with the lamina margins; laminae simple and entire or

rarely denticulate, usually decurrent on the petiole, venation pinnate, surfaces smooth and

rarely completely glabrous. Inflorescences terminal on the main stem or terminal on distal

axillary shoots, paniculate, usually large and with several orders of open distant branch-

ing, basal branches often subtended by small narrow leaves, whitish or pale yellowish, the

flowers usually borne in small bracteate puberulent spikes, each flower subtended by a

basal bract and 2 lateral bracteoles, bracts and bracteoles equal or dissimilar, puberulent

or ciliolate. Flowers bisexual or unisexual, monomorphic or dimorphic, sessile or sub-

sessile, perianth of 5 equal imbricate parts, narrowly oblong to lanceolate, often puberulent

distally, stamens usually 5 (4, 3) united near the base to form a shallow cup, ovary

narrowed at the base, style short with 2 or 3 slender or deltoid stigmas. Fruit an inde-

hiscent utricle loosely or tightly enclosed by the persisting perianth and bracteoles, con-

spicuous tufts of whitish hairs often developed from the base of the perianth abaxially;

seed cochleate-orbicular, thin- or thick-lenticular in cross section, smooth, pale to very

dark in color, often shiny-lustrous.

Iresineis a genus of perhaps 40 species closely allied to Pfaffia and greatly in need

of revision. The often unisexual flowers and the variation in size, arrangement,

and pubescence of inflorescences, floral bracts, and floral parts all contribute to

making specific delimitation difficult. The tufts of hairs in female and fruiting

flowers develop from between the base of the perianth parts and the floral bracts,

not from the interior of the perianth as in Pfaffia. Two species, /. diffusa (formerly

/. celosia) and /. calea, are very common in secondary growth throughout Central

America.

KEY TO THE SPECIES OF Iresine

la Flowers unisexual and only 0.5-1.5 mm long, whitish, subtended by bracts 0.3-1 mm

long; inflorescences to over 50 cm long but usually less than half as broad, branches

of the inflorescences mostly alternate, very common herbs or weak shrubs of ever-

green and partly deciduous formations from near sea level to 2,600 m elevation

/. diffusa

Ib Flowers 1.2-2 mm long, unisexual or bisexual, subtended by bracts 0.3-2 mm long;

breadth of the inflorescence usually more than half the length, distal branches of the

inflorescence opposite or alternate; herbs, shrubs, clambering shrubs or lianas . . . 2a

2a Spikes and flower clusters densely puberulent, branches of the inflorescence densely

sericeous, usually opposite throughout 3a

2b Spikes and flower clusters sparsely puberulent, branches of the inflorescence glabrous

to sparsely and minutely puberulent, usually alternate on the primary divisions of the

inflorescence; stigmas slender 4a

3a Common plants of the Pacific slope and Central Highlands from 0-1,800 m

elevation; inflorescences usually with opposite branching; flowers unisexual and

dimorphic, whitish, stigmas linear; laminae with 4-8 pairs of major secondary

veins /. calea

3b Rare plants of wet forests formations at middle elevations; inflorescence-

branching strictly opposite; flowers apparently bisexual, drying pale yellowish

brown, stigmas rounded; laminae with 7-12 pairs of major secondary veins

/. costaricensis

4a Leaves linear-lanceolate to ovate-lanceolate, with 3-8 pairs of major secondary veins;

flowers bisexual; slender stemmed herbs and subshrubs (rarely clambering), on the

seasonally dry Pacific slope below 800 m elevation /. angustifolia

4b Leaves ovate to ovate-lanceolate, with 6-13 pairs of major secondary veins; flowers

functionally unisexual; clambering shrubs and vines of wet forest formations between

ca. 1,000 and 2,000 m elevation . . . . /. arrecta

IRESINE

diffusa

IRESINE

arrecta

FIG. 30. Amaranthaceae: some species of Iresine.

171

172 FIELDIANA: BOTANY

Iresine angustifolia Euphrasen, Beskr. St. Barthel. 165. 1795. /. elatior Rich., in

Willd., Sp. PI. ed. 4, 766. 1805. Figure 30.

Herbs or shrubs with slender stems, usually erect, 0.5-1.5 (2) m tall, bisexual, leafy

internodes 1-14 cm long, 1-4 mm thick, glabrous or very sparsely and minutely puber-

ulent, usually with 4 or 6 longitudinal ridges, the older nodes often thickened and some-

times with tufts of hairs in the leaf axils. Leaves opposite or subopposite, petioles 5-25 mm

long, 0.5-1 mm thick, with adaxial ridges continuous with the lamina margins, glabrous;

laminae 2-12 (17) cm long, 0.5-3.5 (6) cm broad, linear-lanceolate to lanceolate or narrowly

ovate-lanceolate, tapering gradually to the acute or acuminate apex, tapering gradually to

the acute or obtuse base, margins entire and decurrent on the petiole, the laminae drying

thin chartaceous, smooth, glabrous or sparsely puberulent with short very thin and incon-

spicuous hairs, with 3-8 pairs of major secondary veins. Inflorescences terminal on the

main stem or axillary from distal branches, paniculate, 10-50 cm long, branches of the

inflorescence usually subtended by progressively smaller leaves, the flowers borne on

small pedunculate spikes or in small subsessile fascicles, flower-bearing rachis puberulent,

basal bracts ca. 0.5 mm long, lateral bracteoles ca. 1.5 mm long and broadly ovate, trans-

lucent and drying pale yellowish brown, slightly mucronate; flowers bisexual, sessile,

perianth parts 1.0-1.8 mm long, acute at the apex. Fruit loosely enclosed in the persisting

perianth, long (3-4 mm) white hairs often developed from between the base of the peri-

anth parts and bracteoles; seed cochleate-orbicular, 0.6-0.8 mm in diameter, lenticular in

cross section, black or very dark brown, shiny-lustrous.

Uncommon plants of secondary growth, forest edges, and open sites, from near

sea level to 500 (800) m elevation on the seasonally dry Pacific side of Costa Rica;

possibly flowering throughout the year, but with most all of our collections made

from January to April. The species ranges from Mexico and the West Indies to

northern South America.

Iresine angustifolia is recognized by the open much-branched inflorescences with

flowers closely congested in quite separate pedunculate spikes, the broad lateral

bracteoles twice as large as the basal bracts and equaling the perianth in length,

the very narrow leaves both within and beneath the inflorescences, and the

restriction to our seasonally dry Pacific lowlands. This species has been collected

only a few times in Costa Rica in the areas around the Gulf of Nicoya and near

the border with Nicaragua.

Iresine arrecta Standley, Publ. Field Mus. Nat. Hist., Bot. Ser. 18:1550. 1938.

Figure 30.

Clambering shrubs or woody lianas to 7m high, apparently unisexual, leafy internodes

3.5-18 cm long, 1.5-7 mm thick, very minutely (0.2 mm) puberulent but soon becoming

glabrous, longitudinally striate with usually more than 8 poorly defined longitudinal

ridges, terete or obscurely 4-angIed in cross section, nodes thickened in age and with an

interpetiolar line or ridge. Leaves opposite or subopposite, petioles 6-30 mm long, 1-1.5

mm thick, broadly sulcate above with 2 adaxial ridges continuous with the lamina margins;

laminae (4) 7-18 cm long, (1) 2-8.5 cm broad, ovate to ovate lanceolate, or lanceolate (and

alternate) in the inflorescence, tapering gradually to the acute or less often acuminate apex,

rounded and obtuse to subtruncate at the base, slightly decurrent on the petiole, margins

entire, the laminae drying thin-chartaceous to chartaceous and smooth, very sparsely and

minutely (0.2 mm) puberulent or glabrous, the midvein prominent above with 6-13 pairs

of major secondary veins. Inflorescences terminal on the main axes or terminal on distal

axillary shoots that often arise at right angles from the main stem, paniculate with numer-

ous but distant glabrous or sparsely and minutely puberulent branches, to 30 cm long, the

flowers in small (3-10 mm) sessile or very short-pedunculate spikes, bracts of the spikes

and flowers sparsely puberulent, floral bracts 1-1.5 mm long, broadly ovate cuspidate,

lustrous and translucent; male flowers with perianth 1.3-2 mm long, anthers ca. 0.5 mm

long, elliptic, pistillode well developed but lacking stigmas; female flowers with perianth

1.2-1.6 mm long, ovary turbinate or obovoid, ca. 1 mm long, style very short with 2 long

BURGER: FLORA COSTARICENSIS 173

(0.5 mm) stigmas. Fruit closely enclosed by the persisting perianth, subtended by tufts of

villous or lanate whitish hairs 1-3 mm long arising from the base of the perianth abaxially;

seed cochleate-orbicular, ca. 0.8 mm broad, lenticular in cross section, very dark brown

and highly lustrous.

Rarely collected clambering or climbing plants of wet evergreen montane

(premontane and lower montane rain) forest formations between 1,200 and 1,800

m elevation; flowering material has been collected from late November to early

March. The species is known only from central Costa Rica.

Iresine arrecta is distinguished by its climbing habit, long slender petioles, thin

leaves often rounded at the base and with numerous secondary veins, and

separate little spikes of functionally unisexual sparsely puberulent flowers. The

species is similar in some respects to /. costaricensis (q.v.) and /. argentata (Mart.)

Dietr. of northern South America, but both of those species have densely seri-

ceous inflorescence branches with bisexual flowers. Presently, /. arrectais known

only from near Zarcero, Alajuela (A. Smith A 617 and H 249, the type), near El

Muneco, Cartago (Standley & Torres 51090), and above San Isidro del General,

(Skutch 2563).

Iresine calea (Ibariez) Standley, Contrib. U.S. Natl. Herb. 18:94. 1916. Gom-

phrena latifolia Mart. & Gal., Bull. Acad. Roy. Sci. Bruxelles 10, pt. 1:349. 1843.

Achyranthes calea Iba'nez, Naturaleza 4:79. 1879. /. latifolia Benth. & Hook., Gen.

PL 3:42. 1880, not Dietr. 1839. Figure 30.

Shrubs or clambering and vinelike, 1.5-4 (5) m tall, unisexual, leafy internodes 2.5-13

cm long, 1.5-5 mm thick, sparsely or more often densely short (0.2-0.5 mm) hirsutulous

with whitish or very pale yellowish brown hairs, becoming glabrous and the nodes thick-

ened, irregularly longitudinally striate when dry, the interpetiolar line or ridge usually

obscure (sometimes absent). Leaves opposite, petioles 4-15 mm long, sulcate above with

2 adaxial ridges continuous with the lamina margins: laminae 1.5-8 (11) cm long, 1-4

(7) cm broad, ovate or occasionally ovate-lanceolate, tapering to an acute apex, usually

rounded or obtuse at the base, margins entire and decurrent on the petiole, laminae drying

chartaceous and smooth, usually densely hirsutulous or sericeous beneath with thin

whitish hairs ca. 0.5 mm long and more sparsely puberulent above, midvein prominent

above and with 4-8 pairs of major secondary veins. Inflorescence paniculate, 10-50 cm

long, the open divergent branches densely puberulent, the spikes separate (2-10 mm)

along the rachis and sessile or pedunculate, the male and female spikes quite different in

appearance; male spikes with floral bracts ca. 0.5 mm long and clearly exceeded by the

narrow perianth, the perianth parts 1.5-2 mm long, narrowly oblong, blunt and puber-

ulent apically, often lustrous in the lower half abaxially, stamens united only at the base

and not forming a conspicuous cup, pistillode usually slender and translucent; female

spikes with the flowers hidden by much larger (1-1.5 mm) broadly ovate bracts, with

dense tufts of tomentulous hairs from the bottom of the perianth further obscuring the

flowers, perianth parts ca. 1.5 mm long and narrowly lanceolate, ovary ca. 0.7 mm long,

style with 2 slender stigmas ca. 0.5 mm long. Fruit apparently tightly enclosed within the

persisting perianth and bracts; seeds not seen.

Shrubs or clambering plants of low second growth and thickets from near sea

level to 1,300 (1,800) m elevation on the Pacific slope and Central Mountains in

Costa Rica; flowering in the dry season, from December through April. This

species ranges from Mexico to the eastern edge of the Meseta Central in Costa

Rica along the Pacific slope and Central Highlands of Central America.

Iresine calea is distinguished by its generally shrubby habit, dense whitish

puberulence on young stems, undersides of leaves, and inflorescences, and the

rather different bracts subtending the morphologically distinct male and female

flowers. The different appearance of male and female spikes and the difficulty

174 FIELDIANA: BOTANY

of finding female flowers among bracts and hairs are also noteworthy. This

species has not been collected from the Caribbean slope below 1,200 m elevation;

it appears to prefer the seasonally dry habitats of the Pacific side of Central

America and is especially common in the Meseta Central. Names applied to this

species in central and southern Central America include: Algodoncillo, Chivito,

Chivo, Comenternero, Coyontura, Flor de corona, Siete Pellejos, and Tacuquelite,

Iresine completa Uline & Bray (Bot. Gaz. 21:349, 1896) was originally described

from San Pedro Sula, Honduras, and is distinguished by its larger (2.5 mm)

bisexual short-pedicellate flowers and narrow elliptic-lanceolate leaves glabrous

on the upper surface except above the midrib. I have seen no authentic material

of this species nor any specimens from our area which clearly fit the description.

See the discussion by Duke in the Flora of Panama (Ann. Missouri Bot. Card.

48:33, 1961). This species is related to /. costaricensis and /. hassleriana on the basis

of the short thick stigmas.

Iresine costaricensis Standl., Contrib. U.S. Natl. Herb. 18:94. 1916. Figure 30.

Clambering shrubs and lianas to 15 m high, bisexual, leafy internodes (1.5) 5-12 cm

long, 2-6 mm thick, terete, densely sericeous with short (0.5 mm) appressed-ascending

brownish hairs, becoming glabrescent and the nodes thickened, stems often becoming

contracted above the node and interpetiolar line after drying. Leaves opposite, petioles

6-17 mm long, sulcate above, with adaxial ridges continuous with the lamina margins;

laminae 7-18 on long, 3-7.5 cm broad, elliptic-ovate, to elliptic-oblong or broadly elliptic,

short-acuminate (in the type) to long-acuminate, with a narrow apex 4-10 mm long,

obtuse at the base and decurrent on the petiole, entire but drying slightly undulate, the

laminae drying stiffly chartaceous, and smooth, sparsely puberulent above and more

densely below with thin straight appressed hairs ca. 0.5 mm long, with 7-12 pairs of major

secondary veins. Inflorescences terminal on the main stem or on lateral branches, pan-

iculate to 50 cm long and with strictly opposite branching, the flowers in small (3 mm thick)

densely strigilose spikes or clusters, the flower cluster sessile or less often pedunculate,

bracts at the base of the spike ca. 1 mm long, these and some of the floral bracts (ca. 0.5

mm) densely hirsutulous-strigilose, bracteoles subtending the flowers ca. 1 mm long and

glabrous abaxially, very broadly ovate, tufts of long (1-2 mm) straight white hairs devel-

oping from the base of the flower within the bracteoles; flowers sessile, apparently bisex-

ual, perianth parts ca. 1.5 mm long, oblong, obtuse at the apex, puberulent, filaments

slender and united near the base to form a short cup, style very short with a thickly lobed

stigma. Fruiting material not seen.

Rarely collected lianas or shrubs from wet evergreen montane forests and

presently known only from the Caribbean slope at about 625 m elevation and

above the General Valley at 1,800 m elevation; flowering material has been

collected in December and March. The species is only known from central Costa

Rica.

Iresine costaricensis is recognized by its stiff leaves with up to 12 pairs of major

secondary veins, the inflorescences with consistently opposite divisions, young

flower buds covered with a dense tomentum of pale yellowish brown hairs, and

the short capitate stigmas. The very broad, almost glabrous bracteoles are usu-

ally only found with well-developed flowers. Iresine costaricensis was originally

known only from two collections made by Adolfo Tonduz (12919 and 13183, the

type) near Tucurrique, Cartago. These two collections appear to have been from

woody climbing plants. A recent collection (Burger & Liesner 7057) from an erect

shrub from above San Isidro del General is tentatively placed here. This latter

collection has narrow elliptic-oblong leaves that are short-acuminate and the

inflorescences, bracts, and perianth parts are not as puberulent as those in the

BURGER: FLORA COSTARICENSIS 175

Tonduz collections. Both collections have the unusually short thick stigmas.

This species is closely related to I. hassleriana Chodat, originally described from

Paraguay. Our material has smaller perianth parts, a shorter staminal tube, and

a shorter more rounded stigma than do the flowers of type material of /. has-

sleriana (Hassler 3429 F). The illustration of /. hassleriana in Flora of Panama is

somewhat intermediate, but closer to the material from Paraguay. At present, it

seems best to maintain /. costaricensis as a separate and distinct species closely

related to /. hassleriana.

Iresine diffusa Humboldt & Bonpland ex Willd., Sp. PL ed. 4, 4:765. 1806,

CelosiapaniculataL., Sp. PL 1:206. 1753. Iresine celosia L., Syst. Veg. ed. 10,1291.

1759, nomen illeg. /. celosioides L., Sp. PL ed. 2, 2:1456. 1763, nomen illeg. /.

paniculata (L.) O. Ktze., Rev. Gen. 2:542. 1891, not /. paniculata Poir. Figure 30.

Herbs, subshrubs, or shrubs to 3 (5) m tall, unisexual, the rootstock and lower stem

usually becoming woody, stems erect or clambering, leafy internodes 1-15 cm long 1-6

mm thick, occasionally glabrous but more often sparsely puberulent with short (0.1-0.5

mm) thin crooked hairs, internodes usually with 4 or 6 longitudinal ridges, the nodes

slightly thickened below the leaf base and with an interpetiolar line or puberulent ridge,

stems often narrowed just above the leaf bases when dried. Leaves opposite or sub-

opposite, petioles 0.4-5 (8) cm long, sulcate above with 2 narrow adaxial ridges continuous

with the lamina margins; laminae 2-16 cm long, 1-10 cm broad, elliptic ovate to broadly

ovate or lanceolate (small and linear on the inflorescence), tapering gradually to an acu-

minate or acute apex, acute to obtuse or rounded at the base and decurrent on the petiole,

margins entire, the laminae drying thin chartaceous and smooth to the touch, usually

sparsely puberulent beneath with short (0.1-1 mm) thin hairs, the midvein prominent

above with 3-9 pairs of major secondary veins arising at angles of 40°- 70°. Inflorescences

extremely variable in form and size, essentially terminal on the main or lateral axes,

paniculate arrangements with the ultimate divisions made up of short spikes with 5-35

crowded flowers (becoming separate in fruit) the distal inflorescence branches usually

alternate and often subtended by small lanceolate to linear leaves becoming short-linear or

bractlike near the ultimate branches, the overall inflorescence from 5-50 cm long and

whitish in color, the flowers subtended by subequal bracts and bracteoles 0.3-1 mm long,

broadly ovate and concave, mucronate, translucent to pale yellowish or white; flowers

sessile, unisexual, the perianth 0.7-1.5 mm long, stamens 5, united near the base to form

a short tube, ovary obovoid, style very short with 2 (3) slender stigmas. Fruit loosely

enclosed within the persisting perianth, long (1-2 mm) straight hairs sometimes formed

between the bracts and the perianth; seeds cochleate-orbicular, lenticular in cross section,

0.5-0.8 mm long, very dark brown and lustrous.

Plants of secondary growth in open sunny or partially shaded sites in both

very wet evergreen formations and, less commonly, in seasonally very dry

deciduous formations from near sea level to 2,600 m elevation; in Costa Rica,

some plants can probably be found in flower throughout the year, but almost all

of our fertile collections have been made between October and May, with a great

majority of the collections made in December, January, February, and March.

The species ranges throughout the American tropics.

Iresine diffusa, formerly called Iresine celosia, is one of Costa Rica's most com-

mon and conspicuous weeds. The species is recognized by the usually large,

white, many-branched paniculate inflorescences with very small unisexual

white flowers, stems with longitudinal ridges and unusual nodes, and thin

opposite leaves. The plants are most often collected in evergreen and partially

deciduous formations below 1,500 m elevation. There is considerable variation

regarding the size and arrangement of flowers and bracts in the material placed

under this name. It may be that more than one specific entity is included under

176 FIELDIANA: BOTANY

the broad interpretation adopted here. Material previously placed under the

name I. acicullarisStandl., I. frutescenceMoq., and 1. spiculigeraSeub. is included

here, following James Duke in the Flora of Panama (Ann. Missouri Bot. Card.

48:35, 1961). Common names used in central and southern Central America

include: Adorno de nino, Camron, Coyontura de polios, Hierba de goto, Siete pellejos,

Taba de gueguecho, Velo de gueguecho, and Velo de princesa.

PFAFFIA Martius

REFERENCE: O. Stutzer, Die Gattung Pfaffia. Repert. Spec. Nov. Regni Veg.

Beih. 88:1-46. 1935.

Herbs, shrubs, or climbers, densely puberulent to glabrescent, stems often with long

internodes (in ours) and interpetiolar ridges or lines, often with a tuft of hairs in the leaf

axil; stipules absent. Leaves opposite and simple, petiolate or sessile, entire and pinnately

veined. Inflorescences varied, heads, fascicles, or solitary flowers often on opposing spike-

like axes on terminal stems that bear reduced leaves forming compound panicles; flowers

bisexual or sometimes reduced and nonfunctional, radially symmetrical, small and sessile,

subtended by a bract and 2 lateral bracteoles, bract and bracteoles subequal, puberulent,

flowers often with an unpleasant odor, greenish white, perianth of 5 subequal parts, often

with 3 prominent longitudinal veins on the outer perianth parts, usually puberulent and

with tufts of straight whitish hairs from the base within, persisting but not enlarging in

fruit, stiff and dry, stamens 5, united below to form a short cup, the cup and filaments with

or without appendages, anthers 2-thecous, ovary ovoid to obovoid, 1-locular, with 1 ovule

borne on an elongated funicle from a basal placenta, style very short and bearing a thick

2-lipped stigma. Fruit a thin- walled capsule (utricle), usually obscured by the persisting

perianth, bracts, and tufts of hairs; seeds lenticular, reniform to cochleate-orbicular, red-

dish brown (in ours).

A poorly defined neotropical genus of probably fewer than 20 species, closely

related to Iresine and Gomphrena and intermediate between them in some re-

spects. The relatively few collections and their distribution in Costa Rica may

indicate that our species has been introduced. Our plants are recognized by their

clambering or climbing stems with long internodes and stiff opposite entire

leaves and compound inflorescences with puberulent little flowers with a stiff

dry perianth that bears tufts of whitish hairs from the base adaxially (between

the perianth and the sexual parts). This species and other members of the genus

are easily confused with species of Iresine, especially I. arrecta and /. costaricensis,

in our flora.

Pfaffia grandiflora (Hook.) R. E. Fries, Ark. Bot. 16, no. 12:10. 1921. Iresine

grandiflora Hooker, Icon. PI. 2; tab. 102. 1837. Gomphrena paniculatavar. hookeriana

Seub., in Martius, Fl. Brasil. 5, pt. 1:192. 1875. Hebanthe hookerianus Hemsley,

Biol. Cent. Amer. 3:19. 1882. Pfaffia hookeriana (Hemsl.) Greenman, Publ. Field

Columbian Mus., Bot. Ser. 2:330. 1912. P. hookeriana forma glabriuscula

Suesseng., Repert. Spec. Nov. Regni Veg. 35:332. 1934. Figure 27.

Woody climbers to 3 m long, leafy internodes 2-15 cm long, 1-5 mm thick, sparsely

puberulent to glabrescent with slender ascending hairs 0.5-1.5 mm long, more densely

puberulent at the nodes and in the leaf axils. Leaves opposite, petioles 3-15 mm long, with

lateral margins often forming an adaxial sulcus in the lower half; laminae 3-12 cm long,

1.5-6 cm broad, ovate to lanceolate or elliptic, acute to acuminate at the apex, rounded to

obtuse at the base, margin entire or slightly undulate, decurrent on the petiole, the

laminae drying very stifly chartaceous and pale or dark in color, smooth above and below,

with stiff slender appressed (strigose) hairs on both surfaces, with 3-5 pairs of major

secondary veins. Inflorescences usually compounded panicles at the ends of branches or

BURGER. FLORA COSTARICENSIS 177

lateral stems, made up of spikelike axes 1-5 cm long, the axes often opposite and sub-

tended by small leaves, the flowers usually solitary, sessile and subtended by a bract and

2 bracteoles 1-2 mm long and 1-2 mm broad at the base; flowers bisexual or nonfunctional,

perianth of 5 subequal parts, 2-3.5 mm long, 1-2 mm broad, with short thin whitish hairs

on the outer surface and thin silky hairs 3-8 mm long borne from the base of the inner

surface, stamens 1.5-2.5 mm long, forming a short (0.5 mm) tube at the base and with

anthers ca. 0.5 mm long; pistil with short style and broad 2-lobed stigma. Fruit a utricle

ca. 2 mm long, included within the persisting perianth and bracts and obscured by the

long thin whitish hairs from the interior of the perianth parts; seeds lenticular, reddish

brown, ca. 1.5 mm broad.

Uncommon climbing or clambering plants of wet or seasonally dry evergreen

formations and collected in flower and fruit in February, March, and April in

Costa Rica. Most all of our collections come from between 900 and 1,400 m

elevation around the Meseta Central with only a very few from lower areas. The

species ranges from Mexico through Central America to Peru and the Guianas.

Pfaffia grandiflora is recognized by the clambering stems with opposite thin

entire leaves, compound inflorescences with sessile flowers, and long silky hairs

from within the perianth which obscure stamens, pistil, and fruit.

PHILOXERUS R. Brown

Perennial herbs or subshrubs, prostrate, procumbent, or erect, bisexual, glabrous or

puberulent; stipules absent. Leaves opposite and simple, sessile and often clasping the

stem at their base, entire and pinnately veined, usually semisucculent. Inflorescences

solitary, terminal or axillary, short- to long-pedunculate, often subtended by a pair of

small leaves, the flowers sessile or subsessile and densely crowded in short spikes or

capitula, flowers subtended by a bract and 2 bracteoles, the bract and bracteoles char-

taceous and subequal; flowers bisexual and radially symmetrical, with 5 imbricate sub-

equal perianth parts, the inner 2 usually narrower than the outer 3, acute at the apex,

stamens 5, united near the base to form a very short tube, the tube lacking pseudo-

staminodial appendages, anthers 2-thecous, with introrse dehiscence, ovary with 1 locule

and 1 ovule borne on a long funicle, style short and with 2 (3) stigmas. Fruit an orbiculate

indehiscent utricle; seed lenticular to cochleate-orbicular, smooth, the embryo annular.

The genus is most distinctive among our Amaranthaceae, with its very narrow

semisucculent leaves, short dense spikes, and prostrate habit near seashore

environments. Here considered part of the genus Philoxerus, our species has

been placed in the genus Caraxeron Vaillant ex Rafinesque by Mears (Taxon 29:88,

1980).

Philoxerus vermicularis (L.) R. Br., Prodr. 416. 1810. Gomphrena vermicularis

L., Sp. PI. 224. 1753. Caraxeron vermicularis (L.) Rafinesque, Fl. Telluriana 3:38.

1837. Figure 39.

Semisucculent herbs from a woody base, usually procumbent or prostrate and some-

times forming loose mats, leafy internodes 1-8 cm long, 1-4 mm thick (dry), terete but

becoming longitudinally ridged when dry, glabrous but with thin whitish hairs in the axils

of the leaves, succulent and greenish in life. Leaves opposite and decussate, semi-

succulent, clearly differentiated from the stem at the base but not articulate, a petiole not

clearly differentiated and the lamina narrowed to a clasping base; laminae (1) 2-5.5 (8) cm

long, 2-8 (14) cm broad, linear to narrowly oblanceolate, usually broadest above the

middle, rounded to obtuse at the apex but often with a small mucronate tip, gradually

narrowed to the base, slightly expanded at the base and clasping the stem, margin entire,

smooth and glabrous above and below, venation obscure. Inflorescences solitary and

terminal or less often axillary, short-spicate to capitate, usually very short (0.5-1 cm)

pedunculate above a reduced pair of foliage leaves, flowering portion of the spike 6-18 (28)

mm long, 8-10 mm thick, rachis obscured by the flowers and bracts but with thin whitish

178 FIELDIANA: BOTANY

hairs, flowers sessile or subsessile, subtended by a short (1 mm) abaxial bract and 2 lateral

narrow bracteoles ca. 2 mm long; flowers with the perianth 3-3.5 mm long, dull white in

life and becoming pale straw colored when dry.

A species of sandy and rocky seashores on both the Pacific and Caribbean

shores of Central America; probably flowering throughout the year in our area.

The species ranges from Florida, Texas, and Mexico, through Central America

and the West Indies to Ecuador and Brazil; this species is also found on the west

coast of Africa.

Philoxerus vermicularis is immediately distinguished by its seaside habitat,

succulent prostrate stems, very narrow opposite leaves, and dense spikes with

scarious whitish bracts. These plants appear to be quite uncommon on the

beaches of Central America. Only three collections are known from our area:

Pittier 12692 from Puerto Limon, /. Leon 494 from Caldera, Puntarenas, and von

Wedel 1948 from Bocas del Toro, Panama.

PLEUROPETALUM Hooker f.

Herbs or shrubs, the stems sometimes climbing, plants bisexual, stems glabrous or

sparsely and minutely puberulent and becoming glabrous; stipules absent. Leaves alter-

nate in a spiral or occasionally distichous, simple and petiolate, pinnately veined, thin in

texture. Inflorescences mostly solitary, terminal or axillary, racemes or compound pan-

iculate arrangements usually made up of racemose branches and often with reduced

foliage leaves, glabrous or minutely puberulent; flowers bisexual and radially symmetrical,

pedicellate to subsessile, subtended by 1 bract and 2 bracteoles, these often close together

beneath the perianth and simulating a second smaller perianth whorl, the bracts and

perianth parts often with the same stiff texture and color, perianth of 1 whorl of 5 free

oblong somewhat concave subequal sepals (tepals), imbricate in bud, stiff and persisting

in fruit but not enlarging, green to yellowish or red in fruit, stamens 5-8, united near the

base to form a short filament-tube or cup, free portion of the filaments slightly flattened,

anthers 4-thecous, pistil 1, ovary subglobose, 1-Iocular with many ovules on slender stalks

from a central basal placenta, style 1 and very short with 3-8 small stigmas. Fruit at first

slightly succulent and berry-like but becoming dry and capsule-like, opening irregularly

or with circumscissle dehiscence in the lower half, subtended by the stiff perianth parts;

seeds laterally flattened and somewhat lenticular, cochleate-orbicular to reniform-

orbicular in outline, borne on slender funicles, the testa lustrous black; embryo annular.

A neotropical genus of perhaps three species ranging from central Mexico to

Peru and with an endemic species on the Galapagos Islands. These plants are

distinguished by their thin leaves on slender petioles, the flowers with what

appear to be two whorls of stiff straw-colored (dry) perianth parts, the stamens

forming a short cup at their base, the slightly fleshy capsule-like fruit that often

opens around the middle or lower half, and the shiny black seeds borne on

slender stalks from a basal placenta. The distinctive perianth becomes very stiff

when dry, pale yellowish brown and with many longitudinal ridges, resembling

the glumes of grasses.

la Seeds 1.5-2.5 mm long, larger perianth parts usually 3-4 mm long, inflorescence a

compact panicle with rounded contours; plants erect and few-branched, 500-1,900 m

P. sprucei

Ib Seeds 1-1.5 mm long, larger perianth parts usually 4-5 mm long, inflorescences open

racemes or open panicles with racemose branches; plants often climbing or clam-

bering over others, 0-900 m elevation P. pleiogynum

Pleuropetalum pleiogynum (O. Ktze.) Standley, J. Wash. Acad. Sci. 13:369.

1923. CelosiapleiogynaO. Kuntze, Rev. Gen. 541. 1891. P. standleyi Suessenguth,

Repert. Spec. Nov. Regni Veg. 44:41. 1938. Figure 29.

BURGER: FLORA COSTARICENSIS 179

Shrubs, erect or arching over other plants, to 3 m tall or climbing to 5 m high, leafy

internodes 2-6 cm long, 1-4 mm thick, glabrous, weakly longitudinally ridged. Leaves

alternate in a spiral, glabrous, petioles 0.6-2 (4) cm long, about 1 mm thick, with a narrow

adaxial sulcus in the basal half, decurrent on the stem; laminae (3) 5-15 cm long (1) 2-5

cm broad, elliptic-ovate to narrowly elliptic, narrowly elliptic-oblong or lanceolate, gradu-

ally narrowed to the acuminate (acute) apex, attenuate at the base, margins entire and

decurrent on the petiole, the laminae drying chartaceous and usually dark in color,

smooth and glabrous on both surfaces, with (3) 4-6 (8) pairs of major secondary veins.

Inflorescences racemes or panicles of racemes, axillary or terminal, to 30 cm long, the

individual racemes 1-6 cm long, the rachis often with a zigzag form, glabrous or very

minutely puberulent, pedicels 0.5-2 mm long, becoming 5 mm long in fruit, subtended by

an ovate bract 1-2 mm long with thin membranous edges; flowers ca. 4-6 mm long,

subtended by short (2 mm) broad-based bracteoles, perianth ca. 4 mm long, with usually

more than 20 longitudinal ribs, pale yellow to red, ovary usually with 5 or 6 stigmas. Fruit

a capsule 5-7 mm long, opening by circumscissle dehiscence, upper part cream and the

lower part orange in color; seeds 1-1.5 mm long, lustrous black, usually more than 30 per

capsule.

Plants of forest edges and stream sides in wet evergreen formations from near

sea level to 900 m elevation, but only rarely found above 500 m in Costa Rica;

flowering and fruiting collections have been made mostly between August and

April. The species ranges from western Costa Rica to central Panama.

Pleuropetalum pleiogynutn is recognized by the open racemes or panicles of

flowers with stiff persisting perianth parts, the many longitudinal ridges on the

perianth, and the circumscissle dehiscence of the slightly succulent capsule. The

arching, clambering, or climbing stems in partly open sites further distinguish

this species.

Pleuropetalum sprucei (Hook, f.) Standl., N. Amer. Fl. 21:96. 1917. Melano-

carpum sprucei Hook, f., in Benth. & Hook., Gen. PI. 3:24. 1880. P. costaricense

nom. nud., in Hemsl., Biol. Centr. Am. Bot. 3:12. 1882. P. costaricense Wendl. ex

Hook, f., Bot. Mag. pi. 6674. 1883. P. tucurriquenseDonn. Sm., Bot. Gaz. 61:387.

