Flora of Guatemala

UNIVERSITY OF

ILLINOIS LIBRARY

AT URBANA-CHAMPAIGN

BIOLOGY

3 1 1984

' 3 --,'

•IS

•y
'

fo, 1-3.

FLORA OF GUATEMALA

PAUL C. STANDLEY

AND

LOUIS 0. WILLIAMS

MYRS1NACKAE ^

CYKUS LONGWORTH LUNDELL

The Library of the

JAN 1 8 1979

FIELDIANA: BOTANY
VOLUME 24, PART VIII, NUMBERS 1 AND 2

Published by

CHICAGO NATURAL HISTORY MUSEUM
FEBRUARY 24, 1966

FLORA OF GUATEMALA

PART VIII

FLORA OF GUATEMALA

PAUL C. STANDLEY

The Late Curator of the Herbarium

LOUIS 0. WILLIAMS

Chief Curator, Botany, Chicago Natural History Museum

MYRSINACEAE

by

CYRUS LONGWORTH LUNDELL

Director and Chief Research Scientist
Texas Research Foundation

FIELDIANA: BOTANY

VOLUME 24, PART VIII, NUMBERS 1 AND 2

Published by

CHICAGO NATURAL HISTORY MUSEUM
FEBRUARY 24, 1966

In Memory of
JOHN DONNELL SMITH

1829 - 1928

Who, with reason, may be said to be
the Father of Central American Botany

Library of Congress Catalog Card Number: 48-3076

PRINTED IN THE UNITED STATES OF AMERICA
BY CHICAGO NATURAL HISTORY MUSEUM PRESS

-FB

V •'

Araliaceae
Umbelliferae

CONTENTS

Families Included in Part VIII, Number 1

PAGE

1
21

Cornaceae .......... 67

Garryaceae ......... 70

Families Included in Part VIII, Number 2

Clethraceae ......... 74

Pyrolaceae .......... 81

Monotropaceae ........ 86

Ericaceae .......... 88

Theophrastaceae ....... 127

Myrsinaceae ......... 135

Primulaceae ......... 200

Plumbaginaceae ....... 207

ILLUSTRATIONS

FIGURE PAGE

1. Aralia humilis 3

2. Dendropanax arboreus 6

3. Oreopanax lacnocephalus 15

4. Sciadodendron excelsum 20

5. Arracacia annulata 28

6. Arracacia bracteata 30

7. Centella asiatica 33

8. Coaxana purpurea 35

9. Donnellsmithia guatemalensis 40

10. Enantiophylla heydeana 42

11. Eryngium cymosum 45

12. Hydrocotyle mexicana 51

13. Micropleura renifolia 54

14. Myrrhidendron donnellsmithii 55

15. Spananthe paniculata 63

16. Tauschia steyermarkii 65

17. Cornus disciflora 68

18. Garrya lauriflora 71

19. Clethra mexicana 77

20. Clethra oleoides 78

21. Chimophila maculata 82

22. Monotropa coccinea 87

23. Arbutus xalapensis 90

24. Arctostaphylos cratericola 93

25. Befaria guatemalensis 96

26. Cavendishia guatemalensis 100

27. Disterigma humboldtii 102

28. Empedoclesia brachysiphon 103

29. Gaultheria odorata 107

30. Leucothoe mexicana 109

31. Macleania insignis Ill

32. Pernettya ciliata 113

33. Satyria meiantha 117

34. Sphyrospermum majus 119

vii

35. Vaccinum lundellianum 124

36. Deherainia smaragdina 129

37. Jaquinia aurantiaca 131

38. Ardisia paschalis 148

39. Ardisia Tuerkheimii 154

40. Ardisia verapazensis 155

41. Gentlea tacanensis 158

42. Gentlea Vatteri 159

43. Parathesis columnaris 167

44. Parathesis cubana 169

45. Parathesis membranacea 174

46. Parathesis oblanceolata 176

47. Parathesis papillosa 177

48. Parathesis rufa 180

49. Parathesis sessilifolia 181

50. Parathesis Skutchii 183

51. Parathesis subcoriacea 185

52. Parathesis tomentosa 188

53. Rapanea myricoides 190

54. Stylogyne laevis 194

55. Synardisia venosa 196

56. Yunckeria amplifolia 199

57. Anagallis arvensis 202

58. Centunculus minimus 204

59. Lysimachia steyermarkii 205

60. Samolus ebracteatus 206

61. Plumbago scandens 210

Vlll

Flora of Guatemala ' Part VIII, Number 1

UMBELLIFLORAE

The Umbelliflorae were treated in Engler & Prantl's Natiirliche
Pflanzenfamilien, by Harms, as three families — Araliaceae, Umbel-
liferae and Cornaceae. The order is characterized by the tendency
for the flowers to be borne in determinate umbels, either simple or
compound (or sometimes apparently in heads) ; by the reduction and
simplification in the floral structure; and by epigyny. The carpels
are reduced to two. — The Araliaceae are mostly woody and tropical ;
the Umbelliferae are mostly herbaceous, with a few suffrutescent
members, widely distributed over the world but most abundant in
temperate regions of the northern hemisphere. The Cornaceae, the
smallest family of the order, is made up mostly of trees or shrubs,
with occasional small suffrutescent plants. It is most abundant in
temperate regions of Asia and North America but extends into the
southern hemisphere.

Many plants of economic importance are found in the order.
Food and condiment plants are abundant, especially in the Umbel-
liferae. However, none are of great importance as crops in Guate-
mala or in Central America. Trees are common in the Araliaceae
but the wood is of poor quality and little, if at all, used in our region.
A few ornamentals of this order, mainly Araliaceae, are found in
Guatemala.

ARALIACEAE1

Reference: Smith, Albert C., Araliaceae, North American Flora
28B:3-41. 1944.

Trees or shrubs, sometimes woody vines, rarely herbs, sometimes epiphytic,
glabrous or usually pubescent, the pubescence often of branched hairs; leaves
usually (ours) alternate, simple, lobed, palmately lobed, pinnatifid or ternately
compound or decompound; stipules adnate to the petioles, connate and produced
into a coriaceous ligulate sheath, or hardly distinct or none; flowers in umbels or

1 The field work and travel by the junior author involved in the preparation of
this number of the Flora of Guatemala has been made possible through a grant from
National Science Foundation, which we are happy to acknowledge.

2 FIELDIANA: BOTANY, VOLUME 24

heads and in solitary racemose, corymbose, paniculate, or usually umbellate in-
florescences; bracts subtending the branches of the inflorescence small, usually
soon deciduous; pedicels continuous or articulate; flowers perfect or unisexual,
usually small and greenish; calyx tube cupular or annular, adnate to the ovary,
erect or spreading, sometimes wanting; petals 3-10, generally 5, valvate or imbri-
cate, deciduous, sometimes connate and calyptriform; stamens as many as the petals
and alternate with them, rarely more; anther ovate or oblong or subglobose, dorsi-
fixed, straight or recurved, didymous, the cells longitudinally dehiscent; disc epigy-
nous, explanate or conic, the margin entire or undulate; ovary inferior, 1-many-
locular; the styles as many as the locules, erect or recurved, sometimes connate;
the stigmas terminal or decurrent on the inner surface; ovules solitary in the cells,
pendulous from the apex of the cell, anatropous; fruits baccate or usually drupa-
ceous, the exocarp usually fleshy, the endocarp divided into 1-many distinct, osse-
ous, crustaceous, cartilaginous, or rarely membranaceous nutlets; seeds laterally
compressed or triquetrous in cross-section, the testa thin; endosperm copious,
cartilaginous or carnose; embryo minute, the cotyledons ovate or oblong, the rad-
icle superior.

A family of about 70 genera (nine in North America according to
Smith, loc. cit.) and 600-800 species. Widely distributed in tropical
regions, a few in temperate areas. One other genus, Schefflera, with
several species is in southern Central America and Panama. The
species in Central America are mostly in the mountains.

Leaves pinnately or ternately compounded or decompounded.

Petals in bud valvate; cultivated plants rarely flowering Nothopanax.

Petals in bud imbricated; native plants usually flowering.

Flowers mostly 5- (4-6)-merous Aralia.

Flowers 7-12-merous Sciadodendron.

Leaves simple, lobate or palmately compounded.

Woody vines with lobate leaves; cultivated, rarely flowering Hedera.

Trees or shrubs with simple or palmately compounded leaves; native plants

usually flowering.

Flowers in heads, sessile or short pedicellate, polygamodioecious; petiole with-
out an obvious ligule Oreopanax.

Flowers in umbels, perfect or polygamomonoecious.

Styles and locules 2 or 3 Didymopanax.

Styles and locules 5-9, usually 5 Dendropanax.

ARALIA [Tournef.] L.

Perennial herbs or (ours) shrubs or small trees, glabrous or pubescent; leaves
alternate, pinnately or ternately compounded or decompounded, the leaflets ser-
rated; inflorescence with two or more umbels, radiating or laxly corymbose or in
large compound racemes, the bracts small; the pedicels articulate with the calyx
and the apex often tumid; flowers polygamomonoecious, 5-6-merous, glabrous;
the calyx obconic or cupuliform, truncate, repand, lobed or minutely dentate;
petals imbricated, white or green, obtuse or inflexed-acuminate; anthers mostly
oblong, straight; disc almost flat, fleshy; styles 4-6, connate at their bases, free
above and/or even to the base; fruit baccate, often laterally compressed, nutlets
orbicular to oblong, compressed, crustaceous or harder, as many as the styles.

FIG. 1. Aralia humilis. A, Branch; >£ natural size. B, Fruit; about X 5.
C, Flower; about X 5, from an unpublished Sesse and Mocino drawing.

4 FIELDIANA: BOTANY, VOLUME 24

A genus most common in temperate regions in America, only the
following is found in Central America. There are perhaps 30-40
species, mostly in eastern Asia.

Aralia humilis Cav. Icon. PI. 4: 7, t. 313. 1797. Tacamajaca
(Jalapa); sombrillero (fide Aguilar).

Moist or dry, brushy slopes, often in pine-oak forest, sometimes
on limestone, 500-1,900 meters; Zacapa; Chiquimula; Jalapa; Juti-
apa; Sacatepe"quez; Quiche"; Huehuetenango. Southern New Mexico
and Arizona; Mexico; Honduras.

An unarmed shrub or small tree, commonly 1-3 meters high, sometimes a tree
of 5 meters with a trunk 30 cm. in diameter, the branches brown, terete, pilose with
coarse, simple or sparsely branched hairs; leaves large, pinnate or bipinnate, decid-
uous, long-petiolate, leaflets numerous, sessile or short-petiolulate, membrana-
ceous, oblong-ovate to broadly ovate, 2-8 cm. long, acuminate, rounded or shallowly
cordate at the base, serrate or crenate, sparsely or densely pilose with coarse,
simple or sparsely branched, white hairs, often glabrate on the upper surface; um-
bels few or often numerous, usually paniculate, glabrous or somewhat pubescent,
the flowers 5-merous, numerous, greenish, on very slender but stiff pedicels about
2 cm. long; petals 2.5-3 mm. long; fruit black-purple, 3-7 mm. in diameter.

This is not a common plant of Guatemala, but it is plentiful
locally in the drier regions, most often in forests or scrubby thickets
of Quercus.

EXCLUDED SPECIES: Aralia sololensis Donn.-Sm. Bot. Gaz. 56:
58. 1913. = Sambucus sp.

DENDROPANAX Decaisne & Planchon

Shrubs or trees, glabrous; leaves simple, petiolate, entire or often lobate, often
highly variable on the same plant or even branch; inflorescence of a lateral or ter-
minal umbel or occasionally many umbels (or heads) in racemes or umbels; pedun-
cles subtended by small bracts, sometimes bracteate and articulate above the base,
often swollen distally into a fleshy receptacle; flowers 5-9-merous, perfect or polyga-
momonoecious; calyx obconic or cupuliform, usually dentate; petals usually 5 (6-8),
cucullate, valvate; stamens as many as the petals, inflexed; disc fleshy, short-conic,
confluent with the styles; the styles nearly free or usually connate to the middle
or beyond, the stigmas sessile or short stipitate; fruit globose or ovoid, usually
5-sulcate when dry, black or dark purple, nutlets compressed or subtrigonous.

Perhaps 50 species in tropical America and Asia, — a few addi-
tional ones are in southern Central America and Panama. The wood
of this genus is grayish, light in weight but firm and tough, and easily
worked. In consistency it is similar to Liriodendron. The wood is
not used in Central America, so far as we know. Many authors have

STANDLEY AND WILLIAMS: FLORA OF GUATEMALA 5

used the name Gilibertia Ruiz & Pavon for this genus. For a discus-
sion of the use of the generic name see A. C. Smith in Tropical Woods
66: 1. 1941.

Umbels 3-20 per inflorescence.

Inflorescence slender, the peduncles (3-) 5-7 cm. long, the pedicels 10-20 mm.
long D. leptopodus.

Inflorescence comparatively compact, peduncles 1-5- (7) cm. long, the pedicels
to 8- (12) mm. long.

Peduncles striate or smooth, bracts, if present, rarely connate; common species.

D. arbor eus.

Peduncles rugose, bracts usually connate into a small coriaceous cupule.

D. oliganthus.

Umbels 1 per inflorescence D. schippii.

(See also rare forms of D. arboreus).

Dendropanax arboreus (L.) Dene. & Planch. Rev. Hort. 4, 3:
107. 1854; Smith, No. Am. Fl. 28B: 16. 1944. Aralia arborea L.
Syst. Nat. ed. 10. 967. 1759. Gilibertia arborea Marchal, Bull. Soc.
Bot. Belg. 30: 281. 1891. G. stenocarpa Donn.-Sm. Bot. Gaz. 55:
435. 1913 (type from Barranco Eminencia, Guatemala, J. D. Smith
2666}. G. Smithiana I. M. Johnston, Contr. Gray Herb. 70: 81.
1924 (type from Cubilgiiitz, Alta Verapaz, Tuerckheim II. 407).
G. concinna Standl. Trop. Woods 18: 30. 1929 (type from Honey
Camp, British Honduras, Lundell 115}. D. concinnus Lundell, Field
& Lab. 13: 11. 1945. Mano de ledn; sac-co (Alta Verapaz); chonte
(Quezaltenango) ; mazorco (Suchitepe'quez) ; matapalo; mano de mico;
xup (Pete"n, fide Lundell).

Moist or wet, usually mixed forest or thickets, often in second
growth, sometimes in hedges, 1,500 meters or lower, or very rarely
at higher elevations; Pete"n; Alta Verapaz; Baja Verapaz; Izabal;
Zacapa; Chiquimula; Jutiapa; Santa Rosa; Escuintla; Guatemala;
Suchitepe'quez; Solola; Retalhuleu; Quezaltenango; San Marcos;
Huehuetenango; Quiche*. Mexico; British Honduras to El Salvador
and Panama. West Indies; South America.

A glabrous tree, rather sparsely branched, sometimes 20 meters high but often
only a shrub, the crown spreading, the trunk as much as 60 cm. in diameter, the
bark medium brown or grayish brown, slightly scaly or almost smooth; leaves on
long or short petioles, chartaceous, very variable in size and outline, those on
sterile or juvenile branches often deeply trilobate, those of older or fertile branches
ovate to elliptic or obovate and entire or undulate; umbels of flowers usually numer-
ous (or rarely solitary) and disposed in a short or elongate raceme, the peduncles
short or elongate; flowers few or numerous in the umbel, yellowish or greenish
white, the pedicels mostly 5-10 mm. long; fruit black or dark purple, very juicy,
6-8 mm. in diameter.

6

FIELDIANA: BOTANY, VOLUME 24

Known in British Honduras by the names "white gumbolimbo,"
"potatowood," and "sac-chacah" (Maya); "palo de agua" (Hon-
duras); "tamalcoahuite" (Veracruz); "mano de sapo" (Veracruz);
"palo bianco" (Veracruz). This is a very common small tree of
Guatemala and of many other parts of Central America, often spring-
ing up in abundance in second growth. The tree has a wide range
and exhibits great variation, not only in foliage but in characters of
inflorescence. It seems difficult, if not impossible, to coordinate or
to limit definitely these variations, with the result that there have

FIG. 2. Dendropanax arboreus.
fruit, X 3.

A, Flowering branch; X

B, Immature

STANDLEY AND WILLIAMS: FLORA OF GUATEMALA 7

accumulated in recent or earlier years many names that apparently
must be relegated to synonymy. It is, of course, possible that fur-
ther collection may make it possible to divide the material now re-
ferred to D. arboreus, but it seems more probable that the result will
be further reductions.

Dendropanax leptopodus (Donn.-Sm.) A. C. Smith, Trop.
Woods 66: 3. 1941. Gilibertia leptopoda Donn.-Sm. Bot. Gaz. 57:
421. 1914.

Moist or wet, mixed, mountain forest, 950-2,500 meters; Izabal;
Alta Verapaz (type from Coban, Tuerckheim 1^166} ; El Progreso;
Izabal; Zacapa; Chiquimula; Suchitepe"quez; Solola; Quezaltenango.
Mexico (Chiapas) ; Honduras.

A glabrous shrub or tree, sometimes 9 meters high, with slender branches;
leaves on long or short, slender petioles, chartaceous or membranaceous, lanceolate
or elliptic-lanceolate, mostly 7-14 cm. long, acuminate or narrowly long-acuminate,
obtuse or acute at the base, slightly paler beneath; umbels terminal or lateral, the
peduncles very slender, often exceeding the leaves, 11 cm. long or shorter, naked
or usually with a few very inconspicuous bracts near the base; flowers pale green,
few or rather numerous, the pedicels very slender, mostly 10-20 mm. long, some-
times shorter in early anthesis; petals 1.5 mm. long, apiculate; fruits about 5 mm.
long, purple or almost black.

One specimen referred to here has deeply trilobate leaves similar
to those of D. arboreus, but generally all the leaves are entire, while
in D. arboreus trilobate leaves sometimes are found even on the flow-
ering branches.

Dendropanax oliganthus A. C. Smith, Trop. Woods 66: 3. 1941.
Gilibertia oligantha A. C. Smith, Brittonia 2: 251. 1936.

Moist or wet, mixed forest, 2,100-2,400 meters, or at lower ele-
vations; Alta Verapaz (type from Finca Sepacuite", 0. F. Cook &
R. F. Griggs 98); Zacapa (Sierra de las Minas).

A glabrous tree of 15 meters, the branches very thick; leaves coriaceous, on
stout petioles 1.5-6 cm. long, oblong or obovate-oblong, 9-15 cm. long, 3-6 cm.
broad, short-acuminate to obtuse, acute or obtuse at the base, entire or nearly so,
the veins prominent and reticulate beneath; umbels many-flowered, on very thick
peduncles 2-2.5 cm. long, these fasciculate or very shortly racemose, conspicuously
bracteate below, articulate above the base; pedicels thick, 3-7 mm. long; flowers
7-9-parted (5-parted in other species); calyx 3-4 mm. broad, denticulate; petals
oblong-deltoid, 2.5-3 mm. long, subacute.

The leaves are pale green beneath when fresh; the petals creamy
white.

8 FIELDIANA: BOTANY, VOLUME 24

Dendropanax schippii A. C. Smith, Trop. Woods 66: 3. 1941.
Gilibertia schippii A. C. Smith, Brittonia 2: 252. 1936.

Known certainly only from the type, British Honduras, Sittee
River, in forest on river bank, at sea level, W. A. Schipp 61+1.

A glabrous shrub 1.5 meters high, the branches slender; leaves thick-membra-
naceous, on slender petioles 1-2.5 cm. long, oblong or obovate-oblong, 7-13 cm.
long, 3-5 cm. broad, obtuse or subacute, acute or narrowly cuneate at the base,
remotely callose-denticulate and somewhat undulate; umbels terminal, solitary,
the peduncles 1.5-2.5 cm. long, bracteate above the base, the pedicels in fruit
5-7 mm. long; flowers 10-20 in each umbel; fruit subglobose, 5-7 mm. in diameter.

This may be a form of D. arboreus with inflorescence reduced to
a solitary umbel.

DIDYMOPANAX Decaisne & Planchon

Mostly large trees, the inflorescence and the lower leaf surface generally cov-
ered with a whitish or brownish, minute, often lustrous tomentum; leaves large,
long-petiolate, digitately compound, the leaflets coriaceous, entire, long-petiolulate,
radiately spreading from the apex of the petiole and appearing verticillate; stipules
small, adnate to the petiole; flowers small, greenish, the umbellules usually form-
ing large panicles; bracts minute or none; pedicels not articulate at the apex; mar-
gin of the calyx repand-5-denticulate; petals 5, thick, valvate; stamens 5, the
filaments very short, the anthers ovate, large; disc explanate, the margin undu-
late; ovary 2-celled, the styles distinct, stout, erect, in age recurved-spreading,
stigmatose within near the apex; fruit strongly compressed, broadly subdidymous,
the exocarp thin, carnose; pyrenes strongly compressed laterally, cartilaginous or
hard.

About 20 species, in tropical America. One other Central Amer-
ican species grows in Costa Rica and Panama.

Didymopanax morototoni (Aubl.) Dene. & Planch. Rev. Hort.
4, 3: 109. 1854. Panax morototoni Aubl. Hist. PI. Guiane Fr. 2: 949,
L 360. 1775.

Wet mixed lowland forest at or little above sea level ; Pete"n ; Iza-
bal. Mexico (Oaxaca); British Honduras, along the Atlantic coast
(and southward both coasts) to Panama; South America.

A large tree, commonly 15-30 meters high, the trunk slender, about 30 cm.
in diameter or sometimes thicker, the bark pale gray or pinkish brown, somewhat
scaly, the trunk not buttressed; the crown often umbrella-shaped, rather small;
leaves very large, the petioles commonly 40-60 cm. long; leaflets 7-10, long-petio-
late, thick, elliptic or oblong, 20-40 cm. long, acuminate, rounded or often sub-
cordate at the base, green and glabrous above, densely pale-sericeous beneath,
juvenile leaflets often mucronate-serrate, hispid-pilose above; panicles usually very
large and much branched, with very numerous, small umbellules, the stout pedicels
2-6 mm. long; petals small, white; fruit glaucous, blackish at maturity, 6-7 mm.
broad.

STANDLEY AND WILLIAMS: FLORA OF GUATEMALA 9

Called "mountain trumpet" in British Honduras; known in Oax-
aca by the names "chancarro bianco," "roble bianco," and "mano
de danto."

This is one of the most common and characteristic trees of the
rain forest of the Atlantic lowlands of Central America, where it is
often the most abundant and conspicuous tree of the tallest stratum
of the understory forest. The trees may be recognized from the ground
because of their large size, striking form, and pale under surface of
leaves. They grow rapidly and reproduce abundantly. The wood
is grayish or pale brownish throughout, of medium luster, rather light
and soft to moderately heavy and hard, with a specific gravity of
about 0.50; its texture is medium to rather fine, the grain mostly
straight, easy to work, finishing smoothly; not resistant to decay.
In some regions the wood has been used for match sticks and boxes.

HEDERA L. Ivy

Woody vines, usually climbing by aerial roots; leaves simple, often lobate,
coriaceous; stipules none; umbels paniculate, the flowers polygamous, small, the
bracts minute or none, the pedicels not articulate; margin of the calyx entire or
nearly so; petals 5, valvate; stamens 5, the anthers ovate, straight or finally re-
curved; disc convex, thick-conic, the style columnar; ovary 5-celled, the stigmas
sessile at the ends of the style branches; fruit subglobose, juicy; nutlets ovoid-
triquetrous, membranaceous or chartaceous; seed ovoid, the endosperm ruminate.

Three species, or perhaps only one, in Europe and Asia. One is
cultivated for ornament in most temperate regions of the earth.

Hedera helix L. Sp. PI. 202. 1753. Hiedra (often written Yedrd) ;
siempre-verde.

Native of Europe, Asia, and northern Africa, often grown for or-
nament in other regions; planted occasionally in Guatemala, usually
creeping over walls; noted in Guatemala, Coban, Momostenango,
Huehuetenango, Quezaltenango, and elsewhere.

Plants high-climbing or sometimes creeping, with numerous aerial roots; leaves
thick and somewhat fleshy when fresh, mostly 3-5-lobate and about as broad as
long, dark green above, pale or yellowish green beneath, those of flowering branches
usually entire and ovate; inflorescence covered with whitish stellate hairs; fruit
juicy, usually black at maturity.

The Guatemalan climate, at least in the higher regions, seems well
suited to the growth of ivy, which usually grows luxuriantly in the
rather few places where it is planted.

10 FIELDIANA: BOTANY, VOLUME 24

NOTHOPANAX Seemann

Shrubs or small trees, glabrous or somewhat lanate; leaves digitately or pin-
nately compound, the leaflets usually membranaceous, entire, dentate, or incised;
flowers small, umbellate, usually polygamous, the umbels racemose or paniculate,
the bracts small and caducous or none; pedicels articulate below the flower; margin
of the calyx entire or 5-denticulate; petals 5, valvate; stamens 5, the anthers ovate
or oblong; ovary usually 2-celled, the styles distinct, erect or in age recurved, the
stigmas decurrent on the inner surface; fruit laterally compressed, the exocarp
fleshy; nutlets compressed, indurate, often sulcate or rugose.

About 15 species, in the southern hemisphere, mostly natives of
the South Pacific. Several of the species are widely grown for orna-
ment in tropical America, especially as hedge or patio plants. They
are common in the lowlands of Guatemala and other countries of
Central America. The shrubs are well adapted to hedge culture
since usually they are densely branched, always green, and stand
severe pruning. They attract attention because their leaves are often
variegated with white or cream. They and the species of Codiaeum
are almost equally popular for lowland hedges.

Leaves twice pinnate N. fruticosus.

Leaves once pinnate.

Leaflets 11-17, green N. ornatus.

Leaflets 5-7, usually margined or blotched with white N. guilfoylei.

Nothopanax fruticosus (L.) Miq. Fl. Ind. Bat. 1, pt. 1: 765.
1856. Panax fruticosum L. Sp. PL ed. 2: 1513. 1763.

Probably of Malayan origin, but now grown in most tropical
countries; commonly planted in Guatemala, chiefly in the lowlands,
where it and the other species are used for hedges; sometimes grown
at places as high as Guatemala.

All three species are alike except in the form of their leaves, and
the three are about equally common. In N. fruticosus the leaves are
twice pinnate, the leaflets few or numerous, mostly incised-dentate
and green throughout. The leaves often have a plume-like appear-
ance. The shrubs are common in the lowland parks and often are
seen in the cemeteries.

Nothopanax guilfoylei (Cogn. & March.) Merrill, Philip. Journ.
Sci. Bot. 7: 242. 1912. Aralia guilfoylei Cogn. & March. PL Orna-
ment. L 58. 1874.

Probably of Polynesian origin; frequent in cultivation in Guate-
mala.

STANDLEY AND WILLIAMS: FLORA OF GUATEMALA 11

Leaves once pinnate, the leaflets 5-7, large, long-petiolulate, elliptic or broadly
elliptic, obtuse or acute, deep green, usually edged with white, coarsely serrate.

Nothopanax ornatus (Bull.) Merrill, Philip. Journ. Sci. Bot. 7:
241. 1912. Panax ornatum Bull. Cat. 9. 1888.

Probably of Polynesian origin; frequently planted in Guatemala.

Leaves once pinnate, the leaflets 11-17, green, deeply pinnate-lobate, the leaves
often very large and long-petiolate.

Flowers seem to be infrequent on cultivated plants of this genus,
but they are found sometimes. They are so small and inconspicuous
that they are easily overlooked, and severe pruning probably re-
moves most of the flowering branches before the flowers open.

OREOPANAX Decaisne & Planchon

Shrubs or trees, frequently epiphytic, glabrous or tomentose, the pubescence
often of branched hairs; leaves long-petiolate, entire, lobate, or digitately com-
pound, the margins entire or dentate, the leaves often variable in form on sterile
and fertile branches, usually coriaceous; stipules none or small and inconspicuous;
inflorescence terminal or subterminal, bearing heads of flowers in panicles or ra-
cemes; flowers small, polygamodioecious, bracts scale-like, ternate; margin of the
calyx entire or nearly so; petals 4-7, generally 5, valvate; stamens as many as
the petals, short or somewhat elongate, the anthers ovate or oblong; disc expla-
nate; ovary usually 3-5-celled, the styles free or somewhat connate, usually decid-
uous, the stigmas terminal; fruit globose, often sulcate when dried, baccate, the
endocarp membranaceous or indistinct; seeds ovoid-sub triquetrous; endosperm
ruminate or almost simple.

About 80 species, in tropical America, chiefly in mountain re-
gions. A few others occur in southern Central America. Some of
the species constitute an important element of the wet mixed forest
of the mountains of Central and western Guatemala. Because of
their generally handsome, large, and conspicuous deep green leaves
they are of unusually attractive appearance. The often large in-
florescences also contribute to the beauty of the trees. The wood in
this genus is grayish, of medium luster and moderate density, or
occasionally very light and soft, of rather fine and uniform texture,
and straight-grained; its durability is low. So far as we know, no
use is made of the wood in Guatemala or other Central American
countries.

Leaves simple, often lobate.

Leaves all deeply lobate, often peltate, densely pubescent beneath with stellate
stipitate hairs, or sometimes glabrate.

12 FIELDIANA: BOTANY, VOLUME 24

Pistillate flower heads at anthesis about 1.5 cm. in diameter; lobes of the

leaves entire or nearly so O. lachnocephalus.

Pistillate heads at anthesis less than 1 cm. in diameter; lobes of the leaves

usually coarsely dentate or undulate-lobate O. peltatus.

Leaves entire, not lobate, at least on the fertile branches, the leaves of juvenile
or sterile branches frequently lobate, or the leaves of fertile branches rarely
shallowly lobate but the leaves then glabrous.

Leaves finely stellate-pubescent on both surfaces, often glabrate above but
usually rough to the touch O. arcanus.

Leaves usually glabrous, never rough to the touch on the upper surface.
Leaf blades lanceolate or lance-oblong, mostly 2.5-6 cm. broad, usually

more than 3 times as long as broad, acute or obtuse at the base.
Fruits about 4 mm. in diameter; nerves and veins usually conspicuous on

the lower surface of the leaf O. liebmannii.

Fruits 6-7 mm. in diameter; nerves inconspicuous on the lower leaf sur-
face, the veins obsolete O. steyermarkii.

Leaf blades suborbicular to broadly ovate or rounded-obovate, usually
much wider, generally less than twice as long as broad, rounded or
cordate at the base.

Leaves rounded or very obtuse at the apex O. obtusifolius.

Leaves acute or abruptly acuminate at the apex.

Leaves all entire O. capitatus.

Leaves, at least usually part of those on each branch, more or less

lobate, with very acute lobes O. sanderianus.

Leaves digitately compound.

Leaflets mostly 5-12 cm. broad or larger, with sparse or dense, stipitate,
branched hairs on the upper surface; fruiting heads about 4 cm. in
diameter O. echinops.

Leaflets mostly 2.5-4.5 cm. broad, glabrous on the upper surface or bearing
sessile stellate hairs; fruit heads usually less than 2 cm. in diameter.

Leaflets densely stellate-pubescent beneath, the hairs stipitate.

O. langlassei.

Leaflets glabrous beneath or the pubescence of minute sessile stellate hairs.

O. xalapensis.

Oreopanax arcanus A. C. Smith, No. Am. Fl. 28B: 38. 1944.

Moist or wet, mixed forest, 2,100-2,800 meters; endemic; Solola;
Huehuetenango (type from Cerro Pixpix, Steyermark 50645; collected
also above San Juan Ixcoy).

Terrestrial or epiphytic, sometimes 7.5 meters high, the branches thick, gla-
brous; leaves coriaceous, on petioles 4-10 cm. long, broadly oblong-ovate or ovate-
elliptic, mostly 11-16 cm. long and 6-9 cm. broad, acute or abruptly acute or
short-acuminate, rounded or even subcordate at the base, entire, minutely and
sparsely stellate-puberulent on the upper surface or glabrate, lustrous, slightly
paler beneath, rather densely and minutely stellate-puberulent or in age glabrate,
the nerves elevated and conspicuous; inflorescence (perhaps incompletely devel-
oped or abnormal) about 4 cm. long, open, stellate-pilose, the heads rather numer-
ous, long-pedunculate, the pistillate ones 2-3-flowered.

Oreopanax capitatus (Jacq.) Dene. & Planch. Rev. Hort. 108.
1854. Aralia capitata Jacq. Stirp. Am. 89, t. 61. 1763. Tronador;

STANDLEY AND WILLIAMS: FLORA OF GUATEMALA 13

palo de cohetillo; cohetillo; tapacopcd (Huehuetenango) ; c'ab-choh (chuh,
fide Seler) ; pata de Ie6n (fide Seler) .

Moist or wet forest, 2,800 meters or lower; Alta Verapaz; El Pro-
greso; Izabal; Guatemala; Suchitepe"quez; Solola; Quezaltenango;
San Marcos; Quiche"; Huehuetenango. Southern Mexico; Costa
Rica; Panama; West Indies; South America.

A terrestrial tree, or often an epiphytic shrub or tree, sometimes a tree of 15
meters but generally smaller, glabrous outside the inflorescence; petioles very un-
equal in length (as in most other species), sometimes very short, often much elon-
gated; leaf blades chartaceous or coriaceous, broadly ovate to elliptic or rounded,
mostly 10-25 cm. long, acute or abruptly acute or short-acuminate, rounded to
rather deeply cordate at the base, entire, the nerves conspicuous and elevated be-
neath; young leaves apparently not at all lobate; pistillate flower heads few-
flowered, pedunculate, forming dense or rather open, very stout racemes, these
mostly about 9 cm. long, the rachis more or less whitish-pubescent or in age
glabrate; fruits about 5 mm. in diameter, pale green or at maturity white or
nearly so.

With its wide distribution, this is a variable species. More than
that, some of the species treated as distinct, such as 0. sanderianus
and even 0. liebmannii, cannot be separated sharply from it, distinct
as their extremes appear to be. The entire-leaved Oreopanax species
of Central America are most unsatisfactory to treat on this account,
and it is far from certain how many specific units should be recog-
nized. Like other members of the genus, this plant often begins life
as an epiphyte but in the course of time becomes an independent
tree, after the host is killed and decayed.

Oreopanax echinops (Schlecht. & Cham.) Dene. & Planch. Rev.
Hort. 108. 1854. Aralia Echinops Schlecht. & Cham. Linnaea 5: 174.
1830. Castano, huihuixl (Quezaltenango).

Moist or wet, mixed forest, often on the steep sides of barrancos
in rather open places, sometimes on white-sand slopes, 1,100-2,700
meters; Alta Verapaz; Quiche"; Huehuetenango; Quezaltenango; San
Marcos. Southern Mexico.

A terrestrial shrub or tree, commonly 4-9 meters high, usually with a slender
trunk and few thick branches, densely and coarsely pilose or hirsute throughout
with branched, usually long-stipitate hairs, these often or usually yellowish; leaves
digitately 5-foliate, or the uppermost leaves sometimes only lobate, long-petiolate,
the petioles as much as 75 cm. long; leaflets sessile or short-petiolulate, thin, soft,
obovate to oblanceolate-oblong, sometimes 45 cm. long and 22 cm. broad but
usually not more than half as large, gradually or abruptly acuminate or caudate-
acuminate, remotely sinuate-dentate or almost entire; heads very dense and many-
flowered, subglobose, densely pale-tomentose, at anthesis usually 2 cm. in diameter,

14 FIELDIANA: BOTANY, VOLUME 24

in fruit often 4 cm. or more in diameter, forming long simple racemes; bractlets
ong-acuminate; stamens long-exserted; mature heads with very numerous fruits.

A common and conspicuous tree at many places in western Guate-
mala, very much in evidence at some places along the road above
Rodeo in San Marcos. It is highly ornamental and of striking and
unusual appearance, and has been introduced into cultivation in
southern California. The inflorescences are larger and more showy
than those of other species. The leaves of seedlings are simple and
broadly ovate-cordate.

Oreopanax lachnocephalus Standl. Carnegie Inst. Wash. Publ.
461:77. 1935.

Moist or wet forest, 600-1,500 meters; Pete"n (type collected at
Camp 31, British Honduras Boundary, W. A. Schipp 1272); Zacapa;
Jalapa. British Honduras; Honduras.

A tree as much as 18 meters high with a trunk 30 cm. in diameter, the young
branches stout, covered with a rather dense, ferruginous, stellate tomentum; leaves
coriaceous, on slender petioles 15-18 cm. long or more, palmately 5-7-lobate,
about 26-30 cm. broad, openly cordate at the base and sometimes short-peltate,
the lobes about half the length of the blade, long-acuminate, somewhat contracted
toward the base, glabrous above or when young more or less stellate-pilose, fer-
ruginous-tomentose beneath with branched hairs, usually densely so, the venation
elevated and very closely reticulate on both surfaces, the margins of the leaves
remotely serrate or in adult leaves entire or nearly so; heads numerous, globose,
1.5 cm. in diameter at anthesis, very dense and many-flowered, on stout peduncles
1-1.5 cm. long, forming large terminal panicles as much as 20 cm. long, very
densely ferruginous-tomentose; bracts triangular-ovate, acuminate, 7-8 mm. long;
immature fruit oblong, 7 mm. long, glabrous, depressed at the apex.

This is easily distinguished from 0. peltatus when in flower but
the two are often indistinguishable from sterile material. It is pos-
sible that 0. geminatus Marchal, described from Nicaragua, may be
an older name for this species.

Oreopanax langlassei Standl. Contr. U. S. Nat. Herb. 23:
1083. 1924.

Rare in mountain forests or oak-pine forests, 2,100-2,400 meters;
Baja Verapaz; Zacapa. Mexico (Michoacan or Guerrero, Oaxaca).

A shrub or small tree of 4-8 meters, the branches thick, when young covered
with a close subappressed brownish tomentum; leaves on long slender stellate-
tomentose petioles, digitately 5-7-foliolate; leaflets on short or somewhat elongate
petiolules, narrowly lance-oblong or narrowly lanceolate, 10-16 cm. long, 2.5-3 cm.
broad, narrowly long-acuminate, cuneate-attenuate at the base, entire or remotely
serrate, finely stellate-pubescent above or in age glabrate, deep green, somewhat

FIG. 3. Oreopanax lacnocephalus. Flowering branch; X

15

16 FIELDIANA: BOTANY, VOLUME 24

lustrous, paler beneath, rather densely and coarsely pubescent beneath with stipi-
tate branched hairs; staminate heads 5-6 mm. in diameter, dense, many-flowered,
long-stipitate, forming racemes 25-40 cm. long or longer, the branches densely
stellate-pubescent; fruits several in each head, the fruiting heads as much as 1.5 cm.
broad, the individual fruits 5-6 mm. in diameter, glabrous, greenish white.

Very closely allied to 0. xalapensis.

Oreopanax liebmannii Marchal, Bull. Acad. Brux. II. 47: 87.
1879. 0. oligocarpum Donn.-Sm. Bot. Gaz. 16: 194. 1891 (type from
Alta Verapaz, J. D. Smith 1743). 0. microcephalum Donn.-Sm.
Enum. PI. Guat. 6: 71. 1903, nomen. 0. meiocephalum Donn.-Sm.
Bot. Gaz. 37: 210. 1904 (type from Alta Verapaz, J. D. Smith 1743).
Matapalo; cercil (Chiquimula) .

Moist or wet forest, sometimes on white-sand slopes, 1,000-3,000
meters; Alta Verapaz; Baja Verapaz; Zacapa; Chiquimula; Solola;
Quiche* ; Huehuetenango; Quezaltenango; San Marcos. Southern
Mexico; British Honduras; Honduras; Costa Rica; Panama.

A shrub or tree, sometimes 9 meters high with a trunk 20 cm. in diameter,
glabrous except in the inflorescence; leaves chartaceous, on slender, usually long
but sometimes short petioles, mostly oblong-lanceolate or oblong and 8-20 cm.
long, attenuate to the acute base, somewhat paler beneath or almost concolorous,
the nerves evident or conspicuous beneath, slender; staminate heads 3-4 mm. in
diameter, few-flowered, slender-pedunculate, racemose or paniculate, the panicles
sometimes 15 cm. long, usually much shorter, the heads often few but sometimes
very numerous, the branches more or less pubescent or almost glabrous; fruits
usually only 2 in each head, about 4 mm. in diameter, white at maturity.

The Maya name "yaxyulup" is reported from British Honduras.
The species is a variable one in almost every character, and some of
the forms closely approach 0. capitatus. As a rule it may be recog-
nized by its narrow leaves, but there are some forms intermediate
in leaf characters.

Oreopanax obtusifolius L. Wms. Fieldiana, Bot. 31:20. 1964.
Aralia guatimalensis Hort. ex Lemaire, Fl. Serres 3: Misc. 44. 1847,
in synon., nomen illeg. Oreopanax guatemalense Dene. & Planch, ex
Hemsl. Biol. Cent. Am. Bot. 1: 573. 1881, nomen nudum; Sciadophyl-
lum guatamalense Lemaire ex Hemsl., loc. cit., in synon.

Epiphyte or terrestrial in damp or wet lowland forests, 0-200
meters (type from Forest Home, British Honduras, Schipp 1031);
Pete*n; Izabal; Alta Verapaz. Mexico; British Honduras; Honduras.

Large epiphytic shrubs or occasionally trees to 10 m. with a trunk to 10 cm.
in diameter, glabrous except the young branches and inflorescences, with thick

STANDLEY AND WILLIAMS: FLORA OF GUATEMALA 17

terete branches. Leaves broadly obovate to broadly ovate or suborbicular-ovate,
broadly round or obtuse at the apex, rounded to obtuse or subcordate at the base,
chartaceous to subcoriaceous, the blade 8-20 (-35) cm. leng and 3-18 (-25) cm.
broad, the nerves conspicuous, with 3-5 prominent ones arising at the base, sec-
ondary nerves few; inflorescence paniculate, to 25 cm. long and as broad, rachis
stellate-pubescent, becoming glabrescent, subtending bracts deltoid, 2-8 mm. long,
puberulent, the peduncles 2-10 mm. long, the flower heads ovoid, 6-8 mm. across;
calyx-limb truncate; petals narrowly oblong, 1.5-2 mm. long; fruits few in each
head, 5-6 mm. in diameter, creamy-white.

The species is closely related to the more widespread 0. capitatus.

Oreopanax peltatus Linden ex Regel, Gartenflora 11: 170, I.
363. 1862. 0. salmnii Hemsl. Diag. PI. Mex. 16. 1878 (type from
Volcan de Fuego, Salvin). Memo de leon; tronador.

Moist or rather dry forest, often in pine or oak forest, 1,500-2,700
meters; Alta Verapaz; Baja Verapaz; Zacapa; Jalapa; Guatemala;
Sacatepe"quez; Chimaltenango; Quiche1; Quezaltenango; San Marcos.
Mexico.

A shrub or a large tree, the smaller plants simple or sparsely branched, the
trees with a large crown, the young branches densely tomentose with branched
ferruginous hairs; leaves on long slender petioles, coriaceous but when fresh flex-
ible, 15-50 cm. broad, deeply 5-7-Iobate, the lobes broad or narrow, obtuse to
acuminate, sinuate-lobate to sinuate or remotely serrate, coarsely stellate-pilose on
the upper surface with stipitate hairs or in age glabrate, somewhat paler beneath,
usually densely stellate- tomentose when young or even in age; flower heads vari-
able in size, pedunculate, usually forming large and much branched, densely ferru-
ginous-tomentose panicles; fruits few or numerous in each head, black at maturity,
about 6 mm. in diameter, glabrous.