1916. P. calospermum Standl., J. Wash. Acad. Sci. 13:368. 1923. Figure 29.

Small shrubs, (0.5) 1-2 (3) m tall, few-branched and erect, leafy internodes 0.5-5 cm

long, 1.2-6 mm thick, glabrous or minutely and very sparsely puberulent, becoming

longitudinally ridged. Leaves alternate in a spiral, petioles 1-4 (6) cm long, ca. 1.5 mm

broad, with 2 adaxial ridges producing a slight sulcus above, short-decurrent on the stem;

laminae 5-20 (28) cm long, 2-8 (12) cm broad, elliptic to elliptic-oblong or narrowly

elliptic-ovate to lanceolate, tapering gradually to the acuminate apex, tapering gradually

to the attenuate base, margin entire and decurrent on the petiole, the laminae drying thin

chartaceous and usually dark in color, smooth and glabrous above and below or with a few

minute strigillose hairs along the veins, with 7-10 pairs of major secondary veins. In-

florescences terminal or axillary, solitary, usually compact panicles of short racemose

branches, often somewhat flat-topped or a tight convex group (corymbose), 3-9 cm long,

peduncles 1-4 cm long, secondary peduncles and rachis minutely (0.1 mm) and densely

brownish puberulent, pedicels subtended by broad-based bracts ca. 1 mm long, pedicels

1-4 mm long, 2 or 3 small bracteoles usually subtending the flower just beneath the

perianth; flowers ca. 4 mm long, perianth of a single whorl but appearing as 2 whorls

because of the subtending bracteoles, perianth parts free, ca. 3 mm long and 1-1.5 mm

broad, with usually fewer than 20 longitudinal ribs (striations), yellowish green to reddish

at the base, fleshy in life, stamens ca. 2.5 mm long, ovary with usually 3 or 4 stigmas. Fruit

ca. 6 mm long and 5 mm thick, becoming reddish to dark purple, dehiscing irregularly or

circumscissle in the lower half; seeds 1.5-2.2 mm long, usually fewer than 30 per capsule.

Plants of shaded evergreen forest formations from 500 to 1,900 m elevation on

both the Caribbean and Pacific slopes in Costa Rica; probably flowering and

fruiting throughout the year, but with very few collections made in May and

180 FIELDIANA: BOTANY

June and in October and November. The species ranges from central Mexico to

Peru.

Pleuropetalum sprucei is characterized by its usually short erect habit, flowers

usually borne in small dense corymb-like panicles, and relatively large seeds.

The persisting perianth that dries stiff and with many longitudinal ribs and

is subtended by two or three small bracteoles with similar texture is quite

distinctive.

NYCTAGINACEAE

Herbs, shrubs or trees, erect or scandent, bisexual or unisexual, glabrous or puberulent,

with or without spines, sometimes with viscid hairs, the nodes often somewhat thick-

ened, often with dichotomous branching; stipules absent. Leaves alternate, subopposite,

opposite or whorled, simple, petiolate, laminae entire and often with short-linear raphides

visible as projections on the dried surfaces. Inflorescences solitary or several, terminal or

axillary, variously branched but usually with the distal flowers in cymose groupings, with

1-3 bracts or bracteoles subtending each flower, these sometimes resembling a perianth

whorl; flowers bisexual or functionally unisexual, radially symmetrical (in ours), with 1

perianth whorl but often appearing to have 2 whorls because of the sepaloid or petaloid

bracts (as in Bougainvillea) or because of the differentiation between lower sepaloid and

upper corolla-like parts of the perianth (as in Boerhavia), perianth united to form a broadly

campanulate, urceolate to narrowly cylindrical tube, 5-lobed (rarely 3- to 7-lobed) at the

apex or with 5 (3-7) small teeth, involute-plicate in bud or the lobes valvate, base of the

perianth persisting in fruit, distal portion of the perianth persisting or deciduous; stamens

1-10 (many), filaments usually unequal in length and united at the base to form a short

tube or united to the base of the pistil, anthers dorsifixed near the base, included within

the perianth or exserted at anthesis; pistil solitary, often narrowed near the base or

stipitate, ovary with 1 locule and 1 basal stalked ovule, style long to short or absent, stigma

solitary, globose to fimbriate. Fruit usually enclosed within the persisting and protective

perianth to form what is called an anthocarp, becoming fleshy, coriaceous or woody, the

fruit a thin-walled nut that is not usually separable from the enclosing perianth; embryo

strongly curved.

A family of tropical and subtropical plants with most of its species and genera

found in the New World. The flowers and their bracts differ greatly in different

genera, and they are often difficult to recognize as members of the same family.

The perianth tube that is persistent and adherent around the fruit, the often

stipitate ovary with single basal ovule and solitary stigma, the stamens united

at the base to form a short tube, and the occasional elaboration of bracts to

resemble either calyx or corolla are helpful in recognizing this family. The stalked

basal ovule that produces a strongly curved or folded embryo are important

features shared by other members of the Centrospermae (Caryophy Hales). Ex-

cept for a few important ornamentals, the family is of no economic importance;

a number of species have become pantropical weeds.

Two genera of small weedy species with opposite leaves (Allionia with lentic-

ular fruit and dentate winglike margins and Commicarpus with terete fruit with

longitudinal ranks of stipitate glands) are known from Mexico and Guatemala

and in South America, but have never been reported or collected in the regions

between these areas.

Cephalotomandra fragrans Karsten and Triana (including C. panamensis Standley)

looks very much like some of our larger leaved specimens of Neea. However, in

Cephalotomandra, the male perianth forms an open cup and there are more than

20 stamens in each male flower. The genus is not known from Costa Rica, and

considerable doubt exists that a specimen with a collection label from Panama

BURGER: FLORA COSTARICENSIS 181

was actually collected in Panama (see Woodson's treatment in Ann. Missouri

Bot. Card. 48:55-56, 1961).

•

KEY TO THE GENERA OF NYCTAGINACEAE

la Leaves alternate along the stem, none opposite 2a

Ib Leaves opposite along the stem, a few subopposite or alternate leaves occasionally

present 3a

2a Herbs, erect; flowers subtended by small green bracts; uncommon lowland plants

Salpianthus

2b Woody climbers or shrublike; tubular flowers attached to broad brilliantly colored

red, purple, or orange bracts, usually in groups of 3; widely planted for ornament

Bougainvillea

3a Herbs, woody only at the base 4a

3b Trees, shrubs or subshrubs 5a

4a Bracts subtending the flowers inconspicuous; perianth tube with a distinctly

differentiated calyx-like base and corolla-like upper portion, the perianth 1.5-5

mm long Boerhavia

4b Bracts united near the base of the flower to form a 5-parted calyx-like whorl;

perianth tube 1-5 cm long, uniformly corolla-like and brightly colored

Mirabilis

5a Stems frequently armed with spines (but not all stems have spines); small shrubs,

trees, climbers or woody lianas; fruit often borne on very long stalks, with longi-

tudinal ranks of viscid-tipped glands Pisonia

5b Stems never with spines; small shrubs or trees but not climbers; fruit sessile or

stalked, lacking stalked viscid glands 6a

6a Male flowers campanulate with the stamens well exserted from the broad perianth

opening, female flowers with the stigmas exserted at anthesis and the pistil borne on

a short stalk; fruit becoming dark purple or black, longitudinally ridged when dry;

laminae rarely more than 15 cm long, drying dark in color; small to large trees (rarely

shrubs); evergreen forests of the Pacific slope including the Nicoya peninsula, Meseta

Central, General Valley, and Talamanca mountains, 0-2,000 m elevation . . . Guapira

6b Male flowers tubular to ellipsoid, the stamens included within the perianth tube,

female flowers with the stigmas only rarely exserted, pistil sessile or narrowed at the

base; fruit becoming yellow, orange or purple, occasionally longitudinally striate

when dry; laminae to 30 cm long, drying pale or dark; small subshrubs to small trees

(rarely more than 8 m tall); from the highlands and deciduous formations of northern

Costa Rica to the western edge of the Meseta Central, common in the Caribbean

lowlands and less so on the Caribbean slope and Osa Peninsula (apparently absent on

the Talamanca mountains and in the General Valley), 0-1,600 m elevation . . . .Neea

BOERHAVIA Linnaeus

Herbs, annual or perennial, often with a woody base, bisexual, erect or prostrate, often

much branched, stems often with viscous areas on the internodes, gland-tipped hairs

often present. Leaves opposite or subopposite, opposing leaves of a pair often very un-

equal, petiolate, entire or sinuate, raphides often visible as surface irregularities when

dried. Inflorescences terminal, paniculiform to racemeiform with the flowers in small

umbellate, cymose, capitate or racemose arrangements (rarely solitary), 1-3 slender brac-

teoles present beneath each flower. Flowers bisexual, radially symmetrical and very small,

perianth of a single whorl united to form a calyx-like base and a thinner petal-like distal

limb that is shallowly 5-lobed, the upper and lower areas of the perianth tube divided by

a constriction, stamens 1-5, filaments slender and united at the base to form a very short

tube or cup, pistil stipitate, ovary unilocular, style slender, stigma peltate or capitate,

usually exserted. Fruit an anthocarp formed by the tight enclosure of the ovary within the

perianth tube, obovoid to obpyramidal, 3-5 angled (rarely winged), glabrous or glandular-

puberulent.

A genus of weedy plants ranging widely over the tropics and subtropics. Their

MIRABILIS

violacea

FIG. 31. Nyctaginaceae: a cultivated form of Bougainvillea and two species of Mirabilis.

182

SALPIANTHUS

pupurascens

BOERHAVIA coccinea

FIG. 32. Nyctaginaceae: three species of Boerhavia and a species of Salpianthus.

183

(deciduous f PISONIA ___ ^

forest tree) r aculeata (evergreen forest liana)1

FIG. 33. Nyctaginaceae: a species of Guapiraand two species of Pisonia. Note the different

leaf form and habit in P. aculeata.

184

NEEA delicatula

from deciduous

Guanacaste Prov.

NEEA

psychotrioides

sensu stricto

(common form)

FIG. 34. Nyctaginaceae: Costa Rican representatives of the Neea psychotrioides species-

group. These are part of a polymorphic assemblage that may not be reducible to species.

185

NEEA pittieri

A. montane form

(Zarcero, 1500 m)

NEEA amplifolia

(sensu stricto)

FIG. 35. Nyctaginaceae: Costa Rican representatives of the Neea amplifolia species-group.

These are part of a polymorphic assemblage that may not be reducible to species.

186

BURGER: FLORA COSTARICENSIS 187

distribution patterns in Costa Rica give the impression that these plants are not

native elements of the flora. The genus is recognized by its opposite to sub-

opposite leaves which often are very different in size at the same node, the

frequent presence of gland-tipped hairs or viscous areas on the internodes, the

perianth tube that is calyx-like beneath and corolla-like distally, slender fil-

aments united at the base, capitate stigma and fruit resembling an inferior ovary

and broadest near the apex. The following key and species concepts are based

on the treatment of C. D. Adams in the Flowering Plants of Jamaica (1972, pp.

260-261).

la Fruit glabrous and not at all viscid; distal perianth white to pink or lavender; stems and

leaves minutely puberulent or glabrous, the internodes often with glutinous patches

and the leaves usually pellucid-punctate beneath; inflorescence with many vertical

branches and usually lacking leafy bracts; erect plants from a woody rootstock

B. erecta

Ib Fruit minutely viscid puberulent; distal perianth red to dark purple; stems and leaves

minutely puberulent to densely viscid, lacking glutinous patches on the internodes

and pellucid dots on the laminae beneath; inflorescence with spreading branches and

usually with leafy bracts; decumbent to ascending plants 2a

2a Flowers in distal clusters of 3-12, the inflorescences usually small and few branched,

leaves and stems usually densely viscid puberulent B. coccinea

2b Flowers in distal clusters of 2-5, the inflorescences often large and much branched;

leaves and stems often with minute (0.1 mm) and longer (to 1 mm) thin hairs, not

densely viscid puberulent B. diffusa

Boerhavia coccinea Miller, Card. Dictionary ed. 8, Boerhavia no. 4. 1768. B.

caribaea]acq., Obs. Bot. 4:5. 1771.

Decumbent or clambering herbs from a strong rootstock, to 0.8 m tall or ca. 1.2 m long,

leafy internodes 2-12 (15) cm long, 1-5 (8) mm thick (dry), usually densely puberulent

with slender viscid multicellular hairs 0.2-0.8 mm long. Leaves opposite and subequal,

petioles 2-40 mm long, usually densely viscid puberulent; laminae 1.5-5 cm long, 0.7-3.5

cm broad, ovate to ovate-rhomboid or suborbicular, bluntly acute to obtuse or rounded at

the apex, obtuse to cuneate or truncate and rounded at the base, margin entire or slightly

sinuate (undulate), laminae drying thin- to stiff-chartaceous, and usually dark in color on

the upper surface (dry), obscurely and very minutely (0.1 mm) to densely and con-

spicuously viscid puberulent (occasionally almost glabrous), with 3-5 major pairs of sec-

ondary veins. Inflorescences terminal on short lateral branches or terminal on the many-

branched and leafy apex of the plant, flowers in small (3-10 mm broad) clusters of 3-12

on slender ultimate peduncles 3-15 mm long, glabrous to densely viscid puberulent.

Flowers 2-3 mm long, rose-purple to deep red or red-purple, stamens 2, anthers ca. 0.3

mm long. Fruit 3-4 mm long, narrowly obcvoid, ca. 1 mm thick at the thickest part, with

5 prominent longitudinal ribs, surfaces very minutely (0.1 mm) viscid puberulent.

Herbs of recently disturbed or open sandy sites in the lowlands of both the

Caribbean and Pacific coastal plains, ranging to 600 m elevation in northern

Central America; probably flowering throughout the year, but fertile collections

have been made only from June to August and from November to February in

our areas. The species ranges throughout the American tropics to Africa and

Asia.

Boerhavia coccinea is recognized by the small brightly colored flowers tightly

clustered in small groups of usually 4-10, the very short viscid pubescence on

the fruit and usually on other plant parts as well, and the decumbent growth

habit. These plants are frequently found near the seashore and in disturbed sites;

their present distribution in Costa Rica makes it appear that they were not a

native element of the flora. These plants are often placed under B. diffusa.

188 FIELDIANA: BOTANY

Boerhavia diffusaL., Sp. PI. 3. 1753. B. paniculata L. Rich., Actes Soc. Hist.

Nat. Paris 1:105. 1792.

Herbs to ca. 1 m tall, prostrate, decumbent or short-ascending, usually much-branched,

leafy internodes to 20 cm long, sparsely puberulent with very small (0.1 mm) hairs and

longer slender multicellular hairs to 1 mm long, often becoming glabrous. Leaves opposite,

opposing leaves of the same node very different in size, small leaves on short axillary

shoots often present at the widely separate nodes, petioles 3-15 mm long; laminae 1.5-6

cm long, 1-5 cm broad, ovate to ovate-oblong or ovate-rhomboid, bluntly obtuse to

rounded at the apex, obtuse to truncate and rounded at the base, margin undulate-sinuate,

drying chartaceous and with the lower surface often much paler in color than the upper

surface, sparsely and minutely puberulent on both surfaces or with scattered longer hairs

to 1 mm long, often puberulent along the edge, with 3-5 pairs of major secondary veins.

Inflorescences on the open and diffusely branching terminal stems of the plant, small (1

cm or less) leaflike bracts subtending the major inflorescence branches, ultimate peduncles

with 1-5 flowers, very variable in length, glabrous to minutely puberulent. Flowers ca. 2

mm long, pinkish to deep red or purplish in color. Fruit 3.5-4 mm long, narrowly obovoid

to narrowly obtriangular, thickest ( 1 mm) near the apex, with 5 prominent longitudinal

ridges, minutely (0.2 mm) viscid puberulent.

Plants of open recently disturbed weedy sites and sandy or gravelly situations

in open sun on both the Caribbean and Pacific sides of Costa Rica from sea level

to 1,200 m elevation; probably flowering throughout the year (no fertile col-

lections have been made in May, September, October, or December in our area).

The species ranges throughout the American tropics.

Boerhavia diffusa is recognized by the small clusters of red flowers on an open

diffusely branched inflorescence, small viscid fruit thickest near the apex, leaves

of very different size at the same node (both cauline leaves and those of axillary

shoots), and the short stature. Often found along the sides of roads, in vacant

lots in towns, and near the ocean shore, these plants give the impression of being

recent arrivals in Costa Rica. This species is very closely related to B. coccinea, and

it may be that the two are conspecific, but they seem to segregate themselves

quite clearly in our area.

Boerhavia erectaL., Sp. PI. 3. 1753.

Herbs 0.3-1.2 m tall, erect, woody at the base, leafy internodes 2-15 cm long, 2-5 (8)

mm thick, very minutely (0.1 mm) puberulent or becoming glabrous, occasionally with a

viscid area on the internode, nodes with an interpetiolar line or ridge. Leaves opposite,

petioles 2-40 (60) mm long, very minutely puberulent; laminae 1-5 (8) cm long, 0.5-5 (7)

cm broad, ovate to broadly elliptic or triangular, obtuse to bluntly acute at the apex,

abruptly obtuse to truncate or rounded at the base, margin irregularly sinuate, smooth,

sparsely and very minutely puberulent on both surfaces, drying much paler in color

beneath, dark glandular dots often present on one or both surfaces, with 3-5 pairs of major

secondary veins. Inflorescences to 40 cm long, with many slender (0.5 mm dry) opposite

branches, the flowers variously arranged in groups of 2-5, sessile or sometimes an individ-

ual flower on a slender pedicel, inflorescence branches and pedicels glabrous or obscurely

(0.05 mm) puberulent; floral bracts ca. 1 mm long, linear-lanceolate. Flowers 1.5-2.2 mm

long, with a greenish calyx-like base ca. 1 mm long and a thin corolla-like distal portion

0.8-1.2 mm long, pale greenish white to white tinged with pink or pale lavender. Fruit

narrowly obpyramidal, 3-4 mm long and 1.3 mm broad at the apex, tapering gradually to

the narrow base, strongly 5-angled in cross section with 5 longitudinal ridges and grooves,

essentially glabrous.

Weedy plants of open early secondary vegetation in both seasonally very dry

areas and in evergreen formations from just behind the ocean shore to 600 (rarely

to 1,200) m elevation; probably flowering throughout the year, but with most

collections having been made between February and August in Costa Rica.

BURGER: FLORA COSTARICENSIS 189

Widespread in the tropics and subtropics of the Western Hemisphere and in

Africa.

Boerhavia erecta is recognized by the distinctive little glabrous fruit, the diffuse

many-branched inflorescence (in larger specimens), the opposite leaves often

unequal at a node and with irregular outline, and the little flower with a peri-

anth tube that appears to be divided into calyx and corolla. The viscid areas on

the stems and the gland dots on the leaves are also noteworthy, but not always

apparent. This species is very common in our Pacific lowlands, but uncommon

in the Caribbean lowlands where it has been collected only in the general vicin-

ity of Limon.

BOUGAINVILLEA Commerson

REFERENCE: W. T. Gillis, Bougainvilleas of cultivation. Baileya 20:34-41. 1976.

Woody vines, sometimes forming shrubs or small trees, bisexual, glabrous or puber-

ulent, stems often armed with short spines. Leaves alternate and petiolate, laminae entire

and pinnately veined. Inflorescences solitary, borne on modified axillary shoots, usually

consisting of a group of 3 flowers, each flower subtended by a large leaflike bract, the

pedicel of the flower adnate to the midrib of the bract on the inner (adaxial) side and the

bracts surrounding the flowers, the bracts usually 2-4.5 cm long and brilliantly colored.

Flowers bisexual and radially symmetrical, perianth of a single whorl united to form a long

narrow tube with a small star-shaped distal 5-Iobed limb, perianth tube terete or 5-angled,

yellowish to whitish or less often the same color as the bracts; stamens 8-10, filaments

slender and unequal in length, united near the base, anthers included in the tube or a few

exserted; pistil borne on a short stalk, ovary narrowly ellipsoid and unilocular, ovule

solitary, style straight or curved, stigma long and papillose, included within the perianth

tube. Fruit an anthocarp formed by the perianth tube tightly enclosing the fruit and seed,

narrowly ellipsoid, with 5 longitudinal ribs.

A genus of probably 10 to 15 species native to central and southern South

America. These plants are recognized by their woody usually climbing stems,

alternate entire leaves, and brilliantly colored leaflike bracts in groups of three

that include three narrowly tubular flowers. The brilliant bracts which remain

colorful for months and the woody climbing habit which allows the plants to be

grown in a variety of forms over various supports have made these plants

important ornamentals in frost-free areas. The plants rarely set seed and are

propagated vegetatively. Because of their popularity, they are known by a vari-

ety of names: Bogambilla, Bombilla, Bugambilla, Buguenvitia, Flor de Verano, Lus-

trosa, Manto de Jesus, Napoleon, Pomonce, Pompilla, Pritnavera, Santa Rita, Sempre

Lustrosa, Ires Marias, Trinitaria, and Veranera.

These plants are not known as escapes in our area, and because of propagation

by vegetative means, it is difficult to establish species limits. Species descrip-

tions are not provided here (see fig. 31), but the following key, based on Gillis

1976, should be helpful. For a thorough discussion of the cultivars, see L. H.

MacDaniels' article on this subject in Baileya 21:77-100, 1981.

la Perianth tube distinctly angled with longitudinal ribs and with spreading straight

hairs to 1 mm long; stems and leaves villous; rare in Central America

B. spectabilis Willd.

Ib Perianth tube with distinct longitudinal ribs and with upward curving hairs to 0.5 mm

long; stems and leaves glabrous to puberulent; commonly planted in Central

America 2a

2a Leaves broadly ovate, becoming smaller below the inflorescence; bracts deep red or

orange and with obtuse apices, not crinkled; flowers closely grouped; stems with

straight thorns B. buttiana Holtrum and Standley

190 FIELDIANA: BOTANY

2b Leaves narrowly ovate to elliptic, not becoming smaller below the inflorescence; bracts

magenta to purple and with acute apices, often crinkled; flowers usually separate;

stems with curved thorns or thorns absent B. glabra Choisy in DC.

GUAPIRA Aublet

Trees and shrubs, functionally unisexual (dioecious), puberulent in early stages but

often becoming glabrous. Leaves opposite, whorled or occasionally subopposite, often

very variable in size on the same plant or at the same node, petiolate, entire, pinnately

veined. Inflorescences terminal, 1-3, with a single primary peduncle and rachis and

opposite or subopposite branching, densely and minutely brownish or reddish puber-

ulent, becoming succulent in fruit, flowers usually in groups of 2 or 3 on the ultimate

inflorescence branches, and subtended by 1-3 minute bracts. Male flowers obconic, to

campanulate or funnel-shaped, the 5 small distal lobes valvate in bud, green to white or

yellowish, stamens usually 8 (5-10), filaments slender and unequal, united at the base to

the stipe of the pistillode, anthers versatile and exserted well beyond the floral cup. Female

flowers tubular to tubular-campanulate, staminodes with large sterile anthers and in-

cluded within the perianth tube, pistil narrowly ovoid and with a basal stipe adnate to the

filament tube, locule 1, style 1, stigma penicillate or fimbriate. Fruit an anthocarp formed

by enclosure of the ovary within the succulent perianth tube, drupaceous, the free distal

portion of the perianth tube usually persisting (in ours), becoming red or black, longi-

tudinally striate when dry.

A genus of perhaps 50 species ranging from Mexico through Central America

and the West Indies to northern South America, Brazil, Paraguay, and Bolivia

(apparently absent along the Pacific side of South America). These plants are

recognized by their opposite or whorled leaves, terminal inflorescences with

small flowers that appear to be bisexual but are functionally unisexual, perianth

tube subtended by one to three minute bracts, exserted stamens and stigma, and

drupaceous fruit that may appear to be the product of an inferior ovary. These

plants resemble certain members of the Rubiaceae family, but lack stipules, two

perianth whorls, and an inferior ovary. The genus Torrubia has been united with

Guapira, and it is more than likely that Neea should also be submerged under

Guapira.

We have very few collections of this genus from Costa Rica. While our material

appears to represent a single species, there is some question if it is specifically

distinct from G. linearibracteata (Heimerl) Standley of northern Central America

and Mexico (with smaller leaves but larger flowers) and G. standleyanum as

interpreted by Croat in the Flora of Barro Colorado Island (said to grow to 35 m

in height and with puberulent fruit). Considering the perplexing variability in

our material of Neea, it would not be surprising to find similar problems in

Guapira.

Guapira costaricana (Standl.) Woodson, Ann. Missouri Bot. Gard. 48:62.

1961. Torrubia costaricana Standley, Contr. U.S. Nat. Herb. 13:385. 1911.

Small trees 5-10 m tall (rarely shrubs), unisexual, leafy internodes 1-5 (8) cm long, 1.3-4

mm thick with slender brownish multicellular hairs, stems quickly becoming glabrous and

pale grayish in age, little or obscurely lenticellate. Leaves opposite and subequal to very

unequal at a node, often with a whorl of 4 leaves at flowering or branching nodes, usually

drying dark in color, petioles 5-15 (25) mm long, with 2 lateral or adaxial ridges continuous

with the laminae margins; laminae 4-15 (18) cm long, 2-6 cm broad, narrowly elliptic to

elliptic or elliptic-oblong, (occasionally ovate to obovate in the short leaves) tapering to an

acute or short-acuminate apex, acute to obtuse at the base (rounded at the base on the

broad little laminae), entire, smooth on both surfaces, glabrous or becoming glabrous

above and below, drying thin-chartaceous, with 6-10 pairs of major secondary veins, the

venation often somewhat obscure. Male inflorescences 5-10 cm long, primary peduncle

BURGER: FLORA COSTARICENSIS 191

2-6 cm long, peduncle and inflorescence branches minutely brownish puberulent, male

flowers sessile or subsessile, subtended by minute (0.5 mm) bracts, floral cup 2-3 mm long

and 2-3 mm broad at the campanulate apex (dry), filaments unequal, anthers ca. 0.6 mm

long and exserted up to 3 mm beyond the perianth. Female inflorescences to 15 cm long

in fruit, becoming red in color, minutely puberulent; female flowers 2-3 mm long, tubular,

subtended by minute (0.5 mm) bracts, stigmas exserted 0.5-1 mm at anthesis. Fruit

becoming broadly ellipsoid to subglobose, 8 mm long and 5 mm thick (dry), becoming dark

purple or black, with obscure longitudinal ridges.

Rarely collected small trees of evergreen forest formations from (0) 400 to 2,000

m elevation; flowering from February to May in Costa Rica. The species has only

been collected along the Pacific watershed in Costa Rica from the Nicoya pen-

insula and easternmost Guanacaste province to the General Valley and along the

Pacific slopes of the Talamanca mountains. The species ranges from central

Costa Rica into Panama and probably into northern South America.

Guapira costaricana is recognized by the often unequal opposing leaves (occa-

sionally in whorls of 4), the reddish puberulent terminal inflorescences with

very small tubular female flowers and exserted fimbriate stigmas or slightly

larger campanulate male flowers with exserted stamens, and the fruit becoming

dark purple or black on a red inflorescence. These plants are restricted to central

and southern Costa Rica along the Pacific slope and have not been collected in

the same areas as our representatives of Neea, as if they were excluding each

other from the same habitat. The reason for making such a remark is that Neea

has a rather unusual distribution in Costa Rica and, in addition, our species of

Guapira and Neea should probably be united in the same genus. With very dark

fruit on bright red inflorescences, one would expect these plants to be collected

more often; they appear to be restricted to tropical moist premontane wet forest

formations and to flower only in the dry season.

MIRABILIS Linnaeus

Herbs or small subshrubs, perennial but often capable of flowering after only a few

months growth (annual), sometimes woody near the base, usually much branched, the

nodes often thickened, rootstocks often tuberous; stipules absent. Leaves opposite and

simple, petiolate (in ours) or sessile, entire or undulate, pinnately veined. Inflorescences

mostly small cymes or clusters of bracteate flowers in leaf axils or terminal, with 1 or

several flowers subtended by leaflike bracts that are united near the base and form a

5-parted calyx-like involucre (the product of a reduced cyme); flowers bisexual and radially

symmetrical, perianth united and forming a campanulate to funnelform or salverform

tube, the tube short or long, 5-lobed at the apex and induplicate-valvate in bud, the upper

portion deciduous, stamens 3-5, equaling or exceeding the perianth tube, filaments slen-

der and unequal, short-connate at the base, ovary ovoid to globose, style slender with 1

long papillate or capitate stigma. Fruit included within the persisting and slightly enlarged

bracteate (but calyx-like) involucre, achene-like but said to be an anthocarp formed by

adnation with the base of the perianth tube, cylindrical to obovoid, terete, with 5 longi-

tudinal ridges or sulci, often rugose or tuberculate, glabrous or puberulent, constricted just

above the base, mucilaginous when wet; embryo sharply curved.

A genus of perhaps 60 species, nearly all American. One species (M. jalapd) is

now widely naturalized and planted for ornament. The opposite leaves that are

often slightly viscid and triangular in outline, the bracteate calyx-like involucre

subtending one or a few flowers, the showy corolla-like perianth tube usually

bright reddish purple in color, and the unusual fruit distinguish this genus

which can be mistaken for members of the "Tubiflorae" (Gentianaceae,

Solanaceae, etc.). The genus Oxybaphus is no longer considered distinct from

Mirabilis.

192 FIELDIANA: BOTANY

KEY TO THE SPECIES OF Mirabilis

la Perianth tube 3.5 cm long; fruit 7-9 mm long; common plants of weedy and cultivated

sites in evergreen and partly deciduous areas M. jalapa

Ib Perianth tube 0.5-1 cm long; fruit 3-4 mm long; plants restricted to the seasonally dry

Pacific lowlands of central and northern Costa Rica M. violacea

Mirabilis jalapa L., Sp. PI. 177. 1753.

Herbs or subshrubs to 1.3 m tall, annual or perennial, bisexual, much branched, leafy

internodes 1-10 (15) cm long, 1-5 mm thick (dry), usually minutely (0.1-0.5 mm) puber-

ulent with crooked brownish multicellular trichomes, the hairs often dense in longitudinal

grooves and at the thickened nodes, occasionally glabrous. Leaves opposite and usually

equal at a node, petioles 0.8-6 cm long, 0.5-1.5 mm broad (dry), with an adaxial sulcus

or groove, usually sparsely puberulent; laminae 2-10 (14) cm long, 1-6 (8) cm broad,

triangular to ovate-deltoid or ovate, tapering gradually to the acute or acuminate apex,

rounded or abruptly truncate (subcordate) at the base, short-decurrent on the petiole,

margins entire and often ciliolate, the laminae drying membranaceous to thin-

chartaceous, usually sparsely puberulent with scattered hairs 0.3-1 mm long between the

veins and shorter dense hairs on the veins above, usually glabrous beneath, with 3-5 pairs

of major secondary veins. Inflorescence cymose or glomerate in leaf axils or terminal,

occasionally solitary and axillary, peduncles less than 5 mm long; flowers to 7 cm long,

subtended by a sepaloid whorl of 3-5 partly united leaflike bracts 7-15 mm long, green

and sparsely puberulent, the perianth with a long narrow tube and funnelform, to 6 cm

long and 2.5-4 cm broad at the apex, usually deep fuchsia magenta, or purple- red, occa-

sionally yellow, orange, pink, or white; stamens equaling or slightly exceeding the peri-

anth. Fruit 7-9 mm long, ca. 5 mm thick, ellipsoid or ovoid, with 5 thin dark longitudinal

ridges, dark brown and verrucose, with a slight constriction just above the base.

Cultivated plants in gardens and weeds of open early secondary vegetation,

ranging from sea level to about 2,000 m in both evergreen and partly deciduous

formations in Costa Rica; probably flowering throughout the year, but with

collections made primarily from June to January. This species is now widespread

throughout the tropics and subtropics both as an ornamental and a weed; its

origin appears to have been in Mexico.

Mirabilis jalapa is recognized by its tubular, orange, white, or purplish flowers

subtended by a calyx-like involucre of united bracts, opposite leaves with the

larger laminae almost triangular in outline, and the usual association with culti-

vated or disturbed areas. Flowers of different colors are occasionally produced by

the same plant, and a perianth with longitudinally striped colors is sometimes

seen. The species is called Maravilla throughout Central America and "four-

o'clock" in English-speaking countries.

Mirabilis violacea (L.) Heimerl, Beitr. Syst. Nyctag. 23. 1897. Allionia violacea

L., Syst. ed. 10. 890. 1759. Oxybaphus violaceus Choisy, in DC., Prodr. 13, pt.

2:432. 1849.

Herbs, erect, procumbent, or clambering to 1 m tall, annual or with a perennial root-

stock, bisexual, leafy internodes (0.8) 1.5-9 cm long, 0.3-2 mm thick (dry), sparsely to

densely puberulent with thin brownish multicellular or gland-tipped hairs, the trichomes

often restricted to longitudinal grooves. Leaves opposite and equal at a node, petioles

0.5-3 (6) cm long, ca. 0.5 mm broad (dry), glandular puberulent, with 2 abaxial ridges

forming a groove or sulcus above; laminae 1.5-5 (7) cm long, 1-3 (5) cm broad, ovate-

triangular, to broadly ovate or narrowly triangular, tapering gradually to the acuminate

(rarely obtuse) apex, rounded at the cordate to truncate base, margin entire, the laminae

drying membranaceous to thin-chartaceous (sometimes slightly succulent in life), sparsely

puberulent or minutely puberulent only on the veins above, usually glabrous beneath but

with cystoliths conspicuous on the dried undersurface, with 2 or 3 pairs of major second-

BURGER: FLORA COSTARICENSIS 193

ary veins. Inflorescences small terminal or axillary cymes (flowers sometimes solitary and

axillary), usually congested but sometimes to 5 cm long and with peduncles or pedicels

becoming 2 cm long, pedicels thin and densely covered with thin brownish or glandular

hairs; flowers 10-15 mm long and subtended by a calyx-like involucre of broad green

leaflike puberulent bracts 3-4 mm long and enlarging in fruit to 6 mm long, united near

the base and with viscid hairs, perianth tube 5-10 mm long, 5-lobed, purplish or rarely

white, stamens usually 3. Fruit narrowly obovoid or cylindrical, 3-4 mm long, 2-2.5 mm

thick, dark brown, narrowed at the apex and constricted above the base, usually with

paler-colored longitudinal ribs, puberulent.

Small plants of both open early secondary and weedy sites and in shaded

woodland in the seasonally dry deciduous vegetation of the Pacific lowlands in

Guanacaste and northern Puntarenas provinces; flowering collections have been

made from mid- April to August in our area and from March to November in

Nicaragua and Honduras where the plants are found as high as 900 m above sea

level. The species ranges from southern Mexico along the Pacific side of Central

America to the northern half of Costa Rica, Colombia, and Venezuela.

Mirabilis violacea is recognized by its small stature, opposite cordate to truncate

leaves, bracteate involucre united at the base, pink-purplish perianth tube, usual

presence of viscid hairs, and restriction (in our area) to the drier Pacific lowlands.

NEEA Ruiz & Pavon

Small subshrubs to small- or medium-sized trees (rarely more than 8 m tall), unisexual,

usually reddish brown puberulent in early stages, often becoming glabrescent. Leaves

opposite or in whorls of 4 at flowering or branching nodes, occasionally subopposite or

alternate on the same plant, leaves of the same node often differing in size, petiolate,

laminae entire and pinnately veined, often slightly succulent in life. Inflorescences termi-

nal (appearing axillary only with further growth of a subtending axillary twig), usually

solitary at a node (rarely 2 or 3), paniculate or thyrsiform with a single slender peduncle

and usually a single primary rachis with opposite to alternate primary branches, the

flowers often in cymelike distal groups of 3, sessile or pedicellate and with 1-3 minute

bracteoles at the base of the perianth tube, the inflorescence and its branches often bright

red or purple. Male flowers with an ellipsoid to ovoid perianth tube with 5 small perianth

lobes at the narrow apex of the tube, often semisucculent in life, stamens usually 8 (5-10),

included within the perianth tube, filaments unequal in length and fused at the base, a

pistillode present and often of normal size. Female flowers urceolate to tubular, the tube

often constricted beneath the 5 small lobes, usually much smaller than the male flowers,

staminodes present and with normal but nonfunctional anthers, pistil sessile or narrowed

at the base, unilocular with 1 style and a single fimbriate stigma. Fruit a fleshy anthocarp

formed by the succulent perianth tube tightly enclosing the ovary, bright yellow, pink,

deep red or purple, free distal portion of the perianth tube usually deciduous, surface

smooth but the longitudinal ribs often apparent when dry; seed with straight embryo and

contorted cotyledon, little endosperm.

Neea ranges from southern Florida and central Mexico through Central Amer-

ica and the West Indies as far as Bolivia. These plants, with their shrubby or

small tree habit, opposite leaves, and narrow tubular flowers resemble members

of the Rubiaceae family, but that family has stipules, an inferior ovary, and a

calyx and corolla. The terminal inflorescences of Neea often produce dichoto-

mous branching as growth proceeds beyond the flowering node. The genus Neea

may not be sufficiently distinct from Guapira to merit generic rank, in which case

the species listed here would have to be transferred to Guapira (q.v.). I have seen

no reference to the economic use of these plants, though it is said that the fruit

have been used for coloring.

There seem to be no clearly defined species of Neea in Costa Rica. Even the

most unusual plants can be related to more common forms by intermediate

194 FIELDIANA: BOTANY

collections. There may be more species than the two generalized concepts

presented here, but variation is so great among the common forms that there is

no way of clearly delimiting the rarer forms. This troublesome variability ap-

pears to affect all aspects of the plants: habit, leaves, pubescence, inflorescences,

flowers, and fruit. Add to this the problem of relating specimens with male

flowers to those with female flowers or fruit, and one can understand why

identifying species (if such exist in Need) is so difficult. I have illustrated and

written a key to two "species groups"; these can then be further divided by using

the brief subsidiary keys and the illustrations. However, the two "species

groups," treated as two species in a wide sense in the descriptions, have the

same flowering periodicity, the same altitudinal range, and similarly peculiar

distribution patterns within Costa Rica. This seems to indicate that, however

different some of these plants may look from each other, there may be no

barriers to gene flow within the genus in southern Central America and, in

effect, no real species.

KEY TO THE SPECIES GROUPS OF Neea IN COSTA RICA

la Fruit of the lateral inflorescence branches sessile, subsessile or borne on relatively

short thick pedicels shorter than the fruit; leaves becoming quite large (20-35 cm) and

broadly obovate to broadly oblong or broadly elliptic (leaves shorter above 1,200 m

elevation), glabrous or puberulent; small subshrubs, few-stemmed shrubs or small

trees, 1-3 (8) m tall, often found in the deep shade of the forest understory

N. amplifolia s.l.

Ib Fruit of the lateral inflorescence branches borne on slender pedicels, the pedicels often

becoming as long or longer than the fruit (terminal flowers and fruit often sessile);

leaves rarely exceeding 18 cm in length, generally elliptic to elliptic-oblong (sometimes

obovate in smaller leaved individuals), gradually tapering to both apex and base,

usually glabrous; shrubs, small trees or rarely large (20 m) trees, often found in open

secondary growth N. psychotrioides s.l.

Neea amplifolia Donnell Smith, Bot. Gaz. 61:386. 1916. N. orosiana Standl., J.

Wash. Acad. Sci. 15:473. 1925. N. urophylla Standl., Publ. Field Columbian Mus.,

Bot. Ser. 4:203. 1929. N. elegans P. H. Allen, Rain Forests of Golfo Dulce 273 &

409. 1956. Figure 35.

Few-branched subshrubs or small trees, 0.4-2 m tall, unisexual, leafy inter nodes 0-10

cm long, 1-4 mm thick, soon becoming glabrous, smooth and pale colored in later stages.

Leaves usually opposite, petioles 3-30 (65) mm long, with lateral ridges; laminae 8-26 (35)

cm long (2) 5-12 (17) cm broad, obovate to broadly elliptic-oblong, less often broadly

elliptic, oblong or ovate, usually abruptly narrowed to the apex, usually tapering gradually

to the base, margin entire, surfaces smooth, glabrous or minutely puberulent beneath,

with 6-13 pairs of major secondary veins. Inflorescences terminal, rarely more than 6 cm

long in flower (to 10 cm in fruit, peduncle becoming 7 cm long in fruit). Male flowers 5-9

mm long; female flowers 2-4 mm long at anthesis, usually densely puberulent. Fruit

rounded to oblong or ellipsoid, to 1 cm long when dry, usually borne on a pedicel shorter

than the body of the fruit.

Small few-stemmed shrubs of forest shade in moist evergreen formations from

sea level to 1,600 m elevation; flowering from April to November, but fruit have

been collected from January to November. Distribution in Costa Rica is very

similar to that of N. psychotrioides; throughout the Caribbean lowlands, along the

northwestern volcanic chain to San Ramon and Zarcero, uncommon on the wet

Caribbean slope near La Palma and Bajo La Hondura, Muneco, Orosi, and

Turrialba, and it has not been collected in the General Valley or on the Pacific

slopes of southern Costa Rica. The species ranges from Nicaragua to Panama.

BURGER: FLORA COSTARICENSIS 195

Neea amplifolia is recognized by its larger, obovate to very broadly elliptic-

oblong leaves, short stature, small inflorescences, fruit borne on relatively short

pedicels, and shaded forest habitat. There appears to be no way in which this

taxon can be sharply distinguished from N. psychotr hides, and intermediate

plants are found with some frequency. It may be that the plants placed here are

no more than an ecotype found in deeply shaded sites. The two species, N.

amplifolia and N. psychotrioides, share the same peculiar distribution pattern in

Costa Rica, except that N. amplifolia is more common in the drainage area of the

Rio Reventazon and appears to be absent from the Golfo Dulce area. Flowering

periodicity is also nearly identical in the two entities. I believe that the extreme

forms differ so greatly and with sufficient frequency that it may be useful to

retain N. amplifolia at the level of species rank until we have a better idea of what

is actually going on within and between these populations. Neea acuminatissima

Standley with long (ca. 30 cm) leaves that taper very gradually to the acuminate

apex may be a similar species of deeply shaded sites in the Caribbean lowlands

of northern Central America.

Neea pittien 'Standley (Conrr. U.S. Natl. Herb. 13:383, 1911) appears to belong

in this group and is the earliest name available. However, the type is rather

atypical of the assemblage, and its restriction to higher elevations makes its

placement provisional, hence, use of the name amplifolia.

POSSIBLE SEGREGATES OF Neea amplifolia (IN A WIDE SENSE)

la Montane plants above 500 m elevation; leaves often very broad, occasionally obovate;

shrubs 1-3 m tall N. pittien Standley

Ib Lowland plants 2a

2a Small plants (1-2 m tall) of the dark forest understory, leaves often obovate (to

elliptic-oblong), often attacked by leaf miners N. amplifolia sensu stricto

2b Larger plants usually becoming small trees; leaves rarely obovate; uncommon

plants 3a

3a Leaves very broadly oblong or elliptic-oblong, becoming glabrous in age

N. urophylla Standley

3b Leaves elliptic to elliptic-oblong, remaining brownish puberulent in age

N. elegans P. H. Allen

Neea psychotrioides Donnell Smith, Bot. Gaz. 16:199. 1891. N. laetevirens

Standl., Publ. Field Columbian Mus., Bot. Ser. 4:204. 1929. N. belizensisLundell,

Conrr. Univ. Michigan Herb. 7:9. 1942. N. pycnantha Standl., Ann. Missouri Bot.

Card. 30:85. 1943. N. xanthina Standl., loc. tit. 30:86. 1943. N. popenoei P. H.

Allen, Rain Forests of Golfo Dulce 274 & 410. 1956. Figure 34.

Shrubs, small trees or occasionally subshrubs, 1-5 (8) m tall, unisexual, leafy internodes

0-6 (10) cm long, 1-4 mm thick, minute reddish brown tomentulose in early stages but

quickly becoming glabrescent, smooth and terete, the older stems often pale grayish in

color in contrast to the newer stems which often dry very dark. Leaves usually opposite,

often in a whorl of 4 at branching or flowering nodes, occasionally alternate or subopposite

on the same branch, opposing leaves of a node often slightly unequal, petioles 4-30 mm

long, 0.4-1.6 mm thick, with 2 lateral ridges continuous with the lamina margins; laminae

(2) 3-15 (23) cm long, 1.8-8 cm broad, generally elliptic to elliptic-oblong (obovate to ovate

usually only in small-leaved plants of higher altitudes or seasonally very dry areas),

tapering gradually or abruptly to an acute or short acuminate apex, tapering gradually or

abruptly to an acute to obtuse base, often somewhat unequal at the base, margin entire,

surface smooth and glabrous in age (occasionally remaining brownish puberulent be-

neath), with 5-11 pairs of major secondary veins, the midvein often pinkish or reddish in

life. Inflorescences (3) 8-35 cm long, terminal (apparently axillary by continued growth of

196 FIELDIANA: BOTANY

a subtending stem), usually pendulous or becoming so in fruit, often a panicle with a long

primary peduncle and openly dichotomous cymelike distal branches, the primary pedun-

cle to 20 cm long and often exceeding the length of the flowering portion of the in-

florescence, branches of the inflorescence usually minutely (0.1 mm) reddish brown puber-

ulent, floral bracts 0.5-1 mm long, pedicels 0-5 mm long, often equaling or exceeding the

flowers in length. Flowers deep red in bud but becoming yellowish white or greenish

white, male flowers with a floral tube 4-7 (9) mm long, female flowers with a floral tube

2-3 mm long at anthesis. Fruit becoming 6-10 mm long (dry), ellipsoid to subglobose,

ribbed when dry, pink to deep red.

Shrubs and small trees often found in secondary growth and openings in the

forest interior; ranging from sea level to about 1,500 m elevation. The species

appears to flower for much of the year, but we have very few flowering col-

lections from November to January in Costa Rica. The species ranges from

southern Mexico to Panama and probably extends into South America under a

variety of other names. The geographic distribution of this species in Costa Rica

is unusual. It is not uncommon in the northwestern highlands, as far east as San

Ramon and Zarcero, but it has not been collected from the central volcanoes nor

from the Talamanca range. Likewise, though common in the Caribbean low-

lands, it is not known from the General Valley and appears to be quite uncom-

mon in the evergreen forests along the Pacific in southern Costa Rica.

POSSIBLE SEGREGATES OF Neea psychotrioides (IN A WIDE SENSE)

la Leaves usually less than 13 cm long; infructescences usually less than 10 cm long;

Pacific slope of Central America southward to Guanacaste and the Sierra de Tilaran to

the western edge of the Meseta Central, occasional in the Caribbean lowlands in open

sites N. psychotrioides sensu stricto

Ib Leaves or the fruiting inflorescences larger than above; not known from northwestern

Costa Rica or the Pacific slope of northern Central America 2a

2a Peduncles and pedicels long and thin, pendulous from early stages, pedicels often

greatly exceeding the fruit in length, fruit less than 10 mm long; leaves often quite

narrow, rarely collected N. delicatula auct. in herb.

2b Peduncles and pedicels not so thin, becoming pendulous in later stages(?) 3a

3a Trees to 20 m tall, leaves often abruptly acuminate; male flowers to 7 x 4 mm on long

inflorescences, fruit(?) to 2 cm long; rare N. popenoei P. H. Allen

3b Shrubs or small trees, leaves usually tapering gradually or abruptly to the apex;

male flowers smaller; fruit not so long; very common and intergrading with

N. amplifolia N. laetevirens Standley

PISONIA Linnaeus

Shrubs, small trees, or lianas, unisexual, glabrous or puberulent, axillary short straight

or curved spines usually present, younger leaves often developing from the sides of the

spines. Leaves opposite, subopposite, or occasionally in alternate nearby pairs, laminae

decurrent on the petiole, simple and entire, venation pinnate. Inflorescences solitary or

fasciculate, axillary or terminal on highly modified axillary short shoots, the flowers often

borne in cymes but the cymes clustered in umbelliform, capitate, corymbiform, or other

arrangements. Flowers small and radially symmetrical, greenish, unisexual, subtended by

bracteolate pedicels; male flowers with a campanulate perianth tube, the tube with 5

obscure distal lobes or teeth, stamens 6-8 (10), exerted at anthesis, filaments slender,

united near the base to form a small tube and adnate to the base of the pistillode; female

flowers with a narrow perianth tube, staminodes often in the form of a dentate disk adnate

to the narrowed base (stipe) of the pistil, ovary 1-Iocular and with 1 ovule, style narrow

and with a fimbriate stigma. Fruit an anthocarp formed by the union of perianth tube and

pistil, narrowly elongate, terete or 5-angIed in cross section, with longitudinal rows of

stalked glands in 1 or several ranks, the fruit at maturity usually borne on greatly elongated

peduncle and pedicels.

BURGER: FLORA COSTARICENSIS 197

A pantropical genus with species estimated to number between 15 and 50. The

slender coriaceous fruit borne on elongated pedicels and with longitudinal rows

of stalked glands are quite distinctive. These fruits are said to adhere to the

feathers of birds with their viscid glands.

Pisonia fasciculata, described by Standley (Contr. U.S. Natl. Herb. 13:388,

1911), bears fascicles of little subcapitate inflorescences borne on short shoots

and straight spines. The type was collected by C. Wright on the Pacific side of

Nicaragua between 1853 and 1856. This species appears to be a rare endemic of

Central America's seasonally very dry Pacific lowlands, and it might possibly

occur in northwestern Guanacaste.

la Spines usually curved, stems and leaves glabrous or puberulent, laminae often ob-

ovate; fruit 15-20 mm long; shrubs, trees, or lianas in both deciduous and evergreen

formations, 0-1,000 (rarely to 1,800) m elevation P. aculeata

Ib Spines straight, stems and leaves glabrous, laminae usually narrowly elliptic; fruit

8-10 mm long; shrubs or small trees of evergeen montane forests, 1,000-2,200 m

P. silvatica

Pisonia aculeata L., Sp. PL 1026. 1753. P. aculeata var. macranthocarpa Donn.

Smith, Bot. Gaz. 16:198. 1891. P. macranthocarpa Donn. Smith, Hot. Gaz. 20:293.

1895. P. grandifoliaStandl., Contr. U.S. Natl. Herb. 13:391. 1911, not P. grandifolia

Warb., 1891. Figure 33.

Shrubs or small trees to 10 m tall, often with clambering branches, becoming large lianas

in evergreen formations, unisexual, leafy internodes 0.1-6 (12) cm long, 0.7-6 mm thick,

glabrous and reddish brown to densely puberulent or villous and yellowish brown or

grayish, becoming lenticellate, the nodes slightly thickened and often bearing 1 thick

recurved (less often straight) spine from each axil, newer leaves often borne from the side

of the spine. Leaves opposite, subopposite, or occasionally alternate (on the same stem),

the leaves often borne on short shoots in the axils of fallen leaves, petioles 2-40 mm long,

often very variable on the same stem, with 2 adaxial ridges continuous with the lamina

margins; laminae 2-15 cm long, 1.5-6 (9) cm broad, elliptic to obovate or obovate-

orbicular, usually abruptly narrowed to an obtuse, acute or short-acuminate apex, gradu-

ally narrowed to the acute (less often obtuse) base and decurrent on the petiole, margins

entire, the laminae drying thin- to stiff-chartaceous, smooth, glabrous to densely puber-

ulent on both surfaces or often puberulent only along the midvein beneath, with 5-8 pairs

of major secondary veins. Inflorescences solitary and axillary, 2-6 cm long, made up of

little cymes in complex umbel-like or panicle-like arrangements, unisexual, very minutely

puberulent to hirsutulous with trichomes 0.05-0.5 mm long, often viscid, primary pedun-

cles 1-4 cm long, pedicels subtended by small (1 mm) deciduous bracts, both peduncles

and pedicels greatly elongating in fruit; male flowers with a campanulate perianth tube 2-4

mm long and ca. 3 mm broad, stamens 6 and exerted 2-3 mm beyond the perianth, anthers

ca. 0.5 mm long; female flowers with a tubular or narrowly urceolate perianth tube 2-3 mm

long. Fruit borne on open few-branched expanded infructescences with pedicels 1-3 cm

long, the anthocarp 1.5-2 cm long (in ours), 5-8 mm thick, ellipsoid to narrowly obovoid-

clavate, 5-angled (in cross section), with 5 longitudinal ridges, each ridge or angle with

1-3 ranks of stalked viscid glands, surfaces between the ridges densely and very minutely

(0.05 mm) pale grayish or yellowish velutinous.

Shrubs, small trees, or rarely vines in seasonally very dry deciduous for-

mations or becoming large woody lianas in moist or wet evergreen formations,

to about 1,000 (very rarely 1,800) m elevation in our area; flowering and fruiting

from July to March, but with most of our collections made between November

and February. The species ranges from the southern tip of Florida and Mexico,

through Central America and the West Indies, to Ecuador and northern Argen-

tina; the species is also found in central Africa, southern Asia, and the Philip-

pines.

198 FIELDIANA: BOTANY

Pisonia aculeata is recognized by the opposite or subopposite leaves with vari-

able petioles and decurrent lamina bases, the short thick usually curved spines,

the unisexual flowers with small perianth tubes in dense little inflorescences,

and the very unusual fruit on open few-branched infructescences. The species

appears to present a very unusual correlation between life-form and habitat. In

wet evergreen forests, the plants are described as woody lianas reaching the tops

of tall trees, while in deciduous vegetation, the plants are nearly always shrubs

or small trees. Also, the wet forest material is often densely puberulent and has

larger broader leaves (as in Skutch 4848 from the General Valley). We have very

little material representing these wet forest lianas, and it may be that they differ

in other ways and are worthy of taxonomic recognition as a subspecies or

variety. This species is known as Una de Tigre in Central America.

Pisonia sylvaticaStandley, Publ. Field Mus. Nat. Hist., Bot. Ser. 18:425. 1937.

Slender shrubs or small treelets, 1.5-4 m tall, unisexual, often much branched, leafy

internodes 1-6 cm long, ca. 1-2 (4) mm thick, glabrous, pale in color and relatively smooth

with very few lenticels developing, nodes slightly thickened; spines axillary, 0.5-1.5 cm

long, ca. 1.5 mm thick at the base, straight, smooth and sharp-tipped. Leaves opposite,

subopposite or in alternate nearby pairs, petioles 3-10 mm long, ca. 1.3 mm thick, sulcate

above with 2 adaxial ridges continuous with the lamina margins; laminae 4-10 (13) cm

long, 2-4 (5) cm broad, elliptic, narrowly ovate-elliptic, or narrowly oblong-elliptic, taper-

ing gradually to the acuminate or less often acute apex, tip usually blunt, tapering to the

acute base, margin entire and decurrent on the petiole, the laminae drying stiffly char-

taceous, smooth and glabrous on both surfaces, the midvein raised above, the 4-7 pairs

of major secondary veins obscure on both surfaces. Inflorescences axillary and solitary, a

cluster of small cymes on a long peduncle resulting in a capitate appearance in early stages,

peduncles 1-4 cm long, glabrous but the pedicels papillate-puberulent; male flowers ca. 4

mm long and borne on pedicels to 3 mm long, the perianth tube campanulate with the

usually 10 stamens extended 2-4 mm beyond the tube, anthers ca. 0.5 mm long; female

flowers 2-3 mm long (dry), perianth tube minutely papillate puberulent, style and fim-

briate stigma exerted. Fruit borne on pedicels elongating to 3 cm and becoming reddish,

body of the fruit (anthocarp) 8-10 mm long, 3-4 mm thick (dry), narrowly ellipsoid to

ellipsoid-cylindrical, style and stigma persisting and exerted, with 10 longitudinal rows of

single-ranked stalked gland- tipped projections ca. 0.5 mm long.

Shrubs and small trees of evergreen premontane and lower montane moist

forest formations between 1,000 and 2,200 m elevation; flowering and fruiting

collections have been made from October to December and February and March.

This species is known only from the areas near Monteverde in the Sierra de

Tilaran and from near San Ramon and Zarcero on the western edge of the Meseta

Central in Costa Rica.

Pisonia silvatica is recognized by the glabrous parts (except for the flowering

parts of inflorescences), opposite to subopposite leaves with straight axillary

spines, closely clustered flowers on a distinctive peduncle, and the fruit borne

on greatly elongated pedicels and with 10 longitudinal rows of stalked glands.

It is interesting that this endemic has never been found in areas east of Zarcero

where collecting has been extensive; it may occur in the Sierra de Guanacaste

which has been little collected.

SALPIANTHUS Humboldt and Bonpland

Herbs or small shrubs, usually much branched, bisexual, usually glandular puberulent,

at least near the inflorescences; stipules absent. Leaves alternate and simple, laminae

usually decurrent on the petiole, entire. Inflorescences small racemes, cymes or congested

clusters, often in distal paniculate arrangements, terminal or axillary, bracts subtending

BURGER: FLORA COSTARICENSIS 199

the flowers very small and inconspicuous; flowers bisexual and radially symmetrical,

sessile or pedicellate, perianth tube cylindrical to urceolate, subcoriaceous, with 4 or 5

short lobes, covered with gland-tipped hairs (uncinate hairs present in ours), with 4 or 5

longitudinal ribs, not enlarging in fruit, stamens 3 and borne from 1 side of the perianth

tube, filaments often unequal, anthers rounded, ovary sessile and with a slender style,

stigma simple and acute, ovule solitary and basal. Fruit an achene-like utricle, the persist-

ing glandular-pubescent perianth tube tightly enclosing fruit and seed; seeds very smooth

and lustrous.

A genus of a few, principally Mexican, species. Salpianthusis closely related to

Mirabilis among our representatives of the family, but lacks the corolla-like tube

and subtending bracteate involucre. The broad basal triangular leaves, gland-

tipped hairs on distal parts, stiff calyx tube (perianth tube) with hooked hairs,

3 stamens with thin filaments and rounded anthers exceeding the tube, and

shiny black seed tightly enclosed in the persisting perianth tube help to dis-

tinguish our species of this genus.

Salpianthus purpurascens(Cav.) Hook. & Arn., Bot. Beechey Voy. 308. 1837.

Boldoa purpurascens Cav . ex Lagasca, Gen. & Sp. Nov. 10. 1816. B. ovatifoliaLag.,

loc. cit. Figure 32.

Herbs or subshrubs to 1.5 (2) m tall, much branched, leafy internodes 1-10 cm long 1-7

mm thick, glabrous or very minutely puberulent, angulate, the distal inflorescence bearing

stems densely viscid puberulent. Leaves alternate, very variable in size on the same plant

with smaller leaves on the distal flowering stems, petioles 0.3-10 cm long, with lateral

ridges becoming winglike distally and continuous with the lamina margins; laminae 2-20

cm long, 1.5-18 cm broad, ovate to ovate- triangular or rhombic-ovate, tapering gradually

to the bluntly acute or obtuse apex, rounded and truncate to subtruncate above the

cuneate and long decurrent base, margins entire, the laminae drying thin- chartaceous,

very minutely (0.1-0.2 mm) puberulent above, very minutely puberulent or glabrous

beneath, short linear cystolith-like projections on the dried surface beneath, with 3-6

pairs of major secondary veins, the basal secondaries often very prominent and the ve-

nation subpalmate. Inflorescences small (1-5 cm) terminal and axillary panicles of short

racemes or glomerules, peduncles and pedicels covered with small (0.2-0.3 mm) gland-

tipped hairs; flowers sessile or subsessile, perianth tube 3-4 mm long, ca. 1.5 mm in

diameter, narrowly urceolate to tubular, perianth lobes obtuse, perianth covered with both

gland-tipped and sharply hooked (uncinate) trichomes. Fruit tightly enclosed within the

persisting (but not enlarged) perianth ca. 3 mm long; seed ca. 1.5 mm long, suborbicular

in outline, thick lenticular, lustrous black.

Plants of open fields and partly shaded woodlands from sea level to about 300

m elevation in Costa Rica; flowering from June to January in southern Central

America. The species ranges from Mexico along the Pacific side of Central Amer-

ica to Guanacaste Province in Costa Rica, and it also occurs in Cuba and

Venezuela.

Salpianthus purpurascens is recognized by its very broad ovate triangular leaves

usually found at the base of the plant, the small flower clusters on distal stems

with small leaves, the short narrow perianth tube with both gland-tipped and

sharply hooked hairs on its surface, and the shiny black seed tightly enclosed

within the persisting perianth tube. This species was earlier placed in the genus

Boldoa.

PHYTOLACCACEAE

REFERENCES: Katherine Raeder, Phytolaccaceae in Flora of Panama. Ann. Mis-

souri Bot. Card. 48:408-421. 1961. Joan W. Nowicke, Palynotaxonomic study of

the Phytolaccaceae. loc. cit. 55:294-364. 1969.

TRICHOSTIGMA

polyandrum

T. octandrum

RIVINA humilis

10 cm MICROTEA debilis

FIG. 36. Phytolaccaceae: Costa Rican representative of Microtea, Rivina, and Trichostigma.

200

PHYTOLACCA

rugosa

P. rivinoides

1 * * * * * * * * * * * ^

10 cm

Fie. 37. Phytolaccaceae: three species of Phytolacca.

201

1 mm

FIG. 38. Phytolaccaceae and Chenopodiaceae: a species of Chenopodium and species of

Petiveria and Stegnosperma of the Phytolaccaceae.

202

BURGER: FLORA COSTARICENSIS 203

Herbs or succulent subshrubs, erect or clambering, rarely larger woody shrubs or trees,

bisexual (in ours) or less often unisexual, glabrous or sparsely and minutely puberulent;

stipules absent or small and usually inconspicuous. Leaves alternate, simple and entire,

pinnately veined, often with short linear crystals within. Inflorescences terminal, axillary

or leaf-opposed usually solitary, racemose, spicate or compound paniculate with spicate

or racemose branches, bracts and minute bracteoles present or absent; flowers small and

bisexual in ours (unisexual by abortion), radially symmetrical or rarely somewhat bilater-

ally symmetrical, perianth usually of 1 whorl of 4 or 5 (rarely more) perianth parts (sepals

or tepals), free or united near the base, imbricate in bud, usually persisting in fruit,

greenish white to pink or purple, stamens often as many as the perianth parts and

alternate with them or 3 to many and variously arranged, filaments free or united only at

the base, often borne on a hypogynous disk, slender, anthers 2-thecous, dorsifixed or

basifixed, dehiscing laterally; pistil simple or compound, 1-carpellate (unilocular) or of 2

to many carpels free or variously united, each locule with a single basal ovule, stigmas

linear to capitate, sessile or borne on a short style. Fruit various, made up of 1 or more free

or united carpels, fleshy or dry (rarely winged), subtended by the persisting perianth; seed

subglobose to lenticular or reniform, erect, embryo annular or curved, endosperm present.

A family of about 17 genera and 70 to 80 species, with the majority of genera

and species confined to the South American tropics. The flowers are character-

ized by a single perianth whorl that persists in fruit, and stamens alternating

with the perianth parts when of the same number. However, the number of

stamens and carpels varies greatly, occasionally on the same plant. The locules

of the ovary always have a single basal ovule that produces a curved or annular

embryo. The vegetative parts are often semisucculent and glabrous or only very

sparsely puberulent.

The genus Hilleria is not known to occur in Central America or Panama.

Hilleria subcordata Standl. & L. O. Wms., described in Ceiba 3:199 (1953), was

based on a collection from Turrialba (/. Leon 3488), and this is undoubtedly a

specimen of Scutellaria of the Labiatae family with immature flowers. Species of

the genus Hilleria differ from all our representatives of the family in having a

bilaterally symmetrical perianth with 3 united perianth parts and 1 free perianth

part. The plants are similar to Rivina, but with more densely flowered racemes

and a dry somewhat flattened fruit.

Agdestis clematoides Mocino & Sesse ex DC. from Mexico and northern Central

America is similar to our species of Trichostigma, but the pistil is partly inferior

and has three or four stigmas that become recurved; the leaves are usually

subcordate and the inflorescences become white. This species is sometimes

planted as an ornamental climber, but it has not been recorded from southern

Central America. Its common names are Bejuco de ajo and Vomita because of the

plant's unpleasant odor when crushed.

KEY TO THE GENERA OF PHYTOLACCACEAE

la Flowers subsessile on long open spikes; fruit a narrow achene with 4 sharply retrorse

spines at the top, partly enclosed by the dry persisting appressed perianth; usually

found below 500 m elevation Petiveria

Ib Flowers borne on easily seen pedicels in open racemes or panicles; fruit without

spines, usually globose, persisting perianth not ascending appressed 2a

2a Perianth of 5 or 10 parts; ovary with 2-15 stigmas 3a

2b Perianth of 4 parts; ovary with a single stigma 5a

3a Fruit a capsule splitting open with valves, usually with 1 seed; stamens 10; woody

plants of the very dry deciduous lowlands of northern Guanacaste

Stegnosperma

3b Fruit a berry or drupe, not splitting by valves; herbs and subshrubs not known

to occur in drier northern Guanacaste . . . . 4a

204 FIELDIANA: BOTANY

4a Succulent subshrubs 0.3-4 m tall; ovary with 5-15 locules and 5-15 stigmas, fruit

a fleshy berry with 5-15 seeds; stamens 6-30; common plants from 0-3,300 m

elevation Phytolacca

4b Small semisucculent herbs to 0.8 m tall; ovary with 1 locule and 2 stigmas, fruit

small and globose with spiny ridges forming a reticulum on the surface; stamens

usually 5 (3-9); uncommon plants, below 500 m elevation Microtea

5a Herbs or subshrubs with erect stems; stamens 4; fruit becoming bright red . . Rivina

5b Woody climbers or shrubs with clambering stems; stamens 8 or more; fruit usually

becoming black Trichostigma

MICROTEA Swartz

Small annual herbs, erect or decumbent, stems usually somewhat succulent and gla-

brous; stipules absent or represented by minute tubercles. Leaves alternate and usually

small, petiolate or subsessile, laminae simple and entire, glabrous. Inflorescences terminal

and axillary, 1 to several at a node, racemose, spicate or paniculate, with floral bracts and

with or without bracteoles; flowers very small, bisexual, sepals 5 (4), free above a thickened

receptacle, persisting in fruit, stamens 5 (3-9), usually alternating with the perianth parts,

free or united near the base, anthers dorsifixed, thecae globose and dehiscing laterally,

ovary with 1 locule and 1 ovule, style short, stigmas 2 or 3. Fruit fleshy or hard, smooth

to tuberculate or echinate, the pericarp adhering to the seed; seed erect, embryo curved,

endosperm fleshy.

A neotropical genus of nine species, with only one species reaching con-

tinental North America. The plants resemble some small Caryophyllaceae and

slender stemmed Chenopodiaceae.

Microtea debilis Swartz, Prodr. Veg. Ind. Occ. 53. 1788. Figure 36.

Annual herbs, 15-80 cm tall, usually much branched with prostrate or ascending stems,

leafy internodes 0.5-3 (5) cm long, 0.5-2 mm thick and pale greenish gray when dry,

longitudinally ribbed and glabrous, somewhat succulent in life. Leaves alternate and often

distichous, slightly succulent, petioles 2-20 mm long, poorly differentiated from the blade

with lateral margins continuous with the lamina margins; laminae 0.6—4 (8) cm long, 0.5-2

(3.5) cm broad, elliptic to obovate or ovate, bluntly obtuse to acute at the apex, attenuate

at the base, margin entire, the laminae drying thin chartaceous to membranaceous,

smooth and glabrous on both surfaces, the 4-7 pairs of major secondary veins arising at

angles of ca. 40°-60°. Inflorescences terminal or axillary, often 2 (1) at a node, 1.5-6 cm

long, open racemes or occasionally paniculate with 2 racemose axes, rachis 0.1-0.4 mm

thick (dry), glabrous, flowers on short (0.3-2 mm) thin pedicels, subtended by mem-

branous linear bracts ca. 1 mm long; flowers ca. 1 mm long, tepals 5, narrow, borne on the

edge of a short receptacle, white, 0.5-0.7 mm long, stamens ca. 0.4 mm long, ovary ca. 0.5

mm long, glabrous. Fruit globose, greenish, ca. 1.5 mm long, with a reticulum on the

surface formed by interconnected ridges of tubercles, subtended by the persisting perianth

parts; seed black.

Uncommon weedy little plants of the wet evergreen lowland forest formations

between sea level and 100 (300) m elevation on both the Caribbean and Pacific

coastal plains of Costa Rica; probably flowering throughout the year, but col-

lections from our area have only been made from December to May and in

August. The species ranges from Guatemala and the West Indies to Peru and

Brazil.

Microtea debilis is recognized by its small habit, glabrous slightly succulent

parts becoming very thin when dried, alternate leaves with winged petioles,

minute flowers on open racemes, and unusual little fruit with reticulate surface.

This species has only been collected a few times in Costa Rica: from the Carib-

bean lowlands and the Golfo Dulce region. The plants appear to prefer sandy

soils and are often found along stream and river banks.

BURGER: FLORA COSTARICENSIS 205

PETIVERIA Linnaeus

Erect herbs or subshrubs, woody at the base, minutely puberulent or glabrescent, stems

with longitudinal ridges, tissues often with the odor of garlic (Allium sativum); stipules

paired and lateral. Leaves alternate in a spiral or distichous, petioles canaliculate adaxially,

the laminae simple and entire, acute to acuminate. Inflorescences slender elongate spike-

like racemes of distant flowers subtended by short bracts, terminal or axillary and solitary

or several from a node, the flowers very short pedicellate or subsessile; flowers bisexual

and radially symmetrical, small, perianth of 4 free parts, equal and acute, narrow, spread-

ing at anthesis, persisting and erect in fruit, stiff and dry, stamens 4-9, alternate with the

perianth parts or irregularly placed, shorter than the perianth, filaments filiform, anthers

dorsifixed and linear, 2-cleft at apex and base, ovary unilocular with 4 or 6 reflexed spines

near the apex, stigma solitary, sessile and penicillate, ovule erect. Fruit a long narrow

cuneate achene, laterally compressed with 2 slightly flattened carinate sides, apically

2-lobed with 4 or 6 retrorse or recurved spines, pericarp adhering to the seed; seeds erect

and linear.

A New World genus of one species with two varieties: one widely ranging

throughout the warm areas of the hemisphere and the other in Brazil. The plants

resemble certain Amaranthaceae. Petiveriais distinguished from the Amarantha-

ceae because it lacks stamens that are equal in number and opposite to the

perianth parts and because the stamens are not basally united.

Petiveria alliacea L., Sp. PI. 342. 1753. Figure 38.

Herbs or small subshrubs, 0.5-1.5 (2) m tall, base and lower stems woody, leafy inter-

nodes 1-6 cm long, 1.5-4 cm thick, minutely (0.1-0.3 mm) puberulent with slender

ascending pale-colored hairs, longitudinally ridged; stipules ca. 2 mm long and 0.4 mm

broad, persisting, brownish. Leaves alternate in a spiral or distichous, petioles 4-18 mm

long, ca. 1 mm thick, minutely puberulent, with an adaxial sulcus; laminae 5-16 (20) cm

long, 2-6 cm broad, elliptic to elliptic-oblong, short acuminate at the apex (occasionally

acute), acute at the base, the margins entire, the laminae drying thin to stiffly chartaceous,

smooth above and below, sparsely and very minutely puberulent along the veins but often

becoming glabrous, the 5-8 pairs of major secondary veins arising at angles of 40°-70°.