A frequent tree in some regions, and a highly ornamental one.
It has been introduced into cultivation in southern California, and
occasionally is planted for ornament in Guatemala, sometimes as a
pot plant. The tree is of some considerable economic importance in
Guatemala, for the large durable leaves are much used for covering
baskets or food containers, and as wrapping for soap and other arti-
cles. Bunches of them often are offered for sale in the markets for
this purpose. It might be thought that any large leaf would be suit-
able for such purposes but this is untrue, for most leaves either wither
too quickly or else are not sufficiently tough, pliable, and substantial.
Consequently, the leaves of only a small number of plants are used
ordinarily for wrapping purposes; the most important of all in Guate-
mala being Calathea and Musa leaves.

Many of the specimens available are sterile and it is quite prob-
able that some of these belong to 0. lacnocephalus Standl. We have

18 FIELDIANA: BOTANY, VOLUME 24

no fertile specimens from Honduras or Nicaragua and think it likely
that those so determined may be 0. lacnocephalus.

Oreopanax sanderianus Hemsl. Card. Chron. III. 11: 718.
1892. Cohete; tronador; tronador de montana; cohetillo.

Moist or wet, usually mixed, mountain forest, 1,200-3,000 meters;
Alta Verapaz; Chiquimula; Guatemala; Sacatepe"quez ; Chimalte-
nango (type from Volcan de Fuego, Salvin); Quezaltenango; San
Marcos. Mexico (Chiapas).

A large shrub or a small tree, glabrous outside the inflorescence, terrestrial or
most often epiphytic; leaves long-petiolate, subcoriaceous, very variable in size
and shape, truncate to obtuse at the base, or sometimes deeply and narrowly
cordate, usually angulately 3-lobate or sometimes 5-lobate, the lobes mostly shal-
low and triangular, acute or acuminate, often spreading at a right angle to the
costa, the blades mostly 12-25 cm. broad, almost concolorous; inflorescence usu-
ally large and paniculate, with a sparse, more or less appressed tomentum or almost
glabrous, the heads very numerous, pedunculate, rather few-flowered, the mature
fruiting heads 1.5 cm. broad; fruits usually 3-4 in each head, about 5 mm. in
diameter, white at maturity.

A very handsome plant, cultivated in the mountains of Guate-
mala for ornament either as a tree or as a pot plant. The young
leaves when 5-lobate remind one of a large ivy (Hedera) leaf. They
are of a bright and vivid green and usually glossy. The form of the
leaves is more variable than in any other local species.

Oreopanax steyermarkii A. C. Smith, No. Am. Fl. 28B: 36.

1944.

Moist or wet forest; 1,400-2,500 meters; endemic; El Progreso;
Zacapa (type from Sierra de Las Minas, Steyermark 42649); Hue-
huetenango.

Plants epiphytic or terrestrial, sometimes becoming a tree of 12 meters, gla-
brous except in the inflorescence or the young branches somewhat tomentulose;
leaves on long or short, stout petioles, oblong or narrowly oblong, 13-22 cm. long,
4-6 cm. broad, acuminate or short-acuminate, acute or obtuse at the base, thick-
coriaceous, entire, green and lustrous above, the venation not elevated, usually
much paler beneath or glaucescent, the costa stout and elevated, the lateral nerves
almost obsolete, very inconspicuous, the veins obsolete; heads very numerous,
dense, many-flowered, forming sessile panicles 10-14 cm. long, the branches stout,
tomentulose, the tomentum pale and appressed; fruits 2-3 in each head, globose,
7-8 mm. in diameter.

At first glance this appears to be a very distinct species, and fur-
ther collections may prove this to be the case, but it has little to
separate it from some of the forms of 0. capitatus and 0. liebmannii.

STANDLEY AND WILLIAMS: FLORA OF GUATEMALA 19

Oreopanax xalapensis (HBK.) Dene. & Planch. Rev. Hort. IV.
3: 108. 1854. Aralia xalapensis HBK. Nov. Gen. & Sp. 5: 8. 1821.
0. Taubertianum Donn.-Sm. Bot. Gaz. 19: 4. 1894 (type from Barr-
anco de Corona, Guatemala, J. D. Smith 1905). 0. loesenerianus
Harms, Bot. Jahrb. 23: 127. 1896 (type from Volcan de Fuego, Chi-
maltenango, Osbert Salviri). Mano de ledn; mano de tigre; aticuej
(Volcan de Agua); chilil mazorco (Sacatepe"quez) ; mazorco; mata-
gente; m'shal (Volcan de Santa Maria).

Moist or wet, mixed, mountain forest, sometimes on white-sand
slopes, 750-3,100 meters; Alta Verapaz; Baja Verapaz; Zacapa; Chi-
quimula; Santa Rosa; Guatemala; Sacatepe"quez; Chimaltenango;
Suchitepe'quez; Solola; Quiche1; Huehuetenango; Quezaltenango ; San
Marcos. Central and southern Mexico; Honduras and El Salvador
to Panama.

A shrub or usually a tree, sometimes 18 meters high with a trunk 45 cm. in
diameter, glabrous or nearly so except in the inflorescence; leaves on long slender
petioles, digitately 5-9-foliolate; leaflets chartaceous or subcoriaceous, narrowly
oblong-lanceolate to obovate, mostly 10-25 cm. long, acute to long-acuminate,
attenuate at the base, entire or somewhat serrate, green and lustrous above, paler
beneath, when young minutely stellate-puberulent but in age usually glabrous;
flower heads 5-15 mm. in diameter, few-many-flowered, forming short or often
much elongated, terminal racemes, these often 30 cm. long or more; fruits few or
numerous in each head, subglobose, 5-6 mm. long, becoming white and at maturity
black, very juicy.

It is called "Brazil" in El Salvador and is one of the most abundant
and conspicuous trees of the mountain forest of the central and west-
ern mountains, almost the dominant tree in some regions. Fre-
quently, the trees are planted for ornament in the parks. Wherever
the trees are found, there is an abundance of seedling plants. The
earliest leaves of these are simple and broadly cordate-ovate, the
succeeding ones deeply lobate or with 3 leaflets.

SCIADODENDRON Grisebach

Large or medium-sized, glabrous trees, usually with few branches; leaves very
large, deciduous, long-petiolate, twice pinnate, the leaflets membranaceous, short-
petiolulate, serrate; flowers arising on naked branches or at least on old wood, often
when the trees are leafless, umbellate, the umbels long-pedunculate, paniculate,
or subracemose, the pedicels slender, not articulate; margin of the calyx subentire;
petals 10-12, oval-linear, somewhat imbricate in bud; stamens 10-12; ovary sub-
globose, 10-12-celled, the styles recurved in age.

A single species is known.

5. H. Grove

FIG. 4. Sciadodendron excelsum. A, Paul C. Standley (1884-1963) with a
mature leaf, much reduced. B, Leaflet; X J^. C, Inflorescence; about X M-
D, Flower with petals in place; X 5. E, Flower with petals fallen; X 5. F, Fallen
petals; X 5.

20

STANDLEY AND WILLIAMS: FLORA OF GUATEMALA 21

Sciadodendron excelsum Griseb. Bonplandia 6: 7. 1858.

Moist or dry thickets or open lowland forest, usually on dry rocky
banks, 600 meters or lower; Zacapa; Chiquimula; Retalhuleu. El
Salvador to Panama; Haiti; Colombia.

A tree, sometimes 20 meters high, the trunk straight, pale, sometimes a meter
in diameter but usually smaller, the crown generally rather small and spreading,
sparsely branched, the bark smooth or somewhat roughened; leaves very large,
often a meter long, two or more times pinnate-ternate, the leaflets numerous, ovate
or ovate-oblong, mostly 4.5-8.5 cm. long, long-acuminate, coarsely serrate, green
above, paler beneath; umbels many-flowered, the pedicels mostly 5-8 mm. long;
calyx 3 mm. broad; petals about 4 mm. long.

Called "corroncha de lagarto" or "lagarto" in El Salvador. While
often attaining a considerable size, this plant often flowers when only
a shrub of 2-3 meters. In Panama the long petioles are used for
making bird cages. The wood is gray or yellowish gray throughout,
its luster rather low, moderately light but firm, brittle, of rather
coarse texture, mostly straight-grained and easy to work, but perish-
able in contact with the ground. No local uses are made of it, as
far as we know.

UMBELLIFERAE1 Carrot Family

References: John M. Coulter and J. N. Rose, A synopsis of the
Mexican and Central American Umbelliferae, Proc. Wash. Acad.
Sci. 1: 111-159, tt. 3-13. 1900. Hermann Wolff, Umbelliferae, Pflan-
zenreich IV. 228. 1910-27. Mildred E. Mathias and Lincoln Con-
stance, Umbelliferae, No. Am. Fl. 28B: 43-295. 1944-45.

Annual or perennial herbs, very rarely suffrutescent; leaves alternate or rarely
opposite, often all radical, the radical leaves usually larger and more conspicuous
than the cauline ones, the petiole often dilated at the base into a sheath; leaf blades
simple or usually compound or decompound; stipules usually none; inflorescence
almost always umbellate, rarely capitate, the umbels simple or generally com-
pound, often subtended at the base by an involucre of bracts, the umbellules usu-
ally subtended by bractlets; flowers small, regular or nearly so, perfect or often
polygamous-monoecious; petals mostly white, yellow, purple, or greenish; calyx
tube adnate to the ovary, the limb obsolete or of 5 small teeth; petals 5, inserted
on the margin of the calyx, equal or the outer ones larger, erect or spreading, often
incurved at the apex or with an inflexed acumen, the apex of the petal then appear-
ing emarginate, the petals usually imbricate or reduplicate- valvate in bud; stamens
5, the filaments filiform, inflexed in bud; anthers ovoid or subglobose, versatile,
the cells parallel, longitudinally dehiscent; ovary inferior, 2-celled; epigynous disc
bilobate or bifid, various in form, often continuous with the stylopodium (enlarged

1 An alternative name for this family, Apiaceae of Lindley, is sanctioned for
those who wish to be consistent in the termination of family names.

22 FIELDIANA: BOTANY, VOLUME 24

style bases); styles 2, distinct, filiform, straight or after anthesis often recurved;
ovules solitary in each cell of the ovary, pendulous from the apex of the cell, ana-
tropous; fruit inferior, dry, composed of 2 indehiscent 1-seeded merocarps, these
attached by their inner surface or commissure, separating from a usually persistent
carpophore; merocarps bearing dorsally usually 5 costae, 2 marginal ones, 1 dorsal
one, and 2 intermediate ones between the dorsal and marginal ones, the costae
often extended into broad wings; pericarp usually containing oil tubes; seed 1 in
each merocarp, pendulous from the apex of the cell, the inner face flat, concave,
or sulcate.

A large family with some 200, often not very distinct, genera of
which there are more than 90 in North America, best represented in
temperate regions, in the tropics, except for a few weedy species,
almost confined to mountain regions. Probably all the Central
American genera are represented in Guatemala. The family con-
tains a considerable number of important cultivated plants, used for
both food and condiment. Some few "umbels" are poisonous. Plants
of this family are easily recognized, as a rule, by their umbellate in-
florescences and characteristic fruits. Most also have a characteris-
tic odor. The separation and recognition of the genera is often
difficult.

Radical and principal cauline leaves simple.

Flowers capitate, sessile; leaves often with spine-tipped teeth Eryngium.

Flowers not capitate, pedicellate; leaves never with spine- tipped teeth.

Umbels compound; leaves terete, fistulous, septate Ottoa.

Umbels simple or, if compound, the leaves flat, orbicular to reniform or

broadly ovate, never terete.
Petiole with a dense tuft of hairs at the apex. Erect branched herbs with

usually ovate and cordate leaves Spananthe.

Petiole without a tuft of hairs at the apex.

Umbels compound, the umbellules 3-flowered; leaves reniform.

Micropleura.

Umbels simple or proliferous, not normally compound, when proliferous
the leaves peltate.

Petals acute, valvate; leaves peltate or reniform Hydrocotyle.

Petals obtuse, imbricate; leaves reniform Centella.

Radical and principal cauline leaves compound.

Fruit bearing conspicuous, straight or uncinate bristles or spines.

Fruit covered throughout with short uncinate bristles; leaves palmately di-
vided Sanicula.

Fruit with bristles only on the costae; leaves pinnately dissected or ternate-
pinnate.

Fruit linear-clavate, the bristles straight Osmorhiza.

Fruit broad, the bristles usually uncinate or barbate at the apex . . Daucus.
Fruit without either bristles or spines, usually glabrous.
Fruit compressed dorsally (parallel with the commissure) or not at all com-
pressed.
Plants annual; cultivated plants.

STANDLEY AND WILLIAMS: FLORA OF GUATEMALA 23

Flower white or tinged with pink Coriandrum.

Flowers yellow Anethum.

Plants perennial; native plants.

Fruit oblong or almost linear, twice as long as broad or longer.

Fruit almost linear; leaflets glabrous Myrrhidendron.

Fruit oblanceolate; leaflets somewhat pubescent beneath on the veins.

Enantiophylla.
Fruit broader, almost or quite as broad as long.

Stylopodium lacking, face of the seed sulcate; divisions of the leaves

crenate-serrate, not lobate Prionosciadium.

Stylopodium present, depressed to conic, face of the seed plane; divi-
sions of the leaves deeply cleft and dentate Rhodosciadium.

Fruit more or less compressed laterally (contrary to the commissure) or almost

terete.

Plants cultivated, or else native but then annuals.
Bracts of the involucre present and conspicuous.

Flowers greenish yellow; rays 10-20 Petroselinum.

Flowers white; rays 50-60 Ammi.

Bracts of the involucre none or few, small, and inconspicuous.

Flowers yellow; plants very glaucous Foeniculum.

Flowers white or nearly so; plants not conspicuously glaucous.

Ribs of the fruit filiform and very inconspicuous; plants puberulent.

Pimpinella.

Ribs of the fruit well developed; plants glabrous Apium.

Plants native or thoroughly naturalized, perennial.

Leaves pinnate, the leaflets serrate or incised Berula.

Leaves not simply pinnate, usually two or more times compound.

Mericarps of the fruit conspicuously winged. Involucre none or of 1
bract; tall plants with purple flowers Coaxana.

Mericarps not winged.

Umbels simple. Dwarf plants with scapiform stems; flowers white
or tinged with purple Oreomyrrhis.

Umbels compound.
Stylopodium conical.

Fruit cordate, emarginate at the base, somewhat didymous.

Arracacia acuminata.

Fruit not cordate, neither emarginate at the base nor didymous.

Arracacia.
Stylopodium none or depressed.

Plants dwarf, the stems scapiform; umbels with very few rays.

Tauschia.

Plants very large and tall, with leafy stems; umbels with many
rays.

Involucre none; flowers yellow Donnellsmithia.

Involucre present; flowers white Conium.

AMMI L.

Slender, glabrous annuals or perennials from slender tap roots; leaves ternately
or pinnately dissected, petiolate, the ultimate divisions filiform to lanceolate. In-
florescence a terminal or lateral compound umbel; involucral bracts entire or

24 FIELDIANA: BOTANY, VOLUME 24

divided, many; involucels of many entire bractlets; flowers usually white; petals
ovate or obovate with inflexed tips; calyx very small; styles slender, about twice
as long as the stylopodium; fruit oblong to ovoid, laterally compressed, the ribs
acute, oil tubes one at each interval and two on the commissures.

The genus is adventive in America.

Ammi majus L. Sp. PI. 243. 1753. Madre perla; encajes; lluvia
de plata.

Native of southern Europe. Sparingly introduced as an orna-
mental in Guatemala and elsewhere in Central America.

Erect branching annuals often a meter or more tall, glabrous or the upper part
of the plant obscurely puberulent; leaves ternate or usually pinnate, to 20 cm.
long and nearly as broad, the segments lanceolate, serrate to somewhat lacerate,
those of the stem bipinnate with linear divisions; involucral bracts of the inflores-
cences exceeding the rays, the involucels linear-subulate, entire or sometimes
divided, about as long as or shorter than the pedicels; fruit oblong, to about 2 mm.
long and half as broad.

ANETHUM L. Dill

Glabrous erect annuals, strong-scented, often very glaucous; leaves dissected
into linear or filiform segments, these very numerous, entire, the long petioles
dilated below into a broad, nerved sheath; flowers very small, yellow, in large com-
pound umbels; involucre and involucels none; fruit oblong to elliptic, strongly
compressed dorsally, the lateral costae winged, the dorsal ones slender, acute.

Two species are known, native in southeastern Europe and in
Asia.

Anethum graveolens L. Sp. PI. 263. 1753. Eneldo; hinojo.

Native of southeastern Europe; cultivated commonly in Guate-
malan gardens, and rarely found as an escape in corn fields or other
cultivated ground.

Plants usually a meter high or less, often much branched, very pale and glau-
cous throughout, the stems striate; leaves much dissected into very numerous,
almost filiform segments; umbels as much as 15 cm. broad, with several or very
numerous rays; fruits pedicellate, about 6 mm. long and 3 mm. broad.

The plant has a distinctive odor and the fruits a characteristic
flavor. Both the "seeds" and foliage are much used for flavoring
food, in the United States principally and very commonly for giving
their peculiar taste to cucumber and other pickles. Dill is much
grown about Coban and thrives in that wet and cool climate. The
plants often grow very tall, and some of those seen at various places
in Baja Verapaz must have been at least 2.5 meters high.

STANDLEY AND WILLIAMS: FLORA OF GUATEMALA 25

APIUM L.

Annual or perennial herbs, glabrous or nearly so; leaves pinnate or ternate-
pinnate; flowers yellow or white, the umbels compound, usually opposite the leaves
or terminal and in the forks of the branches; bracts of the involucre very few, 1,
or none, the bractlets of the involucels numerous or none; calyx teeth obsolete;
petals ovate, generally acute and concave, or produced into an inflexed acumina-
tion, entire or rarely emarginate; stylopodium depressed or low-conic; fruit ovate
or broader than long, laterally compressed, constricted at the commissure, often
didymous; carpels 5-angulate; carpophore undivided or short-bifid at the apex.

About 15 species, in tropical and temperate regions of both hemi-
spheres. No other species are known from Central America.

Divisions of the leaves linear or nearly so; native plants A. leptophyllum.

Divisions of the leaves cuneate or broadly cuneate; cultivated plants . . A. graveolens.

Apium graveolens L. Sp. PI. 264. 1753. Apio. Celery.

Native of Europe; grown occasionally for food in Guatemala at
middle or rather high elevations; rarely escaping to cultivated ground
(a weed in street at San Juan, Sacatepe*quez, Guatemala) .

A coarse glabrous strong-scented perennial, 30-90 cm. high, sparsely branched;
leaves pinnate, the basal and lower cauline ones long-petiolate, the upper cauline
ones short-petiolate or subsessile; leaf segments 3-5, petiolulate or sessile, thin,
broadly ovate to oval or cuneate, coarsely dentate and often incised; umbels ter-
minal and opposite the leaves, 3-7-radiate, the involucre and involucels small or
none; flowers small, white, on very short pedicels; fruit oval, 1 mm. long, the ribs
slightly winged.

Celery is occasionally cultivated in Guatemala, and elsewhere in
the highlands of Central America for sale in the markets, principally
to Europeans. It rarely does well and the petioles are often pithy.
It is used more as a flavoring material, especially for soups, than as a
vegetable. The horticultural form most common is apparently var.
dulce (Mill.) Pers. although large rooted celeriac, var. rapaceum DC.,
may be grown. Celery is often imported from the United States for
the market of Guatemala City.

Apium leptophyllum (DC.) F. Muell. ex Benth. Fl. Austr. 3:
372. 1866. Helosciadium leptophyllum DC. Prodr. 4: 105. 1830. A.
ammi Urban in Mart. Fl. Bras. II1: 341, t. 91. 1879, non Crantz.
Culantrillo; anis de ratdn; culantro de chucho.

Wet to dry thickets or fields, frequently in pine-oak forest, often
in waste or cultivated ground, 900-2,300 meters; Alta Verapaz; Chi-
quimula; Jalapa; Guatemala; Sacatepe"quez; Chimaltenango; Solola;
Huehuetenango; Totonicapan ; Quezeltenango. Southern United

26 FIELDIANA: BOTANY, VOLUME 24

States; Mexico; Honduras and El Salvador to Panama; West Indies;
South America; Australia.

A very slender, usually erect annual, generally 50 cm. high or less, glabrous,
sparsely or much branched; leaves thrice ternately parted into numerous, linear or
filiform segments, the lower leaves on rather long petioles, the upper ones on short
vaginate petioles; umbels sessile or nearly so, opposite the leaves, 1-3-radiate, the
rays 8-15-flowered, the pedicels 2-7 mm. long, the flowers white; fruit ovoid, 1.5-
2.5 mm. long, slightly compressed laterally; mericarps with 5 prominent ribs, the
carpophore shortly bifid at the apex.

A common but very inconspicuous, often weedy plant.

ARRACACIA Bancroft

Large or small, perennial herbs, glabrous or puberulent, the roots or rhizomes
sometimes tuberous-thickened; leaves often large, pinnate or pinnately decom-
pound, the leaflets usually broad, dentate or pinnatifid; flowers generally white,
small, the umbels compound, with few or numerous rays; bracts of the involucre
usually foliaceous, sometimes none; bractlets of the involucels numerous or few,
undivided; calyx teeth small or obsolete; petals ovate or broader, subentire, the
acumen inflexed; stylopodium conic, the margin undulate; fruit ovate or ovate-
oblong, usually attenuate at the apex, laterally compressed, constricted at the
commissure; merocarps transversely sub terete or 5-angulate, the primary costa
prominulous, the dorsal and intermediate ones usually more elevated; carpophore
2-parted; seed subterete, the face deeply sulcate, the margins commonly involute.

About thirty species or perhaps fewer, chiefly in the mountains
from Mexico to the South American Andes. Only one additional
species is found in Central America. The generic name is a modifi-
cation of the vernacular one used in Colombia and other Latin
American countries. — Neonelsonia Coulter and Rose, Contr. U. S.
Nat. Herb. 3: 307, t. 9. 1895, is here treated as a generic synonym
although it is maintained by Mathias and Constance in North Amer-
ican Flora.

Nodes of the rachis and petiolules pubescent in rings or with annulate callus thick-
enings.

With annulate callus thickenings; leaflets ovate or cordate-ovate, rounded, trun-
cate or cordate at the base A. annulata.

With rings of pubescence; leaflets linear-lanceolate to ovate-lanceolate, narrowed
to the base.

Leaflets mostly 1.5 cm. broad or less and linear or linear-lanceolate; stylopo-
dium depressed and relatively indistinct; fruits ovoid. .A. donnellsmithii.

Leaflets mostly 1.5 cm. broad or mostly broader, lanceolate to ovate-lanceo-
late; stylopodium prominent; fruits ovoid-cordate A. acuminata.

Nodes of the rachis glabrous and smooth.

Plants cultivated for their edible (yellow) roots A. xanthorrhiza.

Plants native and found only wild.

STANDLEY AND WILLIAMS: FLORA OF GUATEMALA 27

Stylopodium depressed and indistinct; plants in flower less than 1 m. tall.

A. vaginata.
Stylopodium evident, conic; plans in flower more than a meter tall.

Leaflets mostly deeply lobate, usually glaucous or glaucescent beneath.

A. atropurpurea.
Leaflets mostly serrate or dentate (a few rarely lobate), green beneath.

A. bracteata.

Arracacia acuminata Bentham, PI. Hartw. 187. 1845. Neo-
nelsonia ovata Coulter & Rose, Contr. U. S. Nat. Herb. 3: 307, t. 9.
1895. N. acuminata Coulter & Rose ex Drude in Engler & Prantl,
Nat. Pflanzenf. 3, Abt. 8: 167. 1898.

Moist or wet mixed mountain forests, 1,400-2,900 meters; El Pro-
greso; Sacatepe"quez; Chimaltenango; Solola; Quezaltenango; San
Marcos. Mexico; Costa Rica; Colombia to Peru in the Andes.

Large coarse herbs, commonly erect and 1-1.5 meters tall, sometimes more
elongate and as much as 3 meters long, then decumbent and trailing, the stems
hollow, often purplish, glabrous, leafy; basal leaves very large, mostly 30 cm. long
or more, long petiolate, the leaflets numerous, lanceolate to ovate-lanceolate, usu-
ally 4-6 cm. long or larger and 2-4 (4.5) cm. broad, thin, bright green, acuminate
or caudate-acuminate, coarsely and unevenly serrate and often lobate or lobulate,
usually somewhat scabrous or at least somewhat roughened beneath on the pale
veins; umbels few, on long peduncles, compound, often to 16 cm. broad, the rays
very numerous, slender, unequal, glabrous; pedicels slender, glabrous, 18 mm. long
or shorter, usually very unequal; bractlets of the involucre sometimes 5 cm.
long but usually shorter; fruit as broad as or usually broader than long, 4-5 mm.
long and 5-7 mm. broad.

A common and showy plant in the wet forests of the mountains of
the Western Guatemalan highlands, often in great abundance. The
range of the species is curious. It is distributed in Mexico and Guate-
mala and appears again in Costa Rica and then in the Andes from
Colombia to Peru. It has been segregated as Neonelsonia but seems
too near to A. donnellsmithii and to some Andean species to main-
tain it distinct from Arracacia.

Arracacia annulata L. Wms. Fieldiana, Bot. 31:46, fig. 9 1964.

Cool mountain forests, 1,650 to 3,100 meters; Huehuetenango
(type from Sierra de los Cuchumatanes, Steyermark 48965). Mexico
(Vera Cruz and perhaps Puebla).

Large, coarse, essentially glabrous suffrutescent perennial herbs to 1.5 m. or
more tall. Leaves deltoid-ovate in general outline, ternate-pinnate, or ternate-
bipinnate, 10-35 cm. long (excluding petiole) and as broad or broader, petiole to
15 cm. long and sheathing at the base, all junctures of petiolules with rachis and
petioles with rugose annulate callus thickenings, usually also at the junction of

28

FIELDIANA: BOTANY, VOLUME 24

FIG. 5. Arracacia annulata. A, Leaf from upper part of stem; X K. B, Ma-
ture fruit; X 5. C, Annulate callus thickening on petiole; X 5, D, Annulate
callus thickening at the base of a terminal leaflet; X 5.

the petiolule with the leaflet; the uppermost leaves much reduced, occasionally to
a sheathing bract; leaflets ovate to cordate-ovate, acuminate, the base round, trun-
cate or shallowly cordate, serrate-dentate except at the base, 3.5-9 cm. long and
2-4.5 cm. broad (those on upper reduced leaves much smaller), the petiolule on
lateral leaflets 0-1 cm. long, that (rachis?) on terminal leaflets to 3.5 cm. long;
inflorescence cymosely branched, the umbels few-many; peduncles of fertile, ma-
ture umbels 3-5 cm. long; involucre of few short, soon deciduous bracts or none;
involucel of several linear or filiform bractlets 2-5 mm. long and deciduous with
age, fertile rays 2-20, spreading, the pedicels 2-10 mm. long; flowers purple; petals
oblong, ± 0.5 mm. long; sepals broadly ovate, ± 1 mm. long; stylopodium nar-
rowly conic, the styles slightly reflexed, the stigmas capitate, the whole 1.5-2.5
mm. long; the fruit narrowly lanceolate, about 7 mm. long and 1.5-2 mm. broad,
the ribs prominent, subalate on young fruit.

Easily distinguished from the other species of the genus by the
rugose annulate callus thickenings at the juncture of rachis and
petiolules.

STANDLEY AND WILLIAMS: FLORA OF GUATEMALA 29

Arracacia atropurpurea (Lehm.) Benth. & Hook, ex Hemsl.
Biol. Cent. Am. Bot. 1 : 564. 1880. Pentacrypta atropurpurea Lehm.
Ind. Sem. Hort. Hamb. 17. 1828. A. luxeana Coult. & Rose, Bot.
Gaz. 18: 55. 1893 (type from San Miguel Uspantan, Quiche", Heyde &
Lux 3354). A. humilis Rose, Contr. U. S. Nat. Herb. 8: 336. 1905
(type collected between Totonicapan and Los Encuentros, C. & E.
Sekr2352).

Moist or wet, mixed forest, or often with pine, Cupressus, Abies,
or Juniperus, sometimes on limestone, 1,800-3,700 meters; Zacapa;
Jalapa; Sacatepe'quez (Volcan de Agua); Chimaltenango; Huehuete-
nango; Quezaltenango; San Marcos. Central and southern Mexico;
Costa Rica; Panama.

A perennial herb, sometimes a meter high but usually much lower, erect or
sometimes decumbent, glabrous throughout or nearly so, the stems often purplish
and glaucescent, leafy; leaves long-petiolate or the uppermost cauline ones short-
petiolate, ternate, the divisions pinnate; leaflets petiolulate or sessile, oblong-ovate
to broadly rhombic-ovate, mostly 2-4 cm. long, obtuse or acute, usually glaucous
or glaucescent beneath, thin, most of them deeply lobate and irregularly crenate-
serrate with mucronate teeth; umbels few, long-pedunculate, 10 cm. broad or less,
the rays numerous, very unequal, often scabrous at the base, the pedicels stout,
very unequal; involucre none, the bracts of the involucels few, linear or lanceolate,
entire or laciniate, usually much longer than the flowers; fruit glabrous, about
7 mm. long and 2.5 mm. broad.

Arracacia bracteata Coult. & Rose, Contr. U. S. Nat. Herb. 3:
295. 1895. Cicuta (Jalapa) ; arracacha de monte; chunch-botch.

Moist or wet, mixed, mountain forest, or often in pine or oak for-
est, 2,200-3,150 meters; Jalapa; Guatemala; Sacatepe'quez (Volcan
de Agua); Chimaltenango; Huehuetenango; Quezaltenango; San
Marcos. Southern Mexico; Honduras.

A coarse, somewhat succulent herb, commonly 1-3 meters high, or often lower,
the stems thick, hollow, simple or branched, glabrous; radical leaves very large,
on long slender petioles, biternate, the ultimate segments usually pinnate; leaflets
sessile, thin, oblong or ovate-oblong, mostly 5-12 cm. long, acuminate or long-
acuminate, green beneath, unevenly and closely serrate, rarely some of the leaflets
shallowly lobate, glabrous; flowers white, the umbels sometimes 20 cm. broad,
naked at the base, the rays numerous, very unequal, sometimes scabrous at the
apex; pedicels stout, unequal, often very numerous, glabrous or puberulent; fruit
glabrous, about 6-8 mm. long and 3-4 mm. broad, rounded or emarginate at
the base.

Arracacia donnell-smithii Coult. & Rose, Bot. Gaz. 15: 261,
t. 15. 1890. Sis (Huehuetenango); Solojo (San Marcos).

FIG. 6. Arracacia bracteata. A, Leaf and inflorescence; about X M- B, Fruit;
X 4. C, Apex of fruit; X 8.

30

STANDLEY AND WILLIAMS: FLORA OF GUATEMALA 31

Moist or wet, mixed forest, or more often in the higher belts of
pine forest on the high mountains, sometimes in Cupressus forest or
in wet thickets, 2,400-3,600 meters; Sacatepe'quez (type from Volcan
de Agua, J. D. Smith 2196); Chimaltenango; Huehuetenango; Que-
zaltenango; San Marcos; known only from Guatemala but to be
expected in Chiapas.

A large coarse perennial herb, usually 1.5-3 meters high, almost glabrous, the
foliage bright green, the stems thick, hollow, often purplish; simple or sparsely
branched, leafy; radical leaves very large, 2-3 times ternate, the ultimate divisions
mostly pinnate, the nodes densely pubescent or tomentose; leaflets very numer-
ous, oblong-lanceolate to almost linear-lanceolate, mostly 5-9 cm. long, 0.5-1.5
(2.5) cm. broad, glabrous, green beneath, attenuate-acuminate, sessile and often
decurrent on the rachis, finely and evenly serrate, usually not at all lobate; flowers
white, the umbels usually numerous, sometimes 12 cm. broad, the rays numerous,
stout, unequal, densely puberulent, the pedicels short, unequal, stout, puberulent;
involucre none, the bracts of the involucels lanceolate or linear-lanceolate, green-
ish, longer than the flowers; fruit glabrous, ovate, about 6 mm. long and 4 mm.
broad.

A rather handsome and showy, tall plant, often occurring in abun-
dance throughout or especially at the bases of the highest pine-clad
slopes of the volcanoes. On the Volcan de Acatenango it is found
everywhere as one emerges from the dense wet Chiranthodendron
forest into the open pine forest, where most of the ground is covered
with coarse bunch grasses.

Arracacia vagina ta Coult. & Rose, Contr. U. S. Nat. Herb. 3:
297. 1891. Tauschia vaginata Coult. & Rose, Proc. Wash. Acad. Sci.
1: 137, /. 8. 1900.

At 2,500-2,800 meters; Huehuetenango. Mexico (Michoacan;
Oaxaca) .

Plants perennial from a thick perpendicular root, 20-75 cm. high, erect or de-
cumbent, the stems glabrous or nearly so; lowest leaves twice pinnate or pinnate
and with deeply lobate leaflets, long-petiolate, the petioles broadly sheathing at the
base; leaflets thin, ovate or lance-ovate, coarsely dentate or cleft, acute or obtuse,
cuneate at the base, pubescent on the margins and veins or almost glabrous, green
beneath; peduncles short or elongate, the involucre none or of a few linear bracts;
bractlets linear, acuminate, green, entire, about as long as the flowers; rays of the
umbel 6-25 mm. long, the pedicels 2-4 mm. long, glabrous or minutely puberulent;
petals yellow; fruit ovoid, 5-6 mm. long, the styles long, recurved, the costae stout
and prominent.

This species appears quite different from the other species of
Arracacia in Guatemala and is easily distinguished from them by
having the stylopodium depressed and indistinct rather than quite
evident and conic.

32 FIELDIANA: BOTANY, VOLUME 24

Arracacia xanthorrhiza Bancroft, Trans. Agri. Hort. Soc. Ja-
maica 1825: 5. 1825. A. esculenta DC. Prodr. 4: 244. 1830. Arracacia.

Believed to be native of the mountains of Colombia; planted occa-
sionally for food in mountain regions of other parts of tropical Amer-
ica; cultivated occasionally in Guatemala.

An erect perennial from a tuberous, yellow or white root, this bearing several
exterior tubers, these as much as 20 cm. long and 5-6 cm. thick; stems erect,
50-100 cm. high, glabrous, branched, often streaked with purple; radical leaves
large, with usually 5 ovate-acuminate pinnae, these deeply and irregularly pin-
natifid, or the lowest ones pinnate, the ultimate segments acuminate and serrate,
green beneath; umbels few or numerous; involucre none; bractlets of the involucels
2-8, small, setaceous; fruit oblong, the mericarps with 5 subequal obtuse ribs;
styles slender, almost equaling the fruit.

An extended account of this plant is given in connection with
plate 3092 of Curtis' Botanical Magazine. It is propagated from cut-
tings of the tubers, much like the common potato, and is said to yield
a crop of roots within about four months. It is reported that in the
region of Bogota, Colombia, the root is a rather important and pop-
ular vegetable but apparently it has found little favor in Costa Rica
and Guatemala, where it is sometimes cultivated. We do not recall
having seen it served on the table in Guatemala, although it may be
eaten by the Indians, or perhaps it is used only in soups and not eaten
alone.

BERULA Hoffmann

Glabrous perennial herbs, erect, simple or branched, growing in wet soil or in
shallow water; leaves pinnate, the segments few or numerous, serrate or incised;
flowers small, white, the umbels terminal, compound; involucre and involucels of
several narrow bracts and bractlets; calyx teeth very small; stylopodium conic, the
styles short; fruit subglobose, emarginate at the base, slightly compressed laterally,
glabrous, the ribs very slender, the pericarp thick and corky; face of the seed flat.

The genus consists of a single species.

Berula erecta (Huds.) Coville, Contr. U. S. Nat. Herb. 4: 115.
1893. Sium erectum Huds. Fl. Angl. 103. 1762.

Wet soil along streams, 2,100-2,500 meters; Huehuetenango.
Canada and United States; Mexico; Europe; and Asia.

Plants rather stout, erect, usually less than a meter high, simple or often much
branched, the stems striate; leaves pinnate, long-petiolate, the leaflets 7-19, ovate
to oval or narrowly oblong, 1.5-3.5 cm. long, acute, sessile, deeply serrate, laciniate,
or lobate; umbels numerous, short-pedunculate, 10-20-rayed, the rays short or
elongate; pedicels 3-6 mm. long; bracts and bractlets lanceolate or linear-lanceo-
late, entire; fruit scarcely 2 mm. long, suborbicular, subcordate at the base, the
ribs inconspicuous.

X

33

34 FIELDIANA: BOTANY, VOLUME 24

CENTELLA L.

Perennial, glabrous (ours) herbs, the stems prostrate and rooting, with several
long petiolate leaves at the nodes; leaves petiolate, dentate or undulate, palmately
veined, cordate at the base, the petioles with vaginate bases; inflorescence of sim-
ple, subcapitate, few-flowered umbels (ours), peduncles axillary, about as long as
the leaves or usually shorter, involucral bracts usually two and conspicuous; calyx
teeth none; stylopodium flat or none; styles short, filiform; fruits laterally com-
pressed, prominently costate, oil tubes none.

A genus of some twenty species found in both hemispheres, most
abundant in Africa. The following is the only species found in
America.

Centella asiatica (L.) Urban in Mart. Fl. Bras. 11, pt. 1: 287.
1879. Hydrocotyle asiatica L. Sp. PI. 234. 1753. H. erecta L. f . Suppl.
177. 1781. H. asiatica f. luxurians Donn.-Sm. Enum. PI. Guat. 1:
15. 1889, nomen. Centella erecta Fernald, Rhodora 42: 295. 1940;
Math. & Const. No. Am. Fl. 28B: 59. 1944.

Wet meadows or thickets, stream banks, marshes, sometimes in
wet pine woods, 1,900 meters or less; Pete*n; Alta Verapaz; Jalapa;
Huehuetenango. Eastern and southeastern United States; Mexico;
British Honduras; Honduras. West Indies; South America; Old
World tropics.

Stems rather stout and somewhat fleshy, short or elongate, creeping and root-
ing at the nodes; petioles 3-30 cm. long; leaf blades broadly ovate to orbicular-
ovate, 2-7 cm. long, rounded at the apex, openly and often deeply cordate at the
base, rather thick and fleshy, repand-dentate; peduncles usually shorter than the
leaves, mostly 1-5 cm. long, the umbels head-like, 2-4-flowered, subtended by 2
ovate bracts, the flowers sessile or subsessile; fruit 3-4 mm. long and 3-5 mm.
broad, conspicuously costate and somewhat reticulate.

COAXANA Coulter & Rose

Large coarse perennial herbs, glabrous or nearly so, the stems simple or sparsely
branched, leafy; leaves large, bi-triternate, the leaf sheaths conspicuous and some-
what inflated, the leaflets thin, lobate and serrate; involucre none or of a single
bract; involucels conspicuous, narrow, serrate or incised; umbels compound, many-
radiate, the rays unequal; flowers purple; the calyx teeth obsolete; fruit oblong,
glabrous, laterally flattened, glaucous; stylopodium low-conic; styles long and
slender, recurved; merocarps 3-4-winged; face of the seed plane or concave.

The genus consists of two species. The generic name is an ana-
gram of the word Oaxaca, the name of the state of Mexico in which
the plant was first collected.

FIG. 8. Coaxana purpurea. Habit, slightly reduced. A, Dorsal view of fruit;
about X 5. B, Cross-section of fruit; about X 8. [Illustration from original pub-
lication, courtesy of the Smithsonian Institution.]

35

36 FIELDIANA: BOTANY, VOLUME 24

Coaxana purpurea Coult. & Rose, Contr. U. S. Nat. Herb. 3:
297, t. 5. 1895.

Moist or wet, mixed forest, 2,100-3,000 meters; Chimaltenango;
Solola; Quezaltenango; San Marcos. Mexico (Michoacan, Oaxaca).

Plants a meter high or lower, or in Mexico reported as 2 meters tall, the stems
glabrous, glaucous, often purplish; radical leaves very large, triternate or bipin-
nate, the leaflets ovate to ovate-oblong, very unequal in size and shape, acute,
petiolulate or sessile, irregularly and often deeply lobate and unequally doubly
serrate, with acuminate teeth, paler beneath, the cauline leaves reduced upward,
the sheaths conspicuously inflated; peduncles short or elongate, puberulent at the
base of the umbel; involucre of a single foliaceous bract, or none; the pedicels
5 mm. long or less, glabrous or minutely scaberulous; involucels lanceolate or lin-
ear, much longer than the flowers, green, most of them with a few teeth or lacinia-
tions; fruit about 5 mm. long, each merocarp with 3 or 4 conspicuous broad wings.

CONIUM L. Poison hemlock

Tall glabrous biennial herbs with branched stems from a stout tap-root; leaves
pinnately decompound, the numerous segments pinnatifid and dentate; flowers
small, white, polygamous, the umbels large, compound, many-radiate; bracts and
bractlets of the involucre and involucels numerous, small; calyx teeth obsolete;
petals obovate or cuneate, obtuse and entire or with a short inflexed acumen;
stylopodium depressed; fruit broadly ovate, laterally compressed, more or less
constricted at the commissure; merocarps somewhat 5-angulate, the primary costae
prominent, obtuse, undulate-crenate or smooth, the lateral ones distinct; carpo-
phore undivided; seed deeply and narrowly sulcate on the inner face.

Probably two species, the other one African.

Conium maculatum L. Sp. PI. 243. 1753. Perejil de monte;
perejil de chucho.

Sometimes abundantly naturalized in waste ground, especially in
wet soil, 1,800-2,500 meters; Chimaltenango (Patzicia); Quezalte-
nango. Native of Europe, now widely naturalized around the world.

Plants erect, about a meter high, the stems stout, hollow; lower and basal
leaves petiolate, the upper cauline ones sessile or nearly so, pinnately dissected,
the leaflets very numerous, ovate in outline, thin, incised and dentate; petioles
much dilated and vaginate at the base; umbels 2.5-7 cm. broad, with slender elon-
gate rays, the pedicels filiform, 4-6 mm. long in fruit; bracts and bractlets few,
poorly developed, inconspicuous, entire; fruit 3 mm. long, the costae very conspic-
uous in the dry state.

This is the hemlock famous in the history of Greece and other
Mediterranean countries. It is supposed to be the plant by which
Socrates was put to death, and was employed commonly in the poi-

STANDLEY AND WILLIAMS: FLORA OF GUATEMALA 37

soning of criminals. It has become abundantly naturalized about
Quezaltenango, and there are some very large and luxuriant colonies
of it in the lower part of that city.

CORIANDRUM L. Coriander

Slender annuals, branched, glabrous; leaves petiolate or the upper ones sessile,
the basal leaves with ovate incised segments, those of the cauline leaves linear;
flowers white or tinged with pink; inflorescence of loose, compound umbels, usually
few-rayed; involucre usually none; the bractlets of the involucels few, filiform;
calyx teeth acute, generally unequal; petals usually unequal, those of the outer
flowers of the umbel usually larger, obovate, with an inflexed tip, more or less
bilobate at the apex; stylopodium conic, entire; fruit globose or ovoid, the mero-
carps transversely semiterete, not easily separable, the costae prominulous; carpo-
phore bifid; seed strongly compressed dorsally, orbicular, somewhat concave on
the face.

Two species, native in Europe and western Asia.

Coriandrum sativum L. Sp. PI. 256. 1753. Culantro: Culantro
de Castillo,.

Native of Europe, cultivated commonly in most civilized regions,
and grown frequently in Guatemalan gardens; often naturalized
rather freely in waste and cultivated ground, at 900-2,500 meters;
Alta Verapaz; Zacapa; Santa Rosa; Jutiapa; Sacatepe"quez; San
Marcos, and doubtless other departments.

Plants erect, usually 50 cm. high or less; lowest leaves pinnately divided, the
segments ovate to obovate, variously lobate and dentate, the upper leaves divided
into very numerous, short, linear, pale segments; umbels 2.5-5 cm. broad, with
slender rays, the pedicels 2-4 mm. long, the flowers rather conspicuous and showy;
bractlets of the involucels deciduous; fruit about 4 mm. long, the ribs subacute,
narrower than the intervening intervals.

Coriander is grown in Guatemala as elsewhere for its aromatic
"seeds" of distinctive taste that are used to flavor various articles of
food. The fresh plants often are sold in Guatemalan markets, for
use in soups and meat dishes.

DAUCUS L.

Annual or biennial herbs, usually with harsh pubescence; leaves pinnately de-
compound, the segments small or narrow; flowers small, white, the outer ones often
larger than the inner ones; umbels compound, the outer rays longer and in age con-
nivent above the inner ones, or the rays equal or unequal and all spreading even in
fruit; calyx teeth small and acute or obsolete; petals usually unequal, obovate,
with an inflexed or involute acumen, emarginate, or sometimes entire; stylopodium
depressed or short-conic, sometimes undulate-crenulate on the margin; fruit ovate

38 FIELDIANA: BOTANY, VOLUME 24

or oblong, subterete or more or less dorsally compressed; merocarps semiterete or
dorsally convex, the ribs more or less prominent, all or only the secondary ones
aculeolate or setulose; carpophore entire or bifid; seed semiterete or dorsally com-
pressed, the face flat.

About 50 species, almost all of them in temperate or warm regions
of the Old World, only two species native in America. Only the fol-
lowing are known in Central America.

Outer rays of the umbel in age connivent above the inner ones; cultivated plants.

D. carota.
Outer rays of the umbel widely spreading in fruit; native plants D. montanus.

Daucus carota L. Sp. PI. 242. 1753. Zanahoria. Carrot.

Native of Europe and Asia, cultivated for food in almost all civi-
lized parts of the earth; grown commonly as a vegetable in Guatemala.

Plants biennial, setulose-hispidulous or hirsute throughout, erect, a meter high
or usually lower, branched; basal and lower leaves 2-3-pinnate, the segments lan-
ceolate, dentate or pinnatifid, the upper leaves smaller and less divided; bracts of
the involucre parted into linear or filiform lobes; umbels 5-10 cm. broad, the inner
rays shorter than the outer ones, the slender pedicels 2-4 mm. long; flowers white,
the central one of each umbel usually dark purple; fruit 3-4 mm. long, setulose on
the somewhat winged ribs.

The carrot is a common and fairly popular vegetable in Guate-
mala although the Indian peoples probably use it little. It grows
well at most elevations, but the root is usually quite hard and woody.
Most vegetables of the temperate zone are grown in Guatemala. It
may be noted, however, that parsnips are rare in Central America
and the few which we have seen tended to be excessively woody.

Daucus montanus Humb. & Bonpl. ex Schult. Syst. Veg. 6:
482. 1820. Alfileria (Quezaltenango).

Moist or dry, often rocky, brushy or open fields and hillsides,
sometimes in meadows in Alnus forest, in gravel along streams, or a
weed in waste or cultivated ground, 1,000-3,000 meters, most fre-
quent at middle or higher elevations; Alta Verapaz; Zacapa; Chiqui-
mula; Jalapa; Guatemala; Sacatepe'quez; Chimaltenango; Solola;
Totonicapan; Quiche"; Huehuetenango; Quezaltenango; San Marcos.
Mexico; El Salvador; Honduras; Costa Rica; South America.

Plants sometimes annual but probably enduring often for more than one year,
erect, sparsely or much branched, generally 50 cm. high or less, sparsely or densely
hispidulous; leaves mostly small, divided into very numerous, small, narrow, ob-

STANDEE Y AND WILLIAMS: FLORA OF GUATEMALA 39

tuse and apiculate segments; bracts of the involucre much divided and resembling
leaves; umbels many-flowered, usually rather open, as much as 15 cm. broad but
usually much smaller, the rays stiff, very unequal; bractlets of the involucels linear;
pedicels in fruit usually conspicuously elongate and mostly 5-8 mm. long; fruit
about 5 mm. long, with dense rows of stout pale spines along the winged angles,
the spines barbate at the apex.

A rather common plant in meadows of the highlands, often plenti-
ful in overgrazed areas, since no animals eat the foliage. The burlike
fruits adhere closely to clothing and to the hair of animals, thus aid-
ing in wide dispersal of the seeds.

DONNELLSMITHIA Coulter & Rose

Slender glabrous caulescent perennials arising from slender, somewhat thick-
ened roots, the stems simple or sparsely branched, sparsely leafy; leaves 1-4-ternate
or ternate-bipinnately decompound, the rather few segments linear or broader, en-
tire or variously cleft; inflorescence of loose compound umbels; flowers very small,
pedicellate, yellow or purple, involucre present or none, of linear or filiform bracts;
involucels none; calyx teeth obsolete; stylopodium obsolete; fruit rounded-ovoid,
glabrous, somewhat compressed laterally; merocarps with equal filiform costae, the
intermediate one distant from the lateral ones; seed face deeply and narrowly sulcate.

About 12 species, in Mexico and Central America. Only the fol-
lowing have been found in Central America. The genus was named
for John Donnell Smith, who might, with much justice, be called the
father of Guatemalan botany.

Segments of the leaves linear, 1-3 mm. broad D. peucedanoides.

Segments of the leaves lanceolate or broader, 5-18 mm. broad . . . D. guatemalensis.

Donnellsmithia guatemalensis Coult. & Rose, Bot. Gaz. 15:
15, t. 2. 1890. Tauschia guatemalensis Macbride, Contr. Gray Herb.
56: 33. 1918.

Moist or wet meadows or open, pine or oak forest, 1,200-2,500
meters; endemic; Baja Verapaz (type from Santa Rosa, Tuerckheim
1311); Zacapa; Chimaltenango; Chiquimula; Huehuetenango. Hon-
duras.

Plants slender, glabrous, a meter high or less, glaucescent, erect, simple or
usually sparsely and openly branched above; leaves few, at or near the base of
the stem, the lower ones long-petiolate, the uppermost short-petiolate or subsessile,
2-3 times ternate, the leaflets few, very variable in shape and size, linear-lanceo-
late to lanceolate or rhombic, 2.5-5.5 cm. long, acute to long-acuminate, cuneate
at the base, mostly entire and with revolute margins, sometimes lobulate; umbels
compound, few-rayed, terminating long slender naked peduncles; involucre often
present and consisting of 3-4 trifid bracts, sometimes absent; pedicels 4-6 mm.
long; fruit 2-3 mm. long.

FIG. 9. Donnellsmithia guatemalensis. 1, Habit; X M« 2, Umbellule; about
X 1. 3, Fruit; X 6. 4, Cross-section of fruit; about X 8. [Illustration, drawn
by C. E. Faxon, is from original publication, courtesy of The University of
Chicago Press.]

40

STANDLEY AND WILLIAMS: FLORA OF GUATEMALA 41

Donnellsmithia peucedanoides (HBK.) Mathias & Constance,
Bull. Torr. Bot. Club 68: 122. 1941. Cnidium peucedanoides WK.
Nov. Gen. & Sp. 5: 15. 1821. Smyrnium ? linear e Bentham, PI.
Hartw. 83. 1841 (type Guatemala, Hartweg 578). Eulophus peuceda-
noides Benth. & Hook. Gen. PI. 1: 885. 1867. Museniopsis peu-
cedanoides Coult. & Rose, Contr. U. S. Nat. Herb. 3: 303. 1895.
Perejil de rana (fide Aguilar); valeriana (Huehuetenango).

Usually in rather dry, open, pine or oak forest, 1,000-3,000 meters .
El Progreso; Jalapa; Santa Rosa; Guatemala; Chimaltenango; Qui-
che"; Huehuetenango. Mexico; Honduras; South America (Colombia).

Plants very slender, erect, glaucous, 35-90 cm. high, the stems mostly solitary,
simple or sparsely branched, terete, naked or usually bearing 1-2 leaves below;
basal leaves on very long, slender petioles, 2-3 times ternate, the divisions few or
rather numerous, linear or narrowly linear, 7.5 cm. long or shorter, often revolute,
entire; umbels few, on very long and slender, naked peduncles, compound, the rays
3-5; involucre of 2-3 small 3-fid bracts, these frequently deciduous; fruit glaucous,
2-3 mm. long.

This is rather widely distributed in pine and oak forests at middle
elevations in the mountains, but the plants usually are few and iso-
lated, inconspicuous among the tall grasses with which they usually
grow.

ENANTIOPHYLLA Coulter & Rose

Large coarse perennials, herbaceous or suffrutescent, the stems simple or
branched, leafy, the plants almost glabrous; leaves opposite, very large, the rad-
ical and lower cauline ones long-petiolate, ternately compound, the primary divi-
sions pinnate or bipinnate, the uppermost leaves with short sheathing petioles,
pinnate, the leaflets large, thin, lanceolate or lance-ovate, long-acuminate, serrate;
umbels compound; bracts of the involucre few, elongate-linear, entire; bractlets
few, linear, the flowers long-pedicellate, probably white; calyx teeth obsolete; fruit
oblanceolate-oblong, rather large, somewhat compressed dorsally, glabrous; mero-
carps dorsally compressed, the dorsal and intermediate ribs prominent, acute, the
lateral costae with broad pale wings; stylopodium slender-conic; style stout, erect,
conspicuously thickened and subcapitate at the apex; seed strongly compressed
dorsally, the face concave.

The genus consists of a single species.

Enantiophylla heydeana Coulter & Rose, Bot. Gaz. 18: 56,
t. 5. 1893. Canonera (Sacatepe"quez) .

Brushy or wooded, often rocky and steep hillsides, 1,000-2,050
meters; Chiquimula; Jalapa; Santa Rosa; Guatemala; Sacatepe"quez
(type from Santiago, Rosalio Gomez 788); Chimaltenango; Quezalte-
nango. Mexico (Mexico) ; El Salvador.

FIG. 10. Enantiophylla heydeana. Leaf and inflorescence; about X K- Dor-
sal view of fruit is about X 1^. Outline shows cross-section of fruit but dorsal
wings usually more prominent. [Illustration, drawn by C. E. Faxon, is from orig-
inal publication, courtesy of The University of Chicago Press.]

42

STANDEE Y AND WILLIAMS: FLORA OF GUATEMALA 43

Plants 1-3 meters high, sometimes somewhat woody and with a stout erect
trunk a meter high and 10 cm. thick, the stems hollow, simple or sparsely branched;
lower leaves very large and on long thick hollow petioles, the leaflets lanceolate or
lance-ovate, mostly 5-10 cm. long, thin, sessile, closely and rather finely serrate
with somewhat incurved, mucronate teeth, slightly paler beneath, conspicuously
short- villosulous or puberulent beneath on the veins; inflorescence very large and
usually much branched, the peduncles and pedicels scabrous; umbels many-radiate,
the rays straight or nearly so, spreading, subequal, the pedicels 6-8 mm. long;
bracts and bractlets few, linear or lance-linear, scarious-marginate; fruit 10-12
mm., long and 4 mm. broad, slender-rostrate; carpophore 2-cleft.

In Mexico, the plant is reported to attain a height of 6 meters
which, if true, indicates a much greater size than has been observed
in Central America.

ERYNGIUM L.

References: Hermann Wolff, Pflanzenreich IV. 228: 106-271. 1913.
Mildred E. Mathias and Lincoln Constance, A synopsis of the North
American species of Eryngium, Amer. Midi. Nat. 25: 361-387. 1941.

Biennial or perennial herbs, caulescent or acaulescent, usually glabrous, with
stout taproots or rootstocks; leaves coriaceous or membranaceous, simple and en-
tire to pinnately or palmately lobate or cleft, often with spine-tipped teeth or divi-
sions, the venation parallel or reticulate; petioles vaginate; inflorescence capitate,
the heads solitary, cymose, or racemose; involucre of 1 or more series of entire or
lobate bracts at the base of the head; involucel of entire or lobate bractlets sub-
tending the flowers, these white to purple, sessile; petals ovate to oblong, the tips
inflexed and lobate to fimbriate; sepals ovate to lanceolate, acute or obtuse, entire
or rarely spinose-dentate; stylopodium none; fruit globose to obovoid, scarcely
compressed laterally, variously covered with scales or tubercles; ribs obsolete, the
commissure broad; seed sub terete in cross section, the face plane or slightly concave.

Species about 200, widely distributed in both hemispheres, in
North America most numerous in Mexico. At least one other spe-
cies is found in Central America.

Leaves with parallel veins, narrow spine-tipped lobes E. cymosum.

Leaves with reticulate venation.

Leaves acute to attenuate at the base, the petiole broadly marginate to the base,
the teeth tipped with long sharp stiff spines.

Bracts white or bluish on the inner surface E. carlinae.

Bracts green on the inner surface E. foetidum.

Leaves cordate to obtuse at the base, on long naked petioles, the teeth obtuse or
with very short spines.

Bracts of the involucre lobate or coarsely dentate; leaf blades not cordate at
the base E. scaposum.

Bracts of the involucre entire; leaf blades deeply cordate to subcordate at the
base.

44 FIELDIANA: BOTANY, VOLUME 24

Leaves with conspicuous callous margins, the teeth muticous . . . . E. gracile.

Leaves not callous-marginate, the teeth with short spinose tips.

E. ghiesbreghtii.

Eryngium carlinae Delwar. Eryng. 54, t. 23. 1808. C. reptans
Hemsl. in Hook. Icon. 26, t. 254-3. 1897 (type from Quezaltenango,
Bernoulli & Cario 2745). E. affine Wolff, Fedde Rep. Sp. Nov. 7:
345. 1909. Escorzionera; pinapunta (Jalapa); quixtdn (Huehuete-
nango, perhaps an erroneous name).

Moist or dry meadows often in alpine regions, frequently on dry
rocky slopes or in pastures, sometimes on limestone, 1,000-3,400
meters; most frequent at the higher elevations; Alta Verapaz; Baja
Verapaz; Jalapa; Guatemala; Sacatepe"quez; Chimaltenango; Solola;
Quiche"; Huehuetenango; Totonicapan; San Marcos. Mexico; Costa
Rica; Panama.

Plants perennial from a thick root, the stems few or numerous, prostrate to
erect, usually less than 30 cm. high, the plants glabrous throughout, the stems
simple or branched, stout, naked or bearing only a few leaves; basal leaves usually
persistent in age and forming a rosette flat on the ground, oblanceolate or obovate,
mostly 3-10 cm. long, rounded or obtuse at the apex, long-attenuate to the base
and practically sessile, serrate or laciniate-lobate, the teeth spine-tipped; heads
1-many, ovoid or short-cylindric, usually bearing several large and conspicuous
bractlets at the apex, these equaling or often longer than the flowers, the heads
mostly 1 cm. long or shorter; bracts at the base of the head 8-10, bluish or whitish,
rigid, broadly lanceolate or oblanceolate, 10-15 mm. long, with a few spine- tipped
teeth or rigid lobes.

A common plant in meadows of the higher mountains, persisting
in dry places through the dry season. Guatemalan material is highly
variable and some of it might be referred to related Mexican species
on the basis of the key in Mathias and Constance's account of the
genus. The key characters that they use, particularly the relative
length of bractlets and styles, evidently are not dependable for sep-
aration of species in this group of the genus. Either the close rela-
tives that they recognize in Mexico are not distinguishable from
E. carlinae or else better characters must be found for separating
them. In Alta Verapaz this plant is employed in domestic medicine,
especially as a remedy for diarrhea and for the treatment of some
afflictions peculiar to women. Although E. reptans is keyed by Wolff
and by Mathias and Constance far apart from E. carlinae, examina-
tion of the original illustration and study of the description reveal
no characters by which the two may be separated, despite Hemsley's
glib statement that E. reptans "is not closely related to any species,"
something that could not be said of many species in this genus of
poorly marked entities.

STANDLEY AND WILLIAMS: FLORA OF GUATEMALA 45

Eryngium cymosum Delar. Eryng. 63, t. 31. 1808. E. guate-
malense Hemsl. in Hook. Icon. IV. 8: sub t. 2766. 1903 (type collected
near Hacienda Chancol, Huehuetenango, E. W. Nelson 3654). Es-
corzionera; espino; escarzionera; cardosanto; c'ax (Quezaltenango) .

FIG. 11. Eryngium cymosum. A, Habit; X Y^. B, Flower; X 7.

46 FIELDIANA: BOTANY, VOLUME 24

Moist or dry thickets, meadows, or open hillsides, often in rocky
places, open, pine or Juniperus forest, sometimes on limestone, 1,650-
3,750 meters; El Progreso; Jalapa; Guatemala; Sacatepe"quez ; Chi-
maltenango; Huehuetenango; Totonicapan; Quezaltenango; San
Marcos. Southern Mexico.

A very coarse and stout perennial, glabrous throughout, pale green, usually
1-1.5 meters high, the stems terete, striate, simple below, bearing few or numer-
ous leaves; basal leaves usually numerous, linear or nearly so, mostly 10-30 cm.
long, deeply pectinate-lobate, the lobes narrow, spine-like, very stiff, spreading,
sessile or essentially so, parallel-nerved; inflorescence usually much branched and
bearing numerous heads; bracts at the base of the head 10-12, 3.5 cm. long or
shorter, spreading or reflexed in age, entire, linear-lanceolate, rigid and tipped
by a stout sharp spine, longer than the head; flower heads globose, about 1.5 cm.
in diameter; bractlets subulate, longer than the flowers; sepals acute or very obtuse,
mucronate; petals 2 mm. long; fruit 2.5-4 mm. long, densely squamate.

In their monograph of Eryngium Mathias and Constance consid-
ered E. guatemalense a distinct species, separating it on the basis of
its greater size, obviously not an important or stable character, and
the obtuse rather than acute sepals, a character that is variable
and apparently worthless, to judge from Mexican and Guatemalan
material examined. E. cymosum has been reported from Guatemala
as E. pectinatum Presl. It is a very common and conspicuous plant
in the higher mountains of Los Altos, especially in those areas where
coarse bunch-grass (Epicampes) abounds, and where large numbers
of sheep are pastured. The plant is an offensive one, as well protected
as any thistle (Cirsium), and a great pest in the regions where it
occurs. Obviously, it is left severely alone by grazing animals, but
the sheep probably help to spread it by carrying the small burlike
fruits in their wool. It seems to be most plentiful in those too numer-
ous parts of Guatemala where sheep grazing has been carried to excess.

Eryngium foetidum L. Sp. PI. 232. 1753. Culantro; culantro
extranjero; culantro real; alcapate; escorzonera; samat; xamat (Coban;
Quecchi) ; silantro cimarron (Pete*n, fide Lundell) .

Wet or moist, open banks or pastures, sometimes in waste ground ;
1,300 meters or less; often cultivated in gardens; Pete"n; Alta Vera-
paz; Izabal; Jutiapa; Escuintla. Mexico; British Honduras to El
Salvador and Panama. West Indies; South America; naturalized in
Africa and Asia.

Plants perennial from a cluster of rather thick, fleshy roots, the stems solitary,
simple, naked, or sometimes several, frequently branched and often leafy, the whole
plant glabrous and strong-scented; basal leaves usually numerous and forming a
rosette, narrowly or broadly oblanceolate, mostly 8-20 cm. long, obtuse, attenuate

STANDLEY AND WILLIAMS: FLORA OF GUATEMALA 47

to the base and essentially sessile, crenate or usually finely or coarsely serrate, the
teeth tipped with a short yellowish spine; heads few or numerous, usually about
1 cm. long, cylindric, yellowish green; involucral bracts 5-6, foliaceous, lanceolate,
acute, entire or sparsely spinulose-serrate, sometimes 3 cm. long but mostly shorter;
bractlets at the apex of the head usually large, foliaceous, and forming a conspicu-
ous apical tuft; sepals lanceolate to triangular-ovate, 1 mm. long or less; petals
elliptic-oblong, less than 1 mm. long; fruit globose-ovoid, 2 mm. long, densely
squamate.

Sometimes called "acapate" in El Salvador and Honduras. It is
a weedy plant, widely distributed in Central America, found princi-
pally in waste ground near dwellings. It is well known in the region
because of its use in the kitchen. The fresh plant has a very strong
and most nauseous odor, but when boiled in soups or stews it imparts
to them a delicious flavor, such as could never be expected from the
living plant. The plant is used to some extent in household medicine
in Guatemala.

Eryngium ghiesbreghtii Decaisne, Bull. Soc. Bot. France 20:
21. 1873. Lechugilla (Huehuetenango) .

Moist or rather dry, oak forest, or on open hillsides or fields,
1,000-2,200 meters; Baja Verapaz; Santa Rosa; Chimaltenango;
Guatemala; Quiche"; Huehuetenango. Mexico; reported by Wolff
from Bolivia.

Plants perennial from a cluster of rather thick fleshy roots, glabrous, erect,
75 cm. high or less, the stems solitary or few, somewhat fistulous, striate, simple or
branched below, usually somewhat leafy; basal leaves often numerous, on very
long, slender, rather weak petioles, cordate-triangular, as much as 10 cm. long and
6 cm. broad, acute or subacute, shallowly or deeply cordate at the base, densely
and finely crenate with spinulose-mucronate teeth, 7-9-nerved from the base, the
sinus broad and open, much paler beneath; upper cauline leaves sessile and clasp-
ing; bracts at the base of the head rigid, linear-lanceolate, about 1 cm. long, pale
green with white nerves, acute and spine-tipped, entire or nearly so, conspicuously
parallel-nerved; heads globose or depressed-globose, greenish white, much shorter
than the involucre; sepals triangular-lanceolate, acute or obtuse, short-mucronate,
1.5 mm. long; petals broadly spatulate, deeply excised; styles much longer than
the sepals; fruit compressed-globose, 2 mm. long, densely squamate.

Eryngium gracile Delar. Eryng. 54, t. 24- 1808. E. longirameum
Turcz. Bull. Soc. Nat. Moscou 20, pt. 1: 171. 1847.

Moist or rather dry, oak or pine forest, or in alpine meadows,
1,800-3,300 meters; Jalapa; Chimaltenango; Quiche"; Huehuete-
nango. Southern Mexico.

48 FIELDIANA: BOTANY, VOLUME 24

Plants glabrous, perennial from a cluster of slender, somewhat fleshy roots, the
stems erect, simple or sparsely branched, usually 50 cm. high or less, generally
bearing a few leaves below; basal leaves few, on long slender petioles, ovate-trian-
gular or ovate-oblong, 5 cm. long or less, acute or obtuse, broadly rounded or cor-
date at the base, the margins narrowly cartilaginous-thickened, finely and closely
crenate, the teeth not or scarcely apiculate, 7-9-nerved at the base, paler beneath;
heads usually few, cobalt-blue; subglobose, 5 mm. in diameter; bracts about 10,
subequal, much longer than the heads, elliptic-oblong or spatulate, subobtuse,
mucronulate, entire, 5-nerved, about 1 cm. long, white tinged with blue on the
inner surface; sepals ovate, obtuse or truncate, short-mucronate, petals oblong-
spatulate, shallowly excised.

Eryngium scaposum Turcz. Bull. Soc. Nat. Moscou 20, pt. 1:
172. 1847.

Alpine meadows, 3,150-3,500 meters; Totonicapan; Huehuete-
nango. Southern Mexico; Costa Rica.

Plants glabrous, perennial, arising from a short rootstock bearing numerous
long, somewhat fleshy roots, the stems 10-20 cm. high, usually naked and scape-
like, simple or 2-3-furcate, solitary or several; basal leaves few, usually on petioles
about as long as the blades, the petioles naked or winged; leaf blades elliptic to
lance-oblong, 2-5 cm. long, obtuse or rounded at the apex, abruptly contracted at
the base into the petiole and obtuse to rounded or sometimes acute, usually shal-
lowly and inconspicuously crenate, the teeth muticous, depressed; heads solitary
or 2-3, bluish, subglobose, much shorter than the bracts, 6-8 mm. long; bracts of
the involucre about 8, linear-lanceolate, acute or obtuse, muticous or spinulose-
acute, about 1 cm. long, entire or nearly so, whitish on the inner surface; bractlets
equaling or longer than the flowers; sepals ovate-lanceolate, acuminate or obtuse
and mucronate; petals oblong, deeply excised; styles longer than the sepals; fruit
globose-ovoid, 2.5 mm. long, densely squamate.

FOENICULUM Adanson. Fennel

Erect, biennial or perennial, glabrous, often much branched herbs; leaves often
large, pinnately decompound, the segments linear or capillary; flowers small, yel-
low, in compound umbels; involucre and involucels none; calyx teeth obsolete;
petals obtuse or somewhat retuse; stylopodium large, conic; fruit linear-oblong,
terete or nearly so; mericarps semiterete, compressed dorsally, conspicuously cos-
tate; face of the seed plane or slightly concave.

About four species, natives of the Old World.

Foeniculum vulgare Miller, Gard. Diet. ed. 8. Foeniculum 1.
1768. Anethum foeniculum L. Sp. PI. 263. 1753. Hinojo.

Native of Europe, often cultivated in other parts of the earth;
planted rarely in Guatemala, and uncommon.

Plants erect, perennial, usually much branched, a meter high or less, glauces-
cent; leaves much dissected into numerous long capillary segments, the petioles

STANDLEY AND WILLIAMS: FLORA OF GUATEMALA 49

broad and amplexicaul; umbels large, 9-25-rayed, the rays stout, often glaucous,
2.5-7 cm. long, the pedicels 2-8 mm. long; fruit about 6 mm. long.

In some parts of Europe the plant is grown for its enlarged tender
leaf bases, which are eaten like celery.

HYDROCOTYLE L.

References: J. N. Rose and Paul C. Standley, The Central Amer-
ican species of Hydrocotyle, Journ. Wash. Acad. Sci. 17: 194-197. 1927.
Mildred E. Mathias, The genus Hydrocotyle in northern South Amer-
ica, Brittonia 2: 201-237. 1936.

Small perennials with long slender creeping stems or with rootstocks, often
growing in water or very wet soil, glabrous or pubescent; leaves long-petiolate,
peltate or epeltate, usually more or less orbicular in outline; inflorescence a simple
umbel, sometimes proliferous, or the flowers in interrupted spikes, the peduncles
axillary; involucre none or sometimes present; flowers white, greenish, or yellow,
very small; petals ovate, flat; calyx teeth minute or obsolete; stylopodium de-
pressed to conspicuous and conic; fruit transversely ovate to orbicular, strongly
compressed laterally, the dorsal surface rounded or acute, the lateral costae con-
spicuous, slender, acute, rarely obscure; mericarps orbicular to triangular in cross
section.

About 75 species, widely distributed in tropical and temperate
regions. A very few species besides those listed here are found in
southern Central America.

Leaves not peltate.

Petioles glabrous; leaves deeply lobate H. ranunculoides.

Petioles pilose or puberulent; leaves not lobate.

Peduncles densely puberulent; pedicels usually longer than the fruit, often
several times as long H . mexicana.

Peduncles thinly villous; pedicels equaling or shorter than the fruit.

H. leucocephala.
Leaves peltate.

Umbels 2 or more on each peduncle.

Umbels compound H. bonariensis.

Umbels simple H. verticillata.

Umbels solitary on the peduncle.

Petioles glabrous; pedicels usually much longer than the fruit . . .H. umbellata.
Petioles pubescent; pedicels shorter than the fruit H. pusilla.

Hydrocotyle bonariensis Lam. Encycl. 3: 153. 1789. H. yuca-
tanensis Millsp. Field Mus. Bot. 2: 81. 1900.

In shallow water, 300 meters; Huehuetenango; probably collected
also in some other Guatemalan localities. Southeastern United States;
Mexico; Nicaragua; Costa Rica; Panama. South America.

50 FIELDIANA: BOTANY, VOLUME 24

Plants glabrous, arising from tuberous roots, the stems slender, creeping;
leaves on long slender petioles, peltate, orbicular or nearly so, 1.5-5 cm. in diam-
eter, very shallowly crenate-lobulate; peduncles longer than the leaves, bearing
one or more umbels, these proliferous, many-flowered, the flowers white or yellow;
pedicels 2-20 mm. long, spreading and often reflexed; fruit transverse-ovate, 1-2
mm. long, 2-4 mm. broad, with distinct acute ribs; stylopodium depressed.

Hydrocotyle leucocephala Cham. & Schlecht. Linnaea 1: 364.
1826. H. maxonii Rose in Rose & Standl. Journ. Wash. Acad. Sci.
17: 197. 1927.

Moist or wet meadows or thickets, 1,300-1,600 meters; Alta Vera-
paz. Mexico; Costa Rica. South America.

Stems very slender, creeping, glabrous; leaves on long slender petioles, these
laxly and rather sparsely villous; leaf blades orbicular-reniform, not peltate, 1-5
cm. broad, 9-11-nerved, glabrous or nearly so, very shallowly truncate-crenate or
truncate-lobate; peduncles, at least in age, longer than the leaves, villous, espe-
cially toward the apex; umbels simple, 20-30-flowered, the pedicels 1-3 mm. long,
spreading and reflexed, glabrous; flowers cream-colored; fruit suborbicular, 1 mm.
long, 1-1.5 mm. broad, glabrous, the ribs filiform, acute; stylopodium depressed.

The plant is rare in Guatemala. It is reported as collected at
"Coban" by Tuerckheim, but this locality means little in the case
of Tuerckheim collections. Many of those so labeled undoubtedly
were obtained many miles distant from the city of Coban.

Hydrocotyle mexicana Cham. & Schlecht. Linnaea 5: 208. 1830.
Berro cimarronfynaha Luisa de monte.

Moist or wet, mixed forests, or sometimes in pine or Cupressus
forest, open banks, thickets, usually in rich soil, occasionally in
marshy places, 1,200-3,500 meters; Alta Verapaz; El Progreso; Za-
capa; Jalapa; Jutiapa; Escuintla; Guatemala; Sacatepe"quez; Chi-
maltenango; Solola; Quiche"; Retalhuleu; Huehuetenango; Quezalte-
nango; San Marcos. Mexico; El Salvador to Panama. Northwestern
South America.

Plants small or often much elongate, creeping, often forming dense colonies or
mats, the stems glabrous or hirsutulous; leaves on very long, more or less hirsute
or puberulent petioles, orbicular-reniform, not peltate, 1-5 cm. broad, sparsely or
densely hirsutulous on the nerves and veins, shallowly 7-11-lobate, the lobes cre-
nate; peduncles slender, usually longer than the leaves, bearing a single simple
20-70-flowered umbel; flowers pale green or whitish, the pedicels 1-7 mm. long,
spreading and reflexed, glabrous; fruit suborbicular, 1-1.5 mm. long and broad,
glabrous, the ribs distinct, acute; stylopodium depressed.

This is perhaps the plant reported from Guatemala by Hemsley
as H. bonplandii A. Rich., a South American species. H. mexicana is
very common in dense rich forest at middle elevations, often occur-
ring almost everywhere in such places and in the greatest abundance.

X

i

•S
s

o

o

o

IH

X

X

i

a

N

1— I

V

52 FIELDIANA: BOTANY, VOLUME 24

Hydrocotyle pusilla A. Rich. Ann. Ge"n. Sci. Phys. 4: 167, t. 52,
/. 2. 1820. H. costaricensis Rose in Rose & Standl. Journ. Wash.
Acad. Sci. 17: 195. 1927.

Dense wet mixed forest, or on wet banks, especially along streams,
or on wet rocks, 1,500-3,000 meters; Alta Verapaz; Jalapa; Guate-
mala; Chimaltenango; San Marcos. Mexico; Costa Rica; Panama.
Greater Antilles; South America.

Stems filiform, creeping, often forming dense interlaced mats; leaves on slender
petioles 0.5-3 cm. long, these glabrous to densely retrorse-villous; leaf blades orbic-
ular, peltate, 5-20 mm. broad, glabrous or sparsely villous above, glabrous beneath,
shallowly 5-8-lobate, the lobes crenate; peduncles 5-15 mm. long, glabrous or
retrorse-villous; umbels simple, solitary, 2-6-flowered, the flowers sessile or nearly
so, greenish or pinkish; fruit transverse-elliptic, 0.5 mm. long, 1 mm. broad,
sparsely hirsute or glabrous, the ribs obsolete; stylopodium depressed-conic.

Hydrocotyle ranunculoides L. f. Suppl. 177. 1781.

In mud along the borders of streams, or floating in shallow water,
1,600-2,450 meters; Chimaltenango; Quezaltenango. United States;
Mexico; Nicaragua; Panama; Cuba; South America.

Plants rather stout and succulent, yellowish green, glabrous, the stems floating
or creeping; leaves long-petiolate, not peltate, orbicular-reniform, 1-8 cm. in diam-
eter, conspicuously 5-6-lobate, the lobes crenate or lobulate; peduncles shorter
than the leaves, bearing a single simple umbel, this 5-10-flowered; pedicels 1-3 mm.
long, spreading and ascending; fruit suborbicular, 1-3 mm. long, 2-3 mm. broad,
the ribs obscure; stylopodium depressed.

Hydrocotyle umbellata L. Sp. PI. 234. 1753. Lechugilla; naab
mejen (Pete"n, Maya, fide Lundell) .

Floating in shallow water or creeping in wet soil, often in marshes
or at the margins of streams or ponds, sometimes on gravel or sand-
bars along streams, 2,400 meters or less; Pete"n; Baja Verapaz; Izabal;
Zacapa; Chiquimula; Jutiapa; Santa Rosa; Escuintla; Guatemala;
Sacatepe"quez; Suchitepe"quez; Chimaltenango; Solola; Quiche"; Hue-
huetenango; San Marcos. United States; Mexico; British Honduras
to El Salvador and Panama. South America.

Plants perennial, glabrous, the stems slender or often somewhat fleshy, float-
ing or creeping; leaves long-petiolate, peltate, orbicular, 1-6 cm. broad, crenate or
very shallowly crenate-lobate; peduncles usually longer than the leaves, bearing
a single simple umbel, this many-flowered; pedicels 2-25 mm. long, spreading to
reflexed; flowers greenish yellow; fruit suborbicular, 1-2 mm. long, 2-3 mm. broad,
glabrous, the ribs distinct, obtuse; stylopodium depressed.

Sometimes called "lechuga" in El Salvador. This has been re-
ported from Guatemala as H. prolifera Kellogg.

STANDLEY AND WILLIAMS: FLORA OF GUATEMALA 53

Hydrocotyle verticillata Thunb. Diss. 2: 415, t. 3. 1798.
A species of wide distribution in the United States, represented
in Central America by the following variety:

Hydrocotyle verticillata var. triradiata (A. Rich.) Fernald,
Rhodora 41: 437. 1939. H. polystachya var. triradiata A. Rich. Ann.
Gen. Sci. Phys. 4: 171. 1820. H. racemosa Sesse" & Moc. ex DC.
Prodr. 4: 70. 1830. H. prolifera Kellogg, Proc. Calif. Acad. 1: 15.
1854. H. verticillata var. racemosa Mathias, Brittonia 2: 204. 1936.
Hierba de jute.

In wet soil, 1,500 meters or less; Pete"n; Guatemala. United
States; Mexico; El Salvador; Costa Rica; South America.

A glabrous perennial, the stems slender, creeping; leaves on long slender peti-
oles, peltate, orbicular, 1-6 cm. broad, 8-14-nerved, very shallowly 8-14-lobate;
peduncles slender, bearing a spike of few simple umbels, these 4-15-flowered, the
pedicels 1-10 mm. long, spreading or ascending; fruit oblong-ovoid, 1-3 mm. long,
2-4 mm. broad, the ribs distinct, acute; stylopodium depressed.

MICROPLEURA Lagasca

Herbs, glabrous or nearly so, arising from a cluster of small tuberous roots,
caulescent; leaves long-petiolate, simple, reniform, palmate-nerved, crenate, the
petiole abruptly dilated and scarious at the base; umbels long-pedunculate, irreg-
ularly compound; bracts of the involucre 1-2, small; flowers unisexual, the um-
bellule usually bearing 1 subsessile pistillate flower and 2 pedicellate staminate
ones; calyx teeth obsolete; petals entire, broadly imbricate; disk flat, the styles
filiform to the base; fruit laterally compressed, with a narrow commissure, emargi-
nate at the base; mericarps laterally compressed J carpophore none; seed laterally
compressed.

The genus consists of a single species. It is hardly distinct from
Centella although we follow Mathias and Constance in maintaining
it. The stems are erect or suberect and arise from tuberous roots.
The stems of Centella are usually prostrate and root at the nodes,
lacking tubers.

Micropleura renifolia Lagasca, Ocios Esp. Emigr. 4: 347. 1825.
Hydrocotyle grumosa DC. Prodr. 4: 70. 1830. Centella renifolia Urban
in Mart. Fl. Bras. II1: 347. 1879. Canan (Huehuetenango) ; guani-
machaj (fide Aguilar) .

Wet to rather dry fields, banks, or forest, frequently in pine forest,
1,400-3,100 meters; Baja Verapaz; Chiquimula; Jalapa; Santa Rosa;
Guatemala; Chimaltenango; Quiche"; Huehuetenango. Mexico;
Honduras; Costa Rica; South America.

FIG. 13. Micro-pleura renifolia. A, Habit; X %. B, Fruit; X 9. C, Fruit
as seen from above; X 9.

54

FIG. 14. Myrrhidendron donnellsmithii. Leaf and inflorescence of plant;
about X H- Fruit from side; about X 2. Cross-section of fruit; about X 8.
[Illustration, drawn by C. E. Faxon, is from the original publication. Courtesy
of The University of Chicago Press.]

55

56 FIELDIANA: BOTANY, VOLUME 24

Plants arising from small thick tubers, the stems erect or suberect, short or
elongate, often branched, the internodes much elongate; leaves on long slender
petioles, these usually longer than the blades; leaf blades 2-7 cm. broad, membra-
naceous, broadly rounded, deeply but openly cordate at the base, conspicuously
and evenly crenate except at the base, much paler beneath; inflorescence openly
branched, the umbellules few or numerous, long-pedunculate or almost sessile, the
flowers white, short-pedicellate or subsessile, the peduncles often somewhat pubes-
cent; fruit greenish, about 2 mm. long and 3-4 mm. broad, glabrous, deeply con-
stricted between the mericarps, the styles very short.

MYRRHIDENDRON Coulter & Rose

Large coarse perennial herbs or shrubs, or even small weak trees, simple or
branched, the stems very thick, hollow, the whole plant almost glabrous; leaves
alternate, very large, 3-4 times ternate or ternate-pinnate, the leaflets numerous,
petiolulate or sessile, acuminate, serrate, ovate or oblong-ovate; flowers white, the
umbels compound; bracts and bractlets conspicuous, dentate or lobate; calyx teeth
obsolete; fruit linear or oblong-linear, dorsally compressed, glabrous; mericarps
strongly compressed dorsally, the dorsal and intermediate costae slender, the lat-
eral costae narrowly winged; stylopodium low-conic; seeds strongly compressed,
the inner face plane.

One additional species from the mountains of Panama; others in
the Andes.

Myrrhidendron donnellsmithii Coult. & Rose, Bot. Gaz. 19:
466, t. 32. 1894. In mountain forest, 2,500-3,200 meters; Quezalte-
nango. Costa Rica.

Plants 10 meters high, the stems very thick, more or less woody below; leaves
very large, the lower ones long-petiolate, the upper ones with large broad sheathing
petioles; leaflets numerous, ovate or lance-ovate, mostly 2.5-5 cm. long, acute to
long-acuminate, rather thick and firm, unequally serrate, paler beneath, glabrous
or nearly so, the petiolules with a dense tuft of short stout hairs at the base; bracts
of the involucre few, dentate; bractlets numerous, several-dentate or lobate, scari-
ous-marginate; inflorescence large and much branched, the umbels scabrous-puber-
ulent, the rays numerous, subequal, the pedicels mostly 8-10 mm. long; fruit
10-12 mm. long, often dark purple.

This genus is not known to occur between Guatemala and Costa
Rica. The single collection from Guatemala (Volcan Zunil, Steyer-
mark 34750) seems to belong to Myrrhidendron but better material
is needed to verify whether the Guatemalan plant belongs to the
species given here.

OREOMYRRHIS Endlicher

Reference: Mildred E. Mathias and Lincoln Constance, The Genus
Oreomyrrhis, Univ. Cal. Publ. Bot. 27: 347-416. 1955.

STANDLEY AND WILLIAMS: FLORA OF GUATEMALA 57

Perennial herbs, usually dwarf, pubescent or almost glabrous, the stems short,
scapiform, often cespitose; leaves basal, pinnately dissected or decompound, usu-
ally with very numerous, small segments; umbels simple, solitary at the apex of
the scape; bracts of the involucre numerous, ovate, lanceolate, or linear; calyx
teeth obsolete; petals oblong, acute, incurved at the apex, entire; stylopodium de-
pressed or subconic; fruit oblong or oblong-linear, usually contracted at the apex,
slightly compressed laterally; mericarps subterete, conspicuously 3-costate, the
costae obtuse; carpophore 2-parted; seed subterete, sulcate on the inner face.

About 23 species, in the mountains from Mexico to Patagonia,
and in New Zealand, Australia and Tasmania to Borneo and For-
mosa. No other species is found in Central America.

Oreomyrrhis daucifolia I. M. Johnston, Journ. Arn. Arb. 19:
125. 1938.

Moist or dry, alpine meadows, sometimes on dry rocky slopes,
2,200-3,900 meters; endemic; Huehuetenango (Sierra de los Cuchu-
matanes, the type from Chancol, A. F. Skutch 1263).

Plants as much as 30 cm. high but usually 15 cm. tall or less, arising from a
short thick hard root; stems usually 5-8, simple, stout, naked or bearing 1-2 leaves;
leaves all or mostly basal, 3-pinnate, the rachis puberulent, the pinnae 5-10 pairs,
6-20 mm. long, the ultimate divisions linear or narrowly lanceolate, very numerous
and often densely crowded, 1-3 mm. long, setuliferous at the apex; peduncles much
longer than the leaves, puberulent, especially above; umbels 10-20-flowered, dense;
bracts of the involucre pinnatisect; pedicels unequal, 1-5 mm. long; petals white
tinged with purple; fruit 4-5 mm. long, 1-1.3 mm. broad, broadest below the
middle, glabrous at maturity and purplish.

One of the typical alpine plants of the highest areas of Guatemala.

OSMORHIZA Rafinesque

Perennial herbs, pubescent or glabrous, usually arising from a cluster of rather
slender, tuberous-thickened roots, the stems simple or branched; leaves ternate-
pinnate, the numerous segments pinnatifid and dentate; flowers small, white, the
umbels compound, open, with few rays; bracts of the involucre very few or none,
the bractlets of the involucels numerous, narrow, and deciduous, or none; calyx
teeth obsolete; petals obovate or oblong, with an inflexed acumen; stylopodium
small, conic, entire; fruit oblong-linear or clavate, somewhat narrowed at the apex,
long-attenuate to the base, subterete, constricted at the commissure; mericarps
somewhat 5-angulate, slightly compressed dorsally, thinly ciliate on the low ribs;
carpophore slender, bifid; seed dorsally compressed, broadly sulcate on the face.

Perhaps a dozen species in America and Asia, chiefly in temperate
regions; in the tropics confined to the higher mountains. Only one
species occurs in Central America.

58 FIELDIANA: BOTANY, VOLUME 24

Osmorhiza mexicana Griseb. Goetting. Abh. 24: 147. 1879.
Pimpinela (Quezaltenango).

Moist or wet, mixed, mountain forest, or often in forest of pine,
Abies, or Juniperus, 2,400-3,700 meters; Sacatepe'quez ; Solola; Que-
zaltenango; Huehuetenango. Mexico; Costa Rica.