Inflorescences 1-5 in terminal groups or axillary, slender spikes 10-40 cm long, the flowers

5-15 mm distant on the slender (0.5-1 mm) sparsely puberulent rachis, flowers subtended

by scarious triangular bracts 1-2 mm long; flowers ca. 3-4 mm long, subsessile or on very

short (less than 3 mm) pedicels, puberulent on the lower parts, perianth parts 4, white or

greenish, stiff and scarious, 0.5-1 mm broad, stamens 8 (4, 6), free, to 3 mm long, ovary

with 4 retrorse spines near the apex, stigmas 2. Fruit 6-8 mm long, somewhat flattened

and oblanceolate with a deep apical notch separating 2 lateral projections, each of which

bears 2 (3) retrorse spines 1.5-3 mm long, greenish and longitudinally striate, minutely

and sparsely puberulent, subtended by the persisting appressed bracts and perianth, the

fruit and bracts and perianth appressed against the axis of the inflorescence, ascending.

Plants of weedy habitats in partly open as well as shaded sites in both season-

ally dry and deciduous habitats and in wet evergreen formations from sea level

to 300 (rarely to 1,500) m elevation in Costa Rica; probably flowering throughout

the year, but collected with flowers and fruit in our area from August to February

and in April and June. The species ranges from the southern United States

through the West Indies and Central America to Argentina and Peru.

Petiveria alliacea is recognized by its small weedy habit with woody base,

glabrescent alternate leaves, and long spikes with separate scarious flowers that

resemble grass spikelets. The unusual fruit with four sharp recurved spines at

the apex and vegetative parts that often smell like garlic (Allium sativum) further

distinguish this species. Our material belongs to variety alliaceae as defined by

Nowicke. These plants are easily mistaken for members of the Amaranthaceae

206 FIELDIANA: BOTANY

family, while the inflorescences resemble those of some Acanthaceae and Gram-

ineae. Pseudelephantopus of the Compositae also resembles this species.

PHYTOLACCA Linnaeus

Herbs or subshrubs, rarely trees, bisexual in ours, stems erect or clambering, often

somewhat succulent, glabrous or sparsely puberulent; stipules absent. Leaves alternate in

a spiral, simple, sessile or petiolate, margin usually entire, usually glabrous, pinnately

veined. Inflorescences at first terminal but soon becoming leaf-opposed or extra-axillary,

spikes, racemes or raceme-like panicles, pedicels subtended by narrow bracts and often

with 1 or 2 small bracteoles along the length of the pedicel; flowers small and radially

symmetrical, bisexual (in ours) or unisexual, the perianth of 1 whorl of 5 usually free

tepals, the tepals greenish or white to pink and purple, androecium of 5-30 stamens in 1

or 2 whorls, sometimes differing in number in flowers of the same inflorescence, filaments

free or united at the base and borne on a disk, anthers dorsifixed; ovary usually broader

than long, made up of a whorl of 5-16 partly or completely united carpels, the 5-16 locules

with 1 erect ovule each. Fruit usually a fleshy and juicy berry, broader than long with a

centrally depressed apex, 5- to 16-ribbed or parted; seed compressed, embryo curved.

A genus of worldwide distribution, but with most of the 30-odd species in the

American tropics and a few species reaching the temperate zone. The broad

gynoecium with incompletely fused carpels, variable number of stamens, single

perianth whorl, and solitary erect ovule are distinctive features. The semi-

succulent stems and leaves and frequent presence of bright rose red to purplish

pigments in vegetative parts and fruit are also characteristic. Hybridization has

been suspected between many of the species. Plants of this genus resemble,

superficially, Sphenodea zeylanica Gaertn. (Campanulaceae or Sphenocleaceae)

and some species of Monnina (Polygalaceae).

As can be seen in our illustration, P. icosandra is intermediate between P.

rivinoides and P. rugosa in some respects. Jerold I. Davis, who has studied

Phytolacca material from throughout Central America, believes that the earlier

proposed species concepts can be placed in a linear arrangement, reflecting

morphological transitions and hybridization, as follows: P. rivinoides, P. ico-

sandra sensu stricto, P. purpurascens, P. meziana, P. octandra, P. rugosa, P. cos-

taricensis. This sequence also reflects an altitudinal shift from lowland P. riv-

inoides to P. costaricensis above 3,000 m elevation, with P. rivinoides being most

easily separated from among the group (Davis, personal communication, 1980).

While hybridization and intergradation do exist, it is my impression that the

keys and species descriptions provided here will effectively categorize a large

majority of our Costa Rican collections and that hybrid populations or inter-

mediate individuals are relatively rare in Costa Rica.

The fruit contain saponins and have been used as a substitute for soap in many

areas. The common name Jaboncillo, widely used in Central America, reflects this

usage. Other names used in our area are Calalu and Tinta, the latter referring to

the pigments sometimes used as ink (mostly from the fruit). See the discussion

of uses in the Flora of Guatemala (Fieldiana, Bot. 24, part 4:198, 1946).

la Inflorescence axis glabrous, 15-80 cm long; ovary with more than 10 styles, fruit with

more than 10 ribs, fruiting pedicels 5-18 mm long, styles closely connivent; 0-1,300

(1,800) m elevation P. rivinoides

Ib Inflorescence axis with small scurfy or scalelike hairs; ovary with usually 8 styles, fruit

with usually 8 ribs or seeds, fruiting pedicels less than 6 mm long 2a

2a Inflorescence axis 5-15 (20) cm long, pedicels 3-6 mm long; fruit with the styles 1-2

mm distant from the central apex; (1,500) 2,000-3,300 m elevation P. rugosa

2b Inflorescence axis 10-36 cm long, pedicels 0.3.-3 mm long; fruit with the styles close

BURGER: FLORA COSTARICENSIS 207

together at the apex in the depressed center of the fruit; (500) 1,000-1,500 (1,800) m

elevation P. icosandra

Phytolacca icosandra I ., Syst. Nat. ed. 10. 1040. 1759. Figure 37.

Herbs or stout subshrubs to 1.5 (2) m tall, leafy internodes 3-40 mm long, (1) 2-8 mm

thick, longitudinally ribbed when dry, glabrous. Leaves with petioles 5-30 mm long, ca.

1 mm thick (dry) and with lateral margins continuous with the lamina margins, decurrent

on the stem; laminae (3.5) 5-15 (35) cm long, (1) 2-10 (18) cm broad, elliptic or occasionally

ovate-elliptic, tapering to the sharply acute apex, the tip of thickened tissue drying

yellowish and sharp-pointed, attenuate at the base and decurrent on the petiole, margin

entire, the laminae drying thin chartaceous, smooth and glabrous on both surfaces, the

7-9 pairs of secondary veins arising at angles of ca. 45°-60°. Inflorescences usually be-

coming leaf-opposed (9) 12-25 (36) cm long, the raceme only 5-15 mm broad, elongating

in fruit, peduncle 3-6 cm long, the rachis minutely scurfy-puberulent, drying yellowish

but pink to purplish in life, pedicels 0.3-3 mm long, subtended by linear bracts 3-6 mm

long and ca. 0.5 mm broad; flowers ca. 6 mm broad, perianth parts 2.5-3.5 mm long, white

to pale purple, stamens (8) 16-18 (20), 3 mm long, styles usually 8 (6-10). Fruit becoming

dark blue or black, ca. 8 mm broad and 4 mm long, the styles borne close together within

the apical depression at the top of the fruit; seed lustrous black, lenticular, ca. 3 mm long

and 2.5 mm broad.

Plants of recently disturbed secondary formations in and around the Meseta

Central in Costa Rica; probably flowering throughout the year, but with no

collections having been made in September. The Costa Rican collections come

primarily from between 1,000 and 1,500 m elevation with only a few as low as

500 or as high as 1,800 m. The species ranges from Mexico and the West Indies

to northern South America and Ecuador.

Phytolacca icosandra is recognized by the long narrow racemes with subsessile

flowers, styles remaining close together as the fruit expands, and scurfy in-

florescence axis. This species is quite similar to P. rugosa, but the two seem to be

readily separable in our area. Recent collections made by Robert Wilbur and his

associates from the area of the eastern Meseta Central and western Talamanca

mountains indicate that there may be considerable intergradation between P.

icosandra and P. rugosa between 1,500 and 2,500 m elevation.

Phytolacca rivinoides Kunth & Bouche, Ind. Sem. Hort. Berol. 15. 1848, and

Ann. Sci. Nat. Bot. ser 3, 11:231. 1849. Figure 37.

Herbs or weak stemmed subshrubs 1-2 (5) m tall, leafy internodes (1) 2-7 (11) cm long,

1.5-12 mm thick, glabrous and smooth, often reddish or pinkish in life, developing very

small lenticels. Leaves with petioles (1) 2-7 cm long, slightly sulcate above and with lateral

ridges continuous with the lamina margins and decurrent on the stem, often reddish in

color; laminae 5-17 (21) cm long, 2-7 (9) cm broad, elliptic to ovate in outline, tapering

gradually to the sharp-acuminate apex, acute to obtuse or somewhat rounded and sub-

truncate at the base, margin entire and decurrent on the petiole, the laminae drying

thin-chartaceous, smooth and glabrous on both surfaces, often with small punctations

above, the 6-11 pairs of major secondary veins arising at angles of about 60°. Inflorescence

terminal and often becoming positioned just below an opposing leaf, 15-50 (80) cm long,

the rachis essentially glabrous, peduncle 3-8 cm long, 1.2-2 .4 mm thick, glabrous and

smooth, pedicles 3-6 (13) mm long and subtended by thin lanceolate bracts ca. 3 mm long

and with 1 or 2 minute bracteoles along the length of the pedicel, the rachis and pedicels

smooth and glabrous, often pink to purplish; flowers ca. 5 mm broad (dry), perianth parts

2.5 mm long, white to rose or magenta, stamens 9-17; ovary usually green with 10-16

styles. Fruit borne on pedicles 5-18 mm long, ca. 0.5 mm thick (dry), fruit becoming 6-8

mm broad with 10-16 distinct longitudinal ribs, becoming blue or black when ripe, styles

closely borne together in the depressed central apex; seeds lenticular, ca. 2 mm long, dull

black, difficult to separate from the dry fruit.

208 FIELDIANA: BOTANY

Plants of open disturbed and early successional vegetation in wet and moist

evergreen formations from sea level to 1,300 (1,800) m elevation in Costa Rica;

flowering throughout the year. The species has not been collected below 500 m

elevation in Guanacaste or northern Puntarenas. The species ranges from Mex-

ico and the West Indies southward to Bolivia.

Phytolacca rivinoides is recognized by the smooth glabrous inflorescences that

become unusually long, the fruit with 12 to 16 ribs, and a preference for moist

habitats below 1,300 m elevation. Despite the distinct differences between this

species and the other Costa Rican species, hybridization has been reported with

P. rugosa at about 1,600 m near Cervantes.

Phytolacca rugosa Br. & Bouche, Ind. Sem. Hort. Berol. 13. 1851, and Linnaea

25:296. 1852. P. costaricensis Suessenguth, Bot. Jahrb. Syst. 72:273. 1942. Fig-

ure 37.

Herbs or subshrubs 0.3-2 (4) m tall, the basal stems somewhat woody, leafy internodes

0.5-4 (10) cm long, 2-8 mm thick, glabrous or with scurfy hairs, longitudinally ridged

when dry and the ridges continuous with the decurrent leaf bases. Leaves often closely

approximate at the ends of stems in exposed and windy environments, petioles 0.2-3 (5)

cm long, with thin lateral margins continuous with the lamina margins and decurrent on

the stem; laminae (4.5) 6-12 (19) cm long, (1) 2-4 (6.5) cm broad, narrowly elliptic to

elliptic-oblong, gradually tapering to the acute or sharply acuminate apex, acute at the

base, margin entire and decurrent on the petiole, the laminae drying thin to stiffly char-

taceous, smooth and glabrous or with scattered minute (0.1-0.2 mm) scurfy hairs on the

veins beneath, the 6-8 pairs of secondary veins arising at angles of ca. 45°. Inflorescence

terminal and becoming more or less leaf-opposed, 5-15 (20) cm long, at first with the

flowers clustered in a compact conic raceme but soon elongating, peduncle 2-6 cm long

and at first exceeding the flowering rachis in length, rachis and pedicels usually scurfy-

rugose and drying yellowish but often bright pink or reddish purple in life, pedicels 3-6

mm long, subtended by linear bracts ca. 5 mm long, with 1 or 2 bracteoles along the length

of the pedicel; flowers 4-5 mm broad (dry), perianth parts ca. 3 mm long, pink to reddish,

stamens 6-12 in 1 cycle, 1.3-2 mm long, ovary with usually 8 styles. Fruit 6-10 mm broad

and 2-4 mm high, green becoming dark purple, styles becoming 1-2 mm distant from the

central depressed apex of the mature fruit; seeds lenticular, lustrous black, ca. 3.5 mm long

and 2.5 mm broad.

Plants of recently disturbed and open early successional stages in evergreen

highland forest formations between 1,500 and 3,300 m elevation; flowering and

fruiting throughout the year. The species ranges from Mexico to Colombia.

Phytolacca rugosa is recognized by its relatively broad racemes with scurfy

rachis and distinctly pedicellate flowers and the fruit with clearly separate styles

borne some distance from the central apex. This is the only species of Phytolacca

to be found commonly above 2,000 m elevation in Costa Rica. There may be

hybridization between this species and P. icosandra.

An unusual collection by W. Hatheway (2358) from near the summit of Volcan

Poas has the flowers borne on a superficially racemose inflorescence that is

actually paniculate. The main axis bears short lateral flowering branches with

several flowers, but this collection seems to differ in no other way from typical

P. rugosa.

RIVINA Linnaeus

Perennial herbs or subshrubs, usually woody near the base, glabrous or minutely puber-

ulent; stipules absent. Leaves alternate in a spiral, on slender petioles, simple and entire,

BURGER: FLORA COSTARICENSIS 209

thin in texture, pinnately veined. Inflorescences erect solitary open racemes, terminal or

becoming axillary, pedicels subtended by narrow little deciduous bracts, 1 or 2 minute

bracteoles often present at the apex of the pedicel beneath the perianth, the inflorescence

usually minutely puberulent; flowers bisexual and small, radially symmetrical, perianth of

4 subequal parts in 2 decussate whorls, slightly imbricate in bud, oblong spatulate and

usually rounded at the apex, corolla-like and becoming white, persisting and enlarging

only slightly in fruit, stamens 4, free, alternating with the perianth parts and inserted on

the slightly elevated receptacle (hypogynous disk), slightly shorter than the perianth,

anthers elongate and dorsifixed, extrorse-lateral, pistil simple, ovary with 1 locule and 1

basal ovule, style terminal or subterminal, often curved. Fruit a globose red drupe with

pericarp adherent to the seed, subtended by the persisting perianth; seed lenticular, often

with a covering of short hairs, embryo annular.

A genus of apparently only one somewhat variable species, probably of New

World origin but now naturalized throughout the tropics. The genus is very

closely related to Trichostigma, and some authors have treated the included

plants as congeneric.

Rivina humilisL., Sp. PI. 122. 1753. Figure 36.

Perennial (occasionally annual?) herbs and subshrubs 0.3-1 (rarely 2) m tall, usually

woody at the base and sometimes with the lower stems woody, leafy internodes 1-8 cm

long, 1-3 mm thick, glabrous or more often puberulent with thin whitish haris 0.1-0.5 mm

long, longitudinally ridged when dry and greenish; stipules absent. Leaves alternate in a

spiral, petioles 0.4-4 (8) cm long, ca. 1 mm thick when dry, usually puberulent, with a

slight adaxial groove above; laminae (2) 4-12 (17) cm long, (1) 2-6 (8) cm broad, ovate to

elliptic-ovate or lanceolate, tapering gradually to the acuminate apex, rounded and trun-

cate to rounded and obtuse at the base, margin entire, the laminae drying thin-chartaceous

and usually greenish, smooth above and below, usually glabrous above (except on the

midvein) and sparsely puberulent on the veins beneath, the thin pale-colored hairs ca. 0.2

mm long, linear crystals usually visible as minute (0.3 mm) prominent lines between the

veinlets of dried leaves, the 5-7 pairs of major secondary veins arising from the midvein

at angles of ca. 45°. Inflorescences terminal or axillary, racemes 4-12 (16) cm long (be-

coming longer after anthesis), rachis and pedicels usually with very short (0.2 mm) thin

whitish hairs, bracts 1-2 mm long and deciduous, pedicels 2-4 mm long at anthesis and

up to 8 mm long in fruit, slender and only ca. 0.2 mm thick when dry; flower buds ca.

2 mm long, perianth parts ca. 2 mm long, green becoming white or pinkish white,

filaments 1-1.8 mm long, anthers ca. 0.7 mm long, ovary ca. 1 mm long with a style 0.5

mm long. Fruit becoming a bright red or orange globose fleshy drupe, 4-6 mm in diameter,

subtended by the persistent and usually reflexed or spreading perianth parts (in late

stages), the perianth parts to 4 mm long and 1-1.5 mm broad; the seed often with a

reticulate surface formed by the adherent pericarp, seeds 2-2.5 mm broad, dark brown and

with minute brownish hairs on the surface.

Plants of wet or seasonally dry evergreen formations or in moist situations

(such as riparian understory) in deciduous areas between sea level and 1,200

(1,600) m elevation in Costa Rica; flowering throughout the year, but collected

most often from December to January and June to September. The species ranges

from the southern United States (Texas and Oklahoma to Florida) through

Mexico and the West Indies to South America as far as Argentina; the species

is widely naturalized in the Old World.

Rivina humilis is recognized by its short stature from a woody base, thin ovate

leaves on slender petioles, open racemes, flowers with only four perianth parts

and four alternating stamens, and simple pistil that forms a bright red drupe.

The fruits are said to have been used as a source for red pigment. The plants

resemble some small species of Solanaceae.

210 FIELDIANA: BOTANY

STEGNOSPERMA Bentham

REFERENCES: H. G. Bedell, A taxonomic and morphological re-evaluation of

Stegnospermaceae. Syst. Bot. 5:419-431, 1980. D. J. Rogers, Stegnosperma: A new

species and a generic commentary. Ann. Missouri Bot. Gard. 36:475-477, 1949.

Shrubs or small trees, the branches often drooping, vining or clambering, bisexual,

glabrous; stipules absent. Leaves alternate and simple, petiolate and entire, glabrous, often

slightly succulent and drying stiff. Inflorescence racemose, terminal or axillary, solitary,

the pedicels subtended by a bract and 2 bracteoles; flowers bisexual and radially sym-

metrical, borne on slender pedicels, perianth of 2 whorls, the 5 sepals free and imbricate,

thin, persisting and enlarging slightly in fruit, petals or staminodia 5, free, shorter than the

sepals, whitish, stamens 10, filaments slender but expanded near the base and connate to

form a very short ring (perigynous annulus), anthers basifixed and sagittate; ovary supe-

rior and ovoid or ellipsoid, 3- to 5-Iocular but becoming unilocular with a central column,

ovules basal, as many as the locules or style branches (stigmas). Fruit a globose or ovoid

capsule, opening by 3-5 coriaceous valves, styles persisting; seeds 1-5, cochleate-

ellipsoid, with a smooth dark reddish or black lustrous surface and a white or yellow-

ish aril.

A genus of three species; two species are found only in Baja California and

adjacent Mexico, while the third species ranges from Mexico to Costa Rica and

the Greater Antilles. The capsular fruit, two perianth whorls, and woody habit

distinguish this genus among our representatives of the family. The genus has

only recently been found in Costa Rica. This genus has been elevated to family,

but its position in the Caryophyllales remains the same, close to the putatively

primitive Phytolaccaceae (see Bedell, 1980).

Stegnosperma cubense A. Rich., in Sagra, Hist. Nat. Cuba 10:309; 12; tab. 44.

1845. Trichilia scandens A. Robinson ex Lunan, Hort. Jam. 2:319-320. 1814, an

invalid polynomial (fide Rogers). S. scandens (Lunan) Standley, Publ. Field Mus.

Nat. Hist., Bot. Ser. 23:6. 1943. Figure 38.

Small trees or shrubs with scandent or clambering stems, to 3 m tall or 4 m long, leafy

internodes 2-20 mm long, 1-4 mm thick, smooth and glabrous, pale in color but soon

becoming dark. Leaves alternate in a spiral, petioles 5-12 mm long, deeply sulcate above

with 2 longitudinal adaxial ridges continuous with the laminae margins; laminae 2-8 (11)

cm long, 1-5 (7) cm broad, obovate to orbicular or elliptic, obtuse to rounded at the apex,

obtuse to acute at the base and decurrent on the petiole, margins entire, laminae drying

stiffly chartaceous, smooth and glabrous, with 3-5 pairs of major secondary veins (often

obscure). Inflorescences 3-13 cm long, the flowers closely crowded or separated by as

much as 5 mm on the slender rachis, bract and bracteoles 1-1.5 mm long, pedicels 2-12

mm long; flowers 4-6 mm long (dry), sepals 3-4 mm long, broadly imbricate, petals

included within the sepals, filaments 3-4 mm long, slender and united near the base,

anthers 1.5 mm long, styles and stigmas usually 4 (3-5), ca. 1 mm long. Fruit an ovoid to

subglobose capsule 6-7 mm long, subtended by the persisting perianth and with persist-

ing styles, often with only 1 seed per fruit; seed ca. 5x3 mm, cochleate-ellipsoid in

outline, lustrous black, the surface smooth or slightly irregular.

Small trees or clambering shrubs in seasonally very dry deciduous formations

of the northwestern Pacific lowlands in Costa Rica. This species ranges from

northern Mexico to Guatemala and along the Pacific lowlands to Costa Rica and

from Cuba to Puerto Rico in the West Indies.

Stegnospermum cubense is distinguished by its small rounded leaves, lack of

pubescence, terminal racemes, conspicuous pedicels, small petals hidden by the

sepals, and capsular fruit with shiny black seed. This species and genus were not

known to exist in Costa Rica until the recent intensive surveys of Santa Rosa

National Park in Guanacaste Province. These little trees with drooping branches

BURGER: FLORA COSTARICENSIS 211

or shrubs with clambering branches often form thickets behind sand dunes and

on rocky outcrops in northern Central America. Our specimen (Liesner 2284) has

much larger leaves than most Central American material, and the staminal cup

appears to be less well developed. More collections from Guanacaste are needed

to determine whether the Costa Rican populations merit separate specific or

subspecific status.

TRICHOSTIGMA A. Richard

Shrubs or woody climbers, usually glabrous; stipules minute and caducous or not

apparent. Leaves alternate in a spiral, petiolate, the laminae simple and entire, pinnately

veined, chartaceous when dry. Inflorescences terminal or axillary, solitary, elongate open

racemes, the flowers clearly pedicellate, bracts subtending the pedicels present and cadu-

cous or absent, the pedicels often with small bracts and 2 minute bracteoles; flowers

bisexual and radially symmetrical, perianth of 4 subequal parts in 2 whorls, broadly

imbricate in bud persisting and usually reflexed in fruit, becoming reddish, stamens 8-25,

free, borne on the elevated receptacle (hypogynous disk), filaments slender, anthers dor-

sifixed, longer than wide, pistil simple with sessile or subsessile penicillate stigma, ovary

1-locular and with 1 ovule. Fruit a globose fleshy drupe, pericarp attached to the seed;

seed solitary with annular embryo and curved cotyledons, erect.

A New World genus of three species closely related to and perhaps congeneric

with Rivina (q.v.). The plants are recognized by their climbing or trailing stems

with glabrous leaves, open inflorescences that become reddish or purple in fruit,

flowers with only four perianth parts and eight to 25 stamens, and simple pistil

forming a fleshy black fruit. Like Phytolacca the open inflorescence and its pedi-

cels often become wine red or purplish in color. While differing primarily in

floral characters, the two species found in Costa Rica are apparently most com-

mon in the same restricted area: the Sierra de Tilaran and its Caribbean drain-

age area.

la Flowers with ca. 20 stamens, the filaments less than 0.5 mm long and never persisting

in fruit, perianth becoming 5-10 mm long in fruit; the racemes becoming 10-25 cm

long T. polyandrum

Ib Flowers with usually 10 stamens, the filaments to 3 mm long and often persisting

beneath the fruit, perianth not exceeding 5 mm in length; racemes up to 15 cm long

T. octandrum

Trichostigma octandrum (L.) H. Walter, in Engler, Pflanzenr. IV, 83:109.

1909. Rivina octandra L., Cent. PI. 2:9. 1756. Figure 36.

Scandent shrubs or lianas, growing to a height of ca. 10 m, leafy internodes 5-50 mm

long, 1-5.5 mm thick, glabrous, often with prominent lenticels, grayish when dry. Leaves

alternate in a spiral, petioles 1-5 cm long, 0.5-1 mm thick, deeply sulcate above (adaxially);

laminae 6-15 cm long, 2-7 cm broad, elliptic to narrowly elliptic-oblong or narrowly ovate,

acute to short-acuminate at the apex, acute at the base with the margins shortly decurrent

on the petiole, margins entire, laminae drying chartaceous, smooth and glabrous on both

surfaces, the 6-9 (12) pairs of major secondary veins arising at angles of 45°- 70°, often

loop-connected near the margins but difficult to see. Inflorescences solitary and axillary

racemes, 5-11 cm long, peduncle and rachis glabrous, ca. 0.5 mm thick and longitudinally

striate when dry, peduncle (distance to the first flower) usually 2-3 cm long, pedicels 3-6

mm long and 1-6 mm distant on the rachis, with a linear bracteole ca. 2 mm long halfway

up the pedicel; flowers ca. 4 mm long, perianth parts ca. 4 mm long, white, stamens

usually 12 with anthers ca. 2 mm long borne on persisting filaments 2-3 mm long; pistil

1.5 mm long, the penicillate stigma ca. 0.5 mm long. Fruit becoming 8 mm long, globose,

black, succulent, surface venation not readily visible when dry.

212 FIELDIANA: BOTANY

Plants of wet evergreen forest formations on both the Caribbean and Pacific

sides of Costa Rica between 10 and 800 (1,000) m elevation; fruiting collections

have been made from January through April. This is a wide-ranging species

found from Mexico and southern Florida (U.S.A.) through Central America and

the West Indies to Argentina.

Trichostigma octandrum is recognized by the climbing stems, glabrous parts,

racemose inflorescences with slender rachis and pedicels, numerous stamens

persisting as filaments with the fruit, and black succulent drupes. While rela-

tively few collections of this species have been made in Costa Rica, it is inter-

esting to note that many of these come from the same areas where collections of

the congener, T. polyandrum, have been made: Tilaran and the areas bordering

the San Carlos plain. However, this species has been collected in the Golfo Dulce

area, and T. polyandrum has not. The relationship between these two species

growing in the same areas and the differences in their flowers might make an

interesting study.

Trichostigma polyandrum (Loes.) H. Walter, in Engler, Pflanzenr. IV, 83:112.

1909. Rivina polyandra Loesner, Bot. Jahrb. Syst. 23:123. 1896. Figure 36.

Scandent or erect shrubs 1-3 (5) m tall, the stems usually clambering, leafy internodes,

0.5-16 cm long, 0.8-3 mm thick, glabrous or less often very minutely (0.1 mm) puberulent

with straight erect hairs, becoming longitudinally ridged when dry. Leaves alternate in a

spiral, closely spaced on lateral stems as well as widely spaced on long climbing stems,

petioles (0.6) 1-3 cm long, 1-2.5 mm thick, broadly to narrowly canaliculate above; lami-

nae 7-16 (18) cm long, 2-7 (9) cm broad, narrowly ovate to elliptic-ovate, acuminate at the

apex, obtuse to acute or occasionally rounded at the base, margin entire, the laminae

drying thin-chartaceous, smooth and glabrous above and below or with very minute erect

hairs on the veins beneath, the 5-8 pairs of major secondary veins arising at angles of

60°-80°, arcuate ascending distally and often loop-connected near the margin. In-

florescences terminal or axillary, open racemes of distant flowers 7-15 cm long and be-

coming up to 25 cm long in fruit, peduncle and rachis glabrous or occasionally very

minutely (0.1 mm) puberulent, 0.3-1 mm thick (dry) and reddish or purple after anthesis,

pedicels (3) 6-12 (20) mm long with a small (1 mm) linear bracteole on the basal half, larger

(2 mm) bracts subtending the pedicel sometimes present in early stages and caducous, 2

minute (0.5 mm) bracteoles sometimes present on the distal half of the pedicel; flower buds

2-4 mm long, green to pale greenish white, perianth becoming 6-10 mm long in fruit and

rose to red or purple in color, stamens 20-25 with very short (0.5 mm) filaments and

anthers ca. 1.5 mm long, quickly caducous after the flower opens, pistil 3 mm long and 1

mm thick. Fruit 5-8 mm long and equally broad, broadly ellipsoid or ovoid, becoming

black, succulent but with the veins prominent on the outer surface when dry, subtended

by the 4 persisting and brightly colored perianth parts.

Plants of evergreen wet forest formations of the Caribbean slope and lowlands,

from sea level to 1,000 (rarely 1,500) m elevation; flowering and fruiting col-

lections have been made from January to October in Costa Rica. This species is

especially common in the wetter parts of the Sierra de Tilaran and the Caribbean

lowlands of Alajuela Province; the species ranges from southern Nicaragua to

western Panama.

Trichostigma polyandrum is recognized by its climbing stems, long fruiting

inflorescences with distant and long-pedicellate fruit usually subtended by four

red or purplish perianth parts. This species is more commonly collected than T.

octandrum in our area, perhaps, because it has a longer flowering and fruiting

season.

BURGER: FLORA COSTARICENSIS 213

AIZOACEAE

Herbs or subshrubs (rarely shrubs), annuals or perennials, prostrate or erect, stems and

leaves often succulent, glabrous or puberulent; stipules small and thin or absent. Leaves

alternate, opposite or whorled, simple and entire, equal or unequal at a node, sessile or

petiolate, petioles often with a thin sheathing margin at the base; laminae often fleshy,

sometimes reduced to scales. Inflorescences cymose or the flowers solitary, in the axils of

leaves or terminal, flowers sessile or pedicellate, bisexual (rarely unisexual), small (in

ours), radially symmetrical, perianth usually of 1 whorl of 5 (less often 3 or 4) free or united

sepals (tepals), often with an extension or appendage on the back of the midvein distally,

petals (actually staminodes) present and numerous in some cultivated and introduced

genera but not in our native and naturalized species; stamens 3, 4, 5, 10 to many, alternate

with the sepals when of the same number, filaments free or united into fascicles, rarely

united to the perianth, anthers dehiscing longitudinally; pistil 1, ovary superior (in ours)

or inferior, 1- to 5-locular (rarely 6- to 20-locular), placentation axile, basal, apical, or

parietal but not free central, ovules 1 to many in each locule, styles as many as the locules,

stigmas usually narrow, slender. Fruit a loculicidal or circumscissile capsule or indehiscent

and nutlike or baccate; seeds cochleate to reniform, with mealy endosperm, rarely stro-

phiolate, embryo usually curved.

The family, though a member of the very natural order Centrospermae (Caryo-

phyllales), is not easily defined. Some of its genera appear to form links with the

Caryophyllaceae, Portulacaceae, and Chenopodiaceae. The family is very di-

verse in southern Africa, with Australia a secondary center of diversity. Mem-

bers of the family are rarely met with in the wild in Costa Rica. The unusual

succulent forms from South Africa, with larger many-parted flowers, such as

Mesembryanthemum (sensu lato) and the "flowering stones" are popular with

gardeners and collectors of exotic plants. Tetragonia expansa Murr. (Espinaca or

"New Zealand Spinach") is grown in Central America as a vegetable; the alter-

nate slightly fleshy deltoid leaves are abruptly narrowed to the petiole and

become as much as 12 cm long.

la Leaves whorled or opposite, not very fleshy and not sheathing or clasping the stem;

perianth parts free to the base; fruit a loculicidal capsule splitting longitudinally into

3-5 valves 2a

Ib Leaves opposite and fleshy, sheathing or clasping the stem at their base; perianth

parts united to form a short tube, perianth lobes with small distal appendages; fruit

opening by a transverse slit (circumscissile) with the distal lid or cap coming off, 1-

to 5-locular 3a

2a Plants usually puberulent with slender stellate hairs; leaves oblanceolate to ob-

ovate; flowers sessile or subsessile, perianth with slender distal appendages;

seeds ca. 0.3 mm broad, strophiolate (with an aril-like development at the base)

Glinus

2b Plants glabrous, leaves generally very narrow; flowers on slender pedicels; peri-

anth without appendages; seeds ca. 0.5 mm broad, estrophiolate Mollugo

3a Leaves linear to narrowly obovate, basal sheath sometimes united across the stem but

without appendages, laminae equal or subequal at a node, young stems glabrous;

ovary 3- to 5-locular, styles 3-5, seeds numerous, 0.5-1.5 mm broad Sesuvium

3b Leaves obovate to rounded-spatulate, sheathing at the base and with an appendage

on the united interpetiolar sheath, laminae often very unequal at a node, young stems

often with a longitudinal ridge bearing crooked hairs; ovary 1- or 2-locular, styles 1

or 2; seeds less than 15, 1-2 mm broad Trianthema

GLINUS Linnaeus

Herbs, annual, procumbent or ascending, stems slightly succulent, often much

branched, with simple or stellate hairs; stipules absent. Leaves opposite or whorled by

214 FIELDIANA: BOTANY

condensation, often unequal at a node, petiolate; laminae linear to orbicular, entire. In-

florescences of fasciculate or solitary flowers in the axils of leaves, sessile or pedicellate,

often forming congested verticels; flowers bisexual, radially symmetrical, small and incon-

spicuous, perianth of 1 whorl but sometimes with petaloid staminodes, sepals (tepals) 5,

thin and stiff, free, staminodes 0-20, often divided apically; stamens 3-30, free, filaments

slender and borne from the base of perianth; ovary superior, 3- to 5-locular, with numer-

ous ovules on axile placentae, styles short 3-5. Fruit a loculicidal capsule breaking into 3-5

thin-walled valves; seeds numerous and reniform, smooth or granulate, borne on slender

funicles, strophiolate.

A genus of six to 10 species, four native to Africa and only one in our area.

These small diffuse herbs are easily mistaken for members of the Caryo-

phyllaceae.

Glinus radiatus(R. & P.) Rohrb., in Mart., Fl. Bras. 14, pt. 2:238, pi. 55, f. 1.

1872. Mollugo radiata Ruiz & Pavon, FI. Peruv. 1:48. 1798. Figure 42.

Herbs, prostrate or ascending, to 50 (80) cm tall, bisexual, leafy internodes 0.5 cm long,

0.3-2 (3.5) mm thick (dry), densely to sparsely puberulent with thin whitish stellate hairs.

Leaves opposite or whorled, often unequal, petioles 1-5 (12) mm long, with lateral ridges

continuous with the lamina margins; laminae 2-15 (30) mm long, 2-15 mm broad, elliptic

to obovate or spatulate, usually broadest above the middle, obtuse or rounded at the apex,

obtuse to cuneate at the base, margins entire and decurrent on the petiole, sparsely to

densely puberulent with stellate whitish or grayish white hairs to 1 mm broad, venation

pinnate with 1-3 pairs of major secondary veins. Flowers clustered in the leaf axils, sessile,

or subsessile, densely or sparsely puberulent, perianth parts 4-4.5 mm long, greenish,

with a short slender distal extension ca. 0.7 mm long. Fruit ca. 4 mm long and 2 mm thick,

subtended and enclosed loosely by the persisting perianth; seeds many, 0.3-0.4 mm

broad, reniform, smooth and bright reddish brown, lustrous.

A small weedy plant of open sites and often found on muddy river banks, road

sides, cultivated fields and thickets in the Pacific lowlands (0-600 m) of Central

America. Though the species is said to be common in the wet season, we have

very few collections from Central America, and those were mostly made in the

dry season; flowering material has been collected in April and June. The species

ranges from Mexico to central Costa Rica and from the West Indies to South

America.