Plants erect, a meter high or less, often much branched, rather weak, arising
from a cluster of slender fleshy roots, sparsely pilose with whitish hairs; leaves
rather large, very thin and soft, bright green, the lower ones long-petiolate, the
uppermost cauline leaves short-petiolate, the segments rather numerous, deeply
lobate and irregularly dentate; umbels large, open, with about 5 long slender rays,
the flowers on long slender pedicels; fruit linear-clavate, 1-1.5 cm. long, less than

2 mm. broad.

OTTOA HBK.

Glabrous perennial herbs; leaves mostly radical, entire, terete, fistulose, trans-
versely septate; stems simple, naked or bearing 1-2 leaves; flowers small, white,
usually polygamous, the umbels compound; involucre and involucels none; calyx
teeth obsolete; petals oblong, narrowed into a long inflexed acumen, entire at the
apex; stylopodium thick-conic; fruit oblong, laterally compressed, constricted at
the commissure; carpels somewhat 5-angulate, the central rib obtuse, inconspic-
uous, the lateral ones distinct; seed subterete, the race deeply sulcate.

The genus consists of a single species, which may be recognized
at once by its terete fistulous leaves.

Ottoa oenanthoides HBK. Nov. Gen. & Sp. 5: 20, t. 423. 1821.

Mostly in exposed or shaded, rocky places, especially about the
summits of mountain peaks, sometimes in high pine forest, 3,300-
4,000 meters; Sacatepe'quez ; Huehuetenango; Quezaltenango; San
Marcos. Southern Mexico; northwestern South America.

Plants erect, usually 15-40 cm. high, the stems often very numerous from a
thick, apparently perpendicular, fleshy root; leaves several at the base of each
stem, rather slender, terete, 35 cm. long or shorter; umbels few-rayed, the rays
very unequal, the umbellules few-flowered, the central flower sessile or nearly so,
the others on stout pedicels as much as 5 mm. long; fruit about 5 mm. long and

3 mm. broad.

It is rather strange that this plant has not been found in Costa
Rica, where there are regions well suited to its growth. Its absence
there would lead one to suspect that the North American plant might
be a distinct species, but the collections from Guatemala and Mexico
apparently do not differ from those made in Ecuador.

PETROSELINUM Hoffmann. Parsley

Annuals or biennials with strong-scented foliage, usually glabrous, branched;
leaves ternately bipinnatisect or pinnatisect; flowers small, perfect or polygamous,
yellow or white, in compound umbels; bracts and bractlets of the involucels nar-

STANDEE Y AND WILLIAMS: FLORA OF GUATEMALA 59

row, entire; calyx teeth small and inconspicuous; petals subequal, orbicular or oval,
emarginate, short-unguiculate; stylopodium short-conic, the styles short, erect or
reflexed; fruit subglobose or subcordate-ovoid, more or less constricted at the com-
missure; costae of the mericarps filiform, equal, the mericarps coherent at base
and apex, subpentagonal in cross section, the face flat.

Three species, natives of Europe and other parts of the Mediter-
ranean region.

Petroselinum crispum (Mill.) Nyman ex Kew Hand-List Herb.
PL ed. 3. 122. 1925. Apium Petroselinum L. Sp. PI. 264. 1753. A.
crispum Mill. Gard. Diet. ed. 8. no. 2. 1768. P. hortense Hoffmann,
Gen. Umbell. 163. 1814. P. sativum Hoffmann, op. cit. 177. 1814.
Perejil.

Native of the Mediterranean region, but cultivated in almost all
regions of the earth; planted commonly in Guatemalan gardens.

Plants biennial or annual, glabrous, erect from a slender or fusiform root, the
stems usually several, fistulose, leafy, branched; lower leaves long-petiolate, 2-3-
pinnatisect, the pinnae and pinnules petiolulate, the leaflets ovate-lanceolate to
broadly linear, trifid or lobulate; umbels numerous, long-pedunculate, 10-20-rayed,
the rays subequal; bracts of the involucre few, narrow, shorter than the rays;
bractlets of the involucels 5-6; umbellules 10-15-flowered; fruit ovoid or oblong-
ovoid, 2-3 mm. long.

Parsley is grown commonly in Guatemalan gardens for use in
garnishing and flavoring food. It is sold frequently in the markets
of the larger cities.

PIMPINELLA L. Anise

Chiefly perennial herbs, sometimes annuals, glabrous or often pubescent about
the inflorescence; leaves pinnate or ternately or pinnately decompound; flowers
small, white or yellow, the umbels compound ; bracts of the involucre none or rarely
1-2, the bractlets of the involucels few and small or usually wanting; calyx teeth
generally obsolete; petals commonly narrowed into an inflexed acumen, retuse or
entire; stylopodium pulvinate or broadly or narrowly conic; styles usually elon-
gate, rarely short; fruit ovate or broader than long, more or less compressed later-
ally, usually constricted at the broad commissure; mericarps slightly 5-angulate,
dorsally compressed, the ribs slender and small; carpophore bifid or 2-parted; seed
subterete or dorsally compressed, the face somewhat convex, plane, or slightly
concave.

About 70 species, mostly in the Old World, generally in temper-
ate or warm-temperate regions. None are native in Central America.

Pimpinella anisum L. Sp. PI. 264. 1753. Anis.

Native of the eastern Mediterranean region; grown frequently in
other parts of the earth for its aromatic fruits; cultivated occasion-
ally in Guatemala but uncommon.

60 FIELDIANA: BOTANY, VOLUME 24

Plants annual, erect, branched, minutely puberulent almost throughout; low-
est leaves rounded-cordate and incised, the middle ones trisect, the segments cune-
ate and lobate, the upper leaves sessile, cleft into linear or trifid segments; bracts
and bractlets linear or often none; petals puberulent; fruit minutely appressed-
pubescent, ovoid-pyriform, narrowed at the apex.

The "seeds" are employed for flavoring various kinds of foods,
but they are not popular in Central America.

PRIONOSCIADIUM Watson

Large coarse perennial herbs, glabrous or sparsely pubescent; leaves large, 2-3
times ternate-pinnatifid, the segments usually deeply lobate or dentate; flowers
yellow, the umbels compound; involucre none, the bractlets of the involucels small
and narrow; calyx teeth short but usually equaling or surpassing the depressed
stylopodium; fruit oval-rounded, dorsally compressed, with a broad commissure;
lateral costae of the mericarps winged, the intermediate and dorsal ones acute or
narrowly winged; seeds dorsally compressed, with involute margins; carpophore
2-parted.

About seventeen species in Mexico, one reaching Guatemala.

Prionosciadium thapsoides (DC.) Mathias, Brittonia 2: 245.
1936. Elaeoselinum thapsoides DC. Prodr. 4: 215. 1830. Angelica
mexicana Vatke, Ind. Sem. Berol. App. 2. 1876. P. mexicanum Wats.
Proc. Amer. Acad. 23: 275. 1888. P. seleri Rose, Contr. U. S. Nat.
Herb. 8: 335. 1905 (type from Quezaltenango, the exact locality not
indicated, C. & E. Seler 3106). Axan xemel (fide Seler); cuscuta de
sapo (fide Aguilar) ; pimientillo (Huehuetenango) .

Dry rocky open forest of oak and pine, or on brushy hillsides,
1,600-2,600 meters; Baja Verapaz; Sacatepe"quez; Solola; Quiche";
Quezaltenango; Huehuetenango. Mexico.

Plants coarse and stout, as much as 3 meters high, the stems not hollow, terete,
leafy below and with reduced leaves at the base of the inflorescence; radical leaves
very large, long-petiolate, the blades often 35 cm. long or more and equally broad,
ternate, the divisions pinnately decompound, the leaflets or their lobes cuneately
long-decurrent on the rachis, obtuse, rather thick, crenate-serrate with mucronate
teeth, glabrous above, somewhat puberulent or scabrous beneath on the veins with
short thick hairs; petioles of the lower cauline leaves broadly winged and sheath-
ing; inflorescence usually large and much branched, the branches scabrous; umbels
compound, about 7 cm. broad, with numerous stout unequal rays, the bractlets
lanceolate, scaberulous, the pedicels stout, very unequal; fruit when fully mature
about 13 mm. long and 8 mm. broad, the pale wings as much as 4 mm. broad,
glabrous.

While rather widely scattered in Guatemala, the plant is seldom
if ever plentiful. The large coarse leaves, unlike those of any other

STANDLEY AND WILLIAMS: FLORA OF GUATEMALA 61

local plant, are conspicuous during the dry season. The Indians of
Huehuetenango use the dry fruits for flavoring atol, so evidently they
have the aromatic properties common to most plants of the family.

RHODOSCIADIUM Watson

Perennial herbs, usually glabrous or nearly so; leaves large, the lower ones 2
or more times ternately or pinnately decompound ; inflorescences usually large and
much branched, the umbels compound; involucre none or of 1 or more bracts, the
bractlets linear; flowers often or usually purple; calyx teeth very small; stylopo-
dium depressed-conic, the disk with an undulate margin; fruit oblong or broader,
glabrous, compressed dorsally; mericarps with 3 filiform dorsal costae and with 2
marginal wings; seed flat or concave on the inner surface.

About 13 species, in Mexico and Guatemala.

Leaflets entire to crenate-serrate R. glaucum.

Leaflets serrate to encised or lobed R. diffusum.

Rhodosciadium diffusum (Coult. & Rose) Mathias & Con-
stance, Bull. Torr. Bot. Club 68: 124. 1941. Deanea diffusa Coult. &
Rose, Proc. Wash. Acad. Sci. 1: 155. 1900. Ineldo (a corruption of
Eneldo) .

Rocky hillsides, 1,900 meters; Huehuetenango. Mountains of
central and southern Mexico.

Plants tall, erect, glabrous, the stems hollow, thick, glaucous and purplish;
radical leaves very large, long-petiolate, 4-5 times ternate; leaflets numerous, thin,
2-3.5 cm. long, deeply cleft and dentate; inflorescence very large and much
branched, glabrous, the peduncles short or elongate; rays of the umbel stout,
straight, few or numerous, the pedicels 2-3 mm. long or longer in fruit, stout; in-
volucre none; involucels of few filiform bractlets; mericarps oblong or oval, 12-
14 mm. long, 8-10 mm. broad, cordate at the base, with broad thin wings; margins
of the seed involute, the face concave.

Rhodosciadium glaucum Coult. & Rose, Contr. U. S. Nat.
Herb. 3: 309. 1895; Mathias & Constance, N. Am. Fl. 28B: 215. 1945.
Guatemala (according to Mathias & Constance). Mexico.

Slender plants from a tuberous rootstock, 8-15 cm. tall, glaucous, the foliage
scaberulous. Leaves ovate-deltoid, the blade 5-25 cm. long, biternate or ternate-
pinnate, the leaflets lanceolate to ovate-lanceolate, acute or acuminate, cuneate
to the base, sessile, 2-10 cm. long and 0.5-2 cm. broad, crenate-serrate, slightly
incised, scabridulous on the veins, petiole to 35 cm.; inflorescence widely branched,
the peduncles slender, 2-8 cm. long, involucre none or of 1-2 small bracts, rays 3-7,
1-3 cm. long, about equal; fruit oval, retuse at the apex and at the base, 6-10 mm.
long and 5-8 mm. broad, dorsal ribs thin, the wings as broad or broader than the
body of the fruit, seed face slightly concave to sulcate.

We have seen no Guatemalan material of this species.

62 FIELDIANA: BOTANY, VOLUME 24

SANICULA L.

Reference: Hermann Wolff, Pflanzenreich IV. 228: 48-80. 1913.

Mostly perennial herbs; leaves ternate, palmately parted, or variously pinnate;
flowers small, usually yellow; the umbels simple or compound, often head-like, the
involucre conspicuous or greatly reduced; perfect flowers central, sessile or on very
short pedicels, solitary or several, borne in the same umbels with the few or numer-
ous, pedicellate staminate flowers; sepals conspicuous, free in the fertile flowers,
usually connate in the staminate flowers; petals imbricate, emarginate; stylopo-
dium explanate; styles filiform, longer than the sepals; fruit globose to oblong,
covered with uncinate or straight prickles; carpophore none; mericarps broad or
narrow on the commissural side, usually smooth.

About 40 species, widely distributed in both hemispheres, chiefly
in temperate regions, in the tropics restricted to mountain areas.
Only one species is found in Central America.

Sanicula liberta Cham. & Schlecht. Linnaea 1: 353. 1826.
S. mexicana DC. Prodr. 4: 84. 1830.

Moist or wet, usually dense, mixed, mountain forest, 900-2,500
meters; Alta Verapaz; El Progreso; Zacapa; Chiquimula; Guatemala;
Sacatepe"quez; Chimaltenango; Suchitepe'quez; Quezaltenango; San
Marcos. Mexico; Honduras; Nicaragua; Costa Rica; Panama.
Northwestern South America.

Plants perennial, arising from a short thick rootstock emitting numerous
fleshy-fibrous roots, the stems solitary or usually several, erect, simple or branched,
the whole plant glabrous; basal leaves on very long, slender petioles, cordate-
orbicular in outline, 4-10 cm. broad, palmately 5-parted, the segments petiolulate
or sessile, ovate-lanceolate to obovate-cuneate, acute, cuneate at the base, more or
less incised, lobulate, and crenate-dentate with spinulose-tipped teeth; involucral
bracts 5-8, linear, acuminate, reflexed; staminate flowers few, slender-pedicellate,
the sepals narrowly lanceolate, acuminate, scarcely 1 mm. long; petals broadly
obtriangular; fruit 2 mm. long, pedicellate, about 4 mm. long, densely covered
with uncinate prickles.

A common but inconspicuous plant in the denser mountain forests,
often occurring in abundance. The burlike fruits cling persistently
to clothing.

SPANANTHE Jacquin

Branched herbs, usually annual, glabrous or nearly so; leaves mostly opposite,
on long slender petioles, crenate-dentate, often cordate; stipules scarious, lacerate;
flowers very small, white, the umbels simple, few or numerous and forming leafy
panicles; calyx teeth evident; petals obovate, entire or retuse; stylopodium small,
depressed-conic; fruit dorsally compressed, constricted at the very narrow com-
missure; mericarps compressed dorsally and flat, conspicuously tricostate on the
dorsal surface; carpophore undivided; seed dorsally compressed, the face plane.

The genus consists of a single species.

FIG. 15. Spananthe paniculate. A, Habit; X
X 10. C, Sketch of fruit seen from above; X 10.

, Fruit seen laterally;

63

64 FIELDIANA: BOTANY, VOLUME 24

Spananthe paniculata Jacq. Coll. Bot. 3: 247. 1789. Culantro
de monte (fide Aguilar) ; canutillo (Santa Rosa) ; cola de pavito (Guate-
mala, fide Morales); valeriana (Huehuetenango) ; ocoan (Huehuete-
nango).

Moist or wet thickets, meadows, fields, or open forest, sometimes
in pine, oak, or Liquidambar forest, frequently a weed in waste or
cultivated ground, 350-2,000 meters; Alta Verapaz; Zacapa; Jalapa;
Santa Rosa; Guatemala; Sacatepe"quez ; Chimaltenango; Quiche";
Huehuetenango; Quezaltenango. Mexico; British Honduras to El
Salvador and Panama. South America.

Plants a meter high or less, usually or always annual, often much branched,
the stems green, hollow, the whole plant glabrous or nearly so, green; leaves thin,
mostly opposite, on long or very long, slender petioles, the petiole bearing slender
bristles at the apex, the stipules bristly-ciliate; leaf blades broadly ovate to deltoid
or lance-deltoid, mostly 4-9 cm. long, acute to long-attenuate, deeply cordate to
truncate or rounded at the base, rather coarsely crenate-dentate, paler beneath;
umbels slender-pedunculate, few-flowered, the flowers white, on long slender pedi-
cels; fruit about 2.5 mm. long.

A common weedy plant in many parts of Guatemala and other
Central American countries. It grows mostly in wet soil, sometimes
along ditches, and is most plentiful during the rainy months.

TAUSCHIA Schlechtendal

Plants low, herbaceous, acaulescent or short-caulescent, glabrous or pubescent,
with taproots or tubers; leaves entire to pinnate or ternately decompound, the
petioles vaginate; inflorescence of lax compound umbels, the peduncles longer than
the leaves; involucre usually absent, the bractlets several, linear to ovate, free or
connate, entire or dentate or lobate; flowers yellow, white, or purplish, the petals
with a narrow inflexed apex; calyx teeth evident or obsolete; styles short, spread-
ing or reflxed; stylopodium none; fruit oblong to orbicular, slightly compressed
laterally, glabrous, the ribs prominent or filiform, not winged.

Species 20 or more, all American and mostly in Mexico.

Umbels simple T. oreomyrrhioides.

Umbels compound.

Rachis of the leaves puberulent; ultimate division of leaves oblong-linear.

T. steyermarkii.

Rachis of the leaves not puberulent; ultimate division of leaves narrowly linear.

T. filiformis.

Tauschia filiformis Coult. & Rose, Proc. Wash. Acad. Sci. 1:
137. 1900. Arracacia filiformis Coult. & Rose, Contr. U. S. Nat.
Herb. 3: 296. 1895, nomen subnudum; in Hook. Icon. t. 2429. 1896.

FIG. 16. Tauschia steyermarkii. A, Habit, about natural size. B, Umbellule;
about X 21/2- C, Lateral view of fruit; about X 10.

65

66 FIELDIANA: BOTANY, VOLUME 24

Collected in Chiapas on Volcan de Tacand (perhaps on the Guate-
malan side of the boundary) , at 4,000 meters or lower, and doubtless
occurring in San Marcos. Mexico (Oaxaca).

Plants glabrous, scapiform; leaves basal, 10-20 cm. long, on long slender peti-
oles, 2-3 times ternately pinnatifid, the segments numerous, very small, linear,
acute, mostly entire; scape naked, 6-15 cm. high, bearing a single compound um-
bel, the rays few, slender or stout, unequal, often much elongate; bractlets entire
and linear to dentate or pinnatifid, large and conspicuous, much exceeding the
flowers, the pedicels short; flowers dark purple; mericarps ovate-oblong, purplish,
3-4 mm. long.

Tauschia oreomyrrhioides Mathias & Constance, Am. Jour.
Bot. 39: 652, .fa. 1952.

Open slopes in area of pine-fir forests, Tajumulco volcano (type
San Marcos, Steyermark 35495). San Marcos, endemic but surely
also in Mexico.

Low, shortly caulescent, 8-15 cm. high, from a stout, thickened taproot, gla-
brous throughout; leaves oval to ovate, spreading, 1.5-4.5 cm. long, 1-3 cm. broad,
2-3-pinnate, the ultimate division linear-acuminate, mucronate, 2-7 mm. long,
1 mm. or less broad; petioles slender, 2-7 cm. long; peduncles slender, purplish,
4-6.5 cm. long; involucre of several lanceolate, pinnate bracts 3-10 mm. long, ex-
ceeding the flowers and fruit; rays lacking; pedicels 2-3 mm. long; petals not seen;
styles short, recurved; carpophore bifid to the base, the halves divergent for their
apical one-fourth, forming a V; fruit ovoid, 2-5 mm. long, 2 mm. broad, the ribs
low and obtuse, narrower than the intervals; vittae and seed face not seen.

Tauschia steyermarkii Mathias & Constance, Am. Jour. Bot.
39:653,./ig. 1952.

High, cool meadows, mountains of western Guatemala, alt. about
3,000-3,500 meters; Huehuetenango; Totonicapan; endemic (type,
Sierra de los Cuchumatanes, Huehuetenango, Steyermark 50280).

Acaulescent herbs, 10-15 cm. high, from a stout caudex, the foliage and in-
florescence puberulent; leaves oblong, 3.5-6 cm. long, 1-2 cm. broad, pinnate, the
leaflets ovate, 5-15 mm. long, 1-2-pinnately lobed, the ultimate divisions oblong
to linear, callous-tipped, 1-4 mm. long; petioles 2-5 cm. long; peduncles 6-12 cm.
long, puberulent at the summit; involucre lacking, or of a solitary foliaceous bract;
fertile rays 3, stout, unequal, 5-25 mm. long, puberulent; involucel of several linear,
entire, ciliolulate bractlets 2-3 mm. long, about equaling the flowers but shorter
than the fruit; pedicels 2-4 mm. long; calyx teeth obsolete; petals white; styles
short, spreading; carpophore bifid to the base, the halves approximate; fruit ovoid,
tapering slightly at the apex, 5 mm. long, 3 mm. broad, the ribs prominent, acute;
vittae solitary, rarely 2, in each interval, 2 (rarely 3 or 4) on the commissure; seed
face deeply sulcate.

STANDLEY AND WILLIAMS: FLORA OF GUATEMALA 67

CORNACEAE. Dogwood Family

References: Walther Wangerin, Cornaceae, Pflanzenreich IV. 229.
1910. Harold William Rickett, Cornaceae, North American Flora
286:299-311. 1945.

Trees or shrubs, rarely perennial herbs; leaves opposite or alternate, usually
petiolate, entire or denticulate, usually without stipules; flowers small, perfect or
by abortion unisexual and dioecious, regular, in dichotomous cymes or in racemes,
rarely umbellate or capitate and surrounded by an involucre of petal-like bracts,
mostly white or yellowish; calyx tube adnate to the ovary, the limb 4-5-lobate or
subtruncate, sometimes obsolete; petals alternate with the calyx lobes, 4-5 or none,
valvate or convolute; stamens as many as the petals and alternate with them, the
filaments short or elongate, mostly subulate, the anthers introrse, dehiscent by
lateral slits; disk epigynous and angulate, crenate, or lobate, generally glabrous,
rarely annular or obsolete; ovary inferior, 1-4-celled, the style simple, sometimes
divided almost to the base, or the styles several and free; ovules solitary in each
cell, pendulous from the apex of the cell, anatropous; fruit drupaceous, the stone
1-4-celled, or the fruit rarely containing 2-4 osseous or crustaceous nutlets; seeds
oblong, terete or semiterete, the testa membranaceous; embryo small or elongate,
the radicle terete or subcomplanate, the cotyledons subfoliaceous; endosperm abun-
dant, carnose.

Ten genera are recognized, of wide distribution. Only one genus
is represented in America.

CORNUS L.

Trees or shrubs, rarely perennial herbs (in northern species); leaves opposite
or very rarely alternate, entire; flowers small, perfect or rarely dioecious, in panic-
ulate cymes or umbellate or in involucrate heads; calyx limb turbinate, urceolate,
or campanulate, terete or costate, more or less 4-dentate; petals 4, valvate; sta-
mens 4, the filaments filiform or subulate, the anthers dorsifixed, versatile; disk
pulvinar; ovary 2-celled, the style filiform or columnar, the stigma capitate or
truncate; ovules solitary in each cell; drupe globose to ellipsoid or ovoid, areolate
at the apex, the stone osseous or crustaceous, 2-celled, 2-seeded; seeds oblong, com-
pressed, the testa membranaceous, the endosperm carnose.

About 40 species, chiefly in temperate regions of the northern
hemisphere, a few in mountains of the tropics, and one species in
Africa. In Central America only the following species are known.

Flowers in heads C. disciflora.

Flowers in cymes C. excelsa.

Cornus disciflora DC. Prodr. 4: 273. 1830. C. floccosa Wange-
rin, Fedde Rep. Sp. Nov. 6: 101. 1908. C. disciflora var. floccosa
Standl. Field Mus. Bot. 8: 321. 1931. C. disciflora f. floccosa Rickett,
Bull. Torr. Bot. Club 72: 223. 1945. Palo bolitero (fide Aguilar).

Moist or wet mountain forests or Cupressus forests, mostly 2,000-
3,000 meters; Chiquimula; Chimaltenango; Solola; Quiche"; Huehue-

68 FIELDIANA: BOTANY, VOLUME 24

tenango; Quezaltenango; San Marcos. Mexico; El Salvador; Hon-
duras; Costa Rica; Panama.

FIG. 17. Cornus disci flora. A, Flowering and fruiting branch; X Yz- B, Flower; X 10.

STANDLEY AND WILLIAMS: FLORA OF GUATEMALA 69

Shrubs or small trees 6-22 meters tall, the trunk to 40 cm. or more in diameter
with light brown or grayish bark, somewhat roughened; leaves slender-petioled,
lanceolate to lanceolate-oblong, mostly 6-15 cm. long and 2-6 cm. broad, acute to
long acuminate, acute to the base, glabrous above, pale below (usually) and mi-
nutely pubescent with short, appressed malpighiaceous trichomes, sometimes also
with spreading hairs; inflorescence a long pedunculate head about 1 cm. broad with
12-15 sessile flowers; petals whitish or greenish-yellow, 3-3.5 mm. long; fruit ellip-
soidal, 12-14 mm. long, dark purple at maturity.

Closely related to the attractive C. florida of the United States
and Mexico. The authority for this species is given as Mocino and
and Sesse* by Wangerin and by Rickett. However, De Candolle did
not mention Sesse" and Mocino as authors when the name was pub-
ished. Sesse" and Mocino used the name C. capitata for this plant, a
name which did not appear until 1893. The form with spreading
hairs on the lower surface of the leaf is not known from Guatemala
but may be expected. It is forma floccosa (Wang.) Rickett, but hardly
deserving of a formal name.

Cornus excelsa HBK. Nov. Gen. & Sp. 3: 430. 1818. C. tolu-
censis HBK. I.e.

Moist, mixed or pine-oak forest, often in thickets, sometimes
growing along streams, 1,400-2,700 meters; Jalapa; Escuintla; Gua-
temala; Chimaltenango; Quiche"; Huehuetenango. Mexico; Hon-
duras (fide Rickett).

A shrub or a small tree, sometimes 9 meters high with a trunk 15 cm. in diam-
eter, the branches slender, grayish or fuscous, often purplish or brown, puberulent
at first; leaves ovate or lance-ovate, 5-10 cm. long, long-acuminate, obtuse or
rounded at the base, green above and almost glabrous, the nerves conspicuously
impressed, paler beneath and glabrate, the slender nerves very prominent; flowers
in small cymes, pedicellate; petals white, 3.5-4.5 mm. long; fruit white, turning
pale blue, 5-6 mm. in diameter, very juicy.

GARRYALES

An order containing a single family and a single genus with 12-15
species in temperate and tropical North America, — with the charac-
ters of the family.

The systematic position of the family is open to considerable ques-
tion. Classically the family has been placed among the amentiferous
families, often near Betulaceae. Other students have placed the fam-
ily near the Cornaceae, in the Umbelliflorae, or included it in the
Cornaceae. Recently Hutchinson in the second edition of "The
Families of Flowering Plants" (1959) has placed it in the "Arali-
ales," an order with a circumscription somewhat similar to that called
Umbelliflorae in this flora.

70 FIELDIANA: BOTANY, VOLUME 24

The family is placed here, following the Umbelliflorae, but the
Order Garryales is maintained for it. This position is in part a
matter of convenience and in part the junior author's feeling that a
family so different from the Umbelliferae, Araliaceae and Cornaceae
would best be maintained in a separate order.

GARRYACEAE

Reference: Walther Wangerin, Garryaceae, Pflanzenreich IV. 56a.
1910.

Shrubs or trees, the young branchlets subquadrangular, the elder ones terete;
leaves opposite, petiolate, entire or undulate, penninerved, persistent, coriaceous,
the petioles connate at the base; stipules none; flowers small, dioecious, the stami-
nate pedicellate, the pistillate sessile or nearly so, arranged in ament-like racemes,
these chiefly axillary, the bracts opposite, often connate at the base, the flowers
solitary or ternate in their axils; sepals 4 in the staminate flower, valvate, some-
times coherent at the apex; stamens 4, the filaments free, the anthers basifixed,
elongate-elliptic or linear, introrse, dehiscent by longitudinal slits; sepals none in
the pistillate flower, the ovary ovoid or oblong, 1-celled, inferior, the 2 styles sub-
ulate, erect or recurved, papillose within; ovules 2, pendulous from the apex of the
cell, anatropous; fruit baccate, ovoid or subglobose, tipped with the styles and
with 2-4 small calyx lobes at the base of the styles, 1-2-seeded; seeds ovoid or sub-
globose, with membranaceous testa, the endosperm carnose, abundant; embryo
minute, the cotyledons oblong, the radicle terete.

The group consists of a single genus, with the characters of the
family.

GARRYA Douglas

The genus consists of about 15 species, in North America. Only
the following are known from Central America, but about 7 are re-
corded from Mexico.

Leaves small, mostly 2.5-4.5 cm. long, the nerves impressed on the upper surface;
bracts of the staminate inflorescence very broadly ovate G. corvorum.

Leaves large, mostly 8-12 cm. long, the nerves not or scarcely impressed; bracts
of the staminate inflorescence ovate-lanceolate G. laurifolia.

Garrya corvorum Standl. & Steyerm. Field Mus. Bot. 23: 16.
1943.

On slopes of bluffs or in Juniperus forest on limestone, 3,300-
3,700 meters; endemic; Huehuetenango (type from Che'mal, Sierra
de los Cuchumatanes, Standley 81651).

A shrub or small tree 2-15 m. tall, branches blackish, lustrous; leaves coria-
ceous, oblong or elliptic-oblong, 2.5-4.5 cm. long, 1-1.5 cm. broad, obtuse or sub-
acute at the apex and apiculate, obtuse or narrowly rounded at the base, lustrous

FIG. 18. Garrya laurifolia. A, Fruiting branch; X
C, Staminate flowers; about X 7.

, Mature fruit; X 5.

71

72 FIELD IANA: BOTANY, VOLUME 24

and almost glabrous above, the venation impressed, densely pilose beneath with
whitish, subappressed, rather long hairs, the lateral nerves inconspicuous, the veins
obsolete; staminate racemes 3-3.5 cm. long, short-pedunculate, simple, terminal,
solitary, densely pilose with long whitish hairs, the bracts very broadly ovate,
connate below, 8 mm. long or less, cuspidate-acute.

A single shrub of the species was found at the type locality.

Garrya laurifolia Hartweg ex Benth. PI. Hartw. 14. 1839.
G. laurifolia var. quichensis Donn.-Sm. Bot. Gaz. 54: 237. 1912 (type
from San Miguel Uspantan, El Quiche", Heyde & Lux 3175). Palo de
hueso (the usual name for the tree) ; hediondillo (San Marcos) ; ajruch
(Volcan de Agua) .

Moist or rather dry, mountain forest, often in pine-oak or Cupres-
sus forest, 1,500-3,600 meters; Zacapa; Jalapa; Guatemala; Saca-
tepe"quez; Chimaltenango; Solola; Quiche"; Huehuetenango; Quezal-
tenango; San Marcos. Mexico; Costa Rica.

A small or medium-sized tree, sometimes 12 meters high and probably even
taller, often fruiting when only a shrub of 2 meters, the trunk 50 cm. or less in
diameter, the branchlets at first pilose; leaves coriaceous, on petioles 2 cm. long
or less, usually blackening in drying, lance-elliptic to oblong or narrowly lance-
oblong, mostly 6-15 cm. long or even larger, 2-6.5 cm. broad, obtuse or acute and
apiculate at the apex, acute at the base, in the adult stage glabrous or nearly so
but when young usually appressed-pilose, at least beneath, the nerves and veins
usually prominent on the upper surface; staminate racemes dense, often branched,
the bracts ovate-lanceolate or narrowly lanceolate; bracts of the pistillate racemes
often large and leaflike; fruit black, glabrous, lustrous, globose or ovoid, 8-10 mm.
in diameter.

A characteristic and abundant tree in many of the barrancos of
the central region, especially in Guatemala, Sacatepe"quez, and Chi-
maltenango. So far as we know, no use is made of the wood.

Flora of Guatemala - Part VIII, Number 2

ERIC ALES1

Shrubs, trees, or suffrutescent perennials and (in Monotropaceae) saprophytes
or parasites without chlorophyll. Flowers mostly 5-4-merous; the corolla of dis-
tinct or free petals or the petals lightly adherent in the more primitive families
(and in some genera of Ericaceae) but typically connate nearly to the apex (most
Ericaceae); stamens commonly as many or twice as many as the corolla lobes,
hypogynous or epigynous, often opening by terminal pores or sometimes length-
wise by slits; ovary superior or inferior, with axile placentation; fruit commonly
a capsule or a berry.

The order Ericales is divided by Engler and Gilg into two sub-
orders containing 4 families, of which three are in our area. Standley
indicated a preference for placing Monotropa in a separate family, as
was done in North American Flora. We maintain the family Mono-
tropaceae.— The Lennoaceae were placed near the Monotropaceae by
Standley in his preliminary manuscript. I believe that the Lennoa-
ceae should be placed in the order Tubiflorae, where it was placed
by Engler and Gilg and where most evidence seems to indicate it is
related.

The Clethraceae are usually considered to be the most primitive
of the Ericales, followed by Pyrolaceae, Monotropaceae, Ericaceae,
and Epacridaceae, — the last mostly Australian.

1 The field work and travel by the junior author involved in the preparation
of this number of the Flora of Guatemala has been made possible through a grant
from National Science Foundation, which we are happy to acknowledge.

73

74 FIELDIANA: BOTANY, VOLUME 24

CLETHRACEAE

Reference: Nathaniel Lord Britton, Clethraceae, No. Am. Fl. 29:
3-9. 1914.

Shrubs or large trees; leaves alternate, petiolate, entire or dentate, membra-
naceous or coriaceous, penninerved; flowers perfect, in terminal, simple or branched
racemes; calyx deeply 5-cleft, the segments imbricate, persistent; petals 5, distinct
or slightly united at the base, deciduous, usually or always fimbricate at the apex;
stamens 10-12, the slender filaments short or elongate; anthers sagittate, inverted
in anthesis, the cells opening by apical pores; disk none or obscure; ovary superior,
pubescent, 3-celled, 3-lobate; style 1, the stigma entire or 3-lobate; ovules numer-
ous; fruit a 3-celled capsule, loculicidally 3-valvate. A single genus.

The genus Schizocardia A. C. Smith & Standley described in the
Clethraceae has been placed in the genus Purdiaea of the Cyrillaceae
(cf. Thomas in Journ. Am. Arb. 42: 110. 1961). It will be accounted
for in the supplement of the flora.

CLETHRA L.

Large shrubs or small trees, pubescent, often tomentose; leaves mostly persist-
ent, often coriaceous, entire or dentate; flowers small or medium-sized, racemose,
white, cernuous, the racemes terminal, often spike-like, usually paniculate, the
pedicels bracteate at the base, the bracts caducous; calyx 5-fid, the segments ob-
tuse, imbricate; petals obovate-cuneate, rounded at the apex, often emarginate or
bilobate, frequently erose, imbricate; filaments free or connate at the base; ovary
globose, 3-gonous or 3-lobate; capsule subglobose or 3-lobate, 3-celled; seeds small,
compressed or trigonous, the testa cellular, often produced into a wing; endo-
sperm carnose.

Clethra is found in temperate and tropical America, eastern Asia,
and Madeira. There have been more than 50 species described for
temperate and tropical America though there are perhaps fewer than
half so many.

Since preparing the manuscript for this genus Dr. H. Sleumer of
Leiden has borrowed our material for a monographic study of the
genus. We have studied the returned material. Many of Dr. Sleu-
mer's annotations are accepted here but others are not.

Leaves glabrous or at most with a few peltate scales.
Under leaf surfaces conspicuously lighter colored tl

Under and upper leaf surfaces concolorous C. suaveolens.

Under leaf surfaces conspicuously lighter colored than the upper surfaces.

C. oleoides.

STANDLEY AND WILLIAMS: FLORA OF GUATEMALA 75

Leaves sparsely to densely pubescent or tomentose beneath.

Leaves green below and glabrate or stellate puberulent; flower long-pedicellate.

C. johnstonii.

Leaves densely tomentose or pilose beneath and usually grayish or pale, the
pubescence sometimes very close and fine.

Adult leaves densely pilose or tomentose, the hairs spreading and easily visible,
the pubescence more or less velutinous and soft to the touch; lower sur-
face of leaf usually green, not pale or whitish.

Calyx 4-5 mm. long; pedicels about equal calyx, very stout. .C. licanioides.
Calyx 2.5-3 mm. long.

Pedicels shorter than the calyx C. mexicana.

Pedicels conspicuously longer than the calyx C. johnstonii.

Adult leaves covered beneath with fine, closely appressed whitish or grayish
tomentum, hairs sometimes present on nerves and veins, pubescence not
velutinous; lower surface of leaf blade white or grayish.

Calyx 3.5-4.5 mm. long; pedicels all or mostly longer than the calyx.

C. macrophylla.

Calyx 2-2. 5mm. long; pedicels shorter than the calyx.

Leaves thin; rachis of inflorescence minutely whitish-tomentulose.

C. skutchii.

Leaves coriaceous; rachis of inflorescence brown-puberulent.

C. pachecoana.

Clethra johnstonii Standl. & Steyerm. Field Mus. Bot. 22: 258.
1940.

Type from Volcan de Pacaya, above Las Calderas, Dept. Guate-
mala, Johnston 1255; collected also in the same department, without
definite locality.

Young branchlets densely pubescent; leaves on petioles 1-1.5 cm. long, char-
taceous or subcoriaceous, oblong-oblanceolate or narrowly oblanceolate, 7-11 cm.
long, 2-4 cm. broad, rounded at the apex or broadly obtuse and apiculate, cune-
ately long-attenuate to the base, glabrous above, at least in age, green beneath,
sparsely and minutely stellate-puberulent, appearing glabrous to the naked eye,
the lateral nerves about 17 pairs, oblique, almost straight; racemes slender and lax,
few or numerous, forming a terminal panicle, the slender rachis stellate-pilosulous,
the pedicels divaricate, 5-8 mm. long; calyx 3 mm. long, minutely stellate-tomentu-
lose, the segments lance-ovate, acute, ciliate; ovary densely stellate-tomentose.

Clethra licanioides Standl. & Steyerm. Field Mus. Bot. 23: 17.
1943. ZapotiUo.

Zacapa; Jalapa (type from Potrero Carrillo, 13 miles northeast of
Jalapa, 1,500-1,700 meters, Steyermark 33124) ; Chimaltenango.

76 FIELDIANA: BOTANY, VOLUME 24

A tree of 9 meters, the stout branchlets brown, densely brown-tomentose;
leaves on stout petioles 1-1.5 cm. long, rigid-coriaceous, rounded-obovate, 8-10 cm.
long, 5-7 cm. broad, broadly rounded or truncate at the apex, slightly narrowed to
the obtuse or narrowly rounded base, yellowish green above when dried, sparsely
furnished with minute, depressed, radiately stellate hairs, but to the naked eye
appearing glabrous, green beneath, velvety-stellate-pilose, the lateral nerves about
13 pairs, almost straight, ascending at an angle of about 45 degrees; racemes 25 cm.
long or less, laxly flowered, numerous, the rachis stout, densely brown-tomentose,
the stout pedicels 5 mm. long; calyx 4-5 mm. long, densely stellate-tomentulose,
the segments broadly ovate or oval, very obtuse.

The flowers are fragrant, the petals white, the sepals buff with a
faint tinge of pink. The species is closely related to C. mexicana
A. DC. of which it may be but an extreme form.

Clethra macrophylla Mart. & Gal. Bull. Soc. Acad. Brux. 9:
539.1842. C. hondurensisBritton, No. Am. Fl. 29:7. 1914. C.vicen-
tina Standl. Journ. Wash. Acad. Sci. 13: 353. 1923. C. bimatris
Standl. & Steyerm. Field Mus. Bot. 23: 17. 1943 (type from Entre
Rios, Izabal, Kuylen 151). C. caloneura Standl. & L. Wms. Ceiba
1: 157. 1950. C. molinae Standl. & L. Wms. I.e. 158. C. occidentalis
auth. et sensu Sleumer.

Moist or dry mountain forest or lowland forests, 0-1,800 m.
Izabal; British Honduras; El Salvador; Honduras.

A large shrub or a tree, sometimes 13 meters high with a trunk 25 cm. in diam-
eter but usually lower; leaves short-petiolate, firm-chartaceous or subcoriaceous,
narrowly obovate to narrowly oblong-obovate, broadly rounded to subacute at the
apex, cuneately narrowed to the narrowly obtuse or acute base, deep green above
and almost wholly glabrous, the nerves obscure, white or grayish beneath, covered
with a minute, very dense, closely appressed tomentum, without long hairs on the
nerves, the margins entire or somewhat serrate toward the apex, the lateral nerves
slender but conspicuous beneath; racemes lax, short or greatly elongate, few or
numerous, the slender rachis minutely whitish-tomentulose, the pedicels divaricate,
3-6 mm. long; calyx 3.5-4.5 mm. long, minutely whitish-tomentulose, the segments
ovate or elliptic, obtuse; capsule somewhat depressed, 3-lobate. 6-7 mm. broad,
tomentulose.

The name "nance de cerro" is said to be given to the tree in
Honduras. The white flowers are fragrant.

The species is occasional on the east coast of Mexico, it is to be
found in British Honduras and is common and occasionally abundant
in Honduras. It has been found in Guatemala only in the depart-
ment of Izabal. Dr. Sleumer has annotated much of this material as
C. vicentina Standl. but another large part is annotated C. occiden-

FIG. 19. Clethra mexicana. A, Habit, X Yz- B, Mature capsule and sub-
tending calyx, X 2%.

77

78

FIELDIANA: BOTANY, VOLUME 24

talis (L.) Kuntze. The name "occiden tails" has appeared several
times in literature, — first in Steudel's Nomenclator Botanicus where
it is given as a synonym of C. tinifolia Sw. without authority or
basionym. It next appears in Kuntze's Rev. Gen. PI. 32: 192. 1898,
again without basionym but with authority "OK. (L.)." Both uses
would seem to be illegitimate. It is possible, even probable, that the
acceptable name for this species is C. tinifolia Sw., in the broad sense.

Clethra mexicana A. DC. Prodr. 7: 590. 1839. C. lanata Mart.
& Gal. Bull. Acad. Brux. 9: 538. 1842. C. salvadorensis Britton,
No. Am. Fl. 29: 9. 1914.

Dry mountain forest, commonly in pine-oak woods, 550-2,500 m.
Jalapa; Sacatepe"quez; Guatemala; Escuintla; Quiche"; Huehuetenan-

FIG. 20. Clethra oleoides. A, Habit, X Yz- B, Expanded flower, X 1A.
C, Bud partly opened, X 4. D, Average petal, X 2^. E, Anther, X 20. F, Type
of pubescence from the inflorescence, X ± 20.

STANDLEY AND WILLIAMS: FLORA OF GUATEMALA 79

go; Quezaltenango; San Marcos. Mexico; El Salvador; Honduras
to Panama and doubtless South America.

A large shrub or a tree, sometimes 20 meters tall, usually with a thick trunk
and a not very dense, spreading crown, the young branchlets densely villous-
tomentose; leaves on short stout petioles, usually chartaceous, obovate or oblong-
obovate, mostly about 9-15 cm. long and 3.5-6 cm. broad, obtuse to rounded at
the apex, somewhat narrowed to the acute to narrowly rounded base, entire or
obscurely serrate toward the apex, green above but usually sparsely or sometimes
rather densely stellate-pilose with mostly weak slender hairs, usually rufescent
beneath, densely and softly villous-tomentose; racemes few or numerous, short or
elongate, mostly rather dense, forming a large panicle, the rachis slender or rather
stout, densely stellate-pilose, the pedicels 1-2 mm. long, sometimes slightly longer;
calyx 3 mm. long, pale-tomentulose, the segments broadly ovate, obtuse; capsule
tomentulose, 4 mm. broad.

This is possibly the commonest species in Guatemala, usually
found in rather open oak-pine forest areas. The Guatemalan speci-
mens have usually been called C. salvadorensis and sometimes also
C. lanata Mart. & Gal.

Dr. Sleumer has annotated specimens with both C, lanata Mart.
& Gal. and C. mexicana A. DC. We find it difficult to separate the
two and would suggest that C. lanata, as well as C. quercifolia Lindl.
should be placed here as synonyms.