Glinus radiatus is recognized by its small habit, opposite or whorled leaves,

thin-branched stellate hairs, inconspicuous flowers tightly clustered in the leaf

axils, and minute shiny reddish brown seeds. I have seen only a single collection

from Costa Rica: Brenes 3872 from "Los Liros." These plants are very similar to

some species of Caryophyllaceae.

MOLLUGO Linnaeus

Herbs, annuals or short-lived perennials, stems usually slender and much branched,

often prostrate, not conspicuously succulent, glabrous; stipules minute or absent, decid-

uous. Leaves usually in whorls, rarely opposite, often beginning with a basal rosette;

laminae simple and entire, usually broadest above the middle (often very narrow), gla-

brous. Inflorescences cymose or of solitary axillary flowers forming verticels at each node,

pedicellate; flowers small, radially symmetrical, bisexual, perianth of 1 whorl of 5 equal or

subequal tepals (sepals), imbricate in bud, free to the base; stamens 3-5 (rarely 6-10); ovary

3- to 5-locular, superior, ovules usually numerous on axile placentae, styles 3-5, short and

slender. Fruit a thin-walled loculicidal capsule, breaking into 3-5 valves; seeds few to

many, reniform to cochleate, embryo curved.

A genus of about 20 species, mostly in Africa and Asia, ranging from the

tropics and subtropics into the temperate zone. These plants are very similar in

BURGER: FLORA COSTARICENSIS 215

appearance to some of our species of Caryophyllaceae. Only one species is found

in Central America.

Mollugo verticellata L., Sp. PI. 89. 1753. Figure 41.

Herbs, annuals to ca. 0.5 m tall, prostrate or erect, with much-branched slender stems,

leafy internodes 1-9 on long, 0.2-1 mm thick (dry), glabrous; stipules absent. Leaves 4-6

at a node in whorls or pseudowhorls (by condensation), variable in length and shape

(often on the same plant), petioles 1—4 mm long and little differentiated from the lamina,

slightly expanded at the base; laminae 5-30 (50) mm long, 0.5-4 (10) mm broad, linear to

narrowly oblanceolate, broadest above the middle in larger leaves, acute at the apex,

tapering gradually into the petiole, margins entire and continuous with the lateral margins

of the petiole, glabrous and smooth, venation pinnate but usually obscure. Inflorescences

verticel-like with 3-6 flowers in the axils of leaves (at a node), pedicels (3) 5-15 mm long,

very slender (0.1-0.3 mm) and glabrous; flowers 2-3 mm long, white, perianth parts 2-2.5

mm long, 0.5-0.8 mm broad; stamens 3, to 2 mm long; ovary 3-locular, 1-1.5 mm long,

styles 3. Fruit a loculicidal capsule subtended by the persisting perianth parts, 2.5-3 mm

long, ovoid, thin- walled; seeds 20-30 per capsule, ca. 0.5 mm broad, cochleate to reniform,

ridged along the curved back, dark reddish brown, lustrous.

Plants of open early secondary growth; often found on sandy sites along

streams and in cultivated fields. Mollugo verticellata ranges from sea level to 500

(rarely 1,000) m elevation on both the Pacific and Caribbean slopes in Central

America; flowering throughout the year, but with very few collections made in

August and September. This species has a pantropical distribution, but appears

to be uncommon in southern Central America.

Mollugo verticellata is recognized by its diffuse herbaceous habit, slender stems,

verticels of narrow leaves, and small flowers on slender petioles. This species has

been called Clavellina monies in El Salvador, according to Standley. It is easy to

mistake this species for members of the Caryophyllaceae, such as Arenaria lanu-

ginosa and Spergula arvensis, but these plants are restricted to higher elevations in

Central America.

SESUVIUM Linnaeus

Herbs or subshrubs, prostrate to decumbent or erect, often rooting at the nodes, bisex-

ual, glabrous, stems succulent; stipules absent. Leaves opposite, subopposite or alternate,

simple and succulent, sessile or petiolate, clasping or sheathing at the base with the leaf

bases of opposing leaves united across the node; laminae entire, linear to ovate. In-

florescences of solitary axillary flowers, sessile or pedicellate, subtended by 2 minute

bracteoles; flowers bisexual, radially symmetrical, perianth of 1 whorl (sepals or tepals)

united at the base to the floral cup (calyx cup), 5-parted, triangular, imbricate in bud, with

a dorsal (abaxial) distal appendage; stamens 5-15 or many, filaments free or united at the

base, borne on the floral cup; ovary superior and with 2-5 locules, ovules usually many

on axile placentae, styles 2-5, slender. Fruit a circumscissile capsule, the apical lid coming

off as a single unit; seeds several to many, black, cochleate to annular or reniform.

A genus of about six species: one a pantropical plant of the seashore, one

endemic to the Galapagos Islands, and several restricted to eastern Africa.

Sesuvium portulacastrum (L.) L., Syst. Nat. ed. 10:1058. 1759. Portulaca por-

tulacastrum L., Sp. PI. 446. 1753. Figure 39.

Herbs, perennial, prostrate or suberect, often creeping and rooting at the nodes and

much branched, occasionally forming mats, stems thick and succulent, leafy internodes

(0.5) 3-50 (80) mm long, 1-5 mm thick (dry), glabrous; stipules absent but the opposing

leaf bases sheathing the stem and united. Leaves opposite, simple, petioles 2-6 mm long

but not clearly differentiated from the lamina, with broad thin sheathing base; laminae 1-7

216 FIELDIANA: BOTANY

cm long, 1.5-8 (14) mm broad, linear to oblanceolate, obtuse or rounded at the apex,

margin entire and gradually decurrent on the petiole, smooth and glabrous, succulent and

bright green or reddish in life but drying dark and coriaceous, venation usually obscure.

Flowers solitary in leaf axils, glabrous, pedicels 3-10 (15) mm long, ca. 1 mm thick, flowers

7-12 mm long, perianth lobes ca. 5 mm long and unequal, with a thickened central area

and thin lateral margins, greenish on the outside and pink to reddish on the inside, with

a short (1-1.5 mm) stiff appendage on the back (abaxial) distally, floral cup ca. Va the

length of the perianth lobes; stamens numerous and free, filaments 1.5-3.5 mm long,

anthers 0.5-0.8 mm long; ovary 3-4 mm long, 3-locular (rarely 2 or 4), styles 3 or 4. Fruit

5-11 mm long and 5-6 mm thick, opening by a circumscissile slit, the perianth lobes

persistent; seeds 8-30, black and smooth, 0.5-1.5 mm broad, reniform-cochleate.

Sprawling succulent plants usually found on or near sea beaches, occasionally

found in salt flats and on open sandy or rocky sites from sea level to about 50

m elevation. Probably flowering throughout the year in Central America, but

collected primarily from December through March and June through August.

This species is pantropical, but appears to be uncommon in southern Central

America.

Sesuvium portulacastrum is recognized by its succulent parts, narrow opposite

leaves sheathing the stem at their base and united across the stem, solitary

bright pink flowers with many stamens, and capsules opening by a circular slit.

These plants are rarely found far from the seashore. The species has served as a

vegetable, either eaten raw or cooked as greens. Superficially, this species re-

sembles Philoxerus vermicularis of the Amaranthaceae, with small flowers in

whitish spikes.

TRIANTHEMA Linnaeus

Herbs or subshrubs, annual or short-lived, prostrate to erect, stems succulent, bisexual,

glabrous or puberulent; stipule-like developments sometimes present on the united inter-

petiolar sheathing leaf bases. Leaves opposite or subopposite, simple, the opposing leaves

at a node often unequal, petioles expanded at the base and clasping or sheathing the stem;

laminae linear to orbicular, entire, succulent or semisucculent. Flowers solitary or fascicu-

late in leaf axils, sessile or pedicellate, sometimes united to the leaf base, radially sym-

metrical, bisexual, perianth united at the base to form a perianth tube (floral cup or calyx

tube), 5-lobed, lobes often with a distal abaxial (dorsal) appendage; stamens 5 or 10 to

many, borne on the edge of the perianth tube, alternate with the perianth lobes when of

the same number, filaments free; ovary superior, 1-locular, ovules 2 to many on a parietal

placenta, apex of the ovary truncate or depressed around the style, style 1, linear. Fruit a

circumscissile capsule, seeds cochleate-reniform, with a rough or ridged surface.

A genus of nine species: one pantropical, one in Argentina, and the others in

Africa, Asia, and Australia.

Trianthema portulacastrum L., Sp. PI. 223. 1753. Figure 39.

Herbs, annual, prostrate to decumbent or erect, to 50 cm tall or 1 m long, bisexual,

succulent, often reddish in color, often with alternate branching from the axil of the

smaller leaf of a node, leafy internodes 0.5-10 cm long, 0.5-3.5 mm thick (dry), glabrous

or with a single longitudinal ridge bearing slender crooked hairs on younger parts; sheath-

ing leaf bases of opposing leaves united across the node and with a triangular stipule-like

appendage to 3 mm long borne on the interpetiolar portion of the sheath. Leaves opposite

or subopposite, the leaves of each node usually very different in size (the smaller often half

the size of the larger), petioles 2-20 (30) mm long, conspicuously expanded at the base to

form a thin sheathing base united with that of the opposing leaf and encircling the stem;

laminae 1-4 (7) cm long, 0.5-2 (6) cm broad, broadly elliptic to spatulate in smaller leaves,

to broadly obovate or suborbicular, larger leaves abruptly rounded at the apex and often

BURGER: FLORA COSTARICENSIS 217

apiculate or emarginate, cuneate to obtuse at the base, margin entire, drying stiffly char-

taceous to subcoriaceous, smooth and glabrous, with 2-4 pairs of major secondary veins.

Flowers solitary and sessile in the leaf axils, bisexual, ca. 5-8 mm long, perianth tube

campanulate, perianth lobes ca. 1.5 mm long and 1 mm broad; stamens 10, arising from

the apex of the perianth tube (floral cup), filaments 1.5 mm long, anthers 0.5 mm long;

ovary 1 mm long, irregularly lobed at the apex, style 1. Fruit a capsule surrounded by the

persistent perianth tube and hidden within the sheathing leaf bases, ca. 5 mm long, 4 mm

thick, crested at the apex, with circumscissile dehiscence at the middle; seeds 2-12, 1.5-2

mm broad, reniform-cochleate, reddish brown to black, with a rough surface.

Plants of open disturbed sites, from sandy sea beaches to cultivated ground

and roadsides; ranging in altitude from sea level to 1,200 m, but rarely encoun-

tered above 500 m elevation in our area. Flowering primarily in the wet season,

from June to December, but flowering collections have been made throughout

the year in Central America. The species is pantropical, but has not been col-

lected along the Caribbean coast in southern Central America.

Trianthema portulacastrum is recognized by the succulent stems, unequal op-

posite leaves with united sheathing leaf bases, and small pink flowers barely

exserted beyond the sheathing leaf bases. This species has been called Verdolaga

in Guatemala, according to Standley. The flowers are often difficult to see in

pressed collections, since they are hidden by the sheathing leaf bases and are

adnate, in part, to the petioles.

PORTULACACEAE

Herbs or subshrubs (rarely shrubs), bisexual, usually with succulent stems, glabrous or

puberulent at the nodes; stipules present or absent, often represented by scarious, fim-

briate or tufted developments at the node. Leaves alternate, opposite, or condensed and

basal or pseudoverticellate, simple, sessile or petiolate, the laminae usually thick-

succulent, glabrous and entire, often small. Inflorescence terminal or axillary, paniculate,

racemose, cymose or the flowers solitary; flowers bisexual (rarely unisexual), radially

symmetrical, often small but brightly colored, perianth generally interpreted to be of 2

dissimilar whorls, sepals 2 (rarely 4-8), imbricate, free or united at the base, scarious or

herbaceous, persistent or caducous, usually greenish (sepals sometimes interpreted to be

floral bracts), petals 4 or 5 (6-18), free or united at the base, quickly wilting or rarely

persisting, white to yellow or red; stamens 4 or 5 to many, often as many as the petals and

opposite them, usually borne on a short floral cup, filaments free or united into fascicles,

anthers 2-thecous, dehiscing longitudinally; pistil 1, ovary superior (half-inferior to infe-

rior in Portulaca), at first with locules and axile placentation but becoming unilocular at

maturity and with basal placentation, ovules (1) 2 to many, campylotropous, styles 1-9,

united near the base or free. Fruit usually a capsule, opening by loculicidal or circum-

scissile dehiscence (rarely a nut); seeds (1) 2 to many, slightly flattened, reniform to

cochleate, embryo curved or folded.

A family of about 20 genera and 500 species, best represented in open sunny

habitats in the seasonally dry tropics and subtropics. The family is closely related

to Aizoaceae (especially Sesuvium) and exhibits similarities of ovary structure to

that of the Caryophyllaceae. Our species are characterized by their generally

small stature, fleshy stems and leaves, flowers subtended by two greenish

sepals, four or more colorful petals, fruit that begin to open by a circumscissile

slit, and the dark cochleate seeds borne on a basal placenta.

la Plants with the leaves all condensed at the apex of a single thick rootstock, leafy

internodes not visible, leaves linear (1.5-3 mm broad and up to 7 cm long); plants

found only on the highest paramo, above 3,200 m elevation Lewisia

Ib Plants with conspicuous aerial stems and internodes, leaves rarely linear; not known

to grow wild above 2,000 m elevation 2a

218 FIELDIANA: BOTANY

2a Ovary partly or wholly inferior, fruit opening by a circumscissile slit at the middle or

near the top; flowers sessile or subsessile and subtended by a cluster (involucre) of

leaves; stems usually with tufts of hairs or cilia at the nodes (stipules) .... Portulaca

2b Ovary superior, fruit opening irregularly; flowers borne on slender pedicels in an

open panicle, not subtended by leaves; stems and leaves lacking stipules, glabrous

Talinum

LEWISIA Pursh

Herbs, perennial, aerial stem very short from a thick fleshy rootstock, rootstock simple

or branched. Leaves mostly in a condensed basal cluster at the apex of the rootstock,

simple and entire, somewhat succulent. Inflorescences racemose, paniculate or solitary

and axillary, often subtended by leaflike bracts; flowers bisexual, radially symmetrical,

sepals 2-8, persistent in fruit, petals 4-18, white to pink or red; stamens 5 to many; pistil

1, ovary superior, placenta basal and central, styles 3-8 and fused near the base. Fruit a

thin-walled capsule, dehiscence circumscissile near the base and then splitting upward;

seeds 6 to many, ovate to cochleate, smooth and lustrous.

A genus of 20 species of the southwestern mountains of the United States; one

species extends southward on the tops of the highest mountains through Mexico

to Guatemala and has recently been reported from the Chirripo paramo in Costa

Rica. Species of this genus are often planted as ornamentals in rock gardens.

Lewisia megarhiza (Hemsl.) MacBryde, J. Scott. Rock Card. Club 13:295. 1973.

Calandrinia megarhiza Hemsley, Diagn. PL Nov. 23. 1879. Oreobroma mexicanum

Rydberg, N. Amer. Fl. 21:326. 1932. O. megarhiza (Hemsl.) Standl. & Steyerm.,

Publ. Field Mus. Nat. Hist., Bot. Ser. 23:49. 1944.

Herbs, perennial, the short aerial stem and congested leafy internodes from the top of

a single thick fleshy taproot to 10 cm long and 5-18 mm thick. Leaves all basal and

congested at the apex of the taproot, simple and sessile; laminae 2-7 cm long, 1.2-3 mm

broad, linear and entire, expanded near the base with thin sheathing margins, fleshy,

glabrous. Flowers solitary on axillary pedicels 5-20 mm long, sepals 2, 6-7 mm long, petals

4-6, ca. 1 cm long, white. Fruit an ellipsoid capsule ca. 7 mm long and 4 mm thick, with

very thin translucent walls; seeds numerous, 1.5 mm broad, cochleate, black, smooth and

lustrous.

This species has been collected by Arthur Weston on Costa Rica's highest

mountain, Chirripo Grande. All the specimens seen from Mexico and Gua-

temala were collected above 3,500 m elevation. The large taproot for so small a

plant, the succulent linear leaves, the presence of only two sepals, and the

distinctive fruit and seeds easily distinguish this species.

PORTULACA Linnaeus

By Donna Ford and William Burger

REFERENCE: C. Diego Legrand, Desmembracion del genero Portulaca, I and II,

Comun. Bot. Mus. Hist. Nat. Montevideo vol. 3, no. 31:1-15, 1953 and no.

34:1-17, 1958.

Herbs, annual or perennial, often prostrate, stems succulent glabrous or puberulent;

stipules often reduced to a ridge with conspicuous hairs. Leaves alternate, subopposite or

crowded and pseudoverticellate, simple and entire, sessile or petiolate, usually succulent

and glabrous. Inflorescence of 1 to several flowers terminal on main and lateral stems, often

subtended by closely crowded leaves forming an involucre; flowers bisexual and radially

symmetrical, sessile or subsessile, perianth of 2 whorls, sepals 2, unequal and united near

the base, petals 4-6, white, yellow or reddish, united only near the base, quickly with-

BURGER: FLORA COSTARICENSIS 219

ering; stamens (4-6) 7 to many, borne on the base of the united corolla, filaments free; pistil

solitary, ovary half-inferior to inferior, unilocular, ovules numerous on a basal placenta,

styles 3-9 and united at the base. Fruit a circumscissile capsule, the top coming off as a

single unit (calyptra), thin-walled; seeds usually numerous, cochleate to reniform, smooth

to minutely tuberculate.

A genus of 150-200 species, distributed widely in tropical and subtropical

regions. The small stature, succulent stems, brightly colored flowers subtended

by an "involucre" of congested leaves, and calyptrate thin-walled fruit with

cochleate seeds are distinctive. The genus is well represented in Mexico, and a

number of species range sourthward to Guatemala and Honduras.

la Leaves usually more than 4 mm wide, laminae flat; axillary hairs absent or inconspic-

uous (to 1 mm long) 2a

Ib Leaves usually less than 4 mm wide, laminae often terete; axillary hairs present and

conspicuous, 1-18 mm long 3a

2a Ovary superior to half-inferior; capsule not winged, circumscissile at the middle

or below, operculum hemispherical; seeds black, brown or blue-waxy; flowers

yellow P. oleracea

2b Ovary half-inferior to inferior; capsule with a circular wing, circumscissile at the

middle or above, operculum flat or slightly rounded; seeds gray at maturity;

flowers yellow to red P. umbraticola

3a Petals less than 1 cm long, pink to purple; stamens fewer than 40; small plants of open

weedy sites P. pilosa

3b Petals more than 1 cm long and the flowers more than 2 cm broad, variously colored;

stamens more than 40; cultivated ornamentals and not naturalized in Costa Rica;

P. grandiflora Hooker.

Portulaca oleraceaL., Sp. PL 445. 1753. P. marginataH.B.K., Nov. Gen. & Sp.

6:58. 1823. Figure 39.

Herbs, prostrate or ascending, often with many branches radiating from a single base,

stems succulent and often becoming reddish, leafy internodes 0-5 cm long, 0.5-3 mm

thick (dry), glabrous except at the nodes; stipules present in the form of a minute ridge

with slender whitish hairs to 1 mm long (just below the leaf base). Leaves alternate to

subopposite and opposite, often crowded together at the ends of branches; petioles 0.5-4

(8) mm long, expanded at the base with thin lateral margin; laminae 0.4-4 cm long, 0.3-2

cm broad, spatulate to obovate or obtriangular, bluntly rounded at the apex, tapering

gradually to the cuneate base, margins entire and decurrent on the petiole, smooth and

glabrous, succulent but drying dark and stiffly chartaceous, venation obscure. Flowers

sessile at the ends of branches, solitary or clustered together, subtended by a cluster of

leaves, hairs at the base of the flower inconspicuous, with a short floral cup (perianth

tube), sepals 2, broadly ovate, 3-4.5 mm long, petals 4-6, 3—4.5 mm long, bright yellow;

stamens 6-15, filaments ca. 1.6 mm long; style with 4-6 branches. Fruit ovoid, 4-8 mm

long, 3-4 mm thick, circumscissile below the middle, distal cap 2-4 mm long; seeds 0.6-1

mm broad, cochleate, black, with granular or minutely tuberculate surface.

Plants of open sunny sites, often found along roadsides, on sand and gravel

bars along streams, and in open weedy fields from sea level to 1,500 (2,000) m

elevation on both the Caribbean and Pacific sides of Central America. The

species appears to flower throughout the year, but is most commonly encoun-

tered from October to March. This species has become established all over the

world, in both tropics and subtropics.

Portulaca oleracea is recognized by its small stature, numerous glabrous stems,

succulent little leaves often crowded at the ends of stems and forming an in-

volucre beneath the sessile yellow flowers, and the circumscissile fruit with basal

placentation. The distal flowers on main stems and lateral stems with terminal

flower clusters often form cymelike arrangements. These plants are called

220 FIELDIANA: BOTANY

Verdolaga in Central America ("Purselane" in English) and are often cooked as a

vegetable.

Portulaca pilosa L., Sp. PI. 445. 1753. Figure 41.

Herbs, prostrate or erect, stems succulent, rarely exceeding 20 cm in length, leafy

internodes 0-2 cm long; intrapetiolar stipule present, with tufts of whitish haris 3-10 mm

long from a band of tissue ca. 1.5 mm broad just above the leaf base. Leaves alternate or

crowded together, sessile or with a short (1 mm) petiole; laminae 0.5-2 cm long, linear to

narrowly lanceolate, acute at the apex, gradually narrowed at the base and decurrent on

the petiole, margins entire, the laminae succulent (often terete) in life and glabrous, drying

dark, venation obscure. Flowers sessile and crowded at the ends of stems, 2.5-3 mm long,

subtended by a cluster of leaves forming an involucre, sepals 2-3 mm long, petals 3-5.5

mm long, reddish or purple; stamens 15 to many, filaments red. Fruit a circumscissile

capsule, 3-4 mm in diameter; seeds 0.5-0.7 mm broad, slightly flattened, cochleate in

outline, minutely tuberculate, black.

Plants of open sunny sites and often found on rocky sites or sand along

streams from near sea level to 1,200 m elevation along the Pacific slope of Central

America and in the Caribbean lowlands; flowering primarily from June to

February. The species ranges from the southeastern United States, the West

Indies, and Mexico through Central America to South America.

Portulaca pilosa is characterized by its small succulent stems with unusual

h'mbriate intrapetiolar stipules that give a tufted white-hairy appearance to the

stems. The distal rose red flowers subtended by similar tufted hairs and by a

cluster of leaves are also distinctive. The species has only been collected in the

seasonally very dry areas of the northern Pacific slope in Costa Rica, during the

wet season.

Portulaca umbraticola H.B.K., Nov. Gen. & Sp. 6:72. 1823. P. lanceolata En-

gelm., Boston J. Nat. Hist. 6:154. 1850, non Haw. 1803. P. coronata Small, Bull.

Torrey Bot. Club 23:126. 1896. P. plano-operculata Kuntze, Rev. Gen. 3:16. 1898.

P. denudataPoellin., Repert. Spec. Nov. Regni Veg. 33:159. 1933.

Herbs, 10-50 cm tall, annual, stems prostrate to erect and simple or with open branch-

ing, stems glabrous and green to magenta, roots fibrous; stipules absent or represented by

short (1 mm) whitish hairs at the axils. Leaves alternate along the stems and in whorls of

3-8 beneath the flowers, subsessile or with short (1-2 mm) petioles; laminae 1-3 (5) cm

long, 0.4-1.1 (1.6) cm broad, lower laminae spatulate to obovate, the upper oblanceolate

to oblong, usually rounded at the apex, tapering gradually to the base, glabrous, succulent,

venation obscure. Flowers in terminal clusters, bracteoles 1.5-2 mm long, lanceolate to

deltoid; sepals 2, 3-5 (8) mm long, ovate, petals to 1 cm long, yellow to orange or red,

spatulate or obovate; stamens 7-27, anthers narrow; ovary with an adnate corona, style

5-8 mm long, stigmatic branches 3-8. Fruit a capsule subtended by the persisting leaves

and calyx, 3-5 mm long, usually turbinate, sessile or borne on a pedicel to 4 mm long,

circumscissile at the middle or above, the operculum rounded, 3-4 mm broad and 1-2 mm

high, the coronal wing 0.5-1 mm wide, arising just below the line of dehiscence; seeds

0.7-1 mm broad, gray, cochleate and tuberculate.

Plants of open sunny sites in seasonally dry areas, ranging from near sea level

to 1,200 m in our area, but rarely collected below 600 m in central Central

America. Flowering material has been collected from June to September in Cen-

tral America, along the Pacific slope, and in the Central Highlands. The species

ranges from the southern United States (Arizona to Georgia), Mexico, and the

West Indies through Central America to South America.

Portulaca umbraticola is recognized by its succulent and glabrous stems and

BURGER: FLORA COSTARICENSIS 221

leaves (hairs only at the leaf axils), the terminal flowers subtended by a whorl of

leaves, fruit opening by the circumscissile dehiscence of a slightly domed

(curved) lid or operculum, and the restriction to seasonally very dry vegetation.

This species has only recently been collected in Costa Rica at the Santa Rosa

National Park in Guanacaste province (Liesner & Lockwood 2687). While common

in Honduras between 600 and 1,100 m elevation, we have not seen specimens

from Guatemala. In Honduras the species is called Verdolaga.

TALINUM Adanson

Herbs or subshrubs, annual or perennial, bisexual, stems often succulent, usually gla-

brous; stipules absent. Leaves alternate or subopposite, simple and entire, usually quite

succulent. Inflorescences often cymose, the cymes arranged in compound panicles or

simple and few flowered, flowers rarely solitary and axillary, usually pedicellate; flowers

bisexual, radially symmetrical, sepals 2, opposite and free, usually early deciduous, petals

5 or more, free or connate at the base, deciduous; stamens 5 to many, sometimes united

into antipetalous fascicles, filaments slender; pistil 1, ovary superior, 1- or 3-locular near

the base, placentation basal or free central, styles 3, united near the base. Fruit a capsule,

1- or 3-locular near the base, opening from the top into 3 valves or irregularly; seeds

somewhat flattened, cochleate to reniform in outline.

A genus of probably fewer than 50 species, two of which are found in Cen-

tral America. The genus ranges from temperate and tropical America to Africa

and Asia.

la Plants with succulent stems that do not persist through the dry season; larger leaves

3-6 cm broad; inflorescence a much-branched panicle with many distant flowers;

sepals 3-4 mm long; uncommon plants of the seasonally dry Pacific slope from Mexico

to Costa Rica T. paniculatum

Ib Plants with woody stems that persist through the dry season and are perennial; larger

leaves rarely exceeding 2.5 cm in breadth; inflorescence a few-branched cyme or

raceme; sepals 5-6 mm long; ranging from Mexico to central Nicaragua and not

included in the descriptions: T. triangulare (Jaccl-) Willd. (See note below.)

Talinum paniculatum Qacq.) Gaertner, Fruct. & Sem. 2:219, pi. 128, f. 13.

1791. Portulaca paniculata Jacquin, Enum. PI. Carib. 22. 1760. P. patensL., Mant.

PI. 242. 1771. T. patens (L.) Willd., Sp. PL 2:863. 1800. Claytonia paniculata (Jacq.)

O. Ktze., Rev. Gen. 1:57. 1891. Figure 40.

Herbs, erect to 1 m tall, stems succulent and simple or with a few lateral branches, from

a tuberous base, glabrous, leafy internodes 0.5-10 cm long, 1-6 mm thick; stipules absent.

Leaves alternate, glabrous, petioles 1-15 mm long and not clearly differentiated from the

lamina, with lateral wings continuous with the lamina margins; laminae 2-13 cm long, 1-6

cm broad, elliptic to obovate, abruptly obtuse or rounded at the apex, tapering gradually

to the cuneate base and decurrent on the petiole, margins entire, succulent in life but

drying brittle (chartaceous to subcoriaceous) and dark, the 2-4 pairs of major secondary

veins often obscure. Inflorescences 10-60 cm long, 6-20 cm broad, terminal, paniculate

with a central stem and slender alternate branches, flowers in open cymes on very slender

(0.2 mm, dry) pedicels 1-2 cm long; flowers 3-4 mm long, perianth of 2 whorls and not

persisting in fruit, sepals ca. 2 mm long, petals ca. 3 mm long, yellow or pink; stamens 15

or more, filaments 1.2-2 mm long, anthers 0.3-0.5 mm long; ovary globose, 1 mm long.

Fruit a lustrous yellowish globose capsule 3.5-5 mm in diameter, breaking open irregu-

larly; seeds slightly flattened, cochleate in outline, 1-1.5 mm broad, surface black and

lustrous, minutely reticulate, strophiolate.

Plants of open weedy sites, riverbanks, and thickets from (0) 100 to 1,200 m

elevation along the seasonally very dry Pacific slope of Central America; grow-

ing and flowering in the wet season, from June to December. The species ranges

222 FIELDIANA: BOTANY

from the southern United States, Mexico, and the West Indies through Central

America and Panama to South America.

Talinum paniculatum is recognized by its succulent alternate leaves on few-

branched stems, open paniculate inflorescences with slender stems, small flow-

ers with only two sepals, and capsules opening irregularly to disperse small

black snail-shaped seeds. These plants are rarely collected in Costa Rica and are

probably confined to the northern half of the Pacific slope.

Note added in proof: Talinum triangulate Qacq.) Willd. has recently been collected

by Robert Wilbur near the railroad yards at Puerto Limon, where it appears to

be a recent arrival.

BASELLACEAE

Twining herbs or climbing subshrubs or vines, annual or perennial, bisexual, glabrous;

without stipules. Leaves alternate and simple, petiolate, often cordate, usually slightly

succulent, entire and glabrous. Inflorescences axillary or terminal, spikes, racemes or

panicles, glabrous, small slender bracts present at the base of the pedicels and with 2-4

sepaloid bracteoles; flowers small, bisexual or unisexual, perianth of a single whorl but

often appearing to have 2 whorls because of the sepaloid bracteoles adnate to the base of

the flowers, perianth petaloid and 5-parted, forming a tube or almost free to the base;

stamens 5, opposite the perianth parts and borne at their base, filaments free, anthers

versatile, dehiscing variously; pistil 1 with superior ovary, 1-locular with basal placen-

tation, ovule solitary, short-stalked and campylotropous, style 1-3, stigmas separate or

3-lobed. Fruit an indehiscent utricle, enclosed by the often fleshy perianth or winged

bracteoles (winged sepals of some authors); seeds solitary, small, spherical or flattened.

A small tropical family of five genera. Anredera vesicaria (Lam.) Gaertn., with

fruit bearing wings 4 to 5 mm long, is found in northern Central America.

Ullucus tuberosa Lozano, with potato-like tubers, is an important food plant in the

Andean region of South America. Basella alba L. of the Old World tropics is

cultivated for its fleshy leaves which serve as an excellent spinach-like vegetable.

None of the aforementioned species has been reported in Costa Rica, but Basella

albais cultivated in central and eastern Panama (see treatment of the family in the

Flora of Panama, Ann. Missouri Bot. Card. 66:109-115, 1979). The family is

closely related to the Portulacaceae.

BOUSSINGAULTIA Humboldt, Bonpland, and Kunth

Vines or herbaceous twiners with slender distal stems, bisexual, glabrous, often succu-

lent; stipules absent. Leaves alternate and simple, petiolate, slightly succulent, often

becoming cordate in larger leaves. Inflorescences axillary or terminal, racemes or panicles,

flowers subtended by slender bracts and with 2 sepal-like bracteoles at the apex of the

pedicel; flowers bisexual and radially symmetrical, perianth of 5 petaloid parts whitish but

often turning dark when dry, the sepaloid bracteoles united to the base of the corolla-like

perianth tube; stamens borne on the base of the corolla tube, inflexed in bud, filaments

slender, anthers versatile; ovary ovoid, style 1 with 3 stigmas or the style with 3-lobed

stigma. Fruit a utricle enclosed within the persisting perianth; seed solitary erect.

A genus of about 10 species in tropical America. Three species are found in

northern Central America, but only one of these species has been recorded from

Costa Rica.

Boussingaultia ramosa (Moq.) Hemsley, Biol. Centr. Amer. Bot. 3:27. 1882.

Tandonia ramosa Moq., in DC., Prodr. 13, pt. 2:227. 1849. Dioscorea calyculata

Donnell Smith, Bot. Gaz. 20:295. 1895. Figure 40.

PHILOXERUS

vermicularis

TRIANTHEMA

portulacastrum

SESUVIUM

portulacastrum

FIG. 39. Cnryophyllales: succulent species with opposite leaves usually clasping the stem

at their base. Aizoaceae: species of Sesuvium and Trianthema. Amaranthaceae: Philoxerus

vermicularis. Portulacaceae: Portulaca oleracea.

223

BOUSSINGAULTIA

ramosa

TALINUM

paniculatum

FIG. 40. Caryophyllales: a climber and an erect herb, both with semisucculent leaves.

Basellaceae: a species of Boussingaultia. Portulacaceae: a species of Talinum.

224

ARENARIA

lanuginosa

ssp. lanuginosa

SAGINA

procumbens

SCLERANTHUS

annuus

MOLLUGO

verticellata

FIG. 41. Caryophyllales: herbs with narrow opposite leaves. Aizoaceae: Mollugo verti-

cellata. Portulacaceae: Portulacapilosa. Caryophyllaceae: species of Arenaria, Sagina, Scleran-

thus, and Spergula.

225

ARENARIA

lanuginosa

ssp. guatemalensis ,

STELLARIA

ovata

CERASTIUM

guatemalensis

5 mm

GLINUS

radiata

FIG. 42. Caryophyllales: species with broader opposite or whorled leaves. Aizoaceae:

Glinus radiatus. Caryophyllaceae: species of Arenaria, Cerastium, Drymaria, and Stellaria.

226

BURGER: FLORA COSTARICENSIS 227

Herbaceous or woody climbers, annual or perennial, distal stems often pendulous,

glabrous, leafy internodes 2-6 cm long, 1-4 mm thick; stipules absent. Leaves alternate in

a spiral, distant, glabrous, petioles 6-25 mm long, sulcate above with 2 adaxial ridges

continuous with the lamina margins; laminae 2-6 cm broad, broadly ovate, acute to obtuse

at the apex and with a mucronate tip ca. 3 mm long, obtuse to truncate or cordate (in larger

leaves) at the base, margins entire and decurrent on the petiole, somewhat succulent in life

but drying thin-chartaceous or membranaceous, venation pinnate or subpalmate with 2 or

3 pairs of major secondary veins, smooth and glabrous. Inflorescences slender open ra-

cemes borne on leafless paniculiform axillary or terminal branches, 6-35 cm long, rachis

0.2-0.6 mm thick (dry) and glabrous, with slender linear-lanceolate bracts subtending the

pedicels ca. 2 mm long, pedicels ca. 1.5 mm long; flowers ca. 2 mm long, subtended by a

pair of sepaloid bracteoles ca. 0.6 mm long, perianth petaloid, 2 mm long, broadly imbri-

cate, whitish but becoming dark when dried, united and narrowed above the calyx-

like bracteoles, ovary ca. 1 mm long, ovoid. Fruit apparently quickly caducous, usually

enclosed by the dark perianth and pale-colored subtending bracteoles, with 2 lateral

winglike developments at the base of the perianth ca. 1 mm long, fruit lustrous subglobose

and with a small disk at the apex, ca. 0.7 mm long; seed difficult to extricate from the

tightly enclosing fruit wall.