Clethra oleoides L. Wms., Fieldiana, Bot. 31: 167. 1965. Shiorsh.
Moist mixed forests at 2,500-3,100 m.; Huehuetenango. Mexico.

Small evergreen trees or shrubs to 6 m. or more tall. Young branchlets gray,
terete, glabrous or essentially so; leaves elliptic or elliptic-lanceolate, acute or ob-
tuse at the apex, mostly obtuse at the base, glabrous or rarely with a few peltate
scales, distinctly paler below and gray-green, densely but obscurely punctate-
glandular below, (2-) 3.5-8 cm. long and 1.5-3.5 cm. broad, petiole 3-8 mm. long;
inflorescence a several-flowered terminal raceme or sometimes several racemes in a
fascicle, about as long or slightly longer than the subtending leaves, sparsely
peltate or stellate pubescent, becoming glabrous; flowers slightly cernuous and
unilateral on the rachis; pedicels short, about 3 mm. long; calyx-lobes ovate, acute,
about 3-3.5 mm. long, sparsely stellate or peltate pubescent, ciliate, obscurely
punctate-glandular; corolla white, the lobes free or only slightly adnate at the
bases, the lobes (petals) oblong-ovate to broadly oblong-oval, emarginate and the
truncate apex usually laciniate, 6-7 mm. long and 4-5 mm. broad, not all the same
shape in one flower; stamens 10, lightly attached at the base of the corolla or below
it and shorter than the lobes; pistil about 3 mm. long, trilobate at the stigmatic
apex; mature capsules not seen.

80 FIELDIANA: BOTANY, VOLUME 24

The specimens from Guatemala are sterile but match in all vege-
tative detail the type from Mexico.

Clethra pachecoana Standl. & Steyerm. Field Mus. Bot. 22:
259. 1940. Escobo; zapotillo; xiorsh (Huehuetenango).

Damp or wet, usually mixed, mountain forest, often in Chirantho-
dendron forest on the volcanoes, frequently associated with pines or
Abies, 2,000-3,200 meters; Alta Verapaz; Baja Verapaz; Jalapa;
Guatemala; Sacatepe'quez (type from Volcan de Agua, above Santa
Maria de Jesus, Standlel 65123); Chimaltenango; Solola; Quiche";
Huehuetenango; Quezaltenango; San Marcos. Mexico; El Salvador.

A tree 6-15 meters high or larger with a dense crown and often with a tall
slender trunk, the young branchlets densely ferruginous-tomentulose; leaves small,
short-petiolate, coriaceous, the petioles 6-12 mm. long; leaf blades oblanceolate-
oblong to obovate-oblong, 4.5-7 cm. long, 1.2-3 cm. broad, very obtuse at the apex
or somewhat rounded and apiculate, sometimes subacute, narrowed to the subacute
to truncate base, the margins entire or nearly so in adult leaves, in juvenile ones
often coarsely serrate, glabrous above, somewhat paler beneath and appearing
glabrous but actually covered with an almost microscopic and closely appressed
tomentum, the lateral nerves about 10 pairs, almost straight or somewhat curved;
racemes few or numerous, 10 cm. long or less, densely many-flowered, the rachis
minutely brown-tomentulose, the stout pedicels scarcely more than 1.5 mm. long;
calyx 2.5 mm. long, minutely brownish-tomentulose, oblong-ovate, obtuse; petals
white, obovate, ciliate; capsule depressed-globose, 3-3.5 mm. broad, minutely
stellate-tomentulose.

Leaves of young sterile plants or branches are often much larger
than the adult ones, coarsely serrate, and frequently soft-pilose.

Clethra skutchii Standl. & Steyerm. Field Mus. Bot. 22: 260.
1940.

Moist or wet forest, 1,350-2,000 meters or more, Quezaltenango;
type from Palmar, Skutch 1453; collected also on Volcan de Santa
Maria above Santa Maria de Jesus; endemic.

A tree 10-22 meters tall with a trunk as much as 30 cm. in diameter, the young
branchlets minutely hispidulous with brown hairs; leaves on slender petioles 12-
20 mm. long, firm-membranaceous, oblanceolate-oblong or oblong-obovate, 8-13
cm. long, 3.5-7 cm. broad, rounded to subacute at the apex and often abruptly
apiculate, somewhat narrowed to the cuneate to broadly obtuse base, almost gla-
brous above, sometimes pale-puncticulate, grayish or whitish beneath and covered
with a close and minute, very dense tomentum, the lateral nerves about 14 pairs,

STANDLEY AND WILLIAMS: FLORA OF GUATEMALA 81

almost straight, very oblique; racemes rather dense, 13 cm. long or less, fasciculate
at the ends of the branches, the rachis minutely stellate-pubescent, the stout ped-
icels hardly 2 mm. long; calyx 2.5 mm. long, the segments ovate-oblong, subacute,
recurved, minutely tomentulose; petals white, obovate, ciliate, slightly longer than
the sepals.

Clethra suaveolens Turcz. Bull. Soc. Nat. Moscou 36, II: 230.
1863. C. vulcanicola Standl. Journ. Wash. Acad. Sci. 13: 353. 1923.
C. glaberrima Lundell, Phytologia 1: 219. 1937. C. nubium Standl.
& L. Wms., Ceiba 1: 159. 1950 (type from Montana de los Nubes,
Guatemala, Williams & Molina 15276). C. viridifolia Standl. & L.
Wms. I.e. 161. Ka-ut (Coban, Quecchi).

Occasional in oak-pine woods or broad leaf forest at middle or
high elevations of 1,000-2,400 meters. Alta Verapaz; Baja Verapaz;
El Progreso; Zacapa; Chiquimula; Jutiapa; Guatemala; Huehuete-
nango. Mexico; El Salvador; Honduras; Nicaragua.

Shrubs or usually trees 1-15 m. tall, glabrous or the inflorescence pubescent.
Leaves elliptic or elliptic-lanceolate, acute or acuminate, acute or somewhat obtuse
at the base, glabrous, chartaceous at least when dry, 5-15 cm. long and 1-6 cm.
broad; inflorescence of 1-many terminal racemes shorter or usually longer than the
subtending leaves, tomentulose, pedicels slender and 3-10 mm. long; calyx about
4 mm. long, tomentulose, the lobes ovate-oblong, obtuse or acute; corolla white,
about 5 mm. long, the lobes erose or nearly entire; capsule about 6 mm. broad,
tomentulose.

This is the only Clethra in Central America with completely gla-
brous leaves and is easily distinguished.

Clethra matudae Lundell, based on sterile material from Chiapas,
probably belongs here.

PYROLACEAE

Reference: P. A. Rydberg, Pyrolaceae, No. Am. Fl. 29: 21-32. 1914.

Plants perennial, with creeping scaly rootstocks, the stems short, usually per-
sisting for several years and suffrutescent; leaves coriaceous, alternate but often
crowded and appearing verticillate, petiolate, usually serrate or crenate; stipules
none; inflorescence racemose, corymbose, or subumbellate, the peduncles bearing
several bracts; flowers perfect, rather small; sepals 5, slightly if at all united;
corolla regular or nearly so, the petals 5, distinct, white, pink, or purplish, hypogy-
nous; stamens 10, hypogynous, the filaments often dilated at the base; anthers
2-celled, usually attached at or near the middle, generally produced at the proxi-

FIG. 21. Chimophila maculata. A, Habit, X Vz- B, Flower, X 2^. C, an-
ther, X 5. D, Stigma, X 2^. E, Capsule from below, X 1. Pyrola augustifolia.
F, Habit, X 3^. G, Flower, X2^. H, Anther, X 7>£. I, Style and immature
ovary, X 5. J, Capsule from below, X 1.

82

STANDLEY AND WILLIAMS: FLORA OF GUATEMALA 83

mal end into 2 tubes, opening by round or oblong pores at the ends of the tubes;
ovary subglobose, 5-celled, obtusely 5-angulate, the ovules numerous; styles wholly
united, short or elongate, the stigma 5-papillate or 5-radiate; fruit capsular, obtusely
5-angulate, loculicidally dehiscent; seeds very numerous, with carnose endosperm.

Three genera, in the northern hemisphere. Only the following are
found in Central America.

Capsule dehiscent from above downward, without cobwebby threads; inflorescence
corymbose or subumbellate; leaves coarsely serrate Chimaphila.

Capsule dehiscent from below upward, the edges of the valves connected by cob-
webby threads; inflorescence racemose; leaves finely crenate or serrulate or
entire Pyrola.

CHIMAPHILA Pursh

Plants perennial, suffrutescent, erect, with long creeping rootstocks, the stems
simple or branched; leaves often subverticillate or subopposite, coriaceous, usually
serrate; inflorescence subumbellate or corymbose; flowers few; sepals and petals
each 5; stamens 10, the filaments expanded into a disk near the base, then curved
upward ; anthers ventricose, attached near the middle, the tubes rather long, dis-
tinct; style straight, short; stigma peltate, flat, 5-radiate, without papillae; cap-
sule depressed to ovoid or globose, dehiscent from the apex downward.

About 5 species, in the northern hemisphere. The plants grow
mostly in temperate regions, and in the tropics are restricted to the
higher mountains. Only the following are known in Central America.

Leaves striped or blotched with silver on the upper surface, mostly acuminate;
dilated portion of the filament glabrous or merely ciliolate.

C. umbellata var. mexicana.

Leaves green on the upper surface, acute or obtuse; dilated portion of the filament
villous. . . .C. maculata.

Chimaphila maculata (L.) Pursh, Fl. Amer. Sept. 300. 1814.
Pyrola maculata L. Sp. PI. 396. 1753. C. maculata var. acuminata
Lange, Vid. Medd. 1867: 112. 1868. C. acuminata Rydb. No. Am.
Fl. 29: 31. 1914. C. guatemalensis Rydb. I.e. 32 (type from Santiago,
Sacatepe"quez, Rosalio Gomez 795). C. dasystemma Torr. ex Rydb.
I.e. Hierba de madrono (fide Aguilar).

Mostly in pine-oak forest, 1,300-2,500 meters; Zacapa; Chiqui-
mula; Jalapa; Guatemala; Sacatepe"quez ; Chimaltenango; Solola;
Quich^ ; Huehutenango; Quezaltenango; San Marcos. United States;
Mexico; Honduras; Costa Rica.

84 FIELDIANA: BOTANY, VOLUME 24

Plants 10-20 cm. high; leaves few, in pseudoverticels of 2-4, short-petiolate,
lanceolate or oblong-lanceolate, 3-9 cm. long, acute or acuminate, acute at the
base, serrate, usually coarsely and sparsely so, deep green above and more or less
mottled with silver along the nerves, paler beneath; peduncles much elongate,
1-4-flowered, the flowers long-pedicellate; sepals elliptic or rounded, ciliate; petals
waxy white or pinkish, orbicular, 6-8 mm. long; dilated portion of the filament
villous; capsule depressed-globose, 7-8 mm. long.

This has been reported from Guatemala as C. umbellata (L.)
Barton. When in flower, the plant is a handsome one. It is locally
common in some of the mountain forests of Guatemala.

Chimaphila umbellata (L.) Barton var. mexicana (DC.) L.
Wms. Fieldiana, Bot. 31: 169. 1965. C. umbellata /3 mexicana DC.
Prodr. 7: 775. 1839. C. mexicana Rydb. No. Am. Fl. 29: 31. 1914.
C. umbellata mexicana DC. ex Rydb. I.e. in syn.

Dense and moist forest, 1,600-2,800 meters; Zacapa; Solola;
Huehuetenango. Mexico.

Plants perennial and suffrutescent, erect or decumbent, 30 cm. high or less,
glabrous throughout or nearly so; leaves in whorls of 3-7, oblanceolate, 3-10 cm.
long, acute or obtuse, coarsely serrate or spinulose-serrate, attenuate to the base,
short-petiolate, coriaceous, deep green and somewhat lustrous above, the nerves
impressed, paler beneath; peduncles about 10 cm. long, the flowers few, long-
pedicellate, the pedicels in flower somewhat spreading or recurved; sepals broadly
ovate, about as broad as long; petals creamy white tinged with rose, 7 mm. long;
dilated portion of the filament obovate, ciliolate on the edges or glabrous, the an-
thers 3 mm. long; capsule globose-ovoid, 6-10 mm. in diameter.

This plant as well as several other variations in America seem to
vary slightly from the typical Old World form. The Mexican and
Central American variations would seem to differ from temperate
American forms but to separate them all as species, following Ryd-
berg, seems hardly justifiable.

PYROLA L.

Plants perennial, with rootstocks, the stems very short, bearing a few leaves,
the scapes much elongate, bearing 1 or more small bracts; leaves petiolate, mostly
chartaceous; inflorescence racemose, the flowers usually nutant, short-pedicellate;
sepals 5, slightly united; corolla broadly campanulate, the 5 petals distinct, broad,
white, greenish, or purplish; stamens 10, the filaments subulate, somewhat dilated
at the base, incurved; anthers oblong, dehiscent by pores at the ends of the tubes;
ovary with 5 rounded angles; style straight, or deflexed at the base, then curved
upward and gradually thickened toward the apex; stigma of 5 erect papillae; cap-
sule depressed-globose, umbilicate, 5-valvate, dehiscent from the base upward.

STANDLEY AND WILLIAMS: FLORA OF GUATEMALA 85

About 20 species, or perhaps fewer, in the northern hemisphere
and chiefly in temperate regions; in the tropics found only in the
higher mountains.

Style strongly deflexed at the base; petals pinkish; leaves entire or obtusely and
inconspicuously crenate P. angustifolia.

Style straight; petals greenish white; leaves finely serrate with acute teeth.

P. secunda.

Pyrola angustifolia (Alef.) Hemsl. Biol. Centr. Am. Bot. 2: 283.
1881. Thelaia angustifolia Alef. Linnaea 28: 52, t. 1, fig. 7. 1856.
P. liebmannii Lange, Vid. Medd. 1867: 113. 1868.

Moist and usually dense, pine forest, 2,500-3,800 meters; El Pro-
greso (east of Finca Piamonte) Solola (volcanoes of Atitlan and Santa
Clara); Quiche1; Huehuetenango (Sierra de los Cuchumatanes) ; Que-
zaltenango (Volcan de Zunil). Southern Mexico.

Plants erect, with slender rootstocks, the portion of the stem above ground
short, 1-7 cm. long, bearing a few leaves; leaves glabrous, the petioles slender,
2-7 cm. long; leaf blades elliptic or oval, 2-7 cm. long, subacute to rounded at the
apex, acute and decurrent at the base, chartaceous, lustrous, minutely and re-
motely crenulate or practically entire; scapes 15-30 cm. high, 8-15-flowered, with
2-4 bracts near the base; sepals lanceolate, acuminate, 5 mm. long; petals 7-8 mm.
long, pinkish or purplish; capsule depressed, 8 mm. broad.

It does not seem reasonable that anyone would name this plant
"angustifolia" but the illustration provided by Dr. Alef eld leaves no
doubt about the application of the name to this plant.

Pyrola secunda L. Sp. PI. 396. 1753. Ramischia secunda Garcke,
Fl. Deutschl. ed 4. 222. 1858. Actinocyclus secundus var. elatior
Lange, Vid. Medd. 1867: 115. 1868. Ramischia elatior Rydb. No.
Am. Fl. 29: 29. 1914.

Moist or dry forest, mostly in pine woods, 2,700-3,800 meters;
Sacatepe"quez; Totonicapan ; Huehuetenango; Quezaltenango. Widely
distributed in temperate and arctic Canada and United States; Europe
and northern Asia; in Mexico apparently rare and local.

Plants erect, glabrous, from long creeping rootstocks, the stems above ground
suffrutescent, 1-10 cm. high, bearing 1-4 or often more leaves; leaves thin, persist-
ent, lustrous, on petioles 1-2 cm. long, ovate or elliptic, 1-5 cm. long, usually acute,
finely serrulate; scapes 20 cm. high or less; racemes mostly 6-18-flowered, short or
elongate, the pedicels short, the flowers nutant; sepals oval or elliptic, 1 mm. long,
rounded at the apex; petals greenish white, oblong or elliptic, 4-5 mm. long; cap-
sule subglobose, 4 mm. in diameter.

86 FIELDIANA: BOTANY, VOLUME 24

In many parts of the Rocky Mountains this is a very common
plant, but it is rare in the high mountains of Guatemala, also appar-
ently in Mexico. Ramischia elatior, originally described from Vera-
cruz, seems to differ in no essential respect from northern specimens
of Pyrola secunda, and is scarcely a good variety.

MONOTROPACEAE. Indian Pipe Family

Reference: John K. Small, Monotropaceae, No. Am. Fl. 29: 11-18.
1914.

Saprophytic plants or parasitic on roots of other plants, without chlorophyll,
usually succulent and white to brown or red, the stems solitary or often clustered
from matted roots, mostly simple and scapiform; leaves reduced to scales; flowers
perfect, erect or nutant; sepals 2-6, distinct, often deciduous; corolla white or col-
ored, of 3-6 distinct or partially united petals; stamens 6-12, hypogynous or some-
times inserted on a small glandular disk, the filaments slender or dilated; anthers
2-celled, sometimes with confluent cells dehiscent by valves or pores, sometimes
appendaged; ovary superior, mostly of 4-6 united carpels, 4-6-celled or 1-celled,
sessile; styles united, usually columnar, the stigma discoid, funnelform, or capitate,
often sublobate; ovules numerous; fruit capsular, 4-6-celled, sometimes 1-celled,
often more or less fleshy, generally surrounded by the marcescent perianth, locu-
licidally dehiscent; seeds numerous, minute.

The family is often included in the Pyrolaceae or in the Erica-
ceae. It may be distinguished from both these families by its her-
baceous and saprophytic habit, or parasitic habit. There are perhaps
as many as 8 or 9 not very distinct genera, mostly with one or few
species. One genus extends into tropical America.

MONOTROPA L.

Plants fleshy, with a somewhat waxy texture, white, pink, or red, glabrous or
pubescent, the stems simple; leaves scale-like, approximate or crowded below,
larger and more remote above; flowers solitary or racemose; sepals 2-5; petals 5-6,
dilated above, ciliate, slightly saccate at the base; stamens 10 or 12, included, the
filaments slender, often pubescent; anthers short, transverse, the cells opening by
2 slits; ovary 4-6-celled, the style short and thick, the stigma discoid.

Half a dozen species, in America, Europe, and Asia, mostly in
temperate regions, in the tropics found only in the mountains. Dif-
ferent authors vary greatly in the number of species recognized. Only
the following are known in Central America.

Flowers solitary, terminal M. coccinea.

Flowers racemose, several or numerous M . hypopithys.

Monotropa coccinea Zucc. Flora 15, pt. 2: Beibl. 100. 1832.
M. uniflora var. variegata Andres, Verh. Bot. Ver. Brandenb. 52: 94.
1911 (type from Guatemala, but no collection cited). M. uniflora

STANDLEY AND WILLIAMS: FLORA OF GUATEMALA

87

var. coccinea Domin, Sitzungsb. Boehm. Ges. Wiss. Prag, II Classe,
1915: 5. 1915. Flor de tierra (Chiquimula) .

Mostly in dense, wet or moist, oak, pine, liquidambar, or mixed
forest, 1,000-3,100 meters; Alta Verapaz; Baja Verapaz; Zacapa; Chi-
quimula; Jalapa; Guatemala; Huehuetenango. Central and south-
ern Mexico; British Honduras; El Salvador; Honduras; Nicaragua;
Costa Rica; Panama; Colombia.

Plants 10-20 cm. high, 1-flowered, coral-red or rose-red, blackening when dried,
glabrous, often growing in dense colonies; leaf scales numerous, lanceolate or ovate-
lanceolate, mostly 1 cm. long or shorter; flower nutant in anthesis, erect in fruit;
sepals rose-red, oblong or linear-oblong, 12-14 mm. long, erose above the middle;
petals obovate or spatulate, pinkish white, about equaling the sepals, glabrous,
not ciliate; filaments glabrous; capsule subglobose, about 13 mm. long.

This has been reported from Guatemala as M. uniflora L., and
probably is no more than a variety of that, which in its various forms
is widely distributed in the northern hemisphere. M. uniflora in its
typical state is a white plant, sometimes tinged with pink. It is well
known in the United States under the names "Indian pipe" and
"corpse plant."

B

FIG. 22. Monotropa coccinea. A, Habit of plants, X Yi. B, Portion of flower
showing ovary, stamens, and petals, X 2J^. C, Anther dehisced showing veil-like
tissue of anther, X 5.

88 FIELDIANA: BOTANY, VOLUME 24

Monotropa hypopithys L. Sp. PI. 387. 1753. Hypopitys lati-
squama Rydb. Bull. Torr. Bot. Club 40: 461. 1913. M. hypopitys
var. latisquama Kearney & Peebles, Journ. Wash. Acad. Sci. 29: 487.
1939.

Pine forest, 1,700-2,000 meters; Zacapa (Sierra de las Minas, be-
low Finca Alejandria, Steyermark 29782) . Canada and United States;
Mexico; Europe and Asia.

Plants simple, erect, often forming colonies, more or less pubescent, red, pink,
or yellowish, generally 10-20 cm. high; leaf scales mostly 1 cm. long or shorter,
sessile, the lowest rounded or ovate, the upper ones oval or oblong; flowers in
Guatemalan plants orange-red with yellow tips; sepals 6-8 mm. long, cuneate or
spatulate, ciliate; petals cuneate or oblong, 1 cm. long or shorter, pilose and ciliate;
capsule globose to oval, 6-9 mm. long.

The only known collection from Guatemala is in poor condition.
It is not typical for the species but whether or not it is one of the
varieties of this widespread plant or whether it is undescribed can-
not be easily determined. The spelling of the specific name is that
used by Linnaeus.

There is fair reason to separate this species as a monotypic genus
Hypopithys as has been done by Small and others.

ERICACEAE. Heather Family

References: J. K. Small, Ericaceae, No. Am. Fl. 29: 33-102. 1914.
Albert C. Smith, The American species of Thibaudieae, Contr. U. S.
Nat. Herb. 28: 311-547, tt. 1-19. 1932. Hermann Sleumer, Vaccinioi-
deen-Studien, Bot. Jahrb. 71: 375-510. 1941.

Shrubs, trees or vines, glabrous or pubescent; leaves alternate, opposite, or
verticillate, usually persistent (in our region) and often coriaceous, entire or den-
tate, without stipules; flowers regular, perfect; calyx superior or inferior, 5-4-
lobate, the segments imbricate or valvate; corolla hypogynous or epigynous, reg-
ular or rarely somewhat irregular, persistent or deciduous, with 5-4 lobes or rarely
parted almost or quite to the base, the lobes imbricate, contorted, or rarely valvate
in bud; stamens hypogynous or epigynous, sometimes adnate to the base of the
corolla, usually twice as many as the corolla lobes, rarely of the same number, the
filaments free or rarely connate; anthers 2-celled, basifixed or dorsifixed, 2-lobate at
the base, 2-4-aristate dorsally or muticous, produced above into distinct or united
tubules, these dehiscent by pores or slits; disk usually annular or swollen, crenate
or lobate, often obsolete; ovary superior or inferior, 2-12-celled, the carpels often
free above; style cylindric, filiform or columnar, the stigma peltate or capitate,
often dentate, lobate, or marginate; ovules numerous to 1 in each cell, the placentae
adnate to the interior angle of the cell or pendulous from the apex of the cell; fruit
capsular or baccate, when capsular loculicidally or septicidally dehiscent into as
many valves as there are cells, rarely separating into cocci; seeds 1-many in each
cell, generally minute, angulate or compressed; endosperm carnose, the embryo
axial; cotyledons short, the radicle terete.

STANDLEY AND WILLIAMS: FLORA OF GUATEMALA 89

About 70 genera, the species found in almost all regions of the
earth, in the tropics confined usually to the higher mountains. Five
or six other genera are represented in southern Central America and
Panama.

Ovary superior; fruit capsular or drupaceous.

Corolla of distinct petals; fruit capsular Befaria.

Corolla of united petals.
Fruit capsular, the calyx dry, not accrescent.

Corolla symmetric; native plants Leucothoe.

Corolla asymmetric; cultivated plants Rhododendron.

Fruit drupaceous or baccate, or capsular but then surrounded by a fleshy
accrescent calyx.

Fruit capsular, surrounded by the fleshy accrescent calyx Gaultheria.

Fruit baccate or drupaceous, the calyx neither fleshy nor accrescent.

Fruit drupaceous; cells of the ovary 1-ovulate Arctostaphylos.

Fruit baccate; cells of the ovary few-many-ovulate.

Fruit papillose; large shrubs or trees Arbutus.

Fruit smooth; dwarf shrubs Pernettya.

Ovary inferior; fruit baccate, or rarely dry but then berry-like.

Corolla small, 1 cm. long or usually shorter, campanulate to urceolate or urceo-
late-tubular; leaves all or mostly penninerved.

Fruit dry at maturity and with thin fragile walls; plants epiphytic and pend-
ent, or terrestrial and repent; leaves entire Sphyrospermum.

Fruit juicy at maturity, not with fragile walls.

Pedicels and usually also the calyx subtended by 2 broad bracts; stamens 10;
small shrubs with very small, coriaceous leaves Disterigma.

Pedicels and calyx without bracts or these small and inconspicuous.

Vaccinium.

Corolla usually 1.5-3 cm. long, if smaller the plants epiphytic and conspicuously
3-5-plinerved.

Inflorescences in bud covered with large, usually rose-colored bracts, these
usually persistent in anthesis Cavendishia.

Inflorescences not covered with colored bracts in bud, the bracts not con-
spicuous.

Stamens all of equal length; anther sacs conspicuously granular . . Macleania.
Stamens unequal; anther sacs smooth or nearly so.

Filaments unequal, free or nearly so; inflorescence long-racemose and
conspicuously secund Empedoclesia.

Filaments equal in length, connate for their whole length; inflorescence
short, not secund Satyria.

ARBUTUS L.

Shrubs or trees, the bark usually smooth and peeling off in sheets; leaves alter-
nate, usually on rather long petioles, generally coriaceous, entire or dentate; flowers
perfect, paniculate, the panicles terminal, compound; calyx of ten persistent, the 5
lobes much longer than the tube, equal or nearly so; corolla white or pink, urceolate,

90

FIELDIANA: BOTANY, VOLUME 24

the 5 lobes spreading or recurved, rounded, much shorter than the tube; stamens
10, included, the filaments dilated at the base, usually pubescent, not appendaged;
anthers broad, each with 2 slender awns; disk generally 10-lobate; ovary sessile,
mostly 5-celled, the style columnar or subulate, the stigma subcapitate, obscurely
5-lobate; ovules numerous, attached at the inner angle of the cell; fruit drupaceous
or baccate, globose or depressed-globose, rough-granular.

About 10 species, in North America and western Europe. Only
the following is found in Central America.

Arbutus xalapensis HBK. Nov. Gen. & Sp. 3: 279. 1819. A.
rubescens Bertol. Fl. Guat. 420. 1. 42. 1840 (type from Antigua, Saca-
tepe*quez, Vel&squez). A. varians Benth. PI. Hartw. 77. 1841 (type
from Quezaltenango, Hartweg 542}. Arctostaphylos rubescens Hemsl.
Biol. Cent. Am. Bot. 2: 279. 1881. Arbutus donnell-smithii Small,
No. Am. Fl. 29: 85. 1914 (type from San Lucas, Sacatepe"quez, J. D.
Smith 2190). Madrono; guayabo (Volcan de Santa Maria); Guaya-
billo; encino; canelon (fide Aguilar) ; chulube (Huehuetenango) ; pulca
(Quezaltenango) ; ucA (Volcan de Santa Maria) ; madrdn.

U.NVS

FIG. 23. Arbutus xalapensis. A, Flowering branch, X 2^. B, Flower with
subtending bract and bractlets, X 33^. C, Fruit, X 2. D, Anther from front, X 8.
E, Pistil, style, anther with subtending disk, X 6. F, Stamen, X 8. G, Pubescence
along mid-nerve on undersurface of leaf, enlarged.

STANDLEY AND WILLIAMS: FLORA OF GUATEMALA 91

Brushy hillsides, usually in pine or oak forest, 1,300-2,800 meters;
Baja Verapaz; Zacapa; Jalapa; Guatemala; Sacatepe"quez; Chimalte-
nango; Solola; Quiche"; Huehuetenango; Quezaltenango; San Marcos.
Western Texas; Mexico; El Salvador; Honduras; Nicaragua.

A shrub or tree, commonly 4-9 meters tall, at times probably larger, the bark
smooth, brown, peeling off in large thin sheets, the branchlets reddish, when young
tomentose with brown or whitish hairs and often pilose with long gland-tipped
hairs; leaves rather long-petiolate, persistent for most of the year but often decid-
uous at the end of the dry season, broadly oblong to ovate, oval, or ovate-lanceo-
late, mostly 8-14 cm. long, obtuse or acute, subcordate to rounded at the base,
entire or serrate, green and almost glabrous on the upper surface, usually densely
tomentose beneath with rufous or whitish hairs when young, often glabrate in age;
panicles 3-8 cm. long, many-flowered, the branches and pedicels densely tomentose
and usually with scattered gland-tipped hairs; calyx 4-6 mm. broad, the lobes
broadly ovate, obtuse, tomentulose or glabrate; corolla 6-8 mm. long, white or
pinkish, glabrous; ovary pubescent or glabrous; fruits bright red, globose, 8-12
mm. long.

Called "indio desnudo" in Honduras, in reference to the smooth
copper-colored bark. The usual names in Guatemala are "madrono"
or "madr6n." The name "guayabo" is sometimes given because the
bark suggests that of the guava (Psidium). The species has been
reported from Guatemala as A. glandulosa Mart., a name given to
a Mexican plant that probably is no more than a form of A. xalapen-
sis. There is no reason for supposing that A. donnell-smithii is more
than a casual form of A. xalapensis. Both it and A. rubescens were
described from the same region and are presumed to be synonymous,
although it is not stated whether the ovary of A. rubescens is glabrous
or pubescent. Small separated A. donnell-smithii on the basis of the
glabrous ovary, that of A. xalapensis being pubescent. It is absurd
to suppose that the two species, otherwise exactly alike, occur in the
same part of Guatemala, to be distinguished only by the absence or
presence of a few hairs on the ovary.

ARCTOSTAPHYLOS Adanson

Erect or prostrate shrubs or often trees, glabrous or pubescent; leaves alter-
nate, petiolate, coriaceous, entire or serrate; flowers small, white or pink, in ter-
minal racemes or panicles, nutant, pedicellate, bracteate and bracteolate; calyx
5-parted, persistent, inferior; corolla globose to urceolate or subcampanulate, the
limb 5-lobate, the lobes reflexed, imbricate; stamens generally 10, included, the fila-
ments short, subulate, adnate at the very base of the corolla or sometimes inserted
on the disc and free from the corolla; anthers short, laterally compressed, affixed
dorsally below the apex, 2-porose at the apex, bearing 2 dorsal recurved awns; disc
10-lobate or entire; ovary sessile, 5-celled, the style filiform, the stigma small, ob-
scurely bilobate; ovules solitary in the cells, pendulous from the upper part of the

92 FIELDIANA: BOTANY, VOLUME 24

cell; fruit drupaceous, globose, smooth or granular, containing 5 bony nutlets, or
the putamen osseous and 5-10-celled, 5-seeded or by abortion 1-4-seeded; seeds
pendulous, the funicle short, the testa membranaceous; endosperm carnose.

A genus of about 45 species, two of them widely dispersed in the
northern hemisphere, the others in temperate North America or in
the mountains of tropical North America. One other Central Amer-
ican species is found in Costa Rica.

The separation of Arctostaphylos and Arbutus is not always easy
and it is not impossible that only one genus is represented. An addi-
tional genus, Comarostaphylis Zucc., is sometimes used and some of
our species would belong here if it were maintained. Comarostaphylis
is perhaps intermediate between Arbutus and Arctostaphylos. The
differences between these genera are largely in the fruits. Arbutus
has usually 5-celled fruits which are said to be berries, but would
not seem always to be berries. The seeds are usually several con-
tained in an osseous endocarp or one tending to become indurated.
Comarostaphylis has a 4-5-celled ovary also, the endocarp is very hard
and bony, but with the cells containing a single seed and these grown
together to form a hard bony ovary surrounded by a granular cover-
ing (as in Arbutus) . Arctostaphylos typically has 4-10 seed-like nut-
lets, usually (or always?) with a single ovule; the nutlets may be
quite free in the pulp of the drupe or joined to form a solid stone.
Because the nutlets do sometimes form a solid stone most botanists
have placed Comarostaphylis into the genus Arctostaphylos. How-
ever, the nutlets of Arbutus also form a single stone so this character is
not consistently one that can be said to belong only to Arctostaphylos.

Perhaps the only difference between Arctostaphylos (sens, lat.)
and Arbutus is the single ovule in each cell of the ovary in the former
and several seeds in each cell in Arbutus. However, there seems to
be no other basic difference in flower or fruit structure of Arbutus
and that part of the genus Arctostaphylos sometimes referred to as
the genus Comarostaphylis.

Fruit smooth; prostrate shrubs; leaves small, mostly 2 cm. long or less, entire.

A. cratericola.
Fruit verrucose or granular; tall erect shrubs or small trees; leaves much larger,

serrate to entire.
Leaves densely tomentose beneath when young, some of the pubescence usually

persistent in age, usually entire or merely undulate A. arbutoides.

Leaves glabrous in age, closely and very acutely serrate A. pyrifolia-

Arctostaphylos arbutoides (Lindl.) Hemsl. Biol. Cent. Am.
Bot. 2: 278. 1881. Comarostaphylis arbutoides Lindl. Bot. Reg. 29:
t. 30. 1843. Chilu (Huehuetenango) .

STANDLEY AND WILLIAMS: FLORA OF GUATEMALA

93

Dry or moist, brushy or rocky hillsides, often in pine or oak forest,
1,800-3,000 meters; Zacapa; Jalapa; Baja Verapaz; Guatemala; Chi-
maltenango; Quiche"; Huehuetenango; Quezaltenango (described from
plants grown in London, the seeds collected near Quezaltenango by
Hartweg). Mexico; Honduras; Costa Rica; Panama.

A stout shrub 1-4.5 meters high, the branches terete, brown or blackish, when
young rufous-tomentose, often also glandular-hispid; leaves short-petiolate, cori-
aceous, narrowly oblong to elliptic-oblong or oblanceolate-oblong, mostly 5-9 cm.
long, acute to rounded and mucronate at the apex, acute or attenuate at the base,
usually with strongly revolute margins, glabrous above, at least in age, usually
persistently rufous-tomentose beneath, entire or undulate or on sterile branches
often finely serrate; panicles laxly many-flowered, usually longer than the leaves,
densely and laxly tomentose, often also hirsute; calyx 4-5 mm. broad, the lobes
triangular-ovate or triangular-lanceolate, acute, tomentulose; corolla 6-7 mm. long,
laxly tomentose, white; ovary pilose; fruit globose, glabrous, 5 mm. in diameter,
coarsely granular.

Arctostaphylos cratericola Donn.-Sm. Bot. Gaz. 55: 437. 1913.
A. pungens var. cratericola Donn.-Sm. I.e. 16: 13. 1891. Uvaursi cra-
tericola Abrams, No. Am. Fl. 29: 94. 1914. Abril (Volcan de Zunil).

FIG. 24. Arctostaphylos cratericola. A, Flowering branch, X 2. B, Flower,
X 8. C, Fruit, X 2. D, Ovary, style, and stamen, X ± 8. E, Anther from rear,
X ±12.

94 FIELDIANA: BOTANY, VOLUME 24

Dry or wet, alpine meadows or hillsides, often in thickets of Juni-
perus 2,400-3,800 meters; endemic; Sacatepe"quez (type from crater
of Volcan de Agua, J. D. Smith 2159); Chimaltenango; Huehuete-
nango; Quezaltenango.

A depressed prostrate shrub, the stems a meter long or less, sometimes rooting,
often forming dense close mats over the ground, branched, densely leafy, with ex-
foliating bark, the young branchlets puberulent; leaves short-petiolate, coriaceous,
lustrous, obovate or obovate-elliptic, mostly 1.5-2 cm. long, obtuse or acute and
apiculate, acute or obtuse at the base, glabrous, inconspicuously reticulate-veined,
entire; racemes subsessile, nutant, almost head-like, 6-8-flowered, puberulent;
flowers short-pedicellate, white or pinkish; calyx lobes suborbicular, 2 mm. long,
ciliate; corolla urceolate, 6-7 mm. long; fruit bright red, 5-7 mm. in diameter,
smooth, glabrous.

Skutch reports that the fruit is insipid. The plant is abundant at
many places in the region of Che'mal in the Cuchumatanes, often
forming large colonies among bushes of Juniperus. In general appear-
ance it is much like A. uva-ursi (L.) Spreng. of the United States.

Arctostaphylos pyrifolia (Donn.-Sm.) Standl. & Steyerm. Field
Mus. Bot. 23: 138. 1944. A. arguta var. pyrifolia Donn.-Sm. ex Loes.
Bull. Herb. Boiss. II. 3: 219. 1903. Comarostaphylis pyrifolia Small,
No. Am. Fl. 29: 91. 1914. Madron; shulup.

Dry or moist, open or brushy rocky slopes, often or usually on
limestone, frequently with Juniperus standleyi, Abies, or Pinus aya-
cahuite, 2,500-4,000 meters; San Marcos; Huehuetenango (type from
the region of Todos Santos, Seler 2633); Quiche". Mexico (Chiapas).

A shrub or small tree 1-6 meters high, the branches glabrous; leaves subcori-
aceous, on rather short, slender petioles, yellowish green when dried, oval to oblong
or lance-oblong, mostly 4-8 cm. long, acute or short-acuminate, acute at the base,
finely and evenly spinulose-serrate, glabrous throughout or nearly so, silvery be-
neath when fresh, the costa impressed above, salient beneath, the nerves and veins
often rather prominent and reticulate beneath, the upper surfaces usually lustrous;
panicles lax or dense, the branches tomentulose or glabrate; calyx 4.5-5.5 mm.
broad, the lobes deltoid or ovate-deltoid, finely pubescent, ciliate, acute; corolla
5-6 mm. long, white or tinged with pink, minutely pubescent, or almost glabrous;
fruit globose, 4-5 mm. in diameter, glabrous, densely granulate.

This species may be only a variation of A. arbutoides, with gla-
brous or nearly glabrous and spinose-serrate leaves (except in one
specimen) .

BEFARIA Mutis

References: R. Mansfeld and H. Sleumer, Revision der Gattung
Befaria Mutis, Notizbl. Bot. Gart. Berlin 12:235-276. 1935. W. H.

STANDLEY AND WILLIAMS: FLORA OF GUATEMALA 95

Camp, Studies in the Ericales. A discussion of the genus Befaria in
North America, Bull. Torr. Bot. Club 68: 100-111. 1941.

Shrubs or small trees, glabrous or pubescent, the pubescence often glandular;
leaves alternate, coriaceous, petiolate, entire or nearly so, flat or often with revo-
lute margins; flowers large, purple, pink, or rarely white, in terminal or sub terminal,
few-many-flowered, bracteate or ebracteate racemes or corymbs; calyx campanu-
late, usually 7-parted, the segments persistent, ovate, short; petals usually 7, spat-
ulate, thin, free, somewhat imbricate in bud, usually glabrous, much longer than
the sepals, soon deciduous; stamens inserted on the receptacle, twice as many as
the petals or more numerous, about equaling the petals, the filaments free, elon-
gate, subulate, slightly declinate, generally pubescent below; anthers oblong, dorsi-
fixed, muticous, obliquely 2-pored at the apex; disk obsolete; ovary superior,
depressed-globose, glabrous or nearly so, 7-sulcate, 6-7-celled; style filiform, elon-
gate, exserted, glabrous, the stigma depressed-capitate; ovules numerous in each
cell; capsule depressed-globose, 6-7-celled, many-seeded, septicidally 7-valvate
from the apex, the valves thick-crustaceous, separating from the central axis; seeds
flat, imbricate, elongate.

The genus contains some 25 species, mostly from Andean South
America, and are often difficult to distinguish one from another.
The two species here recognized are perhaps only variations of the
South American Befaria glauca H. & B.

Plants glandular-pubescent on the branches and inflorescence B. mexicana.

Plants glabrous or at least without glandular pubescence B. guatemalensis.

Befaria guatemalensis Camp, Bull. Torr. Bot. Club 68: 110.
1941. Azajarillo.

Pine forest, 1,000-2,400 meters; Zacapa (type from Sierra de las
Minas, Steyermark 29783). Mexico; Honduras.

Shrub or tree 3-9 m. tall, the branches glabrous; leaves oblong-lanceolate, cori-
aceous, glabrous, 5-9 cm. long, 1.5-2.5 cm. broad, short-petiolate, entire, acute or
obtuse, acute at the base, green above, paler beneath but sparsely glaucous; in-
florescence short-racemose or subcorymbose, 6-10-flowered, the rachis and pedicels
glabrous, the pedicels stout, 1-3 cm. long; calyx glabrous, in anthesis 4 mm. broad,
the lobes deltoid, ciliate; petals white, obovate-spatulate, attenuate below, 2-2.5
cm. long and 5 mm. broad, sparsely puberulent at the apex; style slender, 3 cm.
long; capsule blackish, about 8 mm. high.

Befaria mexicana Benth. PI. Hartw. 15. 1839.
At 1,500-1,800 meters; Zacapa (Rio Sitio Nuevo, Sierra de las
Minas). Southern and western Mexico; British Honduras; Honduras.

A shrub or a tree to about 7 m. high, the young branches pubescent with gland-
tipped, spreading, grayish or ferruginous hairs, glabrate in age; leaves short-petio-
late, chiefly oblong-lanceolate, 2.5-8 cm. long, 1-3.5 cm. broad, subacute to
rounded at the apex, acute at the base, entire, more or less tomentose when young,

FIG. 25. Befaria guatemalensis. A, Flowering branch, natural size. B, Cap-
sule from below and from above, X 2>a- C, Anthers from front and back, X 10.
D, Stigma, X 21A-

96

STANDLEY AND WILLIAMS: FLORA OF GUATEMALA 97

often with glandular hairs on the mid-rib beneath, in age glabrous or nearly so,
glaucous or green beneath; inflorescence subcorymbose, many-flowered, densely
glandular-pilose; calyx and hypanthium glandular-pubescent or glabrate, the calyx
lobes ovate-lanceolate, subacute, 4-5 mm. long; petals glabrous, white, 3.5-4 cm.
long.

The single Guatemalan collection is an imperfect one, but if pubes-
cence is dependable as a specific character, it is referable to B. mexi-
cana rather than to B. guatemalensis of the same region.

CAVENDISHIA Lindley

Shrubs, usually epiphytic; leaves alternate, petiolate, 3-5-plinerved or penni-
nerved, coriaceous; inflorescence axillary or terminal, subfasciculate or racemose,
bracteate at the base and when young usually enveloped by large, submembra-
naceous, reddish or pinkish bracts; flowers pedicellate, few-many in each inflores-
cence, the pedicels bearing 2 deciduous bractlets; base of the flower articulate with
the pedicel, the hypanthium short-cylindric or campanulate; calyx erect or spread-
ing, 5-lobate, the lobes triangular or ovate; corolla subcylindric, 5-lobate, the lobes
triangular or oblong; stamens 10, alternately unequal, almost as long as the corolla;
filaments distinct or coherent at the base, alternately unequal, attached to the
anther dorsally near its base; anthers firm or membranaceous, the cells smooth or
slightly granular, the tubules somewhat longer than the cells, opening by elongate
introrse clefts; style filiform, about equaling the corolla; fruits baccate, juicy.

More than seventy species, chiefly in the mountains of tropical
America. Numerous other species occur in Central America and
Panama. The plants are not very common in Guatemala, except
about Coban and in some parts of the mountains of the western high-
lands, where they are plentiful enough to be used for decorations.
They are well suited to this purpose because the flowers last for a
long time, in or out of water. The flowers and the bracts of most
species are strikingly beautiful because of their abundance, their form,
and their brilliant pink coloring. In Costa Rica, where the plants are
abundant in the higher mountains, they are among the most beauti-
ful and showy of all native plants. The fruits are juicy and edible,
in most species at least, and they have an agreeable but rather strong
acid flavor. Some of these plants evidently contain a substantial
amount of essential oil, for when in process of drying in the press,
which requires some time because of the leathery leaves, and espe-
cially when dried by artificial heat, they often give off a very intense,
penetrating, and rather agreeable odor, slightly suggestive of winter-
green. The genus is near its northern limit in Guatemala, and but
few species are found in the Mexican mountains. The species are
most numerous from Costa Rica southward to Ecuador.

98 FIELDIANA: BOTANY, VOLUME 24

Calyx longer than the hypanthium, the hypanthium slightly constricted at the
apex; corolla hirsute with spreading hairs C. callista.

Calyx usually much shorter than the hypanthium, the hypanthium not constricted
at the apex; corolla glabrous or pubescent.

Racemes elongate, the rachis 3 cm. long or usually longer; bracts often shorter
than the pedicels, not covering the inflorescence; corolla 15 mm. long or less.

Corolla 9-10 mm. long C. laurifolia.

Corolla about 15 mm. long C. crassifolia.

Racemes short, the rachis less than 3 cm. long; bracts longer than the pedicels
and usually enveloping the inflorescence; corolla often more than 15 mm.
long.

Flowers glabrous or nearly so C. guatemalensis var. chiapensis.

Flowers uniformly and usually densely pubescent throughout.

C. guatemalensis.

Cavendishia callista Donn.-Sm. Bot. Gaz. 20: 5, t. 2. 1895.
Jolonajtzd (Quecchi).

Epiphytic in wet mountain forest, 350-1,500 meters; endemic;
Alta Verapaz (type collected between Coban and Samac, Tuerckheim
91+1} ; Huehuetenango.

A large shrub, sometimes 1.5 meters tall, sparsely branched, the branches
terete, brownish or cinereous, sparsely setose-pilose or glabrous; leaves on peti-
oles 5-12 mm. long, coriaceous, oblong or ovate-oblong, long-acuminate, truncate
or subcordate at the base, glabrous or nearly so, 5-7-plinerved; inflorescences ter-
minal, racemose, 15-35-flowered, the racemes solitary or geminate, the bracts
numerous, as much as 2 cm. long, rose-red, oblong or ovate; rachis 5-10 cm. long,
glabrous, the glabrous pedicels 6-12 mm. long, each one subtended by a bract sim-
ilar to the basal ones; hypanthium glabrous, 2-2.5 mm. long, slightly constricted
at the apex; calyx erect, glabrous, 6 mm. long, the lobes oblong, subacute; corolla
about 18 mm. long, densely pubescent with rather long, pale, spreading hairs;
stamens subequal, 15 mm. long.

Perhaps the most handsome of the Guatemalan species. It was
seen in cultivation in the finca at Samac, where it grew luxuriantly.

Cavendishia crassifolia (Benth.) Hemsl. Biol. Centr. Am. Bot.
2: 273. 1881. Thibaudia crassifolia Benth. PI. Hartw. 65. 1840.

On trees in wet forest, 300-900 meters; Alta Verapaz; Quiche".
Southern Mexico.

A small or large shrub, the branchlets terete, dark brown, glabrous; leaves on
petioles 5-10 mm. long, oblong-ovate, 7-13 cm. long, 2.5-5 cm. broad, caudate-
acuminate, cuneate or rounded at the base, glabrous or nearly so, 5-plinerved;
inflorescences terminal or axillary, 10-20-flowered, the rachis glabrous, 2-5 cm.
long; pedicels 6-11 mm. long, glabrous or minutely puberulent; hypanthium cylin-
dric or broadly campanulate, glabrous or minutely puberulent, 2 mm. long; calyx
2 mm. long, short-pilose or glabrate, the lobes triangular, acute, 1 mm. long; corolla

STANDLEY AND WILLIAMS: FLORA OF GUATEMALA 99

15 mm. long, glabrous; stamens 12 mm. long; young fruit 6 mm. in diameter, when
mature and fresh doubtless much larger.

Cavendishia guatemalensis Loes. Bull. Herb. Boiss. II. 3: 221.
1903. Clavel georgino.

Usually epiphytic in dense wet forest, but often growing on cliffs
or banks, sometimes erect on wet shaded soil, 300-3,000 meters; en-
demic, so far as known, but to be expected in Chiapas; Alta Verapaz;
El Progreso; Guatemala; Sacatepe"quez; Suchitepe"quez; Quiche"; Hue-
huetenango; Quezaltenango; San Marcos.

A shrub 1-4 meters long, when terrestrial usually erect, most often epiphytic
and then pendent or ascending, the branches often angulate, brownish or grayish,
glabrous or sometimes pubescent; leaves on petioles 5-10 mm. long, oblong or
ovate-oblong, mostly 10-15 cm. long and 3.5-6.5 cm. broad, acuminate, rounded
or truncate at the base, 5-7-plinerved ; inflorescences axillary or terminal, short-
racemose, 10-20-flowered, with numerous large, rose-colored, broadly ovate to ob-
long bracts at the base, the bracts densely pubescent on the outer surface; pedicels
pubescent, 8-12 mm. long, each subtended by a large deciduous bract similar to
the basal ones; calyx and hypanthium pilosulous, the hypanthium broadly cam-
panulate, 2 mm. long, the calyx 3 mm. long, the lobes triangular, acute, 2 mm.
long; corolla 15-16 mm. long, 4-5 mm. broad, densely to sparsely pilose with pale
spreading hairs, white or coral red with a white apex; stamens 11 mm. long; young
fruit subglobose, glabrate, 7 mm. or more in diameter, probably white at maturity.

This is the commonest Cavendishia of Guatemala and now in
need of detailed study. The material we have placed here seems to
be quite variable and perhaps not all one species. This species is
occasionally used to decorate altars and homes on religious holidays.
It has been "collected" from the shrine at Fuentes Georginas in Que-
zaltenango.

Gavendishia guatemalensis var. chiapensis (Brandegee) L.
Wms. Fieldiana, Bot. 31: 169. 1965. C. chiapensis Brandegee,
Univ. Cal. Publ. Bot. 6: 188. 1915 (type from Cerro El Boquer6n,
Chiapas).

Epiphyte on trees in mountain forests or occasionally a terrestrial
shrub, 1,200-2,700 meters; Huehuetenango; San Marcos. Mexico
(Chiapas); El Salvador; Honduras.

Similar to the species except glabrous in all parts.

Called "arrayan" in Chiapas; "flor del Nino" and "flor de mon-
tana" in Honduras.

Cavendishia laurifolia (Klotzsch) Benth. & Hook. Gen. PI. 2:
570. 1876. Polyboea laurifolia Klotzsch, Linnaea 24: 31. 1851. Chup-

100

FIELDIANA: BOTANY, VOLUME 24

alon laurifolium Kuntze, Rev. Gen. 383. 1891. Cavendishia tuerck-
heimii Hoerold, Bot. Jahrb. 42: 323. 1909 (type from Coban, Alta
Verapaz, Tuerckheim 64) . Pie de pdjaro; rok'h-ik'h (Coban, Quecchi) .

Usually on trees in dense wet mixed forest, sometimes terrestrial
in wet shaded places, 250-2,000 meters; Alta Verapaz (type said to
have been collected in Alta Verapaz by Warscewicz); Baja Verapaz;
Guatemala; Quiche*; Huehuetenango. Southern Mexico.

FlG. 26. Cavendishia guatemalensis. A, Flowering branch, X >£• B, Flower,
X 21^. C, Corolla opened to show stamens, X 2J^.

STANDLEY AND WILLIAMS: FLORA OF GUATEMALA 101

A small or rather large shrub, suberect or pendent, the branches slender, terete,
brownish or grayish, sparsely puberulent when young; leaves on petioles 2-4 mm.
long, oblong-ovate to lanceolate, mostly 5-9 cm. long and 2-3.5 cm. broad, long-
acuminate, cuneate to truncate at the base, 5-plinerved; inflorescences terminal or
axillary near the ends of the branches, 10-30-flowered, the rachis 3-14 cm. long,
glabrous; pedicels glabrous, 1-2 cm. long, each subtended by an oblong or obovate
bract 1 cm. long or shorter; hypanthium short-cylindric, 1-2 mm. long, puberulent
or glabrate; calyx 2-2.5 mm. long, the lobes triangular, apiculate, 1 mm. long;
corolla 9-10 mm. long, white with a pink apex or rose-pink with a white tip, 4 mm.
broad, sparsely puberulent or glabrous; stamens 7-8 mm. long.

This is less showy than other local species. It is common in wet
forests of the Coban region, often growing with or near C. guatema-
lensis. The plant is more slender and has smaller leaves and flowers
than the other Guatemalan Cavendishias.

Cavendishia warszewiczii (Klotzsch) Hemsl. Biol. Centr. Am.
Bot. 2: 274. 1881. Proclesia warszewiczii Klotzsch, Linnaea 24: 35.
1851. Chupalon warszewiczii Kuntze, Rev. Gen. 383. 1891. The
type is supposed to have been collected in the mountains of Guate-
mala by Warscewicz. According to Smith, the identification of the
plant described is uncertain.

DISTERIGMA (Klotzsch) Niedenzu
Reference: Albert C. Smith, Disterigma, Brittonia 1: 216-232. 1933.

Low terrestrial shrubs, often dwarf and densely branched, the branches densely
leafy; leaves alternate, coriaceous, small, short-petiolate, the nerves usually very
obscure; flowers 1-6 in the leaf axils, often subsessile, subtended by a few minute
subcoriaceous bracts; pedicels 2-bracteolate at the apex, the bractlets relatively
large and embracing the hypanthium; hypanthium articulate with the pedicel,
campanulate or short-cylindric; calyx suberect, 4-5-lobate; corolla subcylindric or
campanulate-cylindric, 4-5-lobate; stamens twice as many as the corolla lobes,
equal, the filaments liguliform, attached to the anthers dorsally near the base; an-
thers membranous, produced into 2 tubules, dehiscent by oval introrse slits; style
filiform, about equaling the corolla, the stigma truncate; ovary 4-5-celled, the
ovules numerous; disk annular, carnose; fruit a coriaceous berry.

Species about 15, ranging from Guatemala to Bolivia, mostly at
very high elevations. One other North American species has been
described from Panama.

Disterigma humboldtii (Klotzsch) Niedenzu, Bot. Jahrb. 11:
224. 1889. Vaccinium humboldtii Klotzsch, Linnaea 24: 57. 1851.

Epiphytic or terrestrial in dense, moist or wet forest, 900-2,600
meters or perhaps even higher; Alta Verapaz; El Progreso; Zacapa;
Huehuetenango. Costa Rica and Panama; Colombia; British Guiana.

102

FIELDIANA: BOTANY, VOLUME 24

A dense shrub, erect or sometimes prostrate, usually 50 cm. high or less, the
branches stiff, brownish or grayish, with shredded bark, hispidulous; leaves on
petioles 1-2 mm. long, coriaceous, glabrous, broadly ovate to oval or broadly ob-
long, 7-15 mm. long, 4-7 mm. broad, rounded at each end, 3-5-nerved at the base
but the nerves usually obscure, paler beneath; flowers glabrous, 1-3 in the leaf
axils, on pedicels 2.5 mm. long or less, the bractlets ovate, rounded at the apex,

FIG. 27. Disterigma humboldtii. A, Flowering branch, X 1. B, Flower, X 5.
C, Flower opened out to show placement of stamens, X 5. D, Stamen from front,
X 1Y2. E, Stamen outline from side, X 11A.

STANDLEY AND WILLIAMS: FLORA OF GUATEMALA

103

2-2.5 mm. long; hypanthium 1-2 mm. long; calyx 2-2.5 mm. long, the 4 lobes del-
toid, acute; corolla white or pink, glabrous, 6-7 mm. long, the 4 short lobes deltoid;
anthers 2.5-4 mm. long, the tubules longer than the sacs.

The plant appears to be very rare in Guatemala, only three col-
lections of it having been made.

FIG. 28. Empedoclesia brachysiphon. A, Flowering branch, X 14. B, Corolla
opened out to show placement of stamens and relative length of style, X 2. C, An-
thers, front and side, X 10.

104 FIELDIANA: BOTANY, VOLUME 24

EMPEDOCLESIA Sleumer

Glabrous epiphytic shrubs with very stout branches; leaves ovate or oblong-
ovate, very thick, lustrous, penninerved, on very short, thick petioles, minutely
denticulate or almost entire; flowers rather large, red, in long axillary many-flow-
ered secund racemes, the flowers articulate with the pedicels; hypanthium broadly
obconic; calyx suberect, 5-lobate; corolla tubular, shallowly 5-lobate; stamens 10,
much shorter than the corolla, alternately slightly unequal, hirsutulous; anthers
granular, the tubules submembranaceous, very short, distinct, opening by round
terminal pores; style filiform, shorter than the corolla.

The genus consists of a single species.

Empedoclesia brachysiphon Sleumer, Notizbl. Bot. Gart. Ber-
lin 12: 124. 1934.

On trees in wet forest, 1,500-2,600 meters; Alta Verapaz (type
from Coban, Tuerckheim 4196); Zacapa (Sierra de las Minas); Hue-
huetenango (Maxbal).

A stout branched shrub, the branches 7-13 mm. thick, apparently obtuse-
angulate; leaves rigid-coriaceous, thicker than in any other local plants of this
family, oblong-ovate or broadly ovate, 6-14 cm. long, 4-8 cm. broad, on very
short, thick petioles, slightly narrowed to the obtuse or very obtuse apex, broadly
rounded or subcordate at the base, lustrous, subentire or with remote black glands
along the margin and callous-mucronate, the venation not elevated on the upper
surface, the costa prominent beneath, the lateral nerves 6-7 pairs, the veins obso-
lete; racemes stout, 10-20 cm. long, many-flowered, strongly secund, 20-25-flow-
ered, the rachis thick, angulate; pedicels very stout, in anthesis 5-7 mm. long,
subtended at the base by a thick elongate-deltoid bract 2.5 mm. long, bearing at
the base of the calyx 2 similar bractlets; hypanthium and calyx together 9 mm.
long, the lobes deltoid, 2-2.5 mm. long; corolla scarlet, fleshy, 18-25 mm. long,
3.5 mm. broad, the lobes obtuse, 3.5 mm. long; stamens one-third as long as the
corolla, the anthers 4 mm. long, the tubules 1 mm. long.

An unusually showy and handsome plant, apparently rare since
it has been collected but three times. In the Thibaudieae it is note-
worthy for its very thick and rigid, practically sessile leaves, and for
the much elongate, one-sided racemes of brightly colored flowers.

GAULTHERIA L.

References: Hermann Sleumer, Notizbl. Bot. Gart. Berlin 12: 285-
287. 1935. W. H. Camp, Gaultheria, Bull. Torr. Bot. Club 66: 9-21.
1939. Louis 0. Williams, Fieldiana, Bot. 31: 169-175. 1965.

Mostly low shrubs, terrestrial, often with horizontal rootstocks, pubescent or
glabrous, often with gland-tipped hairs; leaves alternate, coriaceous, petiolate,
broad, dentate or almost entire; flowers solitary in the leaf axils or usually in

STANDLEY AND WILLIAMS: FLORA OF GUATEMALA 105

racemes or panicles; calyx inferior, persistent, much longer than the hypanthium,
usually accrescent; corolla campanulate, conic, ovoid, or urceolate, generally whit-
ish or pink, the 5 short lobes spreading or recurved; stamens 10, included, the fila-
ments adnate to the base of the corolla tube; anthers usually 2-awned, opening by
terminal pores; disk 10-lobate; ovary superior, 5-celled, 5-lobate, glabrous; style
columnar, the stigma entire; ovules numerous in each cell; fruit a capsule included
in the fleshy or berry-like accrescent calyx and hypanthium.

Perhaps as many as 50 species, mostly in the mountains of trop-
ical America, a few in temperate North America, and others in Asia,
Australia, and New Zealand. Perhaps one or two others are found
in southern Central America and Panama. As in other genera of the
family, many of the species seem to be vaguely separable, and differ-
ent authors are far from agreement as to how they should be treated.
Gaultheria procumbens L. of northern North America furnishes by
distillation the aromatic oil known as oil of wintergreen, much used
for flavoring confectionery and other articles. Most of all commer-
cial wintergreen oil is obtained actually from Betula rather than from
Gaultheria. The North American species is a small plant, very dif-
ferent in general appearance from the Central American ones. They,
however, exhale a similar odor, especially when drying. It is reported
that in Mexico the branches often are spread on the floors of houses
and churches to perfume them.

Leaves lanceolate-oblong, oblong-ovate to cordate with the bases truncate to cor-
date, mostly more than 5 cm. long.

Corolla pubescent outside.

Young branches, leaves, rachis and inflorescence (or some of them) provided
with hirsute or hirsute-glandular hairs G. odorata.

Young branches, leaves, rachis and inflorescence puberulent or some of them
glabrous G. chiapensis.

Corolla glabrous outside G. sp.

Leaves elliptic, oblong-ovate to ovate, mostly attenuate to the base and less than
3.5 cm. long G. cumingii.

Gaultheria chiapensis Camp, Bull. Torr. Bot. Club 66: 12.
1939.

Moist or wet, mixed forest, 2,000-2,500 meters; Chiquimula; Ja-
lapa. Mexico; El Salvador.

A shrub, sometimes 6 meters high (?), the branches glabrous, brown or when
old blackish; leaves ovate-oblong to ovate or lanceolate, on petioles 3-7 mm. long,
obtuse and apiculate to subacuminate, rounded at the base, mostly 5-10 cm. long
and 2-3.5 cm. broad, serrulate, coriaceous, glabrous above, black-punctate be-
neath; racemes lateral and sub terminal, 20-30-flowered, 5-10 cm. long, the rachis
densely puberulent with eglandular hairs, the pedicels puberulent, 5-9 mm. long;
bracts glabrous; calyx lobes 2 mm. long, acute, puberulent; corolla white, 5-6 mm.

106 FIELDIANA: BOTANY, VOLUME 24

long, puberulent and thinly pilose; filaments dilated at the base, pilosulous, the
anthers 2 mm. long; ovary densely whitish-pilose, the style glabrous or pilosulous;
fruit 5 mm. broad.

Perhaps only a somewhat less pubescent phase of G. odorata.

Gaultheria cumingii Sleumer, Notizbl. Bot. Gart. Berlin 12:
125. 1934.

High mountain meadows or sparse forest, 1,800-3,000 meters;
Guatemala; Solola; Huehuetenango. Mexico.

An erect, often densely branched shrub 2 meters high or less, generally only
25-50 cm. high, the branches rather slender, sparsely glandular-hirsute with ascend-
ing hairs; leaves short-petiolate, coriaceous, elliptic, ovate-oblong to broadly ovate,
mostly 2.5-3.5 cm. long and 0.6-2 cm. broad, acute or subobtuse and apiculate,
rounded or very obtuse at the base, in age glabrous or more or less glandular-
papillate beneath, serrulate, conspicuously reticulate- veined; racemes lateral and
terminal, 5 cm. long or less, lax or dense, the rachis whitish-puberulent and sparsely
glandular-hirsute, the bracts puberulent, glandular-ciliate; pedicels puberulent and
usually glandular-pilose; calyx puberulent or glabrous, the lobes ovate, acute to
abruptly acuminate, glandular-ciliate; corolla 5-6 mm. long, white to pink, wholly
glabrous or somewhat puberulent.

This plant is plentiful in the beautiful high meadows of the Volcan
de Pacaya. Six collections made there, all undoubtedly representing
the same species, exhibit well the great amount of variation that may
be found in a single species of this genus. The pubescence of the
calyx also is highly variable, indicating that this character is of rather
doubtful value in separating species of Gaultheria. Material from
the volcanoes of Solola is similarly variable.

Gaultheria odorata Willd. Ges. Naturf. Freunde Berlin Neue
Schrift. 3: 425. 1801. G. odorata var. mexicana DC. Prodr. 7: 595.
1839. G. hirtiflora Benth. PL Hartw. 66. 1840. G. hidalgensis Loes.
Bull. Herb. Boiss. 2: 552. 1894. G. hartwegiana Klotzsch ex Loes.
Bull. Herb. Boiss. II. 3: 217. 1903 (Hartweg 541, type from moun-
tains near Quezaltenango) . G. lancifolia var. dulcis Camp, Bull.
Torr. Bot. Club 66: 14. 1939. G. tacanensis Camp, I.e. 20.

Open or brushy banks or meadows, often in rocky places, common
in oak, pine, or Cupressus forest, sometimes in rather dense, mixed
forest, rarely on mossy hummocks in open swamps, 800-3,700 meters;
Alta Verapaz; Baja Verapaz; El Progreso; Zacapa; Jalapa; Guate-
mala; Sacatepe"quez; Chimaltenango; Solola; Totonicapan; Quiche";
Huehuetenango; Quezaltenango; San Marcos. Southern Mexico;
El Salvador; Honduras; Costa Rica; northern South America.

STANDLEY AND WILLIAMS: FLORA OF GUATEMALA

107

Commonly a shrub of 0.3-2 meters, sometimes as much as 5 meters high, often
only 30-50 cm. high, sometimes depressed and prostrate or nearly so and forming
mats, the branches pubescent and usually sparsely or densely glandular-hirsute;
leaves short-petiolate, coriaceous, ovate to oblong-ovate, oval-ovate or even cor-
date, mostly 3-6 cm. long, obtuse and apiculate or abruptly short-acuminate,

FIG. 29. Gaultheria odorata. A, Habit, X J^. B, Flower, X 2 H. C, Corolla
dissected showing stamens, X 5. D, Accrescent fruiting calyx, X 5. Stamen,
X 15.

108 FIELDIANA: BOTANY, VOLUME 24

rounded, truncate or cordate at the base, serrulate, when young usually sparsely
hirsute, soon glabrate; racemes 6 cm. long or shorter, dense or lax, many-flowered,
the rachis puberulent and usually also pubescent and with numerous long spread-
ing gland-tipped hairs, the pedicels similarly pubescent; calyx nearly glabrous or
puberulent, also often pubescent with glandular-hirsute hairs, the lobes ovate,
acute or short-acuminate; corolla white or pink, ovoid-urceolate, 5-7 mm. long,
pilose or glandular-pilose, often nearly glabrous; filaments pubescent; capsule de-
pressed-globose, 5 mm. broad.

A common, occasionally abundant and quite variable species in
the Central American highlands. Dr. Camp who specialized in cer-
tain Ericaceae believed G. hartwegiana to be a valid species. How-
ever, we believe it to be only a minor variation of G. odorata. G. hirti-
flora Benth. seems to be but only a more pubescent phase of G. odorata.
G. lancifolia var. dukis and G. tacanensis seem not to depart signifi-
cantly from G. odorata. Sometimes called "pasas" in El Salvador.

Gaultheria sp.

A shrub about 1.5 m. tall with a few glandular hirsute hairs on the stem; the
leaves small and lanceolate or elliptic-lanceolate, acute, attenuate to the base,
mostly 3-5 cm. long; the corolla glabrous.

This specimen is not like any of the others that we have seen but
may well be a further variation of G. odorata. The specimen is not
entirely satisfactory. (Steyermark 4.2815).

LEUCOTHOE D. Don

Shrubs or small trees; leaves alternate, petiolate, coriaceous, entire or dentate;
flowers small, perfect, in racemiform, spiciform, or corymbiform panicles, these
often one-sided and spreading or curved, axillary; calyx stellately spreading, the
lobes usually 5, short and broad, much longer than the tube; corolla white or tinted,
urceolate, much longer than the calyx, the lobes much shorter than the tube; sta-
mens 10, included, the filaments subulate or lanceolate, often pubescent; anthers
oblong to oval or subglobose, the cells not awned but obscurely or conspicuously
2-mucronate and free nearly to base; ovary 5-celled, depressed, vertically lobate,
the style columnar, straight or nearly so, the stigma slightly dilated, somewhat
lobate; fruit capsular, depressed, thick-walled, somewhat 5-lobate, septicidally
5-valvate.

About 30 species, in temperate and tropical regions of America.
Only two are known from Mexico and Central America.

Leaves glabrous beneath L. mexicana.

Leaves white pilosulose beneath L. pinetorum.

STANDLEY AND WILLIAMS: FLORA OF GUATEMALA

109

Leucothoe mexicana (Hemsl.) Small, No. Am. Fl. 29: 57. 1914.
Andromeda mexicana Hemsl. Biol. Cent. Am. Bot. 2: 282. 1881.

Mostly in open pine-oak forest, 1,100-2,500 meters; Baja Vera-
paz; El Progreso; Zacapa; Chiquimula; Santa Rosa. Southern Mex-
ico; British Honduras (San Agustin); El Salvador; Honduras.

A shrub or small tree, usually about 6 meters high, sometimes as much as 13
meters tall, the bark very rough, corky, broken into thick quadrangular plates, the
branches ferruginous or fuscous, glabrous or puberulent; leaves on slender petioles
8 mm. long or shorter, coriaceous, lanceolate or oblong-lanceolate, mostly 5-7 cm.
long, narrowly long-acuminate, usually rounded at the base, entire, lustrous above,
the venation not elevated, slightly paler beneath, the costa slender, prominent, the
lateral nerves and veins very inconspicuous, scarcely if at all elevated; glabrous or
when young somewhat pubescent; panicles corymbiform, axillary, 2-3 cm. long,
few-flowered, puberulent or glabrous; calyx 4-4.5 mm. broad, the lobes deltoid-
ovate or broadly ovate, acute or short-acuminate; corolla glabrous, cylindric-
urceolate, 9-12 mm. long; capsules hard, blackish, 5-7 mm. broad.

Called "pellejo de lagarto" in El Salvador, in allusion to the thick
checkered bark. The species has been reported from Guatemala as
Agarista multiflora G. Don.

FIG. 30. Leucothoe mexicana. A, Flowering branch, X 1. B, Flower, X 5.
C, Dehisced capsule, X 5. D, Stamen from side, X 15.

110 FIELDIANA: BOTANY, VOLUME 24

Leucothoe pinetorum Standl. & L. Wms. Ceiba 3: 54. 1952.
Open pine-oak forests, 1,000-1,600 meters; British Honduras;
Mexico; Honduras.

Shrubs or small trees to 6 meters tall, the bark thick, fissured, corky; branchlets
slender, white pilosulose; leaves lanceolate or lance-ovate, apex acuminate, rounded
or obtuse at the base, entire, green above, densely white pilosulose below, becoming
glabrous, 4-6.5 cm. long and 1.5-2.5 cm. broad; inflorescence axillary, short pe-
dunculate, few-flowered, racemose or subumbellate, pedicels about 8 mm. long,
pilosulose, bracts small, ovate, spreading; calyx 5-lobed, 4 mm. broad and 2.5 mm.
high, the lobes broadly ovate, short-acuminate; corolla white, about 9 mm. long
and 3 mm. broad, contracted at the apex, the tube glabrous outside, the lobes erect,
ovate, obtuse; capsule depressed globose, about 7 mm. broad.

Called "Cachimbo" in Honduras where it is often abundant in
the oak-pine woods. The species is closely related to L. mexicana
and perhaps only varietally distinct.

MACLEANIA Hooker

Shrubs, usually epiphytic; leaves alternate, coriaceous, penninerved or basi-
nerved; flowers rather large, usually red or orange-red, the inflorescence axillary or
terminal, the flowers subfasciculate or racemose, pedicellate, few-many in each in-
florescence, the pedicels bearing 2 deciduous bractlets; hypanthium articulate with
the pedicel, short-cylindric or campanulate; calyx erect-spreading, generally 5-
lobate, the lobes triangular, subacute; corolla subcylindric or elongate-urceolate,
the 5 short lobes triangular, subacute; stamens 10, equal, usually half as long as the
corolla, the filaments distinct or connate, attached to the anther dorsally near its
base; anthers stout, the cells strongly granular, the tubules about equaling the sacs,
laterally connate or fused into a single tubule, opening by elongate, distinct or fused
clefts; style filiform, equaling or longer than the corolla; fruit baccate.

About 30 species, chiefly in the mountains of tropical western
South America. Two others are known from southern Central Amer-
ica and one other from Mexico.

Leaves ovate or oblong-ovate, mostly about twice as long as broad . . . . M. insignis.

Leaves linear-lanceolate or oblong-lanceolate, mostly 4-5 times as long as broad.

M. insignis var. linearifolia.

Macleania insignis Mart. & Gal. Bull. Acad. Brux. 9: 531. 1842.
M. cordata Lem. Fl. Serres L 4: 312. 1848.

Usually on trees in dense wet forest, sometimes on mossy hum-
mocks in open swamps, or terrestrial on rocky shaded hillsides, 1,300-
2,500 meters; Alta Verapaz; Baja Verapaz; El Progreso; Zacapa.
Southern Mexico; British Honduras (Pine Peak, Cockscomb Moun-
tains); Honduras; Nicaragua.

STANDLEY AND WILLIAMS: FLORA OF GUATEMALA

111

An erect or pendent shrub, sometimes 4 meters long, the branches stout, terete,
dark brown, with exfoliating bark; leaves on stout petioles 2 mm. long, coriaceous,
ovate to oblong-ovate, mostly 3-8 cm. long and 1.5-4 cm. broad, obtuse or sub-
acute, subcordate at the base, entire, glabrous, 5-plinerved; inflorescences axillary,
2-3-flowered, the pedicels 8-15 mm. long, glabrous; hypanthium glabrous, 4 mm.
long, somewhat winged; calyx 2.5 mm. long, the lobes apiculate, 1.5 mm. long;
corolla elongate-urceolate, 20-25 mm. long, 5 mm. broad, contracted at the throat,
glabrous or nearly so, usually bright red or orange-red; anther sacs 4-5 mm. long,
the tubule 2-2.5 mm. long; style exserted.

When growing, this is a much more handsome and showy plant
than one would suppose from examination of rather dreary herbarium
specimens. The shrub is particularly abundant in the big open swamp
above Tactic, where it grows on the huge mossy hummocks that fill
much of the swamp. It is probable that during the wetter months
the plants stand in water for long periods.

FIG. 31. Macleania insignis. A branch of the plant, X 1A; a flower, X 1^;
a corolla opened out to show stamens, X 1; an anther from the side, X 2H; tip
of the anther showing dehiscence, much enlarged.

112 FIELDIANA: BOTANY, VOLUME 24

Macleania insignis var. linearifolia (Donn.-Sm.) Standl. & L.
Wms. Fieldiana, Bot. 31 : 172. 1965. M. cordifolia var. linearifolia
Donn.-Sm. Bot. Gaz. 16: 12. 1891. M. linearifolia A. C. Smith,
Contr. U. S. Nat. Herb. 28: 368. 1932.

Epiphytic in dense wet forest or on mossy hummocks in open
swamps, 900-1,300 meters; Alta Verapaz (type from Pansamala,
Tuerckheim 1332). Honduras.

Differs from the species in having leaves linear-lanceolate or oblong-lanceolate,
mostly 5-12 cm. long and 1-2 cm. broad.

This variety is hardly more than an extreme variation of the
above, as Captain Smith treated it.

PERNETTYA Gaudichaud

References: Hermann Sleumer, Revision der Gattung Pernettya
Gaud. Notizbl. Bot. Gart. Berlin 12: 626-655. 1935. W. H. Camp,
Pernettya, Bull. Torr. Bot. Club 66: 22-28. 1939.

Shrubs, usually small and terrestrial, erect or depressed; leaves very small,
alternate, short-petiolate or subsessile, coriaceous, dentate or subentire; flowers
small, cernuous, in small few-flowered racemes at the ends of the branches, the
inflorescences usually little if at all exceeding the leaves; calyx inferior, persistent,
the 5 segments broad, often ciliate; corolla urceolate, ovoid, or subglobose, white or
pink, its 5 lobes very short, recurved; stamens 10, included, the filaments dilated
below; anthers oblong or oval, opening by terminal pores, appendaged at the apex;
ovary 5-celled, generally globose, the style columnar, the stigma minute; fruit
baccate, globose, black or purple-black at maturity.

Species 15 or more, a few in Tasmania and New Zealand, the
others mostly in the mountains or cool lowland regions of South
America, some of the species extending northward into central Mex-
ico. One other is known from southern Central America (Costa Rica,
Panama) . Sleumer considered the Mexican material as representing
a single species, while Camp recognized six from that country.

Leaves mostly 6-9 mm. long; plants prostrate; branches minutely puberulent.

P. saxicola.

Leaves mostly 1-2 cm. long; plants erect or ascending; branches more or less stri-
gose or hirsute P. ciliata.

Pernettya ciliata (Schlecht. & Cham.) Small, No. Am. Fl. 29:
82. 1914. Gaultheria ciliata Schlecht. & Cham. Linnaea 5: 126. 1830.
P. tomasii Camp, Bull. Torr. Bot. Club 66: 25. 1939 (type from Vol-
can de Tacana, MacDougall). Cacalote (San Marcos).

Open, moist or wet or even rather dry mountain savannas or
meadows, rarely in the forest, mostly above 3,000 meters; Sacate-

STANDLEY AND WILLIAMS: FLORA OF GUATEMALA

113

p£quez; Chimaltenango; Huehuetenango; Totonicapan; Quezalte-
nango; San Marcos. Mexico; Nicaragua.

A stout erect or ascending shrub, commonly 25-50 cm. high, usually densely
branched, the bases of the plants sometimes prostrate and rooting, the branches
sparsely or densely hirsute or strigose, sometimes glabrate, densely leafy; leaves
on very short petioles, coriaceous, oblong to lance-oblong, ovate-oblong, or oval,
acute or obtuse, rounded or usually obtuse at the base, crenate-serrate, glabrous
and lustrous above, dull and paler beneath, sparsely setose along the costa or almost

FIG. 32. Pernettya ciliata. A, Habit, X 1. B, Flower, X 4. C, Corolla opened
out to show anthers, X 4. D, Fruit, X 2. E, Anther from front, X 10. F, Anther
outline from side, X 10.

114 FIELDIANA: BOTANY, VOLUME 24

glabrous; flowers short-pedicellate; calyx lobes ovate or ovate-lanceolate, 3-3.5
mm. long; corolla 5-7 mm. long, white tinged with pink; berries globose, 6-8 mm.
in diameter, black at maturity, very juicy.

Called "arrayan" in Chiapas. The species has been reported
from Guatemala as P. ciliaris G. Don, P. pilosa G. Don, and P. cori-
acea Klotzsch, the last being a species of Costa Rica and Panama.
Part of the Guatemalan specimens were examined by Camp who re-
ferred most of them to P. coriacea, a species he does not report from
Mexico. We have referred these specimens to P. ciliata. There is
no reason for supposing that that species refuses to cross the inter-
national boundary between Mexico and Guatemala and so far as we
can determine, the Guatemalan collections and the Mexican ones of
P. ciliata are exactly alike. Whether the ones from Costa Rica and
Panama represent a different species is questionable, and according
to Sleumer, the Costa Rican species is hardly different from the South
American P. prostrata (Cav.) Sleumer, which may well be the proper
name for the Central American plant. Dr. Camp's P. tomasii seems
to have no distinguishing characters. It is from near the Guatemalan
border. P. ciliata is an exceedingly abundant shrub in the high
mountains of western Guatemala, and there are thousands of acres
of it, almost continuous colonies, along the highways above Totonica-
pan and Quezaltenango. It densely covers large areas of moorland
in the sheep regions, to the exclusion of most other vegetation.
Rather strangely, it is rare in Huehuetenango, where similar soil
and climate conditions prevail, and the one fruiting collection made
there is atypical. It is reported to be poisonous to sheep, causing
them to become "drunk." The fruits look like blueberries and are
sweet, very juicy, and agreeable in flavor. The senior author has
eaten them in Costa Rica, before being warned about their prop-
erties, and suffered no ill effects, but Pittier once told him that in
the same country he had been made ill by what he believed were
these same berries.

Pernettya saxicola Standl. & Steyerm. Field Mus. Bot. 23: 139.
1944.

Known only from the type, San Marcos, creeping among crevices
of rocks on summit of the dome, Volcan de Tacana, 4,400 meters,
Steyermark 36110.

A prostrate and repent shrub, the very slender branches scarcely 1 mm. thick,
when young very minutely and inconspicuously puberulent; leaves small, almost
sessile, subcoriaceous, oblong-lanceolate or ovate-oblong, or the lowest ovate,

STANDLEY AND WILLIAMS: FLORA OF GUATEMALA 115

mostly 6-9 mm. long, 3 mm. broad or less, obtuse or subacute, obtuse or rounded
at the base, crenate, glabrous, lustrous above, the costa plane or subimpressed,
slightly paler beneath, the costa prominent, the nerves obsolete; flowers axillary,
solitary, on short thick pedicels, the pedicels not concealed by the bracts; sepals
lance-oblong, about 2.2 mm. long, acuminate, glabrous, not ciliate; corolla white,
ovoid-urceolate, 3 mm. long, glabrous.

PERNETTYA SP. — There is a single specimen from the department
of El Progreso (Steyermark lf.3071} which may represent another spe-
cies. It is a much more robust plant than others known in Central
America or Mexico with leaves much larger than the common P. cili-
ata, mostly 2-3.5 cm. long and elliptic. The specimen is in fruit but
lacks flowers.

RHODODENDRON L.

Shrubs or trees, glabrous or variously pubescent, the branches often verticil-
late; leaves alternate, entire, membranaceous to coriaceous; flowers usually large
and showy, mostly in terminal corymbs or racemes, these few-many-flowered;
calyx of 5 sepals or 5-dentate, sometimes obsolete; corolla usually funnelform or
campanulate, rarely subrotate, the limb more or less oblique, generally 5-lobate,
the lobes imbricate in bud; stamens mostly 5-10, more or less unequal, usually
declinate, the filaments subulate-filiform or short and thick, generally pilose below;
anthers dorsifixed, short or elongate, erect, muticous dorsally, dehiscent by termi-
nal pores; disk usually thick and crenate; ovary 5-20-celled, the style short or elon-
gate, declinate or incurved, the stigma capitate, 5-20-lobate; ovules numerous in
each cell; fruit capsular.

Species more than 400, in temperate or rarely arctic regions,
mostly in Asia, about 25 native in temperate and arctic North Amer-
ica. None are native in America below the southern border of the
United States.

Rhododendron indicum (L.) Sweet, Hort. Brit. ed. 2. 343.
1833. Azalea indica L. Sp. PI. 150. 1753.

Said to be a native of Japan; grown commonly for ornament in
many regions of the earth ; an occasional plant of gardens at middle
elevations in Guatemala, sometimes planted in even the colder in-
habited regions.

A densely branched shrub, usually 2 meters high or less; leaves membranaceous
or thicker, short-petiolate, elliptic-lanceolate to lanceolate or oblanceolate, mostly
2.5-3.5 cm. long, acute or obtuse and mucronate, acute at the base, subentire,
green above, paler beneath, ciliate, sparsely or rather densely appressed-pilose with
long hairs; flowers few at the ends of the branches, pedicellate, white or rose-purple,
5-7 cm. broad; stamens 5.

Occasional in Central American gardens in the cooler highlands.
Standley reports that it grows luxuriantly in Coban.

116 FIELDIANA: BOTANY, VOLUME 24

SATYRIA Klotzsch

Rather large shrubs, usually epiphytic, glabrous or nearly so; leaves alternate,
mostly 3-5-plinerved, short-petiolate, coriaceous, entire; inflorescences axillary or
terminal, few-many-flowered, the flowers fasciculate or short-racemose; flowers
pedicellate, the pedicels 2-bracteolate; hypanthium articulate with the pedicel;
calyx spreading, 5-lobate, the lobes ovate or triangular; corolla subcylindric, shal-
lowly 5-lobate, the lobes triangular; stamens 10, alternately unequal, shorter than
the corolla, often only one-third as long, the filaments connate into a tube, attached
to the anther dorsally near its base, equal; anthers subcoriaceous, alternately un-
equal, the sacs slightly granular, the tubules not clearly differentiated from the
sacs, flaring, opening by broad lateral clefts; style filiform, about equaling the
corolla; fruit baccate.

About 15 species, all American, ranging from southern Mexico
to Bolivia. Two other species are known from Central America.

Inflorescence elongate, the rachis usually 2-4 cm. long S. elongata.

Inflorescence short, the flowers fasciculate or short-racemose, the rachis rarely
more than 1 cm. long.

Corolla 18-28 mm. long S. warszewiczii.

Corolla 10-11 mm. long S. meiantha.

Satyria elongata A. C. Smith, Contr. U. S. Nat. Herb. 28: 521.
1932. Arete de guacamaya (Huehuetenango) .

On trees in dense wet forest, 150-400 meters; Alta Verapaz (type
from Cubilgiiitz, Tuerckheim 7633); Huehuetenango (Ixcan). Mex-
ico (Oaxaca); Honduras; Costa Rica.

A pendent shrub, rather coarse, with elongate branches, the branchlets gla-
brous or when young sparsely puberulent; leaves on petioles 4-5 mm. long, lanceo-
late or lance-oblong, 12-17 cm. long, 3-4 cm. broad, long-acuminate, cuneate at
the base, thick-coriaceous, 3-5-plinerved; inflorescences axillary near the ends of the
branches, racemose, 7-15-flowered; the rachis glabrous or practically so, 2.5-5 cm.
long, the pedicels 12-22 mm. long, glabrous; hypanthium minutely puberulent,
2-2.5 mm. long; calyx thin-coriaceous, 1.5 mm. long, the lobes ovate, apiculate;
corolla deep rose-red below, greenish above, 20-30 mm. long, minutely puberulent,
4 mm. broad, contracted at the throat; young fruit subglobose, 7 mm. in diameter,
capped by the persistent calyx.

Satyria meiantha Donn.-Sm. Bot. Gaz. 47: 256. 1909.

Usually epiphytic in dense wet mixed forest, sometimes terrestrial,
850-1,700 meters; Pete"n; Alta Verapaz (type from Coban, Tuerck-
heim II. 2101); Chiquimula; Huehuetenango. British Honduras
(Camp 35, Pete"n boundary).

A coarse epiphytic shrub or sometimes a terrestrial shrub or small tree as much
as 6 meters high, the branches thick, subterete, pale brown or grayish; leaves on
thick petioles 8-12 mm. long, narrowly lance-oblong to ovate-oblong, 12-22 cm.

STANDLEY AND WILLIAMS: FLORA OF GUATEMALA 117

long, 4-9 cm. broad, acuminate or narrowly long-acuminate, acute or broadly cune-
ate at the base, thick-coriaceous, 5-plinerved, glabrous; inflorescences axillary,
often at naked nodes below the leaves, short-racemose, 6-12-flowered, the rachis
5-10 mm. long or rarely longer, the slender pedicels 6-10 mm. long; hypanthium
2 mm. long, minutely puberulent or glabrate; calyx 1.5 mm. long, minutely puberu-

FIG. 33. Satyria meiantha. A, Flowering section of branch, X 1A. B, Flower,
X 2>2- C, Corolla opened out to show anthers, X 2J^. D, Anthers front and
side, X 5.

118 FIELDIANA: BOTANY, VOLUME 24

lent, the lobes triangular, apiculate; corolla dark rose or bright red, waxy, glabrous,
10-11 mm. long or sometimes somewhat longer, 2.5 mm. broad, the lobes 1 mm.
long; anthers 3-3.5 mm. long; fruit subglobose, becoming white but at maturity
bluish black.

SATYRIA OVATA A. C. Smith (Contr. U. S. Nat. Herb. 28: 524. 1932)
is known from four collections, the type and one other from Nicara-
gua, one from Guanacaste, Costa Rica, and one collected in "Guate-
mala" by Friedrichsthal. It has not been found recently in Gua-
temala, and since some of Friedrichsthal 's "Guatemala" collections
are known to have been gathered farther south, some of them in
Costa Rica, it seems likely that the species should be excluded from
the Guatemalan flora.