Plants of open thickets and climbing over others, from 700 to 2,000 m elevation

along the Pacific slope of Central America; collections with flowers and fruit

have been made from October to February. The species ranges from Mexico to

Costa Rica.

Boussingaultia ramosa is characterized by its glabrous slender distal climbing

stems with alternate slightly succulent leaves, the long pendulous racemes with

little white flowers, the two sepaloid bracteoles at the apex of the pedicel, and

the minute fruit with single seed. This species resembles some species of Di-

oscorea and Polygonaceae, but the five-parted perianth and estipulate leaves

serve to place these slender climbers in their appropriate family.

CARYOPHYLLACEAE

By Richard A. Baker and William C. Burger,

with Cerastium by David Good

Herbs (rarely subshrubs or shrubs), annuals or perennials, plants bisexual (rarely uni-

sexual), glabrous to densely puberulent, nodes often slightly thickened; stipules present

or absent and then the leaf often with a clasping or sheathing base, interpetiolar ridges or

sheaths sometimes present. Leaves simple and undivided, opposite or pseudoverricellate

(rarely alternate), laminae usually entire. Inflorescences often cymose, loosely dichasial to

compact and capitate, many to few flowered or the flowers solitary on distal stems, floral

bracts or reduced leaves often subtending the pedicels, rarely with 1-3 pairs of bracts

immediately subtending the calyx (called an epicalyx) as in Dianthus. Flowers bisexual

(rarely unisexual), radially symmetrical, calyx of 5 or 4 free sepals or united to form a calyx

tube with 5 (4) lobes or teeth, hypogynous or borne on a cupulate hypanthium, persisting

in fruit and becoming dry and stiff; petals 5 or 4 (0), free, entire to deeply bilobed or serrate

at the apex, white to pink or purple and red (the petals usually divided into a narrow

petiole-like claw at the base and a distal expanded blade in genera with united sepals, the

clawed petals often with small ligule-like appendages above the junction of claw and

blade); stamens usually 10 in 2 whorls of 5, often fewer by reduction, often united at the

base but the filaments free for most of their length, anthers 2-celled, dehiscing longi-

tudinally; pistil solitary, often narrowed at the base or borne on a short stalk, ovary 1- to

5-Iocular, often with septa at the base and a single locule in the upper half, placentation

axile, central, free-central or basal, ovules few to many (rarely solitary), styles 2-5, free or

united at the base. Fruit usually a thin-walled capsule (1-seeded indehiscent utricles or

berry-like in a few genera), often borne on a short stalk, usually breaking into as many

septa as there are styles, capsules of flowers with united tubular calyx usually opening only

at the top with small teeth; seeds usually reniform to cochleate or circular, often slightly

flattened on 2 sides, smooth to tuberculate, with or without a small aril-like appendage

228 FIELDIANA: BOTANY

(strophiolate or estrophiolate); seeds usually with a curved embryo enclosing perisperm

and little or no endosperm.

A family of about 75 genera which is most numerous in the cooler regions of

the world and very poorly represented in the lowland tropics. A number of

taxonomically difficult genera include species that have become cosmopolitan

weeds; the names and status of these species will probably continue to give us

difficulty for years to come. The family includes a number of ornamentals that

are important both in gardens and in the cut-flower trade. These herbaceous

plants can usually be recognized by their opposite simple and entire leaves, the

separate petals often cleft or fimbriate at the apex, the thin- walled capsules, and

the seeds usually reniform (kidney-shaped) to cochlea te (snail-shaped).

la Flowers with the sepals free or united only at the base (calyx-tube, calyx-cup or

urceolate hypanthium absent); native and naturalized weedy plants, not grown for

ornamental use 2a

Ib Flowers with the calyx united to form a cup or a tube, or the sepals borne on the apex

of an urceolate hypanthium; introduced weeds or garden plants grown for their

ornamental flowers and inflorescences 8a

2a Stipules present at the leaf base (sometimes falling off early), leaf bases not

clasping or sheathing the stem, interpetiolar lines often present 3a

2b Stipules absent, leaf bases often clasping or sheathing the stem, an interpetiolar

line or sheath often present 5a

3a Leaves opposite and broad; styles united near the base; seeds cochleate and

tuberculate; commonly encountered native plants Drymaria

3b Leaves opposite or whorled or appearing to be whorled by condensation of

short axillary shoots, leaves linear to narrowly obovate; rarely collected

introduced weeds of higher elevations 4a

4a Leaves linear and appearing to be whorled; styles free to the apex of the

ovary; seeds subglobose and with a whitish circumferential margin or wing

Spergula

4b Leaves narrowly obovate or oblong-obovate, opposite or whorled; styles

united for most of their length; seeds semicircular to sublunate and without

a margin or wing Polycarpon

5a Capsule cylindrical and usually somewhat curved, opening by 10 short teeth;

plants often conspicuously puberulent Cerastium

5b Capsule ovoid and straight, opening by 2-8 teeth 6a

6a Seeds reniform-globose, seed coat lustrous and smooth, maroon to black in

color; petals entire; laminae often with a delicate submarginal vein along distal

margins Arenaria

6b Seeds usually cochleate with muricate to tuberculate surfaces, brownish and

dull; petals entire to deeply 2-lobed 7a

7a Petals entire; styles 4 or 5; leaves linear and less than 8 mm long, a submarginal

vein absent Sagina

7b Petals 2-lobed; styles usually 3; leaves usually ovate to cordate and the larger

more than 8 mm long (S. nubigena has small lanceolate leaves but is very rare),

a submarginal vein present or absent Stellaria

8a Sepals narrow and borne at the distal rim of a small (1.5 mm) urceolate hypanthium,

petals absent; leaves linear and rarely more than 12 mm long, with an interpetiolar

sheath at the base; fruit with 1 seed; weeds (rare in Costa Rica) Scleranthus

8b Sepals united for much of their length to form a tube or united halfway to form an

open cup; petals present; fruit with numerous seeds; planted for ornament or an

escaped weed with viscid glandular hairs in Silene gallica 9a

9a Flowers less than 1 cm long, calyx united for half its length; styles 2 or 3; capsules

4-valved, 4-5 mm long; inflorescences open and diffuse, with many flowers on long

slender pedicels; grown for ornament Gypsophila

9b Flowers usually over 15 mm long, calyx united to form a tube with small apical teeth,

calyx tube usually enclosing most of the fruit; capsules opening by small apical teeth,

more than 5 mm long; inflorescences usually few-flowered lOa

BURGER: FLORA COSTARICENSIS 229

lOa Styles 5 or 4; calyx 15-20 mm long; garden plants covered with grayish white

tomentum Lychnis

lOb Styles 2 or 3; plants never covered with dense grayish white puberulence lla

lla Styles 3; calyx 6-16 mm long, with 10 or more thick distinct veins; capsule with 6

short apical teeth; glabrous and with showy flowers or a weed with glandular pubes-

cence distally Silene

lib Styles 2; calyx 15-24 mm long, with many veins, the veins not thickened; capsule

usually with 4 apical teeth; glabrous garden plants Saponaria

ARENARIA Linnaeus

REFERENCE: J. McNeill, The delimitation of Arenaria and related genera

in North America, with 11 new combinations in Minuartia. Rhodora 82:495-

502. 1980.

Annual, biennial or perennial herbs, sometimes with a woody rootstock, stems diffusely

branched, procumbent, repent or ascending and clambering, occasionally forming dense

cushion-like mats, glabrous or puberulent; estipulate. Leaves opposite, sessile or very

short petiolate, linear to narrowly elliptic or ovate, entire, usually clasping the stem at their

base. Inflorescences dichasial, axillary or terminal, or the flowers solitary at the nodes (as

in ours); flowers bisexual, radially symmetrical, small, sepals 5, free or only slightly united

at the base, petals 5 (absent in A. quirosiiand some northern subspecies of A. lanuginosa),

white (in ours), free, apically entire to slightly emarginate or dentate; stamens 10, rarely

5, usually borne on a slightly elevated perigynous disk, anthers versatile, dehiscing

longitudinally, 2-thecous; ovary superior, unilocular, placentation basal with many

ovules, styles usually 3 or 5 (occasionally 2 or 4), free. Fruit a capsule dehiscing by as many

valves as there are styles, valves 1-lobed or bilobed at the apex, thin-walled, unopened

fruit ovoid to globose or oblong; seeds often numerous, subglobose or slightly compressed,

reniform to subcircular in outline, the surface smooth and lustrous (in ours) or rough and

tuberculate, estrophiolate.

A genus of perhaps 200 species, primarily of seasonally cold habitats in the

Northern Hemisphere. Two species occur in Costa Rica, one apparently en-

demic. The Costa Rican species of Arenaria can be distinguished from our other

Caryophyllaceae by the combination of their dark maroon or black, smooth and

shiny, reniform-globose seeds, the lack of stipules, the usually narrow leaves,

the solitary axillary flowers borne on conspicuous slender pedicels, and the

entire petals (when petals are present). The members of this genus have only

rarely been collected below 1,000 m elevation in Costa Rica. One of our species

is polymorphic and is here treated as a single species; see the discussion under

A. lanuginosa.

la Laminae less than 5 mm long, broadly ovate to obovate; flowers lacking petals; prob-

ably restricted to areas above 3,000 m elevation, rarely collected A. quirosii

Ib Laminae usually over 10 mm long (linear to lanceolate or narrowly elliptic-ovate when

less than 10 mm long); flowers with petals but the petals deciduous; a polymorphic

array of plants ranging from 700 to 3,400 m elevation A. lanuginosa

Arenaria lanuginosa (Mich.) Rohrbach, in Mart., Fl. Brasil. 14, pt. 2:274. 1872,

sensu la to. Spergulastrum lanuginosum Michaux, Fl. Bor. Amer. 1:275. 1803. A.

alsinoides Willd. ex Schlecht, Mag. Ges. Naturf. Freund. Berlin 7:201. 1816. A.

nemorosa H.B.K., Nov. Gen. & Sp. 6:35. 1823. A. saxosa A. Gray, PI. Wright.

11:18. 1853. A. guatemalensis Standl. & Steyerm., Publ. Field Mus. Nat. Hist.,

Bot. Ser. 23:50. 1944. A. lanuginosum ssp. saxosa (A. Gray) Maguire, Amer. Midi.

Naturalist 49:498. 1951. A. lanuginosa ssp. guatemalensis (Standl. & Steyerm.) J.

Duke, Ann. Missouri Bot. Card. 48:93. 1961. Figures 41 and 42.

230 FIELDIANA: BOTANY

Herbs, prostrate, decumbent or clambering over other plants, to 1 m tall, perennial from

a taproot, usually much branched from the base, bisexual, leafy internodes 0.2-5 (15) cm

long, 0.3-2 mm thick, usually puberulent with slender whitish hairs 0.1-0.3 mm long,

hairs often confined to longitudinal ridges. Leaves opposite, equal or subequal at each

node, differing greatly in different plants but usually quite similar on the same plant,

sessile or subsessile, petioles to 3 mm long; laminae 4-30 (40) mm long, 1-16 mm broad,

linear to narrowly elliptic or elliptic-ovate, obtuse or acute at the apex, slightly (0.5 mm)

mucronate at the tip, gradually narrowed at the base and clasping the stem, margins entire

and often minutely ciliate; laminae drying thin-chartaceous, smooth and usually puber-

ulent (especially on the midvein) with thin whitish hairs ca. 0.2 mm long, with a single

prominent midvein, secondaries obscure. Flowers solitary and borne from the axils of

distal leaves, pedicels slender, 0.5-4 cm long, usually puberulent, the flowers small, 3-6

mm long, sepals 2-5 mm long, usually with green center and translucent margins, acute

at the apex, persisting in fruit; petals shorter or longer than the sepals, to 8 mm long, thin

and whitish, usually deciduous; stamens shorter than or as long as the sepals, filaments

1-4 mm long; ovary ovoid. Fruit 2.5-6 mm long, ovoid-ellipsoid, valves 2-5, thin-walled

and yellowish, splitting to the base or only halfway, thin-walled and yellowish; seeds

0.6-1 mm broad, slightly flattened, suborbicular to slightly cochleate in outline, reddish

brown (in early stages) to lustrous black, surface usually smooth.

Horizontally spreading or clambering herbs in open sites and in moist forest

understory of montane forest and highlands. The species is rarely found below

1,500 m elevation in Costa Rica, but ranges as low as 500 m in Guatemala; it can

be found on our highest mountains, and flowering collections have been made

throughout the year. The species ranges from the southern United States

through Mexico and Central America to Peru and Brazil.

Arenaria lanuginosa (in a wide sense) is characterized by its opposite subsessile,

narrow, little leaves, sprawling or clambering stems, solitary little flowers on

thin pedicels arising from the axils of distal leaves, the biseriate perianth with

undivided petals (in our area), and the thin-walled capsule with basal placenta.

The broad circumscription of this species avoids the problems of erecting new

species, but such a treatment obscures what may be real differences between

population groups in Central America. The following key was developed by

Richard Baker and recognizes four species-like taxa in Costa Rica. Three are

designated subspecies and are consistent with the treatment by James Duke in

the Flora of Panama; the fourth is unnamed. This key seems to work effectively

for a majority of the specimens encountered in Central America, but it may be

ineffective elsewhere in the range of the species. There are occasional collections

that seem to have intermediate characteristics, and this is consistent with the

experience that others have had with other species of Arenaria. We do not know

if the four entities separated by the key deserve specific rank as they were given

in the Flora of Guatemala; we present this key in the hope that it will suggest

further study.

la Valves of the mature capsule not exceeding or only slightly exceeding the sepals;

sepals glabrous, 3—4.5 mm long, petals as long as or longer than the sepals; laminae

usually less than 2 cm long and 5 mm broad, ciliate along the margin only on the

proximal half of the lamina; styles 2 or 3; plants found above 3,000 m elevation on

Volcan Baru, Chiriqui, Panama ssp. saxosa

Ib Valves of the mature capsule usually exceeding the sepals; sepals usually puberu-

lent 2a

2a Petals longer than the sepals; styles usually 3, capsule valves more than 4.5 mm long;

pedicels (1.5) 2-5 cm long; laminae to 4 cm long and often more than 5 mm broad;

plants usually found only above 1,800 m elevation ssp. guatemalensis

2b Petals equaling or shorter than the sepals; capsule valves shorter than 4.5 mm or, if

longer, the pedicels 1.5 cm long or less; laminae rarely more than 2.5 cm long or 5 mm

broad . . . . 3a

BURGER: FLORA COSTARICENSIS 231

3a Styles 2-4; capsule valves 2-4, usually bilobed and revolute, 3-4 mm long (rarely

longer); sepals usually 2-3 mm long; 500-1,800 (2,400) m elevation . .ssp. lanuginosa

3b Styles 5; capsule valves 5, sometimes bilobed, usually entire and straight, 5-7 mm

long; sepals 2.7-4 mm long; (1,800) 2,000-2,800 (3,300) m elevation, (undesignated)

Arenaria quirosii Standley, Publ. Field Mus. Nat. Hist., Bot. Ser. 22:19. 1940.

Small herbs, the stems to 6 cm tall, erect; internodes 1.5-5 mm long, densely pubescent

with multicellular trichomes to 0.3 mm long. Leaves opposite, sessile, the laminae 2.5-5

mm long, 1-2.5 mm broad, ovate to obovate, apically acute to obtuse and short-acuminate

with the tip often thickened and brownish (dry), the base broadly attenuate, the margin

entire, the lamina drying thick-chartaceous and yellowish to brownish, sparsely puber-

ulent above and below, sparsely to densely ciliate along the midrib below and along the

proximal % of the margin, only the midvein conspicuous. Flowers solitary and axillary, the

pedicels 2-4 mm long, densely pubescent with short multicellular hairs; sepals 5, 2.2-3

mm long (anthesis to fruiting), lanceolate to oblong-ovate, the apex short-acuminate, the

tip often thickened and brownish, margins rather broad and scarious, ciliate along the

midvein; petals absent; stamens 10, the filaments 0.5-0.8 mm long, anthers 0.2 mm long

(anthesis); ovary ovoid, ca. 0.6 mm long (anthesis); styles 3, ca. 0.4 mm long (anthesis) to

1.2 mm long (fruit). Fruit a capsule, ellipsoid, the valves after dehiscence approximately

equaling the sepals (ca. 3 mm); seeds numerous, 0.7 mm broad, reniform-globose, seed

coat smooth and shiny, dark maroon to black.

Arenaria quirosii can be recognized by its very small habit, its small broad

leaves, its short pedicels, and its lack of petals. This species was known only

from the type collection (M. Quiros C. 812 collected on Volcan Irazu in September

1938), but has recently been found on Costa Rica's highest mountain, Chirripo,

by Arthur Weston.

CERASTIUM Linnaeus

By David Good

Herbs, annual or perennial, low and caespitose to tall and erect, often ascending, stems

almost always puberulent, with or without glandular hairs; stipules absent. Leaves op-

posite and sessile or subsessile, a basal rosette present or absent, cauline leaves few to

many, usually shorter than the basal leaves when the latter are present; laminae linear to

elliptic, ovate or spatulate, acute or obtuse at the apex, often clasping the stem at their base

and the opposing leaf bases sometimes united, entire, usually puberulent. Inflorescence

invariably a cyme, either compact or loose, borne on distal stems, usually puberulent.

Flowers very small (2-3 mm) to medium sized (ca. 1 cm), bisexual, radially symmetrical,

sepals 5 (4), free, lanceolate to elliptic, acute, scarious margined, usually pubescent, petals

5 (4, 0), free, usually white, variously bifid; stamens 10 (5, 4), free, shorter than the petals;

ovary superior, carpels and styles 5 (rarely 4 or 3), opposite the sepals. Fruit a capsule,

usually well exserted beyond the persisting sepals at maturity and opening apically by

twice as many teeth as there were styles; 1-locular, thin walled, often curved at maturity;

seeds many, small (0.3-1.2 mm), cochleate to U-shaped or subcircular in outline, some-

what compressed, the surface variously tuberculate.

A virtually cosmopolitan genus of 50 to 100 species of small more or less scruffy

herbs. These plants are confined to cooler habitats and are not found below 1,200

m elevation in Central America. The genus can be confused with any of several

other genera in the Caryophyllaceae, particularly with Stellariaand Arenaria, but

the presence of five styles, considerable pubescence, and the exserted curved

capsules usually set it apart. Flowers of Costa Rican species are all five-parted,

with five sepals, five petals, 10 stamens, and five styles.

la Cauline leaves elliptic to ovate or obovate, usually obtuse to rounded at the apex;

pedicels 1-5 mm long, sepals 2.5-5 mm long; capsules 5-8 mm long, seeds 0.3-0.4

mm in diameter C. glomeratum

232 FIELDIANA: BOTANY

Ib Cauline leaves linear to lanceolate or narrowly elliptic, acute to acuminate at the apex;

sepals 4.3-6.5 mm long; capsules 7-12 mm long 2a

2a Pedicels up to 14-35 mm long; petals divided for % of their length; seeds 0.8-1 mm

in diameter C. guatemalensis

2b Pedicels 2-9 mm long; petals divided for Va of their length; seeds 0.5-0.7 mm in

diameter C. triviale

Cerastium glomeratum Thuillier, Fl. Paris, ed. 2, 226. 1799. C. viscosum L.

(nom. ambig.), Sp. PI. 437. 1753.

Annual plants with stems ascending or erect, sometimes caespitose, 2.5-30 cm tall, very

much branched at the base, little otherwise, branches pilose, leafy internodes usually

longer distally than basally, longer or shorter than the leaves. Leaves ± similar through-

out, perhaps slightly smaller above, basal rosette lacking; laminae broadly elliptic to

orbicular or spatulate, 5-20 mm long, 3-12 mm wide, obtuse, pilose. Basal rosette lacking.

Cymes many flowered, flowers very crowded terminally, pedicels very short, 1-5 mm

long, pilose, sometimes slightly glandular. Sepals lanceolate, 2.8-5.1 mm long, 0.5-1.6

mm wide, acute, scarious margined, pilose, sometimes glandular; petals shorter than or

equal to the sepals, bifid ca. V4 of their length, white; filaments 1.8-2.3 mm long, anthers

0.1 mm long; styles 0.8-1.1 mm long. Capsules 5.0-8.8 mm long, 1.3-1.8 mm wide,

curved; seeds 0.3-0.4 mm in diameter, finely tuberculate.

An introduced species of varied habitats including roadsides, fields, wood-

lands, volcanic rocks, cornfields, gardens, and other areas. In Costa Rica it has

been collected at altitudes of between 1,460 and 2,600 m in the provinces of

Alajuela, Cartage, and San Jose. Native to Europe, Cerastium glomeratum is found

in North America throughout much of the southern and central United States,

in central Mexico, and from southern Mexico (Chiapas) south to Chiriqui Prov-

ince, Panama. Flowering specimens can be found virtually throughout the year.

Cerastium glomeratum is recognized by its usually broadly elliptic-ovate leaves,

its bracts lacking scarious margins, and its inflorescences of very small, very

crowded flowers.

Cerastium guatemalense Standley, Field Mus. Nat. Hist., Bot. Ser. 17:244.

1937. Figure 42.

Perennial plants with stems to 45 cm tall, often much branched basally and above,

branches erect or ascending, densely glandular-pilose, internodes usually more or less

equal except at the base, where shorter. Leaves usually similar throughout or the lower

ones somewhat shorter, basal rosette lacking; laminae linear to lanceolate, widest at the

base, narrowing to an attenuate tip, 15-48 mm long, 2-5 mm wide, glandular-pilose,

particularly on the lower surface. Cymes 3-40 flowered, usually 3-15, bracts not scarious

margined, lowermost (longest) pedicels 14-35 mm long, upper ones shorter, all ± hooked

in fruit, glandular-pilose. Sepals lanceolate, 4.4-6 mm long, 1-1.7 mm wide, acute, scar-

ious margined except at the apex, glandular, petals 6-7.1 mm long, bifid ca. Vs of their

length, white; filaments 3.2-3.5 mm long, anthers 0.3 mm long; styles 1.9-2.1 mm long.

Capsules 7.8-11.8 mm long, 2-2.8 mm wide, curved; seeds tuberculate, 0.8-1 mm in

diameter.

Species of the high (2,100 to 4,400 m) pine and juniper forests on many of the

volcanoes of Guatemala and adjacent Mexico. It is also sometimes found in open

upland meadows. In Costa Rica, only a single collection has been made (Burger

and Gomez P. 8216), and it probably represents a disjunct population. This col-

lection was made at 3,250 to 3,450 m altitude on Cerro Chirripo along the upper

Rio Talari, Prov. San Jose. In the main part of its range, this species has been

collected in flowering condition in January, February, and July through October.

It probably flowers to some degree throughout the year.

BURGER: FLORA COSTARICENSIS 233

Cerastium guatemalense is distinguished by its more or less erect habit and its

open inflorescences of relatively (compared to C. triviale and particularly to C.

glomeratum) large flowers. It is the only native species of Cerastium ever collected

in Costa Rica.

Cerastium triviale Link, Enum. Hort. Berol. 1:433. 1821. C. vulgatumL. (nom.

ambig.), Sp. PI., ed. 2, 627. 1762. C. caespitosum Gilibert (nom. ambig.), Fl.

Lithuan. 2:159. 1781. C. holosteoides Fries (nom. ambig.), Novit., ed. 2, 126. 1823.

C. fontanum subsp. triviale (Link) Jalas, Arch. Soc. Zool. Bot. Fenn. "Vanamo"

18:63. 1963.

Weakly perennial plants, much branched, mostly basally, often caespitose, branches

10-25 cm long, decumbent, often long-pilose, internodes longest just below the in-

florescence, shorter below. Leaves usually similar throughout, basal rosette lacking; lami-

nae ovate to spatulate, 7-30 mm long, 3-10 mm wide, acute, sometimes obtuse, pilose.

Cymes usually quite dense, few to many flowered, bracts with scarious margins, pedicels

short, 2.2-8.4 mm long, long-pilose. Sepals lanceolate, 4.3-6.2 mm long, 1-1.6 mm wide,

scarious margined, acute, pilose, petals slightly shorter to slightly longer than the sepals,

bifid ca. Va of their length, white; filaments 3.7-4 mm long, anthers 0.3 mm long; styles

1.8-2.2 mm long. Capsules 7-11.6 mm long, 2.1-3 mm wide, curved; seeds 0.5-0.7 mm

in diameter, red-brown, tuberculate.

An introduced upland species (in our area 1,400 to 2,400 m) of varied habitats

including fields, roadsides, brushy thickets, woody areas, and even cloud forest.

Native to Europe, this species is found in North America throughout most of the

United States and southern Canada, in central Mexico and in Central America

from southern Mexico (Chiapas) south to Chiriqui Province, Panama. In Costa

Rica it has been collected virtually throughout the year from the provinces of

Alajuela, Cartago, and San Jose.

Cerastium triviale is recognized by its prostrate habit, its fairly broad leaves, its

scarious margined bracts, and its inflorescences of crowded, fairly small (though

larger than in C. glomeratum) flowers. The choice to use the name C. triviale in

this work instead of C. fontanum subsp. triviale is arbitrary. Sound judgment on

the best taxonomic level for this taxon would require study of European material.

DIANTHUS Linnaeus

Herbs, biennial or perennial, stems often stiff and erect, nodes usually thickened,

glabrous or glaucous (rarely puberulent); stipules absent. Leaves opposite and narrow,

sessile, leaf bases of opposing leaves connate and forming a short scarious sheath, laminae

entire or serrulate. Inflorescences terminal and often showy, capitate, cymose or pan-

iculate or the flowers solitary, flowers subtended by 1-3 opposing pairs of bracts. Flowers

radially symmetrical, bisexual, sepals united to form a calyx tube, 5-toothed at the apex of

the tube, with multiple parallel veins; petals 5 or more, usually narrowed at the base to

form a petiole-like structure (clawed), the distal blade of the petal entire to dentate or

fringed, lacking appendages at the base, white to rose, red or purple; stamens 10, free;

ovary superior and unilocular, narrowed at the base or stipitate, styles 2, free. Fruit a

capsule, cylindric to oblong or ovoid, dehiscent by 4 (5) valves or teeth; seeds numerous,

usually flattened and with an oval outline, attached on 1 of the flat faces, smooth to

rugose.

A genus of between 200 and 300 species native to the Old World. The four

species mentioned here are the most showy and popular of the species that may

be found in cultivation in the tropics. Each of these four species represents one

of a total of nine horticultural classes listed in Bailey's Manual of Cultivated

Plants. Bailey's manual lists more than 30 species and varieties grown in gardens

234 FIELDIANA: BOTANY

throughout the world; they generally grow best only at higher elevations in the

tropics. This genus is distinguished from our other ornamental genera by the

presence of one to three pairs of appressed bracts at the base of the calyx tube.

la Flowers crowded into dense round-topped or flat-topped heads, bracts linear and

usually as long as the calyx (sepal tube) D. barbatus

Ib Flowers in loose cymes with 2 to many flowers (note that individual flowers with

multiple petals in D. caryophyllus may appear similar to heads, but each one is enclosed

in a single calyx tube), bracts shorter than the calyx tube 2a

2a Bracts Vi-% the length of the calyx, caudate at the apex; leaves narrow-elliptic to

oblanceolate D. chinensis

2b Bracts Vt-l/3 the length of the calyx, acuminate at the apex; leaves linear 3a

3a Leaf margins and bract margins minutely serrulate to subentire or ciliate, leaves

mostly in a basal tuft and with few stem leaves; flowers not double (with 5

petals) D. plumarius

3b Leaf margins and bract margins entire and glabrous, with prominent stem leaves;

flowers usually double (10 or more petals); an important species in cut-flower

commerce D. caryophyllus

Dianthus barbatus L., Sp. PI. 409. 1753.

Perennial herbs, glabrous, stems erect, 20-60 cm tall. Leaves with lanceolate to elliptic

laminae, 3.5-8 cm long, 0.5-1.5 cm wide, apically acute, the margins usually minutely

(0.1-0.5 mm) serrulate (appearing almost ciliate). Inflorescence a dense terminal cyme that

often appears capitate, floral bracts linear to filiform, usually equalling the calyx in length;

flowers numerous within the cyme and sessile, calyx deeply toothed, the teeth long-

acuminate, distal lamina of the petals dentate or crenate, red, purple, pink, white or

variegated, corolla not usually more than 3 cm broad, rarely double.

Dianthus barbatus is a native of southern. Europe and is representative of the

horticultural group called "Sweet Williams" and Clavel Imperial. They are charac-

terized by having small flowers in bracteate heads. As with the other species

mentioned here, many variants and hybrids with other species are found in

cultivation.

Dianthus caryophyllus L., Sp. PI. 410. 1753.

Glabrous and glaucous herbs, the stems erect and stiff, few branched, with conspicuous

nodes, to 1 m tall. Basal leaves 5-15 cm long, linear and stiff, the margins entire or

minutely serrulate near the base, leaves of the stem becoming smaller distally. In-

florescence cymose, few flowered, floral bracts ca. V* as long as the calyx, abruptly acute,

pedicels subtending the flowers and their involucre of bracts to 3 cm long; calyx 2-3 cm

long, calyx teeth acute, petals usually double (10), rose, purple, or white, corolla 2-9 cm

broad and showy, very fragrant.

Dianthus caryophyllus is native to southern Europe. The "Carnation" (Clavel) is

perhaps the most popular of all the species of Dianthus, cultivated for its showy

and fragrant flowers. The distinctive odor is similar to cloves (Clavero; Syzygium

aromaticum) and cinnamon (Canela; Cinnamomum zeylanicum). In Costa Rica it is

cultivated at the middle and higher altitudes, and it is a very important element

in the ornamental cut-flower trade (see discussion in the Flora of Guatemala,

Fieldiana, Bot. 24, pt. 4:226-227, 1946).

Dianthus chinensis L., Sp. PI. 411. 1753.

Herbs, short-lived perennials, glabrous or nearly so, the stems erect. Basal leaves cadu-

cous, stem leaves 2-8 cm long, 0.5-1 cm wide, narrowly elliptic to oblanceolate, acute at

BURGER: FLORA COSTARICENSIS 235

the apex, usually minutely serrulate along the margins. Inflorescence cymose, few to many

flowered, pedicels to 2 cm long, floral bracts ca. Vi as long as the calyx or slightly longer,

cuspidate to caudate at the apex, often divergent; calyx 1.5-2.5 cm long, calyx teeth acute

to acuminate, petals rarely double, variously colored, the corolla often with concentric or

circular patterns, to 3 cm broad, odorless.

Dianthus chinensis is a native of eastern Asia. The circular patterns of the

broadly rotate corolla have resulted in the common name "Rainbow Pinks" for

these plants. Because of the smaller flowers they have been called Clavellina in

Spanish.

Dianthus plumariusL., Sp. PI. 411. 1753.

Glabrous or glaucous herbs, the stems caespitose, to 30 cm long. Basal leaves 2-10 cm

long, linear, the margins minutely serrulate, stem leaves becoming smaller distally. In-

florescence cymose and few flowered, the pedicels to 4 cm long, floral bracts V4-V3 the

length of the calyx, relatively broad, acute to acuminate at the apex; calyx ca. 2 cm long,

the teeth acute, petals dentate or more commonly deeply fringed, red, pink or white, the

corolla 3-4 cm broad, fragrant.

Dianthus plumarius is native to eastern Europe and Russia. It is representative

of a group called the "Cottage Pinks," characterized by their fragrant flowers and

tufted habit.

DRYMARIA Willdenow

REFERENCE: James A. Duke, Preliminary Revision of the Genus Drymaria, Ann.

Missouri Bot. Card. 48:173-268, 1961.

Herbs, annual or perennial, stems procumbent to erect, occasionally slightly woody at

the base, glabrous or puberulent; stipules present, entire to multilacerate, caducous or

persistent. Leaves opposite (in ours) or appearing verticellate, sessile or petiolate; laminae

broad and flat (in ours) to linear, margin entire or rarely dentate, venation palmate to

parallel. Inflorescences dichasial (in ours), terminal or axillary or with flowers clustered or

solitary at the nodes; flowers bisexual, radially symmetrical, small, sepals 5, free, green

with scarious or membranous margins, lanceolate to ovate, petals 3, 4 or 5 (rarely 0),

normally 2-cleft (bifid) at the apex or distal half and often with small auricles at the base;

stamens 2-5, filaments slightly connate at the base (rarely alternating with staminodes),

the anthers versatile, 2-thecous; ovary superior and narrowed at the base, unilocular,

placentation free central with few to many ovules, styles usually 3 (2) and united for ca.

Vz their length. Fruit a capsule, ovoid to globose, dehiscing by as many valves as there are

styles (usually 3-valved), thin-walled; seeds 1 to many, cochleate (in ours) to reniform-

globose, usually with the surfaces and edges tuberculate, embryo curved.

A genus of 48 species, according to Duke, ranging from the southern United

States to subtropical South America and with two of these species now natural-

ized in Asia and Africa, Australia, and the Pacific. Three species occur in Costa

Rica in areas characterized by evergreen (or largely evergreen) vegetation and the

genus can be found at all elevations.

Vegetatively, the Costa Rican species of Drymaria resemble those of Stellaria,

and the two are occasionally confused. Drymaria differs from Stellaria by having

three-valved capsules, partly united styles, and by the presence of stipules.

However, the stipules are often caducous or difficult to distinguish from the

surrounding trichomes and this feature may not be helpful. Species of Stellaria

with broad leaves are pinnately veined with a more or less distinct submarginal

collective vein. In contrast, Drymaria has one to five parallel or palmate primary

veins and lacks a submarginal collective vein.

236 FIELDIANA: BOTANY

la Flower pedicels covered in part with a minute dense gland-tipped tomentum that is

whitish, petals often smaller than the sepals; stipules multilacerate and usually per-

sistent; common plants from 0-1,600 m elevation D. cordata

Ib Flower pedicels glabrous to sparsely hirsute or villous or glandular, never with a

minute dense whitish or farinose tomentum; petals usually as large or larger than the

sepals; uncommon or rare plants 2a

2a Plants hirsute to glabrate; laminae rarely more than 10 mm broad, drying opaque and

the venation obscure; stipules lacerate with filiform hairlike lobes but caducous; sepals

2-3.5 mm long, elliptic to ovate; plants usually found from 1,000-2,000 m elevation

D. villosa

2b Plants glabrous to sparsely glandular hairy; laminae usually more than 10 mm broad,

drying membranaceous and the venation easily seen; stipules with lanceolate lobes

and persistent; sepals 3-6 mm long, 1,600-3,000 m elevation D. multiflora

Drymaria cordata (L.) Willd. ex Roem. & Schult, Syst. Veg. 5:406. 1819.

Holosteutn cordatum L., Sp. PI. 88. 1753. H. diandrum Sw., Prodr. 27. 1788. D.

diandra Blume, Fl. Nederl. Ind. 62. 1825. Figure 42.

Herbs, annual, prostrate to spreading or erect, to 70 cm long, often rooting at the nodes,

glandular puberulent to glabrate, leafy internodes 0.5-3 cm long, ca. 0.5 mm thick (dry);

stipules to 2 mm long, lacerate, often persisting. Leaves opposite and usually well spaced

along the stem, petioles 2-15 mm long; laminae 5-18 (25) mm long, 5-20 (30) mm broad,

broadly ovate to orbicular or reniform, rounded at the apex and occasionally mucronulate,

abruptly rounded at the base and truncate to cordate, entire, usually glabrous. In-

florescences terminal or axillary, few- to many-flowered cymes (rarely solitary axillary

flowers), bracts 2-3 mm long, narrowly lanceolate, pedicels 2-15 mm long, slender and

usually with a covering (at least in part) of very minute (0.1 mm) whitish glandular

puberulence. Flowers small, sepals 5, 2.5-4 (5) mm long, lanceolate to ovate, acute,

glandular puberulent to glabrous, lateral margins translucent, petals 5, 2-3 mm long,

deeply bifid with linear 1-veined lobes, lacking auriculae (rarely subdentate) at the base;

stamens 2 or 3 (4, 5), filaments 2-2.5 mm long and flattened, anthers suborbicular, 0.2-0.3

mm long; ovary with few to many ovules, style 0.5-1 mm long, 2- or 3-parted. Capsule

ovoid, 1.5-2.5 mm long, breaking into 3 entire valves; seeds 1 to many, 1-1.5 mm broad,

cochleate, dark reddish brown, surface with rows of tubercles.

Plants of open sunny or shaded (early secondary) sites in areas of evergreen

vegetation in the Central Highlands and Caribbean side of Costa Rica from near

sea level to 1,400 (1,600) m elevation; collected with flowers throughout the year

in southern Central America. Our plants belong to subspecies cordata which

ranges throughout the American tropics, from Mexico and Florida through Cen-

tral America and the West Indies to Argentina. Subspecies diandra (Blume) Duke

occurs in tropical Asia, Africa, Australia, and the Pacific.

Drymaria cordata is recognized by the small opposite, often broadly cordate,

leaves that are usually evenly spaced along the slender stems, the unusual

puberulence producing a farinose-like whitish covering on a portion of the

flower pedicels and the small petals. This species has not been collected from

below 1,000 m elevation on the Pacific side of Costa Rica.

Drymaria multiflora Brandegee, Zoe 5:232. 1906.

Herbs, perennial, erect or ascending, to 50 cm tall, much branched (in vigorous plants),

glabrous to stipitate-glandular (rarely sparsely villous), leafy internodes 1-10 cm long,

0.5-1.5 mm thick (dry); stipules 1-2.5 mm long, lanceolate, entire. Leaves opposite and

often widely spaced, petioles 2-20 mm long; laminae 4-22 mm long, 4-25 mm broad,

broadly deltoid-ovate to orbicular or reniform, rounded to emarginate at the apex, obtuse

to truncate at the base, entire, drying membranaceous, the 3-5 primary veins often clearly

visible. Inflorescences terminal, cymose with 3 to many flowers, peduncles to 10 cm long,

often stipitate-glandular. Flowers small, whitish, sepals 5, 3-6 mm long, 1.2-2 mm broad,

BURGER: FLORA COSTARICENSIS 237

ovate to narrowly triangular, often saccate at the base, petals 5, 2.5-7 mm long, equalling

or exceeding the sepals, bifid for % of their length, lobes rounded, with ciliate auricles at

the base; stamens 3-5, 2-3.5 mm long, anthers 0.3-0.5 mm long; ovary globose to ovoid,

styles 1-2 mm long, 2- or 3-parted. Capsule 2-3.5 mm long, ovate, with 4-26 seeds; seeds

ca. 1 mm broad, with long dorsal tubercles and substellate facial tubercles.

Rare plants (in Costa Rica) of shaded sites in montane forest between 1,500 and

3,000 m elevation. Flowering collections have been made from October to Febru-

ary in Central America. The species ranges from Mexico to westernmost

Panama.

Drymaria villosaCham. & Schlecht., Linnaea 5:232. 1830. D. hirsuta Bartl. in

Presl, Rel. Haenk. 2:8. 1831.

Herbs, annual, prostrate or more often ascending, to 45 cm long, sparsely villous to

hirsute with slender translucent hairs to 2 mm long, leafy internodes 0.5-5 cm long, ca.

0.5 mm thick; stipules 0.5-1.5 mm long, slender and usually entire, similar to the tri-

chomes. Leaves opposite and usually evenly spaced along the stems, petioles 1-10 mm

long; laminae 3-15 mm long, 3-15 mm broad, orbicular to reniform or broadly ovate,

rounded to acute at the apex, rounded at the truncate to subcordate base, entire, usually

sparsely hirsute or villous. Inflorescences cymose, terminal or axillary, with 5 to many

flowers (ultimate parts often racemose), peduncles 1-5 cm long, bracts 0.5-1.5 mm long,

pedicels 2-20 mm long, slender and sparsely villous. Flowers whitish, sepals 5, 2-3.6 mm

long, 1-2 mm broad, elliptic to broadly ovate, acute to obtuse at the apex, villous to

glabrescent, margins translucent, petals 5 (rarely reduced or absent), 2-3.6 mm long,

divided distally into 2 parts (bifid) for half their length or more, 4-veined, with caducous

filiform auricles at the base, stamens usually 5, 2-3.5 mm long; ovary ovoid to globose,

style 1-1. 5 mm long, the distal half 3-parted. Fruit an ovoid to ellipsoid capsule, 2-3. 5 mm

long, equalling or exceeding the sepals; seeds usually many, 0.5-0.9 mm broad, cochleate,

tuberculate with the facial tubercles stellate.

Small weedy plants of open sunny sites in evergreen vegetation formations,

primarily between 1,000 and 2,000 m elevation, in Costa Rica. Flowering and

fruiting collections have been made from June through the wet season to March.

This species is represented in our area by ssp. villosa; this subspecies ranges from

Mexico through Central America and along the western slopes of South America

to Peru.

Drymaria villosa ssp. villosa is characterized by its small stature, its open dif-

fused branches and inflorescences, pubescence of slender translucent hairs, the

small broad subcordate opposite leaves well spaced along the stem, and the

slender peduncles and pedicels bearing small 5-parted flowers.

GYPSOPHILA Linnaeus

Herbs or rarely subshrubs, annual or perennial, erect or spreading, usually much

branched, glabrous or glaucous; stipules absent. Leaves opposite, often narrow and sessile

or subsessile, entire. Inflorescences cymose in dichasial or paniculate arrangements (rarely

capitate), with numerous flowers, bracts present at the base of pedicels and not at the apex

(an epicalyx lacking). Flowers small, bisexual, radially symmetrical, calyx forming a tube

or united only near the base and campanulate, 5-toothed and 5-veined, scarious (thin)

between the veins, petals 5, exceeding the sepals, white to rose, usually narrowed at the

base (with a claw) and the expanded blade entire to emarginate, lacking appendages near

the juncture of claw and blade; stamens 10; ovary 1-locular and with many ovules, styles

2 (3), free. Fruit a capsule, ovoid to globose, dehiscing by 4 (6) valves or teeth, sessile; seeds

with flattened faces, subreniform, black and tuberculate.

A genus of about 125 species native to Europe and Asia, with outlying species

in Egypt and Australia. One species is commonly cultivated in Central America,

238 FIELDIANA: BOTANY

while a second species (G. paniculata L.), growing to 1 m tall and with a stout

rhizome, may also be found. Both are called "Baby's Breath/' Gipsofila, or L/o-

visna. The open panicle of small long-pedicelled flowers give a diffuse misty

effect, both in gardens and in flower arrangements. The inflorescences are some-

times sprayed with paint for more colorful effects.

Gypsophila elegans Bieb., Fl. Taur. Cauc. 1:319. 1801.

Annual herbs, stems erect, to 60 cm tall, much branched distally, glabrous. Leaves

sessile, 2-7 cm long, 0.2-1 cm broad, linear-lanceolate to lanceolate, leaf bases clasping the

stem and the opposing leaf bases slightly connate or free. Inflorescences cymose, open and

much branched and with many flowers, pedicels 1-3 cm long, slender or filiform. Flowers

small, calyx 4-5 mm long, calyx tube half the length and campanulate, weakly 5-veined,

calyx lobes acute and with thin margins, petals 2-4 x the length of the calyx, white to red,

emarginate; stamens 4-5 mm long; ovary ca. 2 mm long, ovoid, styles ca. 3 mm long,

filiform. Capsule 3.5-5 mm long, ovoid; seeds 1-1.5 mm broad, tuberculate.

Gypsophila elegans, a native of the Caucasus mountain region, Turkey and Iran,

is locally naturalized in Guatemala and commonly grown in gradens. This spe-

cies is important in the cut-flower trade for its many-branched diffuse in-

florescences for which the name Llovisna is so appropriate.

LYCHNIS Linnaeus

Herbs, annual or perennial, stems erect; lacking stipules. Leaves opposite. In-

florescences cymose, often lax and few flowered or capitate and many flowered. Flowers

usually bisexual, radially symmetrical, without an epicalyx, calyx united to form an ovoid

to obovoid tube, 10-veined, petals 5, with a proximal narrow claw and distal blade, entire

to bifid, emarginate or lacerate at the apex, usually with adaxial appendages at the junction

of claw and blade; stamens 10, filaments united only at the base; pistil short-stipitate,

ovary incompletely 5-locular at the base and 1-locular near the apex, with many ovules,

styles usually 5, free. Fruit a capsule dehiscing by 5 teeth (less often 10), borne on a short

stalk; seeds circular to reniform, surface minutely tuberculate.

A genus of about 35 species of north temperate and arctic areas. One species

is likely to be found at higher elevations in tropical American gardens.

Lychnis coronaria (L.) Desr., Lam. Encyc. 3:643. 1792. Agrostema coronariaL.,

Sp. PI. 436. 1753.

Herbs perennial, erect to 1 m tall, all parts densely white-tomentose (woolly). Basal

leaves spatulate and petiolate, 5-18 cm long, stem leaves sessile and smaller, narrowly

ovate to elliptic, acute at the apex, clasping the stem at the base. Inflorescence a compound

dichasium of only a few flowers, pedicels 4-10 cm long, thick and white-woolly. Flowers

showy, calyx 15-20 mm long (including the calyx teeth), teeth contorted in bud, ca. 5 mm

long, linear, petals ca. 3 cm long, the blade usually equalling the claw, appendages 2.5-3

mm long present at the junction of claw and blade; stamens ca. 14 mm long; styles 5, ca.

3 mm long, free. Capsule ca. 12 mm long, borne on a stipe ca. 2 mm long, teeth spreading

at the apex of the capsule; seeds numerous, less than 1 mm broad.

Lychnis coronaria, the "Rose Campion" or Espanolita, is easily recognized by its

densely white-woolly stems and leaves and its large deep rose- or purple-colored

flowers. This species is native to southeastern Europe and generally found only

in gardens above 1,800 m elevation in Central and South America.

POLYCARPON Loefling ex Linnaeus

Herbs, annual or perennial, stems diffuse or dichotomously branching, usually less than

30 cm long, internodes with minute longitudinal ridges, glabrous or puberulent; stipules

BURGER: FLORA COSTARICENSIS 239

present and scarious. Leaves opposite or whorled, laminae obovate to orbicular, entire.

Inflorescences cymose, with few or many branches, flowers subtended by scarious bracts

and usually pedicellate. Flowers small, bisexual, radially symmetrical, sepals 5, with

keeled or prominent midrib and thin margins, free, petals 5, hyaline and smaller than the

sepals, free; stamens 1-5, filaments united at the base; ovary unilocular, ovules few to

many on a free central placenta, style short and 3-lobed. Fruit a 3-valved capsule, valves

splitting almost to the base, becoming spirally twisted; seed ovoid to lenticular or curved,

embryo incurved or almost straight.

A cosmopolitan genus of temperate regions and cooler tropical highlands,

with about 15 species. A number of species are known from southern South

America, but the genus has not been previously recorded from Central America.

Polycarpon tetraphyllum (L.) L., Syst. Nat. ed. 10, 2:881. 1759. Mollugo tet-

raphylla L., Sp. PI. 1:89. 1753.

Small herbs, annual or rarely perennial (lacking a woody rootstock), stems often much

branched, to 20 cm long, essentially glabrous, internodes 5-15 mm long, with thin and

minutely serrulate ridges; stipules conspicuous, 2-3 mm long, triangular, thin and trans-

lucent, erose to cuspidate with a long slender tip. Leaves opposite or in whorls of 4,

petioles ca. 3 mm long; laminae 0.3-1 cm long, 2-6 mm broad, narrowly obovate to

oblong-obovate, bluntly rounded at the apex, tapering gradually to the attenuate base,

essentially glabrous, secondary venation obscure. Inflorescence with few or many

branches, cymose, floral bracts whitish translucent, 1-2 mm long and resembling the

stipules. Flowers small, 2-3 mm long, sepals 1.5-2 mm long, dark green along the midvein

and somewhat keeled, margins thin and translucent, petals smaller than the sepals,

emarginate; stamens usually 3-5. Fruit less than 3 mm long; seeds ca. 0.5 mm long,

semicircular or slightly lunate in outline, pale orange-brown.

Weedy little plants from Europe often found on sandy or rocky soil. The first

collection recorded for Central America was made in June 1982 on the slope of

Volcan Irazu near Tierra Blanca, Cartago, at an elevation of 2,100 m by Jorge

Gomez-Laurito (8630). The species is naturalized in South America and in

Mexico.

Polycarpon tetraphyllum is recognized by its very small stature, the two or four

small obovate leaves at each node, the conspicuous stipules, the cymose in-

florescences, and the small flowers with slightly "keeled" sepals.

SAGINA Linnaeus

REFERENCE: Garrett E. Crow, A Taxonomic Revision of Sagina (Caryophylla-

ceae) in North America, Rhodora 80:1-91, 1978.

Herbs, annual or perennial, stems procumbent, decumbent or ascending, sometimes

slightly woody at the base, short in stature, stems glabrous or glandular puberulent;

stipules absent. Leaves opposite along the stem, rosettes or tufts of basal leaves may be

present in perennial species, without distinct petioles; laminae of the cauline leaves linear

to subulate, simple and entire, clasping the stem at their base and connate with the

opposing leaf base. Inflorescences of solitary flowers, terminal or in the axils of distal

leaves; flowers bisexual, radially symmetrical, small, whitish; ebracteate; sepals 4 or 5, free,

obtuse and with scarious margins, imbricate, petals 4 or 5 (rarely fewer or none),

free, entire or slightly emarginate; stamens 4, 5, 8 or 10, equalling or twice as many as the

styles, in 1 or 2 whorls, nectaries present at the base of the outer whorl, filaments free,

anthers dehiscing longitudinally and extrorse; ovary superior and unilocular, placentation

free central with numerous ovules, styles the same number as the sepals and alternate

with them, united at the base, becoming recurved. Fruit a many-seeded capsule breaking

into 4 or 5 valves, the valves opposite the sepals and thin- walled; seeds usually 0.3-0.5 mm

broad, reniform to triangular or subglobose, surface smooth to papillate or tuberculate.

240 FIELDIANA: BOTANY

A genus of 15 species, chiefly of the cold temperate zone of the Northern

Hemisphere; centers of diversity are Europe and eastern Asia. Our species is a

Eurasian introduction which has become established at higher elevations in

Mexico, Guatemala, and Costa Rica.

Sagina procumbens L., Sp. PI. 128. 1753. Figure 41.

Herbs, perennial, usually procumbent and very small (less than 10 cm tall), much

branched from the base, stems to 15 cm long, glabrous throughout. Leaves of the stems

opposite and sessile, basal leaves 4-15 mm long, distal leaves 2.5-7 mm long, less than 1

mm broad (dry), linear and entire, apiculate to aristate at the tip, united at the base to form

a short (0.5 mm) scarious sheath. Flowers borne on slender pedicels (3) 5-10 (13) mm long,

perianth usually 4-parted, sepals 4 (5), elliptic to orbicular, 1.5-2.5 mm long, ca. 1 mm

wide, margins white, petals usually 4, 0.7-1.4 mm long, often half as long as the sepals,

orbicular to obovate or elliptic; stamens 4 (8), filaments 1-1.5 mm long, anthers 0.3 mm

long; ovary 1-2 mm long, styles ca. 0.6 mm long. Fruit 2-3 mm long, enclosed by the

persisting sepals which open out as the capsule dehisces; seeds brown, reniform to

obliquely triangular, with a distinct dorsal groove, ca. 0.4 mm broad, smooth.

A weedy species of open, often gravelly or sandy, sites, flowering throughout

the year in Central America at altitudes from 1,100 to 3,700 m elevation. This

species is a native of Eurasia and has only been found around the eastern edge

of the Meseta Central in Costa Rica and along the Interamerican Highway.

Sagina procumbens is recognized by its very small habit and its little linear

opposite leaves forming a sheath at their base, the solitary pedicellate flowers

with sepals larger than the petals and the small capsular fruit with minute

somewhat triangular seeds.

SAPONARIA Linnaeus

Herbs, annual or perennial, the stems erect or spreading; stipules absent. Leaves op-

posite, simple and entire. Inflorescence cymose, loosely flowered to capitate. Flowers

bisexual, radially symmetrical, lacking an epicalyx, calyx united to form a tube with 5 teeth

at the apex, many veined but the veins often obscure, petals 5, longer than the calyx, the

base narrowed into a claw, the expanded distal blade of the petal entire or emarginate,

appendages usually present on the base of the blade above the claw (adaxial surface);

stamens 10, united at the base to form a short tube around the stipe; pistil borne on a short

stipe, ovary 1-locular, many ovulate on a free-central placenta, styles usually 2 (3), free.

Fruit a capsule opening only at the apex with 4 (6) short teeth, ovoid to oblong, smooth;

seeds reniform and tuberculate.

A genus of about 30 species of Europe and adjacent Asia, but most numerous

in the Mediterranean area. The following species is occasionally cultivated in the

highland tropics and is widely naturalized in north temperate North America.

Saponaria officinalisL., Sp. PI. 408. 1753.

Perennial herbs, stems erect and stout but little branched, glabrous, nodes thickened

and with an interpetiolar line formed by the connate opposing leaf bases. Leaves sessile

or subsessile, laminae 5-9 cm long, 1-2.5 cm wide, ovate lanceolate to elliptic, acute at the

apex, attenuate at the base and clasping the stem, entire, with 3 distinct primary veins

impressed above. Inflorescences of opposing few-flowered dichasia clustered toward the

ends of the branches, pedicels 1-3 mm long, their subtending bracts often leaflike. Flowers

showy, calyx 15-25 mm long, tubular, calyx teeth ca. 5 mm long and with a mucronate (1

mm) tip, petal claw 20-27 mm long, petal blade 10-15 mm long, emarginate and white to

pale rose, 2 linear appendages ca. 1 mm long at the base of the blade above the claw;

stamens 10, exceeding the petal claws in length; ovary to 15 mm long and 2 mm thick,

styles 2, ca. 15 mm long, filiform. Capsule narrowly ovoid to ellipsoid, to 20 mm long and

BURGER: FLORA COSTARICENSIS 241

8 mm thick, dehiscing by 4 teeth ca. 0.5 mm long and becoming recurved, the calyx tube

persistent and enclosing the fruit; seeds to 2 mm broad, reniform, surface with concentric

rows of tubercles.

Saponaria officinalisis a native of Europe and may be found in gardens at higher

elevations in Central America. Both the Latin name and the English name

"Soapwort" are derived from the fact that the leaves give a lather when rubbed

with water.

SCLERANTHUS Linnaeus

Herbs, annual, biennial or perennial, procumbent to erect, usually densely branched,

glabrous or minutely puberulent; stipules absent. Leaves opposite, clasping the stem at

their base and with the opposing leaf bases often united to form a short sheath, laminae

entire and narrow, small. Inflorescences dense terminal or axillary cymose clusters (in

ours) or solitary flowers in leaf axils; flowers bisexual (often self-fertilizing), radially sym-

metrical, very small, sepals 5 (4), borne on the apex of an urceolate hypanthium (some-

times interpreted as a calyx tube with free distal lobes), petals absent; stamens 1-10, borne

on the hypanthium (perigynous) just below the sepals, filaments free; ovary superior but

tightly enclosed within the hypanthium, unilocular with usually 1 (2) basal ovule, styles

2, separate. Fruit a 1-seeded utricle enclosed by the hardened hypanthium and persisting

sepals; seed lenticular and smooth.

A genus of perhaps 10 species native to Europe, Asia, South Africa, Australia,

New Guinea, and New Zealand. The small size of the plants and crowding of

flowers and axillary branches with condensed internodes produce a character-

istic appearance similar to that of Spergula arvensis.

Scleranthus annuusL., Sp. PI. 406. 1753. Figure 41.

Herbs, annual, decumbent or ascending to 20 cm tall, much branched, leafy internodes

0-30 mm long, sparsely puberulent. Leaves sessile and clasping the leaf at their base;

laminae 3-10 (20) mm long, linear or subulate, the sheathing leaf base united with the

opposing leaf base to form a very short sheath, entire, glabrous on the surfaces but the

basal margins ciliate, venation obscure. Inflorescence a compact few-flowered cymose

cluster exceeded by the leaflike bracts, flowers subsessile; sepals 5, 1.2-2.5 mm long,

lanceolate with a thick midrib, borne at the apex of the urceolate or deeply cup-shaped

hypanthium, the hypanthium ca. 1.5 mm long, glabrous, with 10 indistinct ribs; stamens

ca. 0.5 mm long; ovary ca. 0.9 mm long and 0.5 mm thick, styles 2, ca. 0.7 mm long, linear.

Fruit enclosed within the hardened hypanthium, 3-5 mm long (including the sepals), the

persisting sepals erect to slightly incurved, ca. 1-2 mm long; seeds 0.7-1 mm long,

ellipsoid to ovoid, pale yellowish, with a small projection at the apex and shallow vertical

depressions on each of the 2 slightly flattened faces.

Small weedy plants of open sites; collected by Brenes in June 1906 from along

the Rio Reventado at 1,500 m and more recently on the slopes of Volcan Irazu at

about 2,400 m, both in the province of Cartago. This species is native to north-

western Europe and is now established in North America from Mexico to Can-

ada and in the mountains of Ecuador.

Scleranthus annuus is recognized by its small stature, narrow little leaves ex-

panded and united across the stems at their base, flowers and fruit in dense little

clusters, and the urceolate hypanthium with five distal sepals tightly enclosing

the fruit.

SILENE Linnaeus

Herbs or small shrubs, glabrous to puberulent; stipules absent. Leaves opposite or

whorled, sessile or petiolate, the sheathing bases connate or free, usually entire and

242 FIELDIANA: BOTANY

pinnately veined. Inflorescences cymes, racemes, dichasia or the flowers solitary, pedicels

bracteate only at the base (an epicalyx absent). Flowers bisexual, radially symmetrical,

calyx tubular with (5) 10-30 veins, ovoid to turbinate or campanulate in form, 5-toothed

at the apex, petals 5, narrowed at the base to form a distinct claw and usually with

appendages at the junction of claw and blade, the blade entire or more commonly lobed

or toothed; stamens 10, usually borne at the apex of the androgynophore, filaments

sometimes united at the base; pistil borne on the androgynophore, ovary 3-locular at the

base but 1-locular above, with many ovules on a free-central placenta, styles 3 (rarely 4 or

5). Fruit a capsule dehiscent only at the apex with 6 (3, 8, 10) short teeth, borne on a

carpophore, calyx persisting in fruit and slightly inflated but little enlarged and often

splitting; seeds reniform to subcircular, slightly flattened on 2 faces, minutely tuberculate.

A genus of nearly 400 species, nearly all from the Old World. Both species

included here are natives of Europe. They are distinguished by the lack of an

epicalyx, three styles, and calyx tube with 10 or more veins.

la Plants glabrous; grown in gardens for the showy flowers S. armeria

Ib Plants densely pubescent; weeds of higher elevations S. gallica

Silene armeria L., Sp. PI. 420. 1753.

Erect, annual, mostly glabrous herbs to 70 cm. Leaves 4-8 cm long, 1-2.5 cm wide,

sessile, lanceolate to ovate, apically acute to acuminate, clasping at the base and sometimes

slightly connate. Inflorescence a crowded compound dichasium, many flowered; pedicels

1-4 mm long (to 20 mm for the terminal flower), ebracteate; calyx 1.4-1.6 cm long, clavate

at anthesis, becoming abruptly expanded in its distal half by the expanding stipitate fruit,

10-nerved, 5-toothed, the teeth rounded; petals 5, blades entire or emarginate, the 2

appendages ca. 2 mm long, linear, the blades pink to purple, the corolla ca. 15 mm in

diameter.

This species is called Llovizna and Espanola in Central America; the English

common name is "Sweet William Catchfly." The glaucous and glabrous leaves

are distinctive.

Silene gallica L., Sp. PI. 417. 1753. S. anglica L., loc. cit. 416.

Herbs, annual, erect or spreading, 15-50 cm tall, villous-puberulent throughout and

glandular-pubescent on the distal portions of the plant, leafy internodes (3) 4-9 cm long,

with short scurfy to longer translucent and crooked hairs (0.3-2.5 mm). Leaves sessile or

short (0-5 mm) petiolate, the lower laminae 3-5 (7) cm long, spatulate, rounded at the apex

and often mucronate, tapering to the connate base, upper laminae smaller and narrower

with acute apices. Inflorescence a terminal 1-sided raceme, rachis glandular-pubescent,

upper pedicels very short (0.5-5 mm), pedicels subtended by a pair of linear leaflike bracts

8-20 mm long. Flowers with calyx tubes 7-10 mm long, becoming 5 mm in diameter in

fruit, with 10 veins, calyx teeth ca. 2 mm long, linear, villous with crooked hairs to 3 mm

long; petals bifid or toothed at the apex, entire below, white to pale purple, exceeding the

calyx by ca. 1 mm, basal appendages oblong, ca. 1 mm long. Fruit a capsule 6-9 mm long,

ovoid, dehiscent by 6 teeth, the teeth short (1 mm), acute, recurved; seeds ca. 1 mm broad,

cochleate with rows of tubercles.

This species is a native of southern and central Europe now widely naturalized

in the New World. It is a plant of open weedy sites and cultivated fields in

Central America. In Costa Rica it is found between 1,600 and 2,400 m around the

Meseta Central; it flowers in the wet season between April and November.

Silene gallica can be distinguished from our other members of the family by the

viscid glandular hairs on distal parts, the one-sided inflorescence, the tubular

calyx, and the capsule opening only at the apex with six small teeth.

BURGER: FLORA COSTARICENSIS 243

SPERGULA Linnaeus

Herbs, annual, usually much branched from the base, the stems decumbent to as-

cending, mostly glandular pubescent, rarely glabrous; stipules present and small. Leaves

opposite or appearing verticellate by crowding of leafy axillary branches with condensed

internodes, simple and entire, sessile or subsessile, linear or subulate. Inflorescences

dichasial and terminal, small bracts present and subtending the slender pedicels; flowers

bisexual, radially symmetrical, small, sepals 5, free and imbricate, petals 5, free, margins

entire, white; stamens 5-10, filaments arising from a hypogynous ridge, free, anthers

dehiscing longitudinally; ovary 1-locular, placentation basal with many ovules, styles 5,

alternate with the sepals, free to the base. Fruit a capsule dehiscing by 5 valves, loosely

enclosed within the persisting sepals, valves thin walled; seeds lenticular to subglobose,

suborbicular in outline, with an acute or winged circumferential margin, surface often

papillose.

A genus of four species native to Europe and Asia; one species has become

widely naturalized in the New World. The verticellate appearance of leaves at

most all the nodes (because of the condensed axillary shoots) is very distinctive.

Spergula arvensis is somewhat similar in appearance and habit to Polycarpaea

corymbosa (L.) Lam. which is known to range from South America into southern

Panama. However, the inflorescences of P. corymbosa are congested, and the

cochleate seeds lack the thin winglike margin and minutely papillose surface

found in Spergula.

Spergula arvensis L., Sp. PI. 440. 1753. Figure 41.

Erect to spreading much-branched herbs to 40 cm tall, stems and foliage glabrous below

but often becoming glandular-viscid on the upper portions, internodes 1.5-6 cm long;

stipules 0.8-1.5 mm long, triangular, acute to obtuse, scarious. Leaves congested at the

nodes in verticellate-like clusters, sessile; laminae 1-3.5 cm long, 0.3-0.8 mm broad (dry),

linear to subulate, entire, minutely puberulent with thin whitish hairs ca. 0.2 mm long.

Inflorescences a compound dichasium with the terminal portions becoming racemose, to

20 cm long, lax, 10- to 30-flowered, pedicels 1-1.5 cm long, slender, usually glandular-

viscid; flowers ca. 3 mm long (dry), sepals 2.5-3.5 mm long, persistent in fruit and

becoming 4 mm long, ovate, acute to obtuse, usually viscid, the margins thin; petals 2-3.5

mm long, as long or slightly shorter than the sepals, entire, often persistent in fruit;

stamens usually 10, 1-1.5 mm long; ovary ca. 2.5 mm long, subglobose, styles 5, ca. 0.5

mm long, curved. Fruit a capsule 3.5-5 mm long, as long as or slightly exceeding the

sepals, ovoid; seeds ca. 1 mm broad and 0.7 mm thick, subglobose, with a narrow whitish

circumferential ridge, the broad surfaces brown to black and usually covered with minute

yellowish clavate papillae.

A native of Europe but now widely naturalized in North America and in the

higher elevations of Guatemala and western Panama. Collections by Jorge

Gomez-Laurito near Tierra Blanca, and by Robert Wilbur along the Carretera

Interamericana, all in the province of Cartago, are the first recorded from Costa

Rica. This species is a weed of open fields and also grows in sandy areas along

rivers and streams, between 1,400 and 3,200 m elevation in Central America. The

majority of Central American collections have been made in January and

February, but the Costa Rican collections were made in June, July, and

December.

STELLARIA Linnaeus

Herbs, annual or perennial, generally with slender procumbent or clambering, repent or

ascending (occasionally pendent) stems, puberulent or glabrous; stipules absent. Leaves

244 FIELDIANA: BOTANY

opposite, sessile or petiolate, often clasping the stem at their base, laminae entire, broad

or narrow, venation usually pinnate and with a slender submarginal vein. Inflorescences

of distal dichasial cymes subtended by reduced sessile leaves (bracts) or the flowers solitary

in axils of distal leaves, a progression of basal petiolate leaves to smaller sessile distal bracts

often present on flowering stems, flowers usually subtended by slender pedicels. Flowers

bisexual, radially symmetrical, small, sepals 4 or 5, free to the base, imbricate, petals as

many as the sepals (rarely absent), 2-cleft (bifid and often giving the appearance of 8 or 10

petals), white, lacking basal appendages; stamens 10 or fewer, free, staminodes absent;

ovary superior and unilocular with basal, central or free central placentation, ovules few

to many, styles 2 or 3 (rarely 4 or 5), free. Fruit a capsule, globose, ovoid, or oblong,

dehiscing by as many or twice as many valves as there are styles, valves thin walled; seeds

(1) few to many, cochleate to reniform or globose, usually reddish brown, the surface

smooth to rugose, muricate, papillate, or tuberculate.

A genus of about 100 species, most abundant in the cooler parts of both the

northern and southern temperate zones and usually at higher elevations in the

tropics. These plants can be mistaken for species of Cerastium or Drymaria, but

our species of Stellaria can be distinguished by the presence of a slender sub-

marginal collecting vein which unites the secondary veins near the edge of the

leaf and runs parallel with the margin in the distal half of the lamina (but this

is not always easy to see).

la Laminae lanceolate to oblanceolate, to 8 mm long, drying stiff; stems conspicuously

4-ridged, ridge continuous with the midvein of the leaf; internodes to 15 mm long;

rare endemics of the eastern volcanoes; 1,600-2,600 m elevation S. nubigena

Ib Laminae lanceolate to ovate, cordate or triangular, to 5 mm long, drying thin and

membranous; internodes usually becoming more than 2 cm long, rarely with 4 longi-

tudinal ridges 2a

2a Sepals and petals 4, sepals to 2.5 mm long, inflorescence a compound dichasium;

stems with ridges that are continuous with midveins of the leaves; leaves often

broadly cordate; 1,500-3,300 m S. irazuensis

2b Sepals and petals 5; stems usually terete and lacking ridges extending into the mid-

vein 3a

3a Inflorescence a compound dichasium; sepals and petals 2-3.5 mm long; leaves trun-

cate to subcordate; mostly between 600 and 2,000 m elevation S. prostrata

3b Flowers solitary in the axils of leaves or of large leaflike bracts 4a

4a Sepals 2.5-4 mm long, petals to 4.5 mm long; petioles 0-5 mm long, laminae truncate

to obtuse at the base; 50-1,800 (2,300) m S. ovata

4b Sepals 4-6 mm long, petals 7-13 mm long; petioles 4-25 mm long, laminae subcordate

to cordate at the base; (1,800) 2,400-3,200 m S. cuspidata

Stellaria cuspidata Willd. ex Schlecht., Mag. Ges. Naturf. Freund. Berlin

7:196. 1916. S. ciliataVahl ex Pers., Syn. 1:503. 1805, not Gilib., 1781. S. limitanea

Standl., Publ. Field Mus. Nat. Hist., Hot. Ser. 22:74. 1940.

Herbs, annual (perhaps perennials), much branched, stems procumbent to ascending,

clambering and pendent, growing over other plants to 2 m high, internodes of the main

stems to 14 cm long, mostly glabrous and becoming pale yellowish, internodes of the

axillary shoots mostly 1-2 cm long, often villous, 0.5-1.5 mm thick (dry). Leaves opposite,

sheathing bases of opposing leaves slightly connate, petioles 4-12 (25) mm long (in ours);

laminae 5-16 (22) mm long, 3-14 (18) mm wide, deltoid to narrowly triangular, acute to

acuminate at the apex and with a mucronate tip, abruptly truncate to cordate at the base

and the basal lobes slightly unequal, margins entire, drying membranaceous to char-

taceous, villous to glabrous, secondary veins usually obscure, the submarginal vein ca. 0.3

mm from the edge. Flowers usually solitary and axillary, often with 1 or 2 flowers at distal

nodes, pedicels (10) 20-40 mm long, sparsely to densely villous; sepals 5, 4-6 mm long,

1.5-2.5 mm broad, narrowly ovate to elliptic, sparsely to densely villous, the margins

membranous or scarious, expanding somewhat in fruit, petals deeply bilobed for over half

their length (and often appearing as 10 petals), 7-13 mm long, white; stamens 4.5-7 mm

BURGER: FLORA COSTARICENSIS 245

long; ovary 2-2.5 mm long, cylindrical, styles usually 3, 2-3 mm long. Capsule 6-11 mm

long, ovoid to ellipsoid; seeds ca. 1 mm broad, cochleate splitting into 3 valves, valves

deeply bifid; seeds ca. 1.1 mm broad, cochleate, the tubercles rounded, reddish brown.

Plants of high montane forest formations from (1,800) 2,400 to 3,200 m ele-

vation (to 4,000 m in Guatemala). Central American flowering and fruiting

collections have been made primarily from December through March (no col-

lections have been seen from April and May). This species, whose name and

circumscription are uncertain, ranges from the southernmost United States and

Mexico through Central America to Colombia and Ecuador.

Stellaria cuspidatais recognized by its long scrambling stems, relatively narrow,

cordate to triangular, leaves, and relatively large flowers borne singly in the axils

of leaves (or large leaflike bracts). The species is sometimes confused with speci-

mens of S. prostrata, with usually broader leaves, smaller flowers, and from lower

elevations; compare that species.