Satyria warszewiczii Klotzsch, Linnaea 24: 22. 1851. Huetilla
de cuaresma.

On trees in moist or wet forest, 500-2,000 meters; Alta Verapaz
(Sepacuite1) ; Chiquimula (Cerro Brujo); Suchitepe"quez (Volcan de
Zunil) ; Quezaltenango (Volcan de Santa Maria) . Mexico (Chiapas) ;
Honduras; Costa Rica; Panama.

A large epiphytic shrub, or sometimes a terrestrial shrub or small tree as much
as 6 meters high, the branches stout, subterete, glabrous, brownish or grayish;
leaves on petioles 8-16 mm. long, lance-oblong to ovate-oblong, 10-23 cm. long,
2-9 cm. broad, acuminate, cuneate or rounded at the base, thick-coriaceous, gla-
brous, 3-5-plinerved; inflorescences often numerous, short-racemose, borne in the
leaf axils or at naked nodes below the leaves, 8-30-flowered, the rachis glabrous,
5-10 mm. long; pedicels glabrous, 15-25 mm. long; hypanthium glabrous, broadly
campanulate, 2 mm. long; calyx thin-coriaceous, 2-3 mm. long, the lobes triangu-
lar, apiculate; corolla glabrous, coral-red below, white above, 18-28 mm. long,
4-5 mm. broad, contracted at the throat, the lobes 1 mm. long; anthers 5-6 mm.
long; young fruit globose, 6 mm. in diameter.

This is a very common shrub in the mountains of Costa Rica.
The juicy, somewhat acid fruits, known there as "muelas," are very
good to eat. In the same country the handsome flowers are called
"colmfflos."

SPHYROSPERMUM Poeppig & Endlicher

Reference: Albert C. Smith, The genera Sphyrospermum and Dis-
terigma, Brittonia 1: 203-232. 1933.

Small slender shrubs, usually epiphytic, sometimes terrestrial, glabrous or
pubescent, the branches often pendent; leaves alternate, short-petiolate, small,
coriaceous, fleshy when growing, obscurely nerved, entire; flowers small, few, soli-
tary in the leaf axils, the pedicels slender, cernuous, with 2 deciduous bractlets at

FIG. 34. Sphyrospermum majus. A, Flowering and fruiting branch, X 1.
B, Flower, X 5. C, Corolla opened to show stamens, X 5. D, Anthers from front
and side, X 11A.

119

120 FIELDIANA: BOTANY, VOLUME 24

the base; hypanthium continuous with the pedicel, subglobose, or obconic; calyx
suberect, 4-5-lobate; corolla subcylindric, shortly 4-5-lobate; stamens as many or
twice as many as the corolla lobes, equal or alternately slightly unequal, the fila-
ments slender, attached to the anther dorsally near the base; anthers membranous,
the 2 tubules equaling or longer than the cells, dehiscent by introrse oval clefts;
style filiform, about equaling the corolla, the stigma truncate; fruit globose or ellip-
soid, baccate, the pericarp smooth, brittle, the cells 4-5; ovules numerous.

About 10 species, mostly in the mountains of tropical America.
Three others are known from southern Central America.

Sphyrospermum niajus Griseb. Fl. Brit. W. Ind. 143. 1859.

On trees in wet mixed forest, or often terrestrial, sometimes on
mossy banks or pendent from rocks or cliffs, 2,600 meters or less,
most frequent in the mountains; Alta Verapaz; Izabal; Zacapa; Chi-
quimula; Huehuetenango; San Marcos. Southern Mexico; Honduras
and El Salvador to Panama; West Indies; Colombia to the Guianas.

Plants usually epiphytic and with creeping or long and slender, pendent
branches, frequently terrestrial and then usually prostrate or nearly so, the branches
very slender, often flexuous, glabrous, puberulent, or short-pilose, grayish or brown-
ish; leaves on petioles 1-2 mm. long, thin-coriaceous, lance-ovate to very broadly
ovate, mostly 1.5-4.5 cm. long, very obtuse to acuminate, rounded or subcordate
at the base, glabrous or nearly so, 3-5-nerved from the base, the nerves conspicu-
ous or obsolete beneath; pedicels 1-2 cm. long, very slender, pilose or glabrate;
calyx and hypanthium pilose with spreading hairs or glabrous, the hypanthium
globose, 1.5-2 mm. long; calyx 1 mm. long, the 4 lobes deltoid or short and apicu-
late; corolla white, 5-7 mm. long, 1.5-2 mm. broad, short-pilose or glabrous; sta-
mens 4, slightly shorter than the corolla, the filaments sparsely villous; anthers
2-3 mm. long; fruit subglobose, pilose or glabrous, 4-6 mm. in diameter, pale blue
or purple.

The plant is very common in wet forest, especially in Alta Vera-
paz. The foliage is rather handsome but the few flowers are small
and inconspicuous.

VACCINIUM L.

Reference: Hermann Sleumer, Die Arten der Gattung Vaceinium
L. in Zentral- und Siidamerika, Notizbl. Bot. Gart. Berlin 13: 111-
140. 1936.

Shrubs or small trees, glabrous or pubescent, mostly terrestrial; leaves mostly
small and persistent, rarely membranaceous, often coriaceous, entire or serrate;
flowers small, white or pink, in axillary or terminal racemes or fascicles, rarely soli-
tary, usually bracteate and minutely 2-bracteolate, the bractlets generally decid-
uous; hypanthium globose to turbinate, terete; calyx shallowly 4-5-(6-) lobate;
corolla urceolate to campanulate, terete, the 4-5 (-6) lobes short; stamens 8 or
10(-12), free or shortly adherent to the tube, the filaments free, short or elongate,

STANDLEY AND WILLIAMS: FLORA OF GUATEMALA 121

generally pubescent; anthers arista te dorsally or muticous, produced into 2 straight
or rarely curved tubules, these dehiscent by terminal pores; disk pulvinate or con-
vex, glabrous or pilose, sometimes lobate or angulate; ovary 4-5-(6-) celled or
partially 8- or 10-(12-) celled, the style straight, the stigma minute and simple or
capitellate; ovules few or numerous in the cell; fruit baccate, subglobose; seeds
small or minute, compressed, ovoid, or reniform, the testa coriaceous, the endo-
sperm carnose.

Species about 140, widely distributed in the northern hemisphere,
in temperate and arctic regions, and in the mountains of the tropics.
Three other species are known from southern Central America. In
the United States the fruits of several species of V actinium, known
as "blueberries," are of commercial importance, and vast quantities
of them are sold for food, often preserved in tins for use outside the
normal season. In recent years improved varieties have been devel-
oped in cultivation, and have been planted on a large scale in north-
ern parts of the United States. The cultivated blueberries represent
a remarkable case of the development within a few recent years of a
native plant as an important cultivated one, and is without a recent
parallel. The Central American species have more or less edible
fruits, much inferior to those of the United States, but the fruits are
seldom eaten. Also important commercially in the United States is
the cranberry, V. macrocarpon Ait., a creeping, very slender plant
grown in bogs. Its large, brilliant red fruits are extremely acid.
They are consumed in great amounts in the United States, especially
in the form of jelly, as a relish for meats, particularly with roast tur-
key at Thanksgiving and Christmas. Cranberries often are impor-
ted into Guatemala from the United States, principally for sale to
North Americans living there.

Leaf blades cordate at the base, mostly 2-5 cm. broad V. matudae.

Leaf blades acute to rounded at the base.

Leaves densely pilose V. lundellianum.

Leaves glabrous or essentially so.

Leaves entire; tubules of anther about half or more the length of the anther.
Tubules % or more of the anther length; corollas about 1 cm. long.

V. poasanum.

Tubules about half the anther length; corolla 5-6 mm. long. . .V. breedlovei.
Leaves serrate or crenate, sometimes inconspicuously so; tubules if present less

than half the length of the anther.
Flowers in elongate slender racemes, most of these much longer than the

leaves.
Racemes bearing numerous persistent leaf like bracts; leaf blades acute

at the base V. stenophyllum.

Racemes with small deciduous bracts, these not leaf like; leaf blades usu-
ally very obtuse at the base V. leucanthum.

Flowers fasciculate in the leaf axils or in very short racemes little longer
than the petioles.

122 FIELDIANA: BOTANY, VOLUME 24

Plants mostly tall shrubs 3-4.5 meters high; leaves large, mostly 2-5.5 cm.
long.

Flowers glabrous V. haematinum.

Flowers densely pubescent V. minarum.

Plants low shrubs, 25-50 cm. high; leaves mostly less than 1.5 cm. long.

Branches glabrous or nearly so; leaves rather thin, broadest above the
middle, acute at the base V. geminiflorum.

Branches pubescent, usually conspicuously and densely so; leaves coria-
ceous, broadest below the middle, rounded at the base.

V. confertum.

Vaccinium breedlovei L. Wms. Fieldiana, Bot. 31 : 173. 1965.

In pine-oak forests, at about 3,000 m., the type from near Sta.
Eulalia, Huehuetenango, Breedlove 8594..

Shrubs or possibly epiphytic shrubs of unknown size; branches terete or ob-
scurely angled, completely glabrous; leaves coriaceous ovate-lanceolate, acuminate,
glabrous, 8-10 cm. long and 4-5 cm. broad. The reticulate venation prominulous
on both surfaces, especially so below, petioles fleshy, 7-10 mm. long; inflorescences
axillary or on defoliated nodes, racemose, few-flowered, 3-4 cm. long; flowers green-
ish, borne on pedicels to about 7 mm. long, articulate with the ovary, these sub-
tended by cucullate triangular bracts about 1 mm. long and usually provided above
the middle with two bracteoles; ovary about 2 mm. in diameter, constricted near
the summit; calyx about 2 mm. long, 5-lobate, the lobes triangular, acute, about
1 mm. long; corolla narrowly campanulate, 5-6 mm. long, the lobes triangular,
acute, about 2 mm. long; stamens 10, lightly adnate to the base of the corolla, the
filaments free from one another and about 3 mm. long, glabrous, the anthers
almost 2 mm. long, obscurely papillate, the tubules about 1 mm. long and 0.2 mm.
in diameter, the pores completely terminal; style about 5 mm. long, the slender
apex stigmatic.

Related to V. poasanum and like it an inhabitant of the high
mountain forests.

Vaccinium confertum HBK. Nov. Gen. & Sp. 3: 265, t. 250.
1818. Pernettia ovata Fernald, Proc. Am. Acad. 36: 496. 1901. Coca-
late (San Marcos); Sco (Quezaltenango) .

Moist open banks or slopes or in open or dense forest, often with
pine or Cupressus, 2,600-4,000 meters; Chimaltenango; Solola; Hue-
huetenango; Totonicapan; Quezaltenango; San Marcos. Mountains
of Mexico.

A densely branched shrub, commonly 25-50 cm. high, rarely as much as a
meter tall, the branches densely puberulent or short-hirsute, rarely glabrous; leaves
often crowded, on very short, thick petioles, thick-coriaceous, oval to oblong-ovate,
6-15 mm. long, very obtuse or subacute, rounded at the base, deep green and lus-
trous above, pale beneath and often papillate (sometimes due to a fungus infection),

STANDLEY AND WILLIAMS: FLORA OF GUATEMALA 123

finely and regularly crenate-serrate, the costa and nerves usually evident and im-
pressed on the upper surface, obsolete beneath, glabrous or nearly so; racemes very
short and few-flowered, or the flowers subfasciculate, the inflorescences usually
much shorter than the leaves; hypanthium and calyx together 3 mm. long, gla-
brous or nearly so, the calyx lobes broadly deltoid, acute; corolla 4-5 mm. long,
commonly deep pink; fruit 5-6 mm. in diameter, blue with a bloom or black.

The fruit is sweet and edible, but little or no use is made of it.
The plant often forms dense low thickets of considerable extent.
Some variation is found in regard to the amount of pubescence on
branches, leaves and even on flowers.

Vaccinium geminiflorum HBK. Nov. Gen. & Sp. 3: 267, t. 252.
1818.

Moist or wet meadows or dry grassy hillsides, often among trees
of Pinus or Juniperus, or among rocks on or near mountain summits,
3,000-4,500 meters; Huehuetenango; Quezaltenango; San Marcos.
Mountains of central and southern Mexico.

A small erect shrub, usually 10-25 cm. high, densely branched, glabrous
throughout or nearly so, the slender branches erect, sometimes minutely puberu-
lent; leaves on very short petioles, oblanceolate or narrowly oblong-obovate, mostly
6-15 mm. long, obtuse or acute, narrowed to the acute base, finely crenate-serrate,
thin, glabrous, smooth and usually lustrous above, slightly paler beneath and
coarsely and prominently reticulate- veined; flowers axillary, solitary or geminate,
on recurved pedicels; hypanthium broadly campanulate, the calyx limb very short,
truncate; corolla broadly urceolate, 4-5 mm. long, deep pink or reddish.

This is one of the typically alpine plants of Guatemala, ascending
to the summits of the highest volcanoes. The leaves are found upon
the plants throughout the year, but the old ones are inclined to fall
at the end of the dry season, when new leaves are unfolding.

Vaccinium haematinum Standl. & Steyerm. Field Mus. Bot.
23: 139. 1944.

Type from Huehuetenango, Cerro Huitz, between Mimanhuitz
and Yulhuitz, Sierra de los Cuchumatanes, 1,500-2,600 meters,
Steyermark 48572. Honduras.

A shrub of 4.5 meters, or sometimes procumbent, the branches subterete, black-
ish brown, puberulent when young or glabrous, with short internodes; leaves rigid-
coriaceous, short-petiolate, lustrous, brownish when dried, on stout petioles 4-5
mm. long, elliptic or oblong-elliptic to rounded-ovate, 2-6.5 cm. long, 1.5-4 cm.
broad, obtuse, rounded to broadly cuneate at the base, obsoletely and remotely
appressed-crenate or subentire, glabrous, the costa plane above or shallowly and
broadly impressed, the nerves and veins obsolete, slightly paler beneath, epunctate,
the costa rather stout and prominent, the lateral nerves about 7 pairs, prominu-
lous, ascending at an acute angle, almost straight or irregular, the veins prominulous

124

FIELDIANA: BOTANY, VOLUME 24

and laxly reticulate; inflorescences axillary, short-racemose, few-flowered, scarcely
more than 1 cm. long, sparsely puberulent, the rachis very short, the pedicels stout,
4 mm. long, the 2 bractlets inserted slightly below the middle of the pedicel, lanceo-
late, appressed; hypanthium sparsely and minutely puberulent or glabrate; corolla
white, 6 mm. long, almost 3 mm. broad; calyx lobes deltoid, acute, 1 mm. long;
fruit cranberry-red, 5 mm. long, glabrous.

FIG. 35. Vaccinum lundellianum. A, Flowering branch, X 1. B, Flower,
X 2 1/2- C, Corolla opened to show stamens and relative length of style, X 2^.
D, Stamen from front, X 7^. E, Stamen from side, X 1%. F, Mature fruit,

STANDLEY AND WILLIAMS: FLORA OF GUATEMALA 125

Vaccinium leucanthum Cham. & Schlecht. Linnaea 8: 524.
1833. Cacj (Huehuetenango) .

Moist or wet forest or thickets, or open wet banks, sometimes in
pine-oak forest, 1,700-3,300 meters; El Progreso; Zacapa; Chiqui-
mula; Huehuetenango. Mexico.

A shrub 1-4.5 meters high, the branches slender or rather stout, brown, terete
or when young somewhat angulate, puberulent or glabrate; leaves lance-oblong to
broadly ovate, very variable in size and shape, mostly 2-4.5 cm. long, acute to
obtuse or rounded and apiculate, rounded at the base, short-petiolate, coriaceous,
crenate-serrate, glabrous or nearly so, green above, often lustrous, pale beneath,
sometimes black-punctate; racemes many-flowered, 3-6 cm. long, dense or usually
lax, puberulent, the bracts small, usually early deciduous, the flowers on long slen-
der pedicels; calyx and hypanthium usually pubescent, the calyx lobes broadly
deltoid, acute; corolla urceolate or subglobose, 3 mm. long, white; fruit black,
5 mm. in diameter.

Guatemalan material referred here is variable and may represent
more than one species. This group of the genus is highly variable
and critical, and the nomenclature and specific limits cannot be de-
cided until all the various entities have been considered together.

Vaccinium lundellianum L. Wms. Fieldiana, Bot. 31: 173.
1965. Malea pilosa Lundell, Am. Midi. Nat. 29: 484. 1943, non
Vaccinium pilosum A. Chev. 1930.

Epiphyte in moist or wet forest, at about 3,000 meters; Huehue-
tenango. Mexico (the type from Mt. Male", near Porvenir, Chiapas).

An epiphytic shrub, pilose throughout with short spreading whitish hairs,
sometimes subscandent, the branches terete, often blackish; leaves on petioles
3-6 mm. long, lanceolate or lance-oblong, 3-5.5 cm. long, 1-2 cm. broad, some-
times slightly larger, narrowed to an obtuse apex, rounded or obtuse at the base,
densely short-pilose on both surfaces; inflorescences numerous, the rachis usually
less than 5 mm. long, the pedicels 8 mm. long or less, bearing 2 narrow bractlets
near the middle; calyx 5-lobate, the lobes narrowly triangular, 1.3 mm. long;
corolla white, 8-10 mm. long, densely short-pilose, the 5 lobes erect or suberect,
obtuse, as much as 3 mm. long; filaments pilose, the anthers attached dorsally at
the middle, the tubules opening by elongate introrse clefts; cells of the ovary usu-
ally 2-ovulate; disk annular; style slender, equaling the corolla; immature fruit
globose, 4 mm. in diameter, densely short-pilose.

This species is the basis of the monotypic genus Malea of Lundell,
but would seem to have no satisfactory characters by which it may
be distinguished from Vaccinium.

Vaccinium matudae Lundell, Phytologia 2: 4. 1941, as V. ma-
tudai (type from Chiapas, Mexico).

126 FIELDIANA: BOTANY, VOLUME 24

Epiphytic shrubs in cloud forest at about 3,000 meters; Quiche";
Huehuetenango. Mexico (Chiapas).

An epiphytic or terrestrial shrub with stout, somewhat angulate branches,
these sparsely hirtellous when young, soon glabrate; leaves practically sessile, rigid-
coriaceous, lustrous, glabrous,, broadly ovate or rounded-ovate, 3-8 cm. long,
mostly 2.5-5 cm. broad, almost rounded to subacute at the apex, shallowly and
narrowly cordate at the base, obscurely and remotely crenate-serrate or subentire,
the venation scarcely prominulous above, prominent beneath and closely reticu-
late; inflorescences racemose, many-flowered, as much as 10 cm. long but mostly
shorter, glabrous or nearly so, the pedicels and rachis deep rose; calyx lobes 5, deep
rose, deltoid, acute or subacuminate; corolla whitish suffused with deep pink, 7-
8 mm. long, glabrous outside, the 5 lobes short, obtuse; filaments pilose, the anthers
3 mm. long, muticous; fruit 5 mm. in diameter.

Because of its large, broad, cordate, very thick and lustrous leaves,
this is very unlike other North American species of the genus. Lun-
dell compared V. matudae with V. selerianum (Loes.) Sleumer, a plant
to which it is not closely related.

Vaccinium minarum Standl. & Steyerm. Field Mus. Bot. 23:
219. 1947.

At 2,000-3,200 meters; endemic; Zacapa (type from middle and
upper slopes of Volcan Gemelos, Sierra de las Minas, Steyermark
43295; also collected between Loma Pichaco and Cerro de los Monos) .

A shrub or small tree 3-4.5 meters high, the branches slender, subterete, fus-
cous or brownish, the young ones often reddish, puberulent, soon glabrate, densely
leafy; leaves rather small, coriaceous, on petioles 3-5 mm. long, somewhat lustrous,
elliptic-oblong, obovate-oblong, or lanceolate, 2.5-3.5 cm. long, 8-15 mm. broad,
obtuse or narrowed to an obtuse apex, acute or cuneate at the base, remotely and
rather inconspicuously appressed-crenate, glabrous, the costa sometimes impressed
above, prominulous beneath, epunctate beneath, the lateral nerves obsolete or
prominulous, the veins obsolete; racemes axillary, few-flowered, less than half as
long as the leaves, the pedicels red, 3-7 mm. long, puberulent, bracteate below the
middle; hypanthium 1.5 mm. long, puberulent, the 5 sepals triangular-acuminate,
1 mm. long, puberulent outside; corolla deep rose, 6 mm. long, 2 mm. broad or
slightly broader, whitish-hispidulous, the 5 lobes ovate, subacute, 1.3 mm. long,
erect.

Vaccinium poasanum Donn.-Sm. Bot. Gaz. 24: 395. 1897.
Wet mixed mountains forest, 1,500-2,400 meters; Zacapa (Sierra
de las Minas). El Salvador; Honduras; Costa Rica; Panama.

A shrub or tree 4.5-10 meters high, glabrous throughout, the bark thin, deep
cinnamon-brown; leaves short-petiolate, coriaceous, mostly oblong-elliptic and
5-9 cm. long, acuminate or long-acuminate, acute at the base, entire, the veins
and nerves conspicuous beneath, the leaves somewhat paler on the lower surface;

STANDLEY AND WILLIAMS: FLORA OF GUATEMALA 127

racemes axillary, umbelliform, several-many-flowered, the pedicels about 1.5 cm.
long, often pendent; calyx very short, repand-dentate; corolla narrowly campanu-
late, pale greenish yellow or sometimes tinged with pink, about 1 cm. long; fruit
subglobose, 6-7 mm. in diameter or larger, pale yellow.

Vaccinium stenophyllum Steud. Nom. Bot. ed. 2. 2: 740. 1841.
V. angustifolium Benth. PI. Hartw. 45. 1840, not V. angustifolium
Ait. 1789.

Oak forest, 1,900 meters; Quiche" (Nebaj, A. F. Skutch 1733}.
Western and southern Mexico.

A shrub or small tree as much as 6 meters high, the branches slender, puberu-
lent or glabrate; leaves short-petiolate, elliptic-lanceolate to linear-oblong, coria-
ceous, lustrous, 2-4.5 cm. long, acute or acuminate, acute at the base, serrulate
or entire, paler beneath; racemes terminal or lateral, usually much elongate, bear-
ing numerous small leaves or more reduced, leaf-like bracts, the rachis and slender
pedicels puberulent or glabrate; calyx puberulent, the lobes deltoid, puberulent,
acuminate; corolla white or pinkish, 5 mm. long, lobate to about the middle, the
lobes triangular, acute, glabrous outside; fruit black, globose, 6 mm. in diameter.

PRIMULALES

The Primulales consist of three families: Theophrastaceae, Myrsi-
naceae and Primulaceae, all of which are represented in Guatemala.
Dr. C. L. Lundell has prepared the treatment of the Myrsinaceae for
this flora. It is the largest and most important family of the order
in Guatemala.

Plants of this order vary from small herbaceous annuals to, more
commonly, trees and shrubs ranging from dry coastal plains to high
mountain forests.

The Primulales have sympetalous corollas, the flowers are normally pentam-
erous; the ovary is superior, unilocular and has free central placentation: the leaves
are usually alternate or sometimes subverticellate or whorled, simple.

THEOPHRASTACEAE

Reference: Carl Mez, Theophrastaceae, Pflanzenreich IV. 236a:
1-48. 1903.

Trees or shrubs; leaves alternate, often crowded and subverticillate at the ends
of the branches, simple, entire or spinose-serrate, often tipped with a rigid spine;
stipules none; flowers mostly perfect and 5-parted, regular, large or small, mostly
white, yellow, or red, the inflorescences generally terminal, racemose or abbrevi-
ated or even reduced to a single flower; calyx inferior, the segments free or very
shortly connate, crenate or ciliate, imbricate, generally rounded at the apex, per-

128 FIELDIANA: BOTANY, VOLUME 24

sistent; corolla gamopetalous, rotate, urceolate, or funnelform, carnose or coria-
ceous, the lobes imbricate; staminodia usually 5, inserted on the corolla tube, some-
times petaloid; stamens 5, opposite the petals, free or connate, the filaments united
wholly or partly with the corolla, the anthers dorsifixed, dehiscent extrorsely by 2
longitudinal slits; ovary ovoid or clavate, 1-celled, the style stout or almost obso-
lete; stigma discoid or rarely conic, entire or irregularly lobate; placenta central,
short-stipitate, the ovules numerous, multiseriate, anatropous; fruit baccate or
drupaceous, sometimes almost dry; seeds numerous or few, surrounded by muci-
laginous pulp, the testa thin; endosperm abundant, smooth, corneous; embryo
straight.

Four genera, confined to tropical America. One other, Clavija,
is represented in southern Central America.

Staminodia minute, gland-like; leaves hirsute beneath; corolla green; flowers soli-
tary or fasciculate Deherainia.

Staminodia petal-like, conspicuous; leaves glabrous; corolla yellow or red, rarely
white; flowers usually racemose Jacquinia.

DEHERAINIA Decaisne

Shrubs or small trees, usually hirsute; leaves short-petiolate, often lepidote,
entire (in Central American species) or spinose-dentate; flowers rather large, per-
fect, usually green, inserted at the ends of the branches; sepals short-connate at
the base, rounded or ovate, ciliolate; corolla short-campanulate, lobate to the
middle, the lobes broad, rounded; staminodia minute, liguliform or triangular, in-
serted at the apex of the corolla tube; stamens 5, inserted on the base of the corolla,
the filaments connate with the corolla; anthers free, dorsifixed slightly above the
base, truncate or caudate at the apex; ovary ovoid, glabrous, contracted into a
style of equal or greater length; stigma broad, disk-like; fruit ellipsoid or ovate-
oblong, large, many-seeded.

Three species are known: the following, one in Cuba, and one
from Chiapas, D. matudae Lundell, which is to be expected in west-
ern Guatemala. It differs from D. smaragdina in having glabrous
leaves in addition to flower characters.

Deherainia smaragdina (Planch.) Dene. Ann. Sci. Nat. VI. 3:
139, 1. 12. 1876. Jacquinia smaragdina Planch, ex Linden, PI. Nouv.
8. 1859.

Mostly in wet limestone forest, 1,100 meters or less; Pete"n; Alta
Verapaz; Huehuetenango. Mexico; British Honduras; Honduras.

A branched shrub, mostly 2-4.5 meters high, sometimes a small tree, the
branches stiff, rufous-hirsute; leaves coriaceous, short-petiolate, elliptic to oblong-
oblanceolate, 8-16 cm. long, 2-5 cm. broad, acute or acuminate, not spine-tipped,
attenuate to the base, glabrous above, or nearly so, rufous-hirsute beneath; flowers
solitary or binate at the ends of the branches, the pedicels rufous-hirsute; flowers 16-
20 mm. long; sepals suborbicular, broadly rounded at the apex, ciliate; corolla lobes

STANDLEY AND WILLIAMS: FLORA OF GUATEMALA 129

FIG. 36. Deherainia smaragdina. A, Flowering branch, X Y^ B, Flower, X 2.
C, Corolla opened out, X 2. D, Fruit, X J^. E, Stamens, front and side, X 4.
F, G, Sepal margin, X 8.

suborbicular, minutely ciliolate; anthers emarginate at each end; ovary glabrous,
ovoid, the style longer than the ovary, slender, the stigma large, capitulate-pulvi-
nate; fruit green, lustrous, ovoid-oblong, about 7.5 cm. long and 3 cm. broad, atte-
nuate-acute, narrowly rounded at the base, glabrous.

The shrub is common in the characteristic thickets on almost bare
limestone in the lower portions of Alta Verapaz. It is unusual in its
large, deep green corollas, which are attractive because of their strange
coloring.

JACQUINIA L.

Shrubs or trees, usually glabrous; leaves alternate or pseudo-verticillate, cori-
aceous, often rigid-coriaceous and armed at the apex with a spinelike cusp or
mucro, sometimes muticous; flowers rather small, perfect, 5-parted, in terminal

130 FIELDIANA: BOTANY, VOLUME 24

or pseudo-axillary racemes, rarely solitary, mostly yellow or orange-red, sometimes
white; sepals free or very shortly connate at the base, imbricate, ovate or orbicu-
lar, ciliate or crenate; petals connate to form a short or elongate tube, the lobes
imbricate, spreading in anthesis, rounded, somewhat coriaceous; staminodia 5, in-
serted at the apex of the tube and resembling corolla lobes, but smaller, generally
crenate and emarginate at the apex; stamens 5, inserted near the base of the corolla,
more or less exserted, the filaments dilated below; anthers dehiscent by extrorse
longitudinal slits, elongate-ovate or subsagittate, dorsifixed slightly above the base,
emarginate, rounded, or rarely acuminate at the apex; ovary glabrous, attenuate
to the usually short style, the stigma obtuse or discoid-capitulate; fruit ovoid or
globose, generally cuspidate, coriaceous or crustaceous; seeds few, ovoid immersed,
in pulp; endosperm cartilaginous; cotyledons ovate, the radicle slender, elongate,
inferior.

About 40 species, in tropical America. A few additional species
are known from Central America and Mexico.

The taxonomy of the genus is not well understood. Mez saw only
a few more than 125 specimens when he revised the genus in 1903
and credited to it 33 species — about two-thirds of these North Amer-
ican. Eleven additional species have been described from North
America since Mez' revision. Mez maintained numerous species on
the basis of anther characters which, at best, seem to be vague and
inconstant. A revision based on the more ample material now avail-
able would seem to be in order.

Leaves not spine-tipped «7. schippii.

Leaves spine-tipped, the spine small or large.

Flowers yellow or white, about 7 mm. long; leaves thin, the apical spine short
and weak J. paludicola.

Flowers orange-red, 8-10 mm. long or larger; leaves usually thick and rigid, the
apical spine generally long and stiff.

Leaves rather conspicuously triplinerved; connective of the anther acumi-
nately produced beyond the cells J. donnell-smithii.

Leaves not triplinerved, 1-nerved or penninerved, often obscurely so.

Leaves very narrowly lanceolate, 1-nerved; connective of the anther acu-
minately produced beyond the cells J. pungens.

Leaves oblong-elliptic to elliptic or obovate, penninerved, the nerves often
obscure; connective of the anther not acuminately produced.

J. aurantiaca.

Jacquinia aurantiaca Ait. Hort. Kew. ed. 2. 2: 6. 1811. J. au-
rantiaca var. latifolia Mez in Urban, Symb. Antill. 2: 451. 1901.
J. albiflora Lundell, Wrightia 2: 60. 1960 (type from Pete"n, Contreras
.45); J. lepidota Lundell, I.e. 61 (type from Pete"n, Lundell 15997).
Chicajam (reported from the Atlantic coast); tzic or chacsic (Pete"n,
Maya) ; naranjillo; zincin (Pete"n, Maya) ; mata-pescado (Zacapa) ; the
name ducuche also reported.

STANDLEY AND WILLIAMS: FLORA OF GUATEMALA

131

FlG. 37. Jaquinia aurantiaca. A, Flowering branch, X 1. B, Flower, X
C, Corolla opened out, X 2}^. D, Stamens, front and side, X 5.

Dry and moist, brushy plains and hillsides, sometimes in moist
lowland forest or on saline flats, 1,100 meters or lower; Pete"n; Baja
Verapaz; El Progreso; Zacapa; Chiquimula; Jutiapa; Escuintla; Re-
talhuleu; Huehuetenango. Mexico; British Honduras to Panama.

A shrub or tree, sometimes 9 meters high with a trunk 15 cm. in diameter, often
very densely branched, the young branchlets sometimes puberulent or tomentu-
lose; leaves subsessile, usually rigid-coriaceous, oblanceolate to obovate-oblong or
obovate, mostly 3.5-8 cm. long, acute or obtuse at the apex and tipped with a long
rigid spine, lineolate beneath when dried, evidently penninerved, the lateral nerves

132 FIELDIANA: BOTANY, VOLUME 24

very slender, often prominulous beneath, the upper surface usually lustrous; ra-
cemes terminal, few-several-flowered, racemose, the pedicels about 6 mm. long,
often thickened; bracts ovate, acute; flowers bright orange-red, 8-10 mm. long; se-
pals serrulate; fruit globose, yellow or orange, 1.5-2 cm. in diameter or even larger,
with a very hard but somewhat fragile shell.

Called "knock-me-back" in British Honduras; "cansic" (British
Honduras, Maya); "muyche" (Yucatan, Maya); "siche," "flor de
San Antonio" (Tabasco); "naranjillo" (Chiapas); "barbasco," "li-
moncillo," "escorpioncillo," "mirra," "espino ruco," "carambolillo,"
"crucillo" (El Salvador). This has been reported from Guatemala
as J. axillaris Oerst. This shrub or tree is particularly plentiful in
the dry lower Motagua Valley, where it occurs in great abundance.
The plants are well protected against large animals, since the spiny
tips of the leaves penetrate the fleshlike needles. The flowers, when
orange-red, are showy and pretty. They often are strung on cords
to make garlands, and preserve their shape and color for a long time.
They are said to have been used by the Mayas for decorating their
temples. The pulverized seeds are said to be administered in Pete"n
to expel or destroy intestinal parasites. Almost throughout the range
of the genus the species of Jacquinia are employed as a barbasco or
fish poison, the crushed roots or fruits being used. In Guatemala the
roots are macerated in water with fresh lime. The latter produces a
foam which aids in spreading the barbasco more thoroughly. J. albi-
flora Lundell seems, at most, to be a variety with light colored flowers
and puberulent branches.

Jacquinia donnell-smithii Mez, Pflanzenreich IV. 236a: 39.
1903.

Moist or dry thickets of the Pacific plains, 300 meters or less; en-
demic; Pete"n; Zacapa; Santa Rosa; Escuintla (type from Santa Lucia,
Heyde & Lux 63^3) ; Retalhuleu.

A glabrous shrub 1-3 meters high; leaves subsessile, rigid-coriaceous or sub-
coriaceous, narrowly elliptic to oblong-lanceolate, 4-10.5 cm. long, 1.5-2.5 cm.
broad, acute or obtuse and tipped with a long slender stiff spine, attenuate to the
base, more or less obviously triplinerved beneath and also penninerved above the
basal nerves; inflorescences terminal, mostly 5-9-flowered, racemose or subcorym-
bose, the stout pedicels about 6 mm. long; bracts minute, ovate, acute; flowers
orange-red, 8-9 mm. long, the sepals minutely crenulate.

This has been reported from Guatemala as J. pungens Gray. Its
validity as a species is questionable. It is probably only a minor
form of J. aurantiaca.

STANDLEY AND WILLIAMS: FLORA OF GUATEMALA 133

Jacquinia paludicola Standl. Field Mus. Bot. 11: 138. 1932.
Wet mixed forest, at or little above sea level; Izabal; Pete"n. Brit-
ish Honduras (type from Forest Home, W. A. Schipp 1028).

A shrub or tree 3-7 meters tall, glabrous, the trunk as much as 8 cm. in diam-
eter; leaves thin, on petioles 3-5 mm. long, pseudo-verticillate at the ends of the
branches, oblong-oblanceolate to ovate-elliptic, mostly 8-12 cm. long and 2.5-
4.5 cm. broad, acute or acuminate, tipped with a short weak spine, attenuate to
the base, penninerved; racemes few-several-flowered, often umbelliform, the slen-
der pedicels 5-14 mm. long; sepals orbicular, 2 mm. long, broadly rounded at the
apex; corolla white or pale yellow, 6 mm. long, the rounded lobes shorter than
the tube; fruit globose, about 2 cm. in diameter.

Jacquinia pungens Gray, Mem. Am. Acad. Sci. n. ser. 5: 325.
1855. Matapeje; luruche.

Dry rocky plains and hillsides, 200-700 meters; Zacapa; Chiqui-
mula; El Progreso. Western and southern Mexico.

A shrub or small tree, glabrous, the trunk sometimes 12 cm. in diameter; leaves
subsessile, rigid, linear-lanceolate, 3-6 cm. long, 1 cm. broad or narrower, acute
and tipped with a long stiff yellowish spine, attenuate to the base, 1-nerved; in-
florescences racemiform, terminal, few-flowered, the pedicels about 7 mm. long,
the bracts small, ovate-acuminate; flowers 7-8 mm. long, orange-red, the sepals
crenulate only at the apex if at all; fruit subglobose, 2 cm. in diameter, several-
seeded, the endocarp woody and thick.

It is doubtful whether the Guatemalan material represents more
than a narrow-leaved variety of J. aurantiaca. The bark is ashy gray,
rough and thin, the inner bark greenish; wood pale brown.

Jacquinia schippii Standl. Carnegie Inst. Wash. Publ. 461 : 78.
1935.

Known only from the type, Jacinto Hills, British Honduras, in
forest, 270 meters, W. A. Schipp 1233.

A tree of 10 meters, the trunk 20 cm. in diameter, glabrous; leaves on petioles
2-4 mm. long, subcoriaceous, oblong or narrowly oblanceolate-oblong, 3-5 cm.
long, 1-2 cm. broad, narrowly rounded or very obtuse and muticous at the apex,
attenuate to the base, somewhat lustrous, the costa obsolete above, the whole
lower surfaces closely and conspicuously longitudinally striate, the margins corne-
ous; flowers short-racemose, the slender pedicels 1 cm. long; calyx in fruit 5 mm.
broad, appressed, the sepals broadly rounded at the apex, broader than long,
minutely erose; fruit subglobose, 1 cm. long, smooth, apiculate.

MYRSINACEAE

by CYRUS LONGWORTH LUNDELL'

Director and Chief Research Scientist
Texas Research Foundation

Reference: Carl Mez, Myrsinaceae, Pflanzenreich IV. 236. 1902.

Trees or shrubs, glabrous or pubescent; leaves alternate, often clustered at the
ends of the branches, entire, crenulate or serrate, glandular-punctate; stipules none;
flowers perfect, or dioecious, usually 5-parted, regular, in terminal or axillary in-
florescences, small, white, pink or green; calyx inferior, the segments free or more
or less connate, mostly ciliate and glandular-punctate, valvate, imbricate, or sinis-
trorsely contorted, persistent; corolla regular, usually gamopetalous, rotate to tub-
ular, petals valvate or dextrorsely, or rarely sinistrorsely, imbricate or contorted,
often quincuncial; stamens as many as the petals and opposite them, the filaments
usually short, sometimes equaling or longer than petals, connate with the corolla
tube or almost wholly free; anthers mostly dorsifixed, sagittate, ovate, elliptic or
linear, dehiscent by introrse slits or by apical pores; ovary globose, ovoid, or cla-
vate, free, sessile, 1-celled; style long or short, the stigma punctiform, capitate,
discoid, or conic; placenta central, usually globose, the ovules numerous or few,
uniseriate or pluriseriate; fruit drupaceous, 1-seeded; seeds with a thin testa, the
endosperm copious, smooth or rarely ruminate; embryo cylindric or curved, the
cotyledons small, the radicle elongate.

Genera about 40, widely dispersed in tropical regions.

The classification of the tribes and genera of the Myrsinaceae is
based primarily on the number of ovules and their arrangement in
the placenta. Most generic characters are weak and those separating
species are often weaker. Fine distinctions are necessary for an un-
derstanding of natural groups.

Flowering material is essential for the identification of genera and
species. In Guatemala, Stylogyne A. DC. and Rapanea Aubl. are dio-
ecious. In Stylogyne, which has been poorly understood, the pistillate
flowers with long slender style and well developed stamens appear
perfect, and superficially resemble those of Ardisia Swartz, but the

1 Assisted by a grant from the Dodge Jones Foundation.

135

136 FIELDIANA: BOTANY, VOLUME 24

staminate flowers have a short style subequaling the abortive ovary.
The ovules are few and uniseriate, which places Stylogyne in the Tribe
Myrsineae. Rapanea with its fasciculate axillary flowers and large
sessile stigma is easy of recognition.

Parathesis Hook, f., which has valvate sepals and petals, is a large
natural genus with Guatemala the center of its highest development.
Yunckeria Lundell cannot be placed without dissection of the pla-
centa, yet the few erect uniseriate ovules mark it as one of the most
distinctive in the family.

Synardisia (Mez) Lundell, a monotypic genus, is notable for its
campanulate or suburceolate corolla. It has glabrous branchlets and
leaves, but the large inflorescences are pubescent with conspicuous
gland- tipped hairs, which aid in identification.

Gentlea Lundell is notable for the slender exserted stamens, and
greenish or white flowers. The greenish hue is unusual in the Myr-
sinaceae.

Ardisia contains a series of natural groups which probably deserve
generic recognition. Most of its species are well-marked in our area.

Inflorescences small, axillary, shorter than the petioles; flowers dioecious, fascicu-
late; style none, the stigma of pistillate flowers large and morchelliform.

Rapanea.

Inflorescences usually large, axillary or terminal, much longer than the petioles,
pedunculate; style evident, short or much elongate, the stigma small, usually
punctiform.

Flowers dioecious; the style of staminate flowers short, subequaling abortive
ovary; the style of pistillate flowers elongate, much exceeding ovary; ovules

few, uniseriate; sepals and petals convolute in bud Stylogyne.

Flowers perfect, the style long and slender.

Sepals and petals valvate; petals pubescent Parathesis.

Sepals and petals imbricate or convolute in bud; petals usually glabrous.
Ovules uniseriate, erect from base of placenta; petals imbricate, about 1 cm.

long Yunckeria.

Ovules pluriseriate, immersed in placenta.

Stamens exserted; filaments long and slender; anthers small, cordate;

corolla greenish or white; flowers corymbose Gentlea.

Stamens included; filaments usually short, if elongate, the flowers either

racemose or spicate; corolla pink or white.

Corolla campanulate or suburceolate, the petals connate up to three-
fourths their length; inflorescences large, densely pubescent with

gland-tipped hairs Synardisia.

Corolla with petals connate at base, nearly free Ardisia.

ARDISIA Swartz

Shrubs or trees; leaves alternate, petiolate or rarely sessile, entire, dentate,
crenate or serrate; inflorescences varied in form, the racemes, umbels and corymbs
paniculate; flowers pedicellate or subsessile, perfect, usually 5-parted, often 4- or

LUNDELL: FLORA OF GUATEMALA 137

6-parted, small, white, pink or lilac, dextrorsely, or rarely sinistrorsely, imbricate
or contorted, often quincuncial; sepals short-connate or nearly free; petals united
at base, sometimes connate almost to middle, recurved or spreading, rarely erect;
stamens free, inserted at or slightly above base of corolla, the filaments usually
short; anthers dorsifixed, mostly elongate and subsagittate, concolor or glandular-
punctate, dehiscent by longitudinal slits or sometimes by apical or subapical pores;
ovary ovoid or subglobose, the style long and slender, often exserted, the stigma
minute, punctiform; ovules few to numerous, pluriseriate; fruit globose or sub-
globose, bearing at the apex the persistent style base, 1-seeded, the endocarp crus-
taceous or osseous; seed globose.

Species numerous, mostly in tropical America and Asia. In North
America, the genus is most abundantly represented in Costa Rica.
One exotic species, perhaps the Asiatic A. crenulata Lodd., has been
observed in cultivation in Guatemala as an ornamental plant.

Flowers in paniculate racemes or spikes; leaves entire.

Sepals and petals sinistrorsely imbricate or quincuncial A. escallonioides.

Sepals and petals dextrorsely imbricate or quincuncial.

Sepals and petals epunctate A. escuintlensis.

Sepals and petals conspicuously punctate.

Corolla tube glandular-lepidote or papillose within, the petals otherwise
glabrous; flowers subsessile or with pedicels usually less than 6 mm.
long.

Flowers spicate, usually subsessile, rarely with pedicels up to 6 mm. long
at base of spike; sepals coriaceous and thickened medially, rugose.