It is difficult to see why these plants were placed under the European S.

nemorum L. Plants of S. nemorum have thicker erect stems usually branched only

in the distal inflorescence, basal leaves twice as large and with winged petioles,

and the flowers are usually borne on distal inflorescence-like stems with greatly

reduced leaves (i.e., bracts). Our use of the name S. cuspidata is consistent with

its usage in the Flora of Guatemala. However, a photo said to be type material

from Ecuador looks rather different from typical Mexican and Central American

material placed here. The Ecuadorean specimen has larger leaves on longer

petioles, and the flowers are on long pedicels, but the sepals appear to be the

same as in our material. A type photograph of S. cilia ta Vahl and collections from

northern South America placed under S. ciliata are very similar to our material;

we follow the treatment in the Flora of Peru by considering all of these as

variants of the same species.

Stellaria irazuensis Donnell Smith, Bot. Gaz. 23:236. 1897.

Herbs, perennial, prostrate or spreading, usually much branched, leafy internodes 1-10

cm long, 0.4-2 mm thick, glabrous, becoming pale yellowish when dry. Leaves opposite,

usually evenly spaced along the stems, petioles 6-25 mm long, sparsely villous with

slender crooked translucent hairs ca. 1 mm long; laminae 10-20 (40) mm long, 7-15 (25)

mm broad, deltoid to triangular or ovate, acuminate to abruptly attentuate at the apex,

rounded at the truncate to cordate base, margin entire, laminae drying membranaceous

and light green, sparsely villous, with 2 or 3 pairs of major secondary veins and a slender

submarginal vein running parallel with the edge of the lamina ca. 0.5 mm from the margin.

Inflorescences to 25 cm long, terminal or appearing to be axillary, of open dichotomously

branching cymes, a pair of bracts at each dichotomy with the basal pair of bracts leaflike

and sessile and the successive bracts becoming smaller, primary peduncle 4-15 cm long,

each succeeding peduncle becoming shorter by l/3-l/2, pedicels sparsely to densely

glandular-villous. Flowers 4-parted, sepals 1.2-2.5 mm long, 0.5-1 mm broad, elliptic,

acute to obtuse, glabrous; petals 4, 2-4 mm long, deeply cleft, the lobes 1.5-3.5 mm long,

white; stamens 4-8, ca. 1.5 mm long, ovules 3-6 on basal placentae. Capsule ellipsoid,

1.5-2.5 mm long, subtended by the persisting sepals and petals, dehiscing by 4 valves;

seeds 1-1.3 mm broad, cochleate, dark reddish brown, with a tuberculate surface.

Plants of wet evergreen montane formations between 1,500 and 2,800 m ele-

vation (to 3,300 m in Guatemala). Flowering collections have been made primar-

ily between June and January, with a few collections made in February and

March. This species ranges from Chiapas, Mexico, to Guatemala, Nicaragua,

Costa Rica, and the westernmost highlands of Panama (we have seen no col-

lections from El Salvador or Honduras).

246 FIELDIANA: BOTANY

Stellaria irazuensis is distinguished by its very small four-parted flowers, the

open inflorescences with long peduncles and widely spaced cymes, and the

deltoid to cordate leaves that dry thin and translucent green. This species is

apparently a close relative of S. venezuelana Steyerm.

Stellaria nubigena Standley, J. Wash. Acad. Sci. 17:250. 1927.

Herbs, perennial, much branched and often forming clumps or mats, stems to 15 cm

long, leafy internodes 5-15 mm long, glabrous, sharply 4-angled (in cross section), drying

pale yellowish. Leaves opposite, decussate and sessile, clasping the stem at their base and

the bases of opposing leaves arising close together; laminae 4-8 mm long, 1-2.5 mm wide,

lanceolate to oblanceolate, apex acute and with a calloused tip, margin entire, blades thick,

stiff, drying yellow to yellow-green, distally glabrous and villous near the base, midvein

continuous with the ridges of the stem, secondary veins inconspicuous, submarginal veins

obscure. Inflorescence few-flowered cymes or the flowers solitary, peduncles 2.5-5 mm

long and subtended by a pair of scarious bracts 1.5-2 mm long, pedicels 4-10 mm long,

glabrous, subtended by a pair of scarious bracts 1-1.5 mm long. Flowers 5-parted, sepals

ca. 2 mm long at anthesis (up to 3 mm in fruit), lanceolate, glabrous, green with a scarious

margin, petals 5, shorter than the sepals (ca. 1 mm at anthesis to 2 mm in fruit), 2-cleft

nearly to the base, white; stamens apparently 8, unequal, the largest to 1 mm long; ovary

ovoid, ca. 1 mm long, styles 3, 0.7-0.9 mm long. Capsule ca. 3 mm long, ovoid, splitting

into 6 valves; seeds ca. 0.7 mm broad, cochleate-reniform, tuberculate, reddish brown.

Stellaria nubigena is endemic to Costa Rica and known only from collections by

Paul Standley on Volcan Turrialba (35344, the type, US) and on the slope of

Volcan Irazu near Las Nubes, San Jose (numbers 38736, 38784, and 38829)

collected in February and March of 1924 between 1,600 and 2,600 m elevation.

This species is easily distinguished from our other species of Stellaria by its stiff

little sessile lanceolate leaves, short internodes,- and four-angled stems.

Stellaria ovata Willd. ex Schlecht., Mag. Ges. Naturf. Freund. Berlin 7:196.

1816. Figure 42.

Herbs, perennial, prostrate or spreading to 60 cm long, much branched from the base,

leafy internodes 0.4-6 cm long, 0.5-1.5 mm thick, glabrous to densely hirtellous, the

trichomes thin and translucent, to 1 mm long, stems green to pale yellowish when dry,

and longitudinally ridged. Leaves opposite and usually evenly spaced along the stem,

petioles 0-5 mm long, with lateral margins continuous with the lamina margins; laminae

5-25 (30) mm long, 7-14 mm wide, ovate to triangular, acute to obtuse at the apex and

with a minute (0.7 mm) mucronate tip, attenuate at the base and decurrent on the winged

petiole, drying yellowish or greenish, hirtellous, with 2-4 pairs of major secondary veins,

submarginal vein 0.2-0.4 mm from the margin and often obscure. Flowers solitary in the

axils of leaves, borne on long (6-25 mm) slender (0.3 mm) ebracteate hirtellous pedicels

(rarely glabrous); sepals 5, 2.5-4 mm long, ovate to obovate, apically acute to obtuse, often

hirtellous, petals 5, 3.5-4.5 mm long, bilobed, lobes to 1 mm long, white; stamens 10, ca.

3 mm long; ovary ovoid 2.5-3.5 mm long, with numerous ovules on a basal placenta, styles

3. Capsule 3-4 mm long, ovoid; seeds 1 mm broad, subreniform to subcircular in outline,

dull reddish brown, with rows of prominent tubercles to 0.2 mm high.

Plants of moist or shaded areas in regions of evergreen vegetation formations

between 50 and 1,800 (2,300) m elevation, having been collected on the Carib-

bean slopes and lowlands, the Central Highlands, the General Valley, and the

Golfo Dulce area in Costa Rica. Flowering collections have been made through-

out the year. The species ranges from Mexico southward through Central Amer-

ica to Venezuela and Peru.

Stellaria ovata is distinguished from other species of the genus by its combina-

tion of nearly sessile ovate leaves, solitary axillary flowers, and lower elevation

habitat.

BURGER: FLORA COSTARICENSIS 247

Stellaria prostrata Baldwin, in Elliott, Bot. South Carol. & Georgia 1:518. 1821.

Herbs, annual or perennial, stems procumbent to subscandent, to 60 cm long, leafy

internodes 1-7 (12) cm long, 0.5-3 mm thick, minutely puberulent to villous with tri-

chomes to 0.5 mm long. Leaves opposite and usually evenly spaced on the stems, petioles

10-25 mm long, with lateral wings, glabrous to sparsely villous along the margins of the

wing; laminae 5-30 (40) mm long, 5-20 (35) mm broad, ovate to deltoid, acute to acuminate

at the apex, truncate to subcordate at the base, slightly decurrent on the petiole, drying

membranaceous and pale green, glabrous or sparsely villous along the entire margins,

with 2-4 pairs of major secondary veins, submarginal vein difficult to see. Inflorescences

dichasial, lax, with few to many flowers, moderately to densely villous or villous-glandular

with hairs 0.1-0.5 mm long, bracts subtending the inflorescence 6-20 mm long and

petiolate (leaflike), each succeeding pair of bracts becoming smaller, often villous. Flowers

5-parted, sepals 2-3.5 mm long (becoming 4.5 mm long in fruit), ovate to elliptic-

lanceolate, glabrous to villous-glandular, petals 5, 2-3.5 mm long, deeply lobed, the lobes

1.3-2.5 mm long, white; stamens 1.5-2 mm long; ovary ca. 1.5 mm long, styles 3 or 4, ca.

1 mm long. Capsule 4-5.5 mm long, ellipsoid to ovoid, splitting into 6 valves, the valves

thin walled and strongly revolute at the tip; seeds 0.7-1 mm broad, cochleate-reniform,

with rows of minute cylindrical capitate tubercles, reddish brown.

Plants of moist and shaded sites in areas of evergreen vegetation formations

between 600 and 2,000 m elevation in Costa Rica (to 4,000 m elevation in Gua-

temala); flowering and fruiting throughout the year, but with most collections

made from December through March. The species ranges from the southern

United States (Texas to Georgia and Florida) through Mexico and Central Amer-

ica to Colombia.

Stellaria prostrata is very similar to S. irazuensism vegetative characteristics, but

it is distinguished from that species by its larger five-parted flowers and capsule

valves revolute at the tip. It has been confused with certain forms of what is here

called S. cuspidata and can be distinguished from that species by the capitate

tubercles on the seeds, the sepals usually less than 4 mm long (at anthesis), the

petals usually equal to the sepals in length, and the strongly revolute capsule

valves. These differences appear to be consistent with ecological differences: S.

prostrata is most commonly found between 1,200 and 1,800 m elevation (extremes

of 700 to 2,000 m), while S. cuspidata is usually found between 2,500 and 3,200

m (extremes of 1,900 to 4,000 m) in northern Central America. (Specimens

identified as S. prostrata by Standley and Steyermark from over 2,000 m elevation

in Guatemala are sterile.) See the discussion under S. cuspidata.

INDEX

The index includes all accepted names (in roman typeface), synonyms (italics ),

common English names (roman) and vernacular names (italics). Page numbers

of illustrations are in boldface. Hyphenated words and multiple words are

alphabetized letter by letter.

Abanico 162

Acelga 139

Acetosella 132

Achyranthes 144

altissima 165

aspera 144, 146

bettzickiana 148

brasiliana 148

calea 173

costaricensis 151

indica 145, 146

laguroides 151

lehmannii 153

megaphylla 151

panamensis 153

prostrata 167

stenophylla 153

williamsii 154

Adorno de nino 176

Aethanthus nodosus 47

Agdestis clematoides 203

Agonandra 15, 28

brasiliensis 29

macrocarpa 15, 28

obtusifolia 29

racemosa 29

Agrostema coronaria 238

Aguacatia 11

Aizoaceae 213, 223, 225, 226

Algodoncillo 174

Allionia 180

violaceae 192

Alternanthera 145, 149, 152

bettzichiana 148, 149

bettzickiana 148

brasiliana 148

caracasana 149, 150

costaricensis 151, 152

ficoidea 153, 156

laguroides 151, 152

lanceolata 153

lehmannii 153

Alternanthera panamensis 153

paronychioides 149, 153

polygonoides 154

pubiflora 152, 154

repens 150

sessilis 155

stenophylla 153

tenella 149, 155

williamsii 154

Amaranthaceae 142-180, 146, 149, 152,

158, 163, 171, 223

Amaranthus 156, 158

caudatus 159

cruentus 158, 159

dubius 157, 158

hybridus 157, 158

subsp. cruentus 159

hypochondriaceus 157

scariosus 158, 159

spinosus 158, 160

tristis 157

viridis 158, 161

Amor seco 162

Andripetalum suaveolens 11

Anredera vesicaria 222

Antidaphne 32, 37

viscoides 32, 37

Antigonon 100, 104

cordatum 106

grandiflorum 101

guatemalense 101, 104

leptopus 104, 106

macrocarpum 101

Ant-plant 137, 138

Apinagia 3

myriophylla 6

Apodantheae 90

Apodanthes 37, 90

caseariae 37, 91

flacourtiae 91

panamensis 91

tribracteata 91

248

BURGER: FLORA COSTARICENSIS

249

Arenaria 225, 226, 229

alsinoides 229

guatemalensis 229

lanuginosa 225, 226, 229

ssp. guatemalensis 226, 229-231

lanuginosum ssp. saxosa 229, 230

nemorosa 229

quirosii 231

saxosa 229

Aristolochia 79, 80, 81, 82

anguicida 81, 82

bicolor 84

bilabiata 82

chapmaniana 85

constricta 80, 83

costaricensis 85

var. zamorensis 85

gigantea 81, 83

grandiflora 81

linearifolia 82

littoralis 84

loriflora 82

maxima 80, 84

odoratissima 81, 84

pandurata 84

pilosa 81, 85

ringens 85

schmidtiana 86

securidata 83

sylvicola 83

tonduzii 80, 85

translucida 80, 86

trilobata 81, 86

veraguensis 80, 87

Aristolochiaceae 79, 80, 81

Baby's Breath 238

Balanophoraceae 93, 94, 95

Banksia 8

Basella alba 222

Basellaceae 222, 224

Batchelor's button 169

Bdallophyton 90, 91, 92

americanum 91, 92

andreuxii 91

bambusarum 91

cerantantherum 91

oxylepis 91

Beets 139

Bejuco de ajo 203

Bellisima 106

Beta 139

vulgaris 139

Bilderdykia 104, 106

aubertii 104, 106

Black Man wood 24

Blandowia myriophylla 6

Bledo 156

Boerhavia 181

Boerhavia coccinea 183, 187

diffusa 183, 188

erecta 183, 188

paniculata 188

Bogambilla 189

Boldoa 199

ovatifolia 199

purpurascens 199

Boria 162

Bougainvillea 182, 189

buttiana 189

glabra 190

spectabilis 189

Boussingaultia 222, 224

ramosa 222, 224

Brandesia lanceolata 153

pubiflora 154

Bugambilla 189

Buguenvitia 189

Calandrina megarhiza 218

Caldasia cayennensis 97

Calalu 206

Callite 138

Campderia floribunda 120

mexicana 120

Camron 176

Canilla de Mula 138

Caraxeron 177

vermicularis 177

Carnation 234

Caryophyllaceae 225, 226, 227

Caryophyllales (families 63-70) 138-247

Casearia 91

Celosia 161, 163

argenta 162, 163

var. cristate 162

paniculata 165, 175

pleiogyna 178

virgata 162, 163

Centrospermae (families 63-70) 138-247

Centrostachys aspera 144

indica 145

Cephalotomandra 180

fragrans 180

panamensis 180

Cerastium 226, 231

caespitosum 233

fontanum ssp. trivale 233

glomeratum 232

guatemalense 226, 232

holosteoides 233

trivale 233

viscosum 232

vulgatum 233

Chamissoa 163, 164

acuminata 165

altissima 163, 165

macrocarpa 165

250

FIELDIANA: BOTANY

Chard 139

Chaunochiton 16, 18

kappleri 16, 18

Chenopodiaceae 138, 202

Chenopodiales (families 63-70) 138-247

Chenopodium 140, 202

album 140

ambrosioides 141, 202

anthelminticum 141

quinoa 140

Chile de perm 129

Chivito 174

Chivo 174

Cladocolea 35, 38

oligantha 35, 39

Clavel 234

Clavel Imperial 234

Clavellina 235

Clauellina mantes 215

Claytonia paniculata 221

Coccoloba 107, 108, 109, 110, 111

acapulcensis 109, 113

acuminata 110, 114

allenii 116

belizensis 111, 114

browniana 113

caracasana 108, 115

cardiophylla 113

changuinolana 116

floribunda 120

guanacastensis 111, 116

hirsuta 114

lehmannii 110, 116

molinae 120

obovata 109, 117

padiformis 111, 119

riparia 117

roseiflora 118

standleyana 115

tuerckheimii 109, 118-119

uvifera 108, 119

venosa 110, 120

wercklei 113

williamsii 116

Colacion 106

Colchbn de nino 148

Comenternero 174

Commicarpus 180

Confite 106

Confitillo 106

Conopholis 93

Coqueta 148

Cordia alliodora 75

Corynaea 95, 96

crassa 95, 96

sphaerica 95, 97

Cottage Pinks 235

Coyontura 174

Coyontura de polios 176

Cresta de gallo 162

Cricamola 24

Crillo 24

Cyathula 146, 166

achyranthoides 146, 166

prostrata 146, 167

Cynomorium cayennense 97

Cytineae 90

Cytinus americanum 91

Danto hediondo 13

Dendrophthora 60, 61

ambigua 61, 64

biserrula 65

costaricensis 61, 64

subsp. poasensis 64

geniculata 65

squamigera 61, 65

terminalis 61, 66

Desmochaeta achyranthoides 166

Dianthus 233

barbatus 234

caryophyllus 234

chinensis 234

plumarius 235

Dioscorea 82

calyculata 222

Drymaria 226, 235

cordata 226, 236

diandra 236

hirsuta 237

multiflora 236

villosa 237

ssp. villosa 237

Dufourea hypnoides 7

trifaria 7

Eremolepidaceae 31, 37

Eriogonum 99

Escambron 124

Escambron negro 124

Espanola 242

Espaholita 238

Espinaca 141, 142, 213

Eubrachion 31

Fallopia aubertii 107

Flor de corona 174

Flor de mano 162

Flor de San Miguel 106

Flor de Verano 189

Flares de palo 50

Flowering stones 213

Four-o'clock 192

Gaiadendron 34, 39

poasense 39

punctatum 34, 39

Gateador 121

BURGER: FLORA COSTARICENSIS

251

Gipsofila 238

Glinus 213, 226

radiatus 214, 226

Golondrina 154

Gomphrena 168

brasiliana 148

decumbens 169

disperse 169

elegans 168

ficoidea 153

filaginoides 168

globosa 168

latifolia 173

paniculata var. hookeriana 176

serrata 149, 169

sessile 155

tuerckheimii 168

vermicularis 177

Grain amaranths 156, 157-159

Grevillea 9, 10

banksii 9

robusta 9, 10

Guapira 184, 190

costaricana 184, 190

linearibracteata 190

standleyanum 190

Gypsophila 237

elegans 238

paniculata 238

Hebanthe hookerianus 176

Heisteria 17, 18

acuminata 18, 19, 20

acuta 23

cocinnea 20

concinna 18, 20

costaricensis 18, 20

cyanocarpa 18, 21

eurycarpa 22

fatoensis 22

kappleri 16

latifolia 22

longipes 21

macrophylla 18, 22

scandens 18, 22

Helosis 94, 95, 97

cayennensis 94, 95, 97, 98

var. cayennensis 98

var. mexicana 97, 98

guyanensis 97

mexicana 97, 98

Hierba de gato 176

Hilleria 203

subcordata 203

Holosteum cordatum 236

diandrum 236

Homalocladum platydadum 121

Hongo 98

Hormigo 138

Howardia costaricensis 85

hoffmanni 84

Hydnora 87

Hydnoraceae 87, 88

Inmortal 169

Insect-plant mutualism 138

Iresine 170, 171

acicullaris 176

angustifolia 171, 172

argenta 173

arrecta 171, 172

calea 171, 173

celosia 175

celosoioides 175

costaricensis 171, 174

diffusa 171, 175

elatior 172

frutescence 176

grandiflora 176

hassleriana 175

latifolia 173

paniculata 175

spiculigera 176

Jaboncillo 206

Jamesbondia costaricensis 151

Lads schiedeanum 6

foeniculacea 4

Langsdorffia 94, 98

hypogaea 94, 99

Lechugon 134

Lengua de caballo 134

Lengua de vaca 134

Lewisia 218

megarhiza 218

Llovizna 238, 242

Lora malodora 13

Loranthaceae (sensu lato) 29,. 30, 33,

34- 37, 57-63

Loranthaceae (sensu stricto) 33, 34- 37

Loranthus aduncus 45

alveolatus 41

amplexicaulis 41

angustifolius 69

calyculatus 47

cansjeraefolius 56

chordocladus 55

leptostachyus 52

marginatus 53

nodosus 47

occidentalis 42

oerstedii 54

orbicular is 55

paniculatus 45

piperoides 75

punctatus 39

pyrifolius 44

auadrangularis 75

auercicola 56

252

FIELDIANA: BOTANY

Loranthus ramiflorus 48

retroflexus 45

schiedeanus 49

spicatus 43

theobromae 45

Lustrosa 189

Lychnis 238

coronaria 238

Manglillo 17

Mono de lebn 162

Manto de Jesus 189

Mann 24

Marathrum 2, 3

allenii 5

cheiriferum 4, 5

flexuosum 6

foeniculaceum 4

indifferens 5

leptophyllum 5

minutiflorum 5

forma allenii 5

forma diversifolium 5

forma indifferens 5

forma intermedium 5

oxycarpum 5

pusillum 5

schiedeanum 2, 6

tenue 2, 6

utile 2, 7

Maravilla 192

Medicinal uses 82, 134, 140-141

Melanocarpum sprucei 179

Mesembryanthemum 213

Microtea 200, 204

debilis 200, 204

Minquartia 15, 23

guianensis 15, 23

Mirabilis 182, 191

jalapa 182, 192

violaceae 182, 192

Mitrastemma 93

Mitrastemon 90, 93

matudai 93

Mitrostemonaceae 93

Mollugo 214, 225

radiata 214

tetraphylla 239

verticellata 215, 225

Mono 162

Mono de reina 162

Monte negro 148

Mosotillo 145

Mozote 145

Muehlenbeckia 104, 121

platyclada 121

tamnifolia 104, 122

volcanica 123

Mulato 138

Napoleon 189

Neea 185, 186, 193

acuminatissima 195

amplifolia 186, 194, 195

belizensis 195

delicatula 185, 196

elegans 186, 194, 195

latevirens 185, 195, 1%

orosiana 194

pittieri 186, 195

popenoei 185, 195, 196

psychotrioides 185, 195, 1%

pycnantha 195

urophylla 194

xanthina 195

Negro 24

Neolacis myriophylla 6

New Zealand Spinach 213

Nispero 24

Notanthera 29

Nyctaginaceae 180, 182-186

Olacaceae 14, 15, 18

Opiliaceae 15, 27

Oreobroma megarhiza 218

mexicanum 218

Oryctanthus 33, 37, 40, 41

alreolatus 37, 41

botryostachys 41

cordifolius 37, 42

florulentus 43

guatemalensis 43

occidentalis 37, 42

spicatus 37, 43

Oxybaphus 191

violaceus 192

Palo de papa 11

Panopsis 9, 10

costaricensis 11

suaveolens 10, 11

Papa 11

Papaturro 112, 120, 121

Papaturro Blanco 116

Papaturro de Playa 121

Papaturro Negro Macho 121

Papaturro Rastero 121

Pasacarne 3

Perico 148

Peristethium leptostachyum 52

Petiveria 202, 205

alliacea 202, 205

Pfaffia 152, 176

grandiflora 152, 176

hooker iana 176

forma glabriuscula 176

Philoxerus 177

vermicularis 177

BURGER: FLORA COSTARICENSIS

253

Phoradendron 61, 62, 63, 66, 67

acinacifolium 63, 68

a/feni'i 64

angustifolium 69

annulatum 62, 69

auriculatum 72

biolleyi 75

ceibanum 75

conzattii 70

cooperi 77

corynarthron 61, 70, 76

var. seibertii 75

crassifolium 63, 71

var. pittieri 71

crispum 64

davidsoniae 70

dichotomum 62, 71

dipterum 62, 72

dodge/ 76, 77

ensifolium 69

falcifolium 76

flavens 63, 73

flavescens 64

gracilispicum 78

herrerense 75

mucronatum 73

navae-helvetiae 77

obliquum 63, 74, 77

paquitanum 75

pergranulatum 76

piperoides 63, 75

quadrangulare 61, 75

auinquenervium 73

rensonii 75

robaloense 68

robustissimum 63, 76

seibertii 75

sonarum 75

sp. aff. undulatum 62

sauamigerum 65

supravenulosum 73

tonduzii 62, 77

undulatum 62, 78

venezuelense 75

woodsonii 73

Phrygilanthus 33

panamensis 55

Phthirusa 33, 35, 44

adunca 45

paniculata 45

pittieri 52

pyrifolia 35, 44

retroflexa 35, 45

Phytolacca 201, 206

costaricensis 208

icosandra 201, 207

meziana 206

octandra 206

purpurascens 206

rivinoides 201, 206, 207

rugosa 201, 208

Phytolaccaceae 199, 200-202, 203

Pilostyles 90, 93

covillei 93

globosa 93

thurberi 93

Pisonia 184, 1%

aculeata 184, 197

var. macranthocarpa 197

fasciculata 197

grandifolia 197

macranthocarpa 197

sylvatica 184, 198

Pleuropetalum 163, 178

calospermum 179

costaricensis 179

pleiogynum 163, 178

sprucei 163, 179

standleyi 178

tucurriquense 179

Podopterus 105, 123

mexicanus 105, 123

Podostemaceae 1

Polycarpaea corymbosa 243

Polycarpon 238

tetraphyllum 239

Polygonaceae 99, 102-105, 108-111

Polygonum 102, 124

acuminatum 102, 125

aubertii 106, 107

aviculare 126

baldschuanicum 106

grandiflorum 101

guatemalense 125

hispidum 102, 126

hydropiper 129

hydropiperoides 102, 127

meisnerianum 102, 128

mexicanum 129

pennsylvanicum 130

persicaria 128

persicarioides 128

platycladum 121

punctatum 102, 128, 129

segetum 102, 129

tamnifolium 122

uvifera 119

volcanicum 123

Pomonce 189

Pompilla 189

Portulaca 218, 225

coronata 220

denudata 220

grandiflora 219

lanceolata 220

marginata2l9

oleracea 219, 223

paniculata 221

patens 221

pilosa 220, 225

plano-operculata 220

portulacastrum 215

254

FIELDIANA: BOTANY

Portulaca umbraticola 220

Portulacaceae 217, 223-225

Primavera 189

Prosopanche 87, 88

americana 89

costaricensis 88, 89

Protea 8

Proteaceae 8

Psittacanthus 34, 46

allenii 48

calyculatus 34, 47

chrismarii 47

dilatatus 48, 49

lateriflorus 48

mayanus 47

nodosus 34, 47

ramiflorus 34, 48

scheryi 34, 48

schiedeanus 34, 49

Purselane 220

Rabo de Chanco 145

Rabo de gato 162

Rafflesiaceae 89, 90, 92

Rainbow Pinks 235

Remolacha 139

Rhaptostylum acuminatum 19

Rheum 130

rhabarbarum 130

rhaponticum 130

Rhubarb 130

Ribbon Bush 122

Rivina 200, 208

humilis 200, 209

octandra 211

polyandra 212

Rosas de pah 50

Rose Campion 238

Roupala 10, 11

borealis 13

complicata 13

glaberrima 10, 12

loranthoides 10, 13

montana 10, 13

var. dentata 14

panamensis 13

Ruibarbillo 132

Ruibarbo 130

Rumex 103, 131

acetosella 103, 132

costaricensis 103, 132

crispus 103, 133

peruanus 133

tolimensis 133

Ruprechtia 105, 134-135

costata 105, 135

cumingii 136

deamii 135

kellermanii 135

nepalensis 103, 134

Ruprechtia obtusifolius 103, 134

pringlei 136

Sagina 225, 239

procumbens 225, 240

Salpianthus 183, 198

purpurascens 183, 199

San Andres 106

San Diego 169

San Miguel 106

Santalaceae 14

Santalales (families 56-58) 14-79

Santa Rita 189

Saponaria 240

officinalis 240

Schoepfia 15, 24

americana 25

arborescens 25

obliquifolia 26

schreberi 15, 25

species A 15, 26

vacciniiflora 15, 25

Scleranthus 225, 241

annuus 225, 241

Sea Grape 120

Sempre Lustrosa 189

Sesuvium 215, 223

portulacastrum 215, 223

scandens 210

Siempreviva 169

Siete pellejos 174, 176

Silene 241

anglica 242

armeria 242

gallica 242

Soapwort 241

Solitaria 122

Spergula 225, 243

arvensis 225, 243

Spergulastrum lanuginosum 229

Spinacea oleracea 142

Spinach 141, 142

Spinach-like vegetables 217, 220-222

Spinacia 141

Stegnosperma 202, 210

cubense 202, 210

scandens 210

Stellaria 226, 243

ciliata 244, 245

cuspidata 245

irazuensis 245

limitanea 244

nemorum 245

nubigena 246

ovata 226, 246

prostrata 247

venezuelana 246

Struthanthus 35, 36, 50

aff. dichotrianthus 57

belizensis 55

BURGER: FLORA COSTARICENSIS

255

Struthanthus cansjeraefolius 56

chordocladus 55

costaricensis 36, 52

guatemalensis 43

leptostachyus 35, 42

macrostachyus 58

marginatus 36, 53, 57

oerstedii 36, 54

oliganthus 39

orbicularis 36, 55

panamensis 33, 35, 55

papillosus 57

polystachyus 52

quercicola 36, 56, 57

retroflexus 45

rotundatus 36, 57

woodsonii 35, 58

Sweet William 234

Sweet William Catchfly 242

Trichilia scandens 210

Trichostigma 200, 211

octandrum 200, 211

polyandrum 200, 211

Trinitaria 189

Triplaris 108, 136

cumingiana 137, 138

macombii 137

melaenodendron 105, 137

Tristicha 2, 7

hypnoides 7

Tsu-Kra 27

Turi-Svan-Kra 138

Ullucus tuberosa 222

Una de Tigre 198

Uva 120

Uva de Playa 120

Tabaco 138

Tabaco de Monte 138

Tabacon 127, 138

Taba de gueguecho 176

Tacuquelite 174

Talinum 221, 224

paniculatum 221, 224

patens 221

triangulate 221, 222

Tandonia ramosa 222

Tapeworm Plant 122

Telanthera bettzichiana 148

Tenia 122

Terciopelo 162

Tetragonia expansa 213

Thonningia 98

Tinta 206

Tobaquillo 127

Torrubia 190

costaricana 190

Tres Marias 189

Trianthema 216, 223

portulacastrum 216, 223

Vegetables 134, 141, 142, 156, 217, 220, 221

Velasquezia melaenodendron 137

Velo de gueguecho 176

Velo de princesa 176

Veranera 189

Verdolaga 217, 220, 221

Viscaceae 57-63

Viscum angustifolium 69

cordifolium 42

crassifolium 71

dichotomum 71

ensifolium 69

/toens 73

mucronatum 73

squamigerum 65

Vomi'ta 203

Ximenez, Francisco 27

Ximenia 15, 26

americana 15, 27

Families of seed plants known or expected to occur in Costa Rica and adjacent ai

numbered according to the sequence oiEngler'sSv//a6u,s der Pflanzenfamuien, edi

11, reworked by L. Diels (1936).

areas

tion

1 Cy cad no

3 Podocarp

iceae

5 Pinaceae

6 Cupressaceae

7 Cm-:

8 Typhaceae

• mogetonaceae

10 Najadaceae

11 A!

12 Butom.i'

13 Hydroch;u

14 Triuridaceae

15 Gramineae

16 Cypenu

ilmae

* lanthaceae

19 Araceae

20 Lemnaceae

iceae

23 Eriocaulaceae

24 Bromeliaceae

25 Commelinaceae

26 Pontederiaceae

27 Juncaceae

28 Liliaceae

29 Haemodoraceae

30 Amaryllidaceae

.•lloziaceae

32 Dioscoreaceae

33 Indaceae

34 Musaceae

35 Zingiberaceae

36 Cannaceae

37 Marantaceae

38 Burmanniaceae

39 Orchidaceae

40 Casuarinaceae

41 Piperaceae

42 Chloranthaceae

43 Lacistemaceae

44 Salicaceae

45 Garryaceae

46 Myncaeeae

47 Juglandaceae

48 Batidaceae

49 Betulaceae

50 Fagaceae

51 Ulmaceae

52 Moraceae

53 Urticaceae

54 Podostemonaceae

55 Proteaceae

56 Olacaceae

57 Opiliaceae

58 Loranthaceae

59 Aristolochiaceae

60a Hydnoraceae

60b Rafflesiaceae

61 Balanophoraceae

62 Polygonaceae

63 Chenopodiaceae

64 Amaranthaceae

tagmaceae

66 Phytolaccaceae

67 Aizoaceae

68 Portulacaceae

69 Basellaceae

70 Caryophyllaceae

72 Ceratoph -

73 Ranunculaceae

74 Berberidaceae

75 Menispermaceae

tgnoliaceae

\nonaceae

82 P>

incl t

iceae

91 Brunei!:

92 Cunoni.

uelidaceae

94 Re

95 Connaraceae

96 I.egumn.

97 Kramenaceae

98 Oxalidaceae

99 Geraniaceae

i opaeolaceae

101 Linaceae,

incl. Humiriaceae

102 Erythroxylaceae

103 Zygophyllaceae

104 Rutaceae

105 Simarubaceae

106 Burseraceae

107 Meliaceae

108 Malpighiaceae

109 Trigoniaceae

110 Vochysiaceae

111 Polygalaceae

112 Dichapetalaceae

113 Euphorbiaceae

114 Callitrichaceae

115 Buxaceae

116 Coriariaceae

117 Anacardiaceae

118 Cyrillaceae

119 Aquifoliaceae

120 Celastraceae

121 Hippocrateaceae

122 Staphyleaceae

loacinaceae

124 Hippocastanaceae

125 Sapindaceae

126 Sabiaceae

127 Balsaminaceae

128 Rhamnaceae

129 Vitaceae

130 Elaeocarpaceae

131 Tiliaceae

132 Malvaceae

•nmbacaceae

't-rculiaceae

135 Dilleniaceae

136 Actinidiaceae

137 Ochnaceae

138 Caryocaraceae

139 Marcgraviaceae

140 Quiinaceae

141 Theaceae

142 Guttiferae,

incl. Hypericaceae

143 Elatinaceae

i-ae

\aceae

146 Cochlospermaceae

Violaceae

149 Turner:

150 Passifloi

ceae

152 Loasaceae

154 Cactiiceae

155 Thymelaeaceae

156 Elaeagnaceae

157 Lythraceae

.ceae

163 Melastomataceae

164 Onagraceae

165 Halorrhagaceae

eae

167 Umbelliferae

168 Cornaceae

169 Clethraceae

'lotropaceae

171 Pyrolaceae

172 Ericaceae

i-ophrastaceae

174 Myrsinaceae

175 Primulaceae

176 Plumbaginaceae

177 Sapotaceae

178 Ebenaceae

mplocaceae

180 Styracaceae

181 Oleaceae

182 Lopaniaceae

••ntianaceae

184 Apocynaceae

. lepiadaceae

186 Convolvulaceae

187 Polemoniaceae

188 Hydrophyllaceae

189 Boraginaceae

190 Verbenaceae

191 Labiatae

192 Solanaceae

193 Scrophulariaceae

194 Bignoniaceae

195 Pedal iaceae

196 Martyniaceae

197 Orobanchaceae

198 Gesneriaceae

199 Lentibulariaceae

200 Acanthaceae

201 Plantaginaceae

202 Rubiaceae

203 Caprifoliaceae

204 Valerianaceae

205 Dipsacaceae

206 Cucur'oitaceae

207 Campanulaceae

208 Compositae

UNIVERSITY OF ILLINOIS-URBANA

30112028176011

Field Museum of Natural History

Roosevelt Road at Lake Shore Drive

Chicago, Illinois 60605-2496

Telephone: (3 1 2) 922-9410

UNIVERSITY OF ILLINOIS-URBANA