A. densi flora.

Flowers racemose, pedicels usually 4-6 mm. long, sometimes up to 1 cm.
long; sepals thin A. revoluta.

Corolla with conspicuous glandular, yellow or orange, star-shaped eye with-
in; pedicels elongated toward base of raceme, 1-2.5 cm. long, rarely
reduced A. paschalis.

Flowers in paniculate umbels, corymbs, or corymbiform racemes.

Flowers in corymbiform racemes (see also A. paschalis); pedicels elongated,
rigid, usually 1-3 cm. long; inflorescences glabrous.

Sepals entire, with conspicuous round plate-like glands on dorsal surface.

A. verapazensis.
Sepals ciliate, without glands on dorsal surface.

Corolla 9-12 mm. long; corolla tube 3.5-5 mm. high, glabrous on outer sur-
face; leaves subentire, coriaceous or subcoriaceous A. sexpartita.

Corolla 7-9 mm. long; corolla tube about 2.5 mm. high, pubescent at
base on outer surface with gland-tipped hairs; leaves entire, thin,
chartaceous A. cucullata.

Flowers in umbels or corymbs, the corymbs sometimes elongated after anthesis.
Inflorescences furfuraceous, at least at base.

Leaves sessile or nearly so, the petioles broadly marginate to the attenuate
narrowly rounded base; inflorescences narrowly paniculate, with pri-
mary branches less than 2 cm. long A. apoda.

Leaves petiolate; inflorescences broadly paniculate, usually pyramidal.

Leaf blades thin, large, with abundant black glands beneath; anthers
2.3-2.6 mm. long, black-lineate dorsally, longitudinally dehiscent.

A. nigropunctata.

138 FIELDIANA: BOTANY, VOLUME 24

Leaf blades usually chartaceous, sometimes coriaceous, inconspicuously
punctate; anthers 2.5-4 mm. long, concolor, dehiscent by apical pores.

A. compressa.

Inflorescences not furfuraceous, either glabrous or puberulent, often glandular-
puberulent.

Leaves pectinate-dentate with very close, spreading, subulate teeth.

A. pellucida.

Leaves entire, crenulate, dentate, or appressed-serrulate, the teeth not sub-
ulate.

Pedicels of flowers rigid, 3-10 mm. long; leaves acute or subacuminate;
flowers usually corymbose.

Leaf blades thin, broad, mostly elliptic, 5.5-9.5 cm. wide, the margin
usually dentate or appressed-serrulate A. Schippii.

Leaf blades subcoriaceous, narrow, mostly oblanceolate-oblong, 2.5-5 cm.
wide, the margin entire or subentire A. rarescens.

Pedicels of flowers filiform, weak, usually 1-2.5 cm. long, rarely shorter;
leaves acuminate, mostly caudate-acuminate; flowers often umbellate.

Filaments glabrous; branchlets rufous-hirtellous (see also A. nigrescens).

A. hirtella.
Filaments glandular-puberulent.

Branchlets hirsute-tomentose with short red hairs A. nigrescens.

Branchlets glabrous or minutely puberulent.

Anthers cuspidate, up to 9 mm. long A. Mitchellae.

Anthers merely apiculate, 3-5.5 mm. long.

Leaves glabrous; branchlets usually glabrous. . . . A. Tuerckheimii.

Leaves with petioles and costa puberulent beneath; branchlets
ferruginous-puberulent.

Petals ovate, 5.5-6 mm. wide A. Carlsonae.

Petals lanceolate-oblong, 3-4 mm. wide A. erythrocarpa.

Ardisia apoda Standl. & Steyerm., Field Mus. Bot. 23: 219. 1947.
Wet forest, 900 meters or less; Izabal (endemic, type from Cerro
San Gil, J. A. Steyermark 41923).

A tree 6 meters tall, with thick branches, densely ferruginous-furfuraceous,
glabrescent, densely leafy at the ends; leaves very large, sessile or nearly so, sub-
chartaceous, entire, oblanceolate, up to 40 cm. long, 9.5 cm. wide, acuminate,
narrowly attenuate to the base, the base narrowly rounded, densely ferruginous-
furfuraceous beneath at first and at base above, rather sparsely so at maturity,
densely punctate, paler on lower surface, the costa elevated below, the primary
lateral veins very slender, inconspicuous, obscurely reticulate; inflorescences
terminal, very slender, 1- or 2-branched, narrowly paniculate, up to 12.5 cm.
long, the primary branches short, up to 2 cm. long, ferruginous-furfuraceous;
flowers corymbose, the fruiting pedicels up to 1 cm. long; sepals free, ovate-orbicu-
lar, 1.5 mm. long, ciliolate, black-punctate; fruit subglobose, glabrous, black and
shiny at maturity, 8 mm. in diameter.

A. apoda closely resembles A. Wedelii Lundell, a species described
from Panama and found also in Costa Rica and Nicaragua. The
narrowly paniculate slender inflorescences, longer pedicels and the in-
conspicuous venation of the leaves are minor features which appear

LUNDELL: FLORA OF GUATEMALA 139

to distinguish the Guatemalan tree. Flowers of A. apoda are un-
known. Entirely unlike any other of the genus in Guatemala, the
species is easy of recognition because of its large sessile leaves.

Ardisia Carlsonae Steyermark, Ceiba 4: 301. 1955.

Known only from the type; Mexico, around Los Arcos, Los Lagos,
3 miles northwest of Rancho San Jose" and 34 miles southeast of
Comitan, Chiapas, 1,600 meters, Margery C. Carlson 1744.

A shrub up to 2 meters high, the branchlets slender, at first ferruginous-puber-
ulent; leaves with petioles 3-5 mm. long, nearly glabrous; leaf blades narrowly
elliptic, 4-6.5 cm. long, 1.5-2.3 cm. wide, apex abruptly acuminate, base acute, the
margin crenulate above, conspicuously black-punctate, the costa prominent and
minutely puberulent beneath, otherwise glabrous; inflorescences terminal, laxly
subcorymbose, about 12-flowered, much shorter than the leaves, up to 5 cm. in
diameter; pedicels 1.2-2.3 cm. long, sparsely puberulent, almost glabrous; flowers
5-parted; sepals short-connate at base, ovate, 2.8-3 mm. long, 1.5 mm. wide, sub-
obtuse, ciliate, conspicuously punctate; petals almost free, connate only at base,
ovate, 10 mm. long, 5.5-6 mm. wide, subobtuse, densely punctate; filaments 2 mm.
long, glandular-puberulent; anthers narrowly linear, 5.5 mm. long, with apex ob-
tusely apiculate; ovary ovoid, glabrous; style slender, 6.5 mm. long; dry fruit 7 mm.
in diameter.

The type at Chicago has been misplaced, or lost, and I know the
species only from description. Steyermark (I.e.) notes its affinity:
"From A. Mitchellae Johnston, to which it is probably most closely
related, it differs in the obtusely short-apiculate, not cuspidate, an-
thers, white petals, with finer and relatively fewer dark punctations
of the leaves, and sparsely puberulous to nearly glabrous pedicels."
Possibly its wider petals, described as 5.5-6 mm. in width, are its
chief distinctive feature, for the petals of both A. Mitchellae and
A. erythrocarpa Lundell are oblong and usually about 3 mm. wide,
rarely up to 4 mm. wide. Both A. Carlsonae and A. Mitchellae have
larger flowers than A. erythrocarpa.

Ardisia compressa H.B.K. Nov. Gen. & Sp. 3: 245: 1818. A.
belizensis Lundell, Contr. Univ. Mich. Herb. 7: 38. 1942 (type from
Stann Creek Valley, British Honduras, Percy H. Gentle 3500) . Sirasil
de montana; ixbambul; hueso bianco; cerezo morado; capulin (Guate-
mala) ; male blossom-berry grape; bird berries (British Honduras) .

Moist or wet forest, often in second growth, frequently in pine-
oak forest, 2,500 meters or less; Pete"n; Alta Verapaz; Bajo Verapaz;
Izabal; Zacapa; Chiquimula; Jalapa; Santa Rosa; Escuintla; Guate-
mala; Sacatepe"quez; Chimaltenango; Suchitepe"quez; Quezaltenango;

140 FIELDIANA: BOTANY, VOLUME 24

San Marcos. Mexico; British Honduras to El Salvador and Panama;
northern South America (type from Venezuela).

A shrub or small tree up to 7 meters tall, 15 cm. in diameter, the branchlets
slender, at first furfuraceous-lepidote; the leaf buds furfuraceous-lepidote; leaves
variable, with short usually stout narrowly marginate petioles rarely over 1 cm.
long, often much shorter, the petioles furfuraceous-lepidote beneath at first; leaf
blades variable, lanceolate, oblong, elliptic-oblong, elliptic, oblanceolate or obovate,
6-20 cm. long, 3-9 cm. wide, acuminate or subabruptly acuminate, the acumen
acute to obtuse, base acutish and decurrent, usually entire, sometimes inconspicu-
ously crenulate or denticulate, usually chartaceous, sometimes coriaceous, glabrous,
costa elevated beneath, the primary lateral veins slender, the veins obscure to con-
spicuously reticulate; inflorescences terminal, reddish, furfuraceous at base, very
sparingly furfuraceous above at first, glabrous early, paniculate, 2- or 3-pinnate,
the panicles very variable in size, 5-15 cm. long, usually many-flowered, open;
pedicels slender, 4-9 mm. long, rarely shorter; flowers corymbose or subumbellate,
glabrous; calyx pink, sepals dextrorsely imbricate, almost free, ovate-elliptic, ovate
or oval, 1-2 mm. long, apex rounded, minutely erose or ciliolate, conspicuously
orange-red punctate; petals dextrorsely contorted, oblong, oblong-elliptic or ob-
lanceolate-oblong, 5-7 mm. long, united 1-1.5 mm. at base, rather sparsely orange-
red punctate; stamens 3-5.5 mm. long; filaments 1-2 mm. long, sometimes longer;
anthers linear or lanceolate-linear, 2.5-4 mm. long, erect, dorsifixed about 0.5 mm.
above base, dehiscent by apical pores, concolor; ovary ovoid; style slender, sub-
equaling petals; placenta ovoid; ovules pluriseriate, numerous, immersed; fruits
(immature) shallowly pitted; mature fruits subglobose, 6-8 mm. in diameter,
purple-black.

One of the commonest shrubs of forest at low and middle eleva-
tions, A. compressa is quite variable over its wide range. A diagnostic
characteristic, present in all the variable populations, is the furfura-
ceous internode at the base of the inflorescence. The indument per-
sists through fruiting.

The ripe fruit is eaten by children as well as by birds.

Ardisia cucullata Lundell, Wrightia 3: 26. 1962.
In wet forest, 2,540 meters or less; Mexico (type from Fraylesca,
near Siltepec, Chiapas, E. Matuda 5201).

A tree, up to 7 meters tall, the branchlets rather slender, minutely papillate-
puberulent in leaf axils, otherwise glabrous; the leaf buds ciliate; leaves with rather
stout narrowly marginate petioles 8-15 mm. long, the petioles inconspicuously lepi-
dote above; leaf blades oblong-elliptic or oblanceolate-oblong, 12-20 cm. long,
4-7.5 cm. wide, base acutish and decurrent, apex acute or subabruptly acuminate,
chartaceous, glabrous, entire, the young leaves conspicuously black-punctate mostly
with small rounded glands, costa elevated beneath, nearly plane above, reticulate-
veined on both surfaces; inflorescences terminal, broadly pyramidal, up to 12 cm.
high, 15 cm. wide, the racemes corymbiform, paniculate, glabrous; pedicels slender
but rigid, 1-3 cm. long, the longest at base of racemes; flowers 5-merous, 7-8 mm.

LUNDELL: FLORA OF GUATEMALA 141

long at anthesis; calyx 4-4.5 mm. long, the sepals quincuncial or imbricate, united
at base, ovate-elliptic or ovate-oblong, narrowed and rounded at apex, ciliate,
lepidote-papillose at base within, conspicuously black-punctate, the medial glands
linear; corolla 7-9 mm. long, the petals united at base into tube about 2.5 mm.
long, the tube pubescent outside at base with gland-tipped hairs, lepidote-papillose
within, the petals symmetrical, black-punctate, multilineate medially, the two
outer petals larger, elliptic-lanceolate, obtuse at apex, the three inner petals cucul-
late apically; stamens 4-6 mm. long, attached about 1.5 mm. above base of tube;
filaments stout, enlarged below, 2-2.5 mm. long, densely pubescent with short
gland-tipped hairs; anthers oblong-triangular, 3-4.5 mm. long, epunctate, dorsi-
fixed about 1 mm. above base, apex rounded and mucronate; ovary ovoid, black-
punctate, glabrous, tapering into the slender style 6-7 mm. long; placenta acicular
at apex; ovules pluriseriate, numerous.

The long very slender pedicels, the elongated subcorymbiform
racemes, smaller flowers and thinner leaves are features by which
A. cucullata differs markedly from A. verapazensis Donn.-Sm. and
A. sexpartita Lundell, the species with apparent affinity.

Ardisia densiflora Krug & Urb. Notizbl. Bot. Gart. Berl. 1: 79.
1895 (type from Blue Mountains, Jamaica, W. Harris 5227). A. spi-
cigera Donn.-Sm. Bot. Gaz. 27: 434. 1899 (type from Comitan, Chi-
apas, E. Seler 2678). A. Gentlei Lundell, Field & Lab. 13: 11. 1945
(type from Toledo District, British Honduras, Percy H. Gentle 4561) .
Caraso berries; wild spice (British Honduras) .

Mostly in wet forest, sometimes in swampy places or pine-oak
forest, 1,800 meters or less, usually at low elevations; Pete"n; Zacapa.
Mexico (Veracruz, Oaxaca, Chiapas); British Honduras; Honduras.

A shrub or tree, sometimes up to 25 cm. in diameter and 12 meters high, the
branches stout; the short terminal internodes at first minutely ferruginous-tomen-
tose with closely appressed tomentum, glabrescent early; the leaf buds finely fer-
ruginous-tomentose; leaves with stout marginate petioles usually less than 1 cm.
long, sometimes up to 1.5 cm. long; leaf blades obovate-elliptic, oblanceolate or
oblong-elliptic, 7-20 cm. long, 2.5-8 cm. wide, apex usually obtuse, sometimes
acutish, base acutish and decurrent, entire, glabrous, coriaceous or subcoriaceous,
drying pallid, costa impressed above, elevated beneath, the veins slender and ob-
scure; inflorescences terminal, spicate-paniculate, up to 15 cm. long, 2-3-pinnate,
sparingly ferruginous-tomentulose at first, glabrescent; flowers sessile or subsessile,
the pedicels rarely up to 6 mm. long at base of spikes; flowers 5-6 (7) mm. long at
anthesis; sepals quincuncial, free almost to base, broadly ovate or suborbicular,
1.7-2 .4 mm. long, rounded apically, ciliate, coriaceous and thickened medially,
rugose, punctate, the margin scarious, glandular-lepidote within at base, otherwise
glabrous; petals dextrorsely imbricate or quincuncial, oblong, 5-5.5 mm. long,
united at base, the tube about 2 mm. long, apex rounded, black-punctate medially,
lineate, glandular-lepidote within tube, glabrous otherwise; stamens usually sub-
equaling petals, rarely exserted or included, up to 6 mm. long; filaments slender,
up to 4.5 mm. long, coalescing at base, attached medially in corolla tube; anthers

142 FIELDIANA: BOTANY, VOLUME 24

ovate-triangular, rather thick, epunctate, 1.5-2 mm. long, acute or apiculate, dor-
sifixed about one-third above base; ovary glabrous; style up to 6 mm. long; ovules
numerous, pluriseriate, immersed; fruit subglobose, drying 6-7 mm. in diameter,
black and shining at maturity.

The fruits are reported to be acid-sweetish and edible (Steyer-
mark 42899). In both continental and Jamaican collections, fruits
are often malformed and enlarged from disease. Since the species
appears to be rather rare, possibly this has contributed to the sparse
population.

A. densiflora is closely related to A. revoluta H.B.K. but a very
distinctive species. Flowers in some specimens are pedicellate, and
this has led to confusion. The thick rugose calyx helps to distinguish
even fruiting material of A. densiflora.

Ardisia erythrocarpa Lundell, Wrightia 2: 59. 1960. Chalche
(Pete*n); Chilich (Huehuetenango).

Wet forest, 300 meters or less; Pete"n (type from San Luis, C. L.
Lundell 16267); Huehuetenango. British Honduras.

A slender shrub or small tree up to 3 meters high, the branchlets slender, rather
sparsely puberulent with minute red hairs, glabrescent; leaves with slender, mi-
nutely puberulent petioles 3-8 mm. long; leaf blades lanceolate-elliptic, elliptic or
oblanceolate, 3-10 cm. long, 1.3-4.3 cm. wide, apex acuminate, the acumen ob-
tusish, base acute, membranaceous, decurrent, the margin crenulate above or
subentire, at first puberulent above and along costa beneath, the veins slender;
inflorescences lateral and terminal, minutely puberulent, subcorymbose, few-flow-
ered, rarely reduced to 1 flower; pedicels slender, 8-15 mm. long, usually less than
12 mm. long; flowers 5-merous, white; sepals free, lanceolate, ovate or lanceolate-
oblong, 2-4 mm. long, acutish, ciliate, punctate; petals connate at base, lanceo-
late-oblong, 6-9 mm. long, 3-4 mm. wide, obtuse and laterally emarginate at apex,
conspicuously punctate, glandular-puberulent within above base of filaments; sta-
mens up to 5 mm. long; filaments stout, 1.5-2 mm. long, glandular-puberulent,
inserted at base of corolla; anthers linear, only slightly enlarged at base, 3-4 mm.
long, apiculate, dorsifixed about 0.5 mm. above base; ovary glabrous; style slender,
about 5 mm. long; placenta ovoid; ovules 8 or more, pluriseriate, immersed; fruit
bright red, depressed globose, drying 6-8 mm. in diameter.

Ardisia escallonioides Schlecht. & Cham. Linnaea 6: 393. 1831.
Cyrilla paniculata Nutt. Amer. Journ. Sci. 5: 290. 1822, not Ardisia
paniculata Roxb. 1814. A. Pickeringia Torr. & Gray, in DC. Prodr.
8: 124. 1844. Icacorea paniculata (Nutt.) Sudw. Gard. & For. 6:
324. 1893. Uva (Huehuetenango); hullaba (British Honduras).

Usually in low wet or dry forest, 900 meters or less, often near sea
level; Pete"n; Huehuetenango. Southern Florida; Mexico (Tamauli-

LUNDELL: FLORA OF GUATEMALA 143

pas, San Luis Potosi, Veracruz, Oaxaca, Chiapas, Campeche, Yuca-
tan); British Honduras; West Indies (Bahamas, Cuba, Hispaniola).

A shrub or small tree, sometimes up to 12 meters high, the bark thin, scaly,
light gray or white, the young branchlets rather slender, finely pubescent with
minute appressed ferruginous tomentum; leaves with petioles usually about 5 mm.
long, but ranging from 3-10 mm. long, pubescent with minute ferruginous tomen-
tum; leaf blades oblanceolate, obovate, elliptic or oblong, 5-10 (18) cm. long, 2-4
(6.5) cm. wide, apex obtuse or acute, base acute, chartaceous or subcoriaceous,
entire, glabrous or nearly so, minutely pitted, usually drying pallid and paler be-
neath, costa elevated beneath, the veins very slender and inconspicuous; inflores-
cences terminal, paniculate, 2-3-pinnate, 5-12 cm. long, very minutely puberulent
or finely ferruginous-tomentulose; pedicels (2) 4-6 (9) mm. long; flowers racemose-
paniculate, fragrant, white or pink, at anthesis up to 8 mm. long including exserted
style; sepals quincuncial or sinistrorsely imbricate, ovate-elliptic, 2 mm. long or less,
rounded at apex, ciliate, conspicuously black-punctate with lines and small round
glands; petals up to 6 mm. long, usually sinistrorsely imbricate, rarely quincuncial,
connate up to one-third at base, oblong-elliptic, conspicuously lineate, papillose-
puberulent within at base; stamens up to 4 mm. long; filaments slender, up to 2.5
mm. long; anthers erect, lanceolate-triangular or rarely ovate, 1.2-2 mm. long,
dorsifixed 0.3-0.5 mm. above base, eglandular, acute or apiculate, rarely cuspidate;
ovary glabrous, ovoid; style rather stout, up to 6 mm. long, conspicuously ex-
serted; ovules numerous, pluriseriate, immersed; fruit globose, usually 4-5 mm. in
diameter, rarely larger.

In our area the species is quite uniform, but some Pete"n collec-
tions have leaves approaching those of A. revoluta H.B.K. in size.
The sinistrorse sepals and petals, along with the stout exserted style,
make this racemose-flowered species easy to recognize.

In its northeastern Mexican range, the species has small flowers
with cuspidate anthers scarcely 1.2 mm. long, differences possibly
of varietal importance. The pedicels vary considerably in length
throughout the range of A. escallonioides, but they are possibly short-
est in collections from San Luis Potosi and Tamaulipas. In the
Bahamas, specimens with fruits up to 1 cm. in diameter may repre-
sent another variety.

Ardisia escuintlensis Lundell, Wrightia 3: 98. 1964.
Known only from the type, Dept. Escuintla, 1942, Jose Ignacio
Aguilar 1679.

Branchlets slender, terete, glabrous; leaves with slender petioles 1 cm. long;
leaf blades oblanceolate or narrowly oblanceolate-elliptic, 7.5-10 cm. long, 2.8-
3.5 cm. wide, apex obtusely subacuminate, base acute, glabrous, entire, charta-
ceous, slightly paler beneath, the costa slightly impressed above, nearly plane,
elevated beneath, the veins very slender, inconspicuous; inflorescences terminal,
bipinnately paniculate, shorter than leaves, glabrous, the flowers racemose at ends
of branches; bracts thin, deciduous, the basal up to 1.5 cm. long, conspicuously

144 FIELDIANA: BOTANY, VOLUME 24

punctate with very small elevated reddish glands; pedicels 3-5 mm. long; sepals 5,
free or essentially so, ovate-lanceolate, about 2 mm. long, glabrous, obtusish, suben-
tire, membranaceous, epunctate or sometimes with one or more small glands; petals
dextrorsely imbricate, short connate at base, ovate-lanceolate, 4 mm. long (in
buds), membranaceous, epunctate, glabrous; stamens (in buds) 3 mm. long, at-
tached at base of corolla; filaments rather stout, about 1.5 mm. long; anthers
dorsifixed about two-fifths above base, sagittate, narrowly triangular and caudate-
acuminate, about 2.75 mm. long (in buds), epunctate, longitudinally dehiscent;
ovary ovoid, glabrous; style slender, short, about 1.5 mm. long, the stigma puncti-
form; ovules numerous, pluriseriate.

The only specimen available is a single flowering twig with flower
buds, but the species is very well marked. A. escuintlensis has mem-
branaceous sepals and petals, long caudate-acuminate anthers dehis-
cent longitudinally, and inflorescences paniculate with flowers in
short racemes. Although entirely different in appearance, it is prob-
ably related to A. revoluta H.B.K.

Ardisia hirtella Lundell, Wrightia 3: 98. 1964.
Wet cohune ridge, about 750 meters (type from British Honduras,
Retire in El Cayo District, C. L. Lundell 6302).

Arborescent shrub, 3 cm. diameter, 3.5 m. high, branchlets slender, hirtellous
with red hairs; leaves petiolate, the petioles red puberulent, 7-15 mm. long, the
acute base of blade decurrent on petiole; leaf blades membranaceous, lanceolate,
8.5-14 cm. long, 3-4.7 cm. wide, apex obtusely acuminate, base acute, margin
obscurely crenulate above, puberulent on both surfaces at first, glabrescent above ;
inflorescences terminal, corymbose or corymbose-umbellate, multi-flowered, up to
7 cm. in diameter, red puberulent; pedicels slender, 1-1.5 cm. long, red puberu-
lent; flowers 5-merous; calyx red puberulent, 3-3.5 mm. long, the sepals lanceolate,
conspicuously ciliate, obtusish; corolla 7-10 mm. long, pinkish, the petals dex-
trorsely imbricate, asymmetrical, laterally emarginate at apex, lanceolate-oblong,
3-4 mm. wide, minutely ciliate, rather sparsely glandular puberulent within at
base, otherwise glabrous, united at base; filaments stout, up to 2 mm. long, gla-
brous; anthers linear-lanceolate, up to 4 mm. long, obtuse and apiculate at apex,
opening by apical pores, attached to filaments above the subsagittate base; ovary
glabrous; style slender, about 6 mm. long.

Ardisia Mitchellae I. M. Johnston, Contr. Gray Herb. 81: 96.
1928.

Wet forest, 600 meters or less, usually near sea level (type from
Honduras, vicinity of Tela, Elizabeth R. Mitchell 66).

A shrub or small tree up to 5 meters high, the branchlets slender, minutely
puberulent; leaves with slender puberulent petioles 3-10 mm. long; leaf blades
elliptic, obovate-elliptic or oblanceolate-elliptic, 6-13 cm. long, 2.5-5.5 cm. wide,
apex abruptly and broadly acuminate, base acute and decurrent, margin crenulate
from about the middle, chartaceous, paler beneath, at first puberulent along the

LUNDELL: FLORA OF GUATEMALA 145

midvein on both surfaces, the midvein elevated beneath, nearly plane above, the
veins slender, obscure on upper surface; inflorescences strictly terminal, laxly sub-
corymbose, few-flowered, up to 6 cm. long, densely puberulent with minute red
glandular hairs; pedicels very slender, up to 2 cm. long, puberulent, flowers 5-
merous, up to 1.2 cm. long at anthesis; sepals short-connate at base, broadly ovate
or ovate-orbicular, up to 3 mm. long, acute, puberulent, ciliate, densely punctate;
petals (only mature buds available) dextrorsely contorted, oblong, up to 11 mm.
long, 4 mm. wide, united about 1 mm. at base, minutely ciliolate, glandular-
puberulent within above base of filaments, otherwise glabrous, conspicuously punc-
tate; stamens up to 9.3 mm. long (in buds of type), inserted near base of corolla;
filaments thick, about 2 mm. long, glandular-puljerulent; anthers linear, up to

9 mm. long (in buds of type), apex cuspidulate, the cusp shallowly bifid, not punc-
tate; ovary ovoid, glabrous; style slender, placenta ovoid, apically acicular; ovules

10 or more, pluriseriate, immersed; fruit bright red, globose, drying 8-10 mm. in
diameter.

In the type, which has only mature flower buds, the petals exceed
a centimeter in length and these are subequaled by the stamens. The
large anthers, up to 9 mm. long, are distinctly cuspidulate and the
cusp is bifid. Apparently the abnormally large flowers and anthers
of the type represent an extreme development of the species. No
collections from British Honduras and Guatemala are definitely refer-
able to A. Mitchellae, but the species should be found in the Atlantic
lowlands.

Ardisia nigrescens Oerst. Vid. Medd. Kjoebenhavn 130. 1861.
A. Donnell-Smithii Mez, Bull. Herb. Boiss. II. 3: 235. 1903 (type from
Cubilquitz, Alta Verapaz, H. von Tuerckheim 7766). A. multilineata
Mez, Repert. Sp. Nov. 16: 311. 1920 (based on same type as A. Don-
nell-Smithii Mez). A. nigrescens Oerst. var. Donnell-Smithii (Mez)
Lundell, Wrightia 3: 99. 1964. "Coc-ishilil" (Kekchi in Quiche").
Uva de paloma; blossomberry (British Honduras).

Wet forest, 350 meters or less; Pete"n; Alta Verapaz; Huehuete-
nango. Mexico (San Luis Potosi, Veracruz, Oaxaca); British Hon-
duras.

A shrub or small tree up to 8 meters high, the trunk 10 cm. or less in diameter,
the young branchlets slender, densely hirsute-tomentose with matted dark red
hairs; leaves with canaliculate petioles (3) 5-10 (15) mm. long; leaf blades oblong,
lanceolate, oblong-elliptic, elliptic, oblanceolate or obovate-elliptic, 5-18 cm. long,
2-7.5 cm. wide, apex subabruptly acuminate or acuminate, base acute to obtuse or
rounded, decurrent, subentire, serrulate or crenulate-dentate, chartaceous, glabres-
cent above except at base and along the midvein, pilosulous beneath, glabrescent,
the costa elevated beneath, shallowly impressed above and often subbullate,
the veins slender, obscure or rather conspicuous; inflorescences terminal, usually
small, broadly paniculate, 1- or 2-branched, 3-7 cm. long, densely ferruginous-
hirtellous; pedicels slender, 6-18 mm. long; flowers subcorymbose, 7.5-10.5 mm.

146 FIELDIANA: BOTANY, VOLUME 24

long at an thesis; sepals short connate at base, ovate, ovate-oblong or lanceolate-
oblong, 2.3-2.8 mm. long, slightly keeled, hirtellous, long-ciliate, densely punctate;
petals dextrorsely contorted, oblong-elliptic, up to 10 mm. long, connate about 2
mm. at base, glandular-puberulent above filaments at base, minutely ciliolate,
glabrous otherwise, densely punctate, lineate, obliquely emarginate at apex; sta-
mens inserted below middle of corolla tube, 5-5.5 mm. long; filaments stout, about
1.5 mm. long, humped above, pubescent with short glandular hairs; anthers erect,
linear, subsagittate, up to 4.5 mm. long, dorsifixed about 0.75 mm. above base,
epunctate, apiculate; ovary ovoid, glabrous; style slender, 5-5.5 mm. long; ovules
10 or more, pluriseriate, immersed in placenta; fruits subglobose, drying up to
8 mm. in diameter.

With the reduction to varietal status of A. Donnell-Smithii Mez,
the enigma of A. nigrescens Oerst. is resolved. The small-leaved
shrubs of San Luis Potosi, Veracruz and Oaxaca have their counter-
part in British Honduras (W. A. Schipp 287 from Middlesex). The
plants with large mostly subentire or obovate leaves, usually rather
obtuse or rounded at base and with venation rather conspicuous are
typical of A. Donnell-Smithii, but these intergrade into the smaller
leaved types.

With extremes of variability in the foliage of the species, it does
not seem that the large-leaved A. Donnell-Smithii should be accorded
specific rank, but varietal status can be justified. The entire A. ni-
grescens complex has flowers very much alike. The uniformity of the
stamens is particularly remarkable. The pubescence of A. nigrescens
and its variety consists of rather stiff dark red hairs and these are
usually dense on the younger growth. The undersurface of the leaves
is pilosulous.

The leaves vary considerably in size with the greatest reduction
shown in Sesse, Mocino, Castillo & Maldonado 679 (F) . The largest
leaves are represented by the type collection of A. Donnell-Smithii.
A collection from Quiche" (Harry Johnson 228) has large leaves with
crenate-dentate margin typical of the small-leaved plants of Mexico.

This shrub is very ornamental, its attractive flowers ranging from
white or pink to lilac, and its bright red berries are shiny.

Ardisia nigropunctata Oerst. Vid. Medd. Kjoebenhavn 127.
t. 2. 1861. A. pulverulenta Mez, Pflanzenreich IV. 236: 88. 1902 (type,
Friedrichstahl 613). A. chontalensis Mez, Pflanzenreich IV. 236: 90.
1902 (type from Laguna de Chiriqui, Panama, John Hart 136) . Sira-
sil (Izabal) ; blossomberry, high ridge blossom berry, blossomberry grape,
asib (British Honduras).

LUNDELL: FLORA OF GUATEMALA 147

Wet forest, 1,600 meters or lower, mostly at low elevations; Alta
Verapaz; Izabal. British Honduras; Honduras; Nicaragua; Costa
Rica; Panama.

Shrub, or small tree up to 8 meters high, the branchlets minutely and densely
ferruginous-lepidote at first; leaves with short marginate petioles up to 1 cm. long;
leaf blades oblong-elliptic, obovate-elliptic or oblanceolate, 15-30 cm. long, 5-12
cm. wide, rarely larger, usually acuminate, base acute and decurrent, thin, margin
usually entire, sometimes denticulate, minutely and sparsely lepidote, dotted be-
neath with abundant black glands, the costa and primary lateral veins elevated
beneath, often slightly impressed above; inflorescences terminal, many-flowered,
open, large and pyramidal, often longer than the leaves, 2-4-pinnate, minutely
lepidote or furfuraceous-lepidote; pedicels sparsely lepidote, 3-5 mm. long; flowers
5 mm. long, the calyx lepidote at base, otherwise glabrous, conspicuously black-
punctate with elevated glands; sepals dextrorsely imbricate, broadly ovate, 1.6-
2 mm. long, acutish, minutely erose-ciliate, sparsely lepidote at base, glabrous
otherwise; petals dextrorsely imbricate, united about 1 mm. at the lepidote base,
ovate-lanceolate, slightly asymmetrical, often erose; stamens shorter than petals,
3.75 mm. long; filaments slender, 1.5-1.75 mm. long; anthers dorsifixed one-fifth
above base, lanceolate-oblong, 2.3-2.6 mm. long, with black line dorsally, apex
apiculate or subulate, longitudinally dehiscent; ovary ovoid, glabrous; style slen-
der, up to 4.5 mm. long; placenta ovoid, apiculate, the ovules pluriseriate, 14-17;
fruit globose, 5-6 mm. in diameter.

Ardisia paschalis Donn.-Sm. Bot. Gaz. 19: 5, t. 1. 1894 (type
from Cuyotenango, Suchitepe"quez, J. Donnell Smith 2465). Icacorea
paschalis (Donn.-Sm.) Standl. in Standl. & Cald. Lista Salvador 168.
1925. A. dichropetala Standl. Trop. Woods 37: 29. 1934 (type from
La Libertad, Dept. Comayagua, Honduras, J. P. Edwards P-621).
A. Matudai Lundell, Lloydia 4: 55. 1941 (type from La Palma, Ta-
basco, Mexico, E. Matuda 3295). Xilil (Pete*n); Chilil; Capulin;
sirasil de pava; morrito.

Moist or wet forest, 2,500 meters or lower; Pete"n; Alta Verapaz,
El Progreso; Izabal; Jutiapa; Santa Rosa; Escuintla; Guatemala;
Sacatepe"quez; Suchitepe'quez; Quezaltenango. Mexico (Chiapas,
Tabasco); British Honduras; Honduras; El Salvador.

A shrub or small tree, sometimes 9 meters high, the branchlets rather stout,
glabrous; leaves with short, narrowly marginate petioles up to 1.5 cm. long; leaf
blades obovate, oblanceolate, elliptic or oblong-elliptic, 7-25 cm. long, 4.5-10 cm.
wide, obtuse, rounded, acute or broadly acuminate at apex, acute and decurrent
at base, chartaceous to subcoriaceous, glabrous, drying pallid, densely punctate
along the margin, entire, costa elevated beneath, nearly plane above, the veins
very slender; inflorescences terminal, usually lax and open, up to 20 cm. long, 25
cm. wide, rarely congested, many-flowered, the racemes bipinnate-paniculate,
corymbiform with the lower pedicels usually elongated; pedicels rather slender,
straight or reflexed, mostly 1-1.5 cm. long, rarely only 0.5 cm. long, sometimes up
to 2.5 cm. long at base of racemes; flowers 8-12 mm. long at anthesis; sepals sub-

FIG. 38. Ardisia paschalis. A, Branch with inflorescence; X 3^- B> Calyx
opened out, and pistil; X 2^. C, Open flower, showing star-shaped eye at base
of corolla; X 2J^. D, Stamen, dorsal view; X 5.

148

LUNDELL: FLORA OF GUATEMALA 149

orbicular, broadly ovate or oblong-ovate, 2-3.5 mm. long, rounded at apex, ciliate,
sparingly to densely punctate, rarely epunctate, glabrous; corolla white or yellow-
ish with glandular, yellow or orange, star-shaped eye within, otherwise glabrous,
the petals elliptic or oblong-elliptic, up to 1.1 cm. long, sparingly to conspicuously
punctate, sometimes epunctate or nearly so, united at base into a short tube; sta-
mens large, 5-7.5 mm. long, attached about 1 mm. above base of tube; filaments
stout, usually short, sometimes up to 3 mm. long; anthers large, thick, narrowly
triangular, 3-6 mm. long, dorsifixed near base, apiculate, the apex usually punctate
with several small glands, dorsally sparingly punctate or epunctate, before anthesis
cohering in cone, longitudinally dehiscent; ovary glabrous, style slender, 6-7.5 mm.
long; ovules numerous, pluriseriate, immersed; fruit subglobose, 8-10 mm. in
diameter when dry, usually purple-black at maturity.

A. Lindenii Mez and A. Karwinskyana Mez, described from Mex-
ico, appear to be referable here, but I have not seen type material of
either. Both are separated from A. paschalis largely on the degree
of punctation of the calyx and corolla. A. dichropetala Standl. and
A. Matudai Lundell are in the same category.

There appears to be some correlation between punctation and
elevation, those plants growing at lower altitudes having less punc-
tate or epunctate flowers. Other differences are of the same minor
degree.

E. Matuda 16724 from Escuintla, Chiapas has fruits which are
reported by the collector to be milky-white.

The flowers of this species are singularly large and showy for the
genus, and they are reported to be very fragrant. They are used for
decorating the altars of churches during Easter week. Both flowers
and leaves are used for wreaths. The leaves dry pallid, a distinctive
characteristic of the plant.

Ardisia pellucida Oerst. Vid. Medd. Kjoebenhavn 130, t. 2.
1861. A. pectinata Donn.-Sm. Bot. Gaz. 12: 132. 1887 (type from
Pansamala, Alta Verapaz, H. von Tuerckheim 942). A. myriodonta
Standl. Journ. Wash. Acad. Sci. 17: 13. 1927 (type from Barro Colo-
rado Island, Panama, P. C. Standley 40848) . A. pellucida Oerst. var.
pectinata (Donn. Sm.) Lundell, Wrightia 3: 99. 1964. Tapacajete
(Veracruz) .

Wet forest, 1,650 meters or less, mostly at low elevations; Pete"n;
Alta Verapaz; Izabal; Escuintla; Suchitepe"quez ; Huehuetenango.
Mexico (San Luis Potosi; Veracruz, type from Pital, F. M. Liebmann
29. C); British Honduras; Honduras; Panama; Colombia.

Usually a simple shrub up to 2.5 meters, sometimes a tree of 7 meters, the
branchlets thick, apically densely glandular-puberulent and usually villous, the
glandular hairs reddish; leaves large with petioles 0.5-2.5 cm. long; leaf blades

150 FIELDIANA: BOTANY, VOLUME 24

oblanceolate or oblong-elliptic, mostly 20-40 cm. long, 6-12 cm. wide, sometimes
larger, apex short-acuminate, base attenuate and decurrent, very thin, membrana-
ceous, at first minutely puberulent beneath, especially along the veins and very
sparsely lepidote, glabrous otherwise, minutely and densely puncticulate, the mar-
gins closely and finely pectinate-dentate with short subulate teeth, the costa large
and elevated beneath, the primary lateral veins slender but conspicuous; inflores-
cences terminal, very variable in size, 3.5-20 cm. long, paniculate, 2-3-pinnate,
densely and minutely glandular-puberulent, and with interspersed villous hairs,
mostly at base; flowers corymbose, rose-purple, about 5 mm. long, the pedicels;
4-11 mm. long, papillose; sepals almost free, dextrorsely imbricate, ovate or lan-
ceolate, 2-3 mm. long, apex acute, acuminate or subulate, usually densely orange-
red punctate, sometimes obscurely and sparsely punctate, papillose-puberulent,
margin erose and ciliolate; petals dextrorsely contorted, united about 1.5 mm. at
base, 5-6 mm. long, lanceolate, acuminate, minutely puberulent on outer surface,
sparsely punctate or epunctate, minutely glandular-puberulent within at base,
margin obscurely ciliolate; stamens 2-4 mm. long; filaments short, 0.3-0.75 mm.
long; anthers erect, narrowly lanceolate-triangular, 2-3.75 mm. long, attenuate-
acuminate, dorsifixed 0.1-0.2 mm. above base, not punctate; ovary ellipsoid, usu-
ally black-punctate; style 2-2.5 mm. long; placenta ovoid, ovules pluriseriate, 10
or more, immersed; fruits depressed-ovoid, about 6 mm. in diameter, purple-black
when ripe.

The flowers are unusually variable in size, but uniform in their
characteristics except for punctation. The typical A. pellucida plant
has leaves, sepals and ovary densely black-punctate with small
rounded elevated glands. A. pellucida var. pectinata is sparingly
punctate, and in this feature quite distinctive. An intermediate form
is represented by W. A. Schipp S-559 from British Honduras which
has the sparingly punctate leaves of A. pectinata but the cuspidate
densely punctate sepals of A. pellucida. A. myriodonta differs only
in its marginally punctate leaves, having the sepals and ovary of
A. pellucida.

Ardisia rarescens Standl., Field Mus. Bot. 4: 248. 1929. Cerezo
morado (Quezaltenango) .

Wet, mixed forest, 1,300-2,700 meters; San Marcos; Quezalte-
nango. Mexico (Chiapas, type from Cerro del Boqueron, C. A.
Purpus7Q32).

A shrub or small tree, up to 10 meters tall, the branchlets rather slender, very
minutely glandular-puberulent, glabrescent, with leaves usually crowded at apex;
leaves with narrowly marginate stout petioles 5-11 mm. long; leaf blades oblan-
ceolate or oblanceolate-oblong, 7-15 cm. long, 2.5-5 cm. wide, apex short acumi-
nate, the acumen obtuse, base subcuneate and decurrent, margin entire or subentire,
subrevolute, subcoriaceous, slightly paler beneath, glabrous, punctate, costa ele-
vated beneath, prominent, nearly plane and paler above, primary lateral veins very
slender; inflorescences terminal, panicles broadly subpyramidal, up to 18 cm. long,

LUNDELL: FLORA OF GUATEMALA 151

3-4-pinnate, laxly multiflowered, the branches spreading, slender, minutely glan-
dular-puberulent, glabrescent with age; pedicels slender, 5-10 mm. long, finely
glandular-puberulent; flowers subumbellate, 5-5.5 mm. long at anthesis; sepals
free, dextrorsely imbricate, ovate-lanceolate, 2 mm. long, acutish, punctate, finely
and sparsely glandular-puberulent, ciliate with gland-tipped hairs; petals dex-
trorsely contorted, elliptic, 5 mm. long, united 1 mm. at base, minutely glandular-
puberulent at base on outer surface, otherwise glabrous, sparsely punctate, the
glands mostly small and round, apex obtuse-rounded, laterally notched; stamens
3 mm. long; filaments 1-1.2 mm. long, rather stout, sparsely glandular-puberulent
at base, glabrous above; anthers erect, oblong or ovate-oblong, 2-2.4 mm. long,
subtruncate and apiculate at apex, dorsifixed about 0.6 mm. above base, not punc-
tate, dehiscent apically; ovary ovoid, glabrous; style 4 mm. long; placenta ovoid,
apiculate; ovules 12, pluriseriate, immersed, enclosed; fruits depressed-globose,
about 7 mm. in diameter, dark purple.

The wide, paler, nearly plane midvein of the upper leaf surface
is distinctive. In leaf form and pubescence of inflorescences, the spe-
cies in fruit can be readily confused with Gentlea micrantha (Donn.-
Sm.) Lundell.

Ardisia revoluta H.B.K. Nov. Gen. & Sp. 3: 246. 1818. A. sco-
paria Mez, Pflanzenreich IV. 236: 82. 1902. A. scopulina Brandegee,
Zoe 5: 215. 1905 (type from vicinity of Culiacan, Sinaloa, Mexico,
T. S. Brandegee). Silacil; cerecilla; mora; morita; ciracil.

Wet to rather dry forest or thickets, often along streams, some-
times in pine-oak forest, 250-1,700 meters; Baja Verapaz; Guate-
mala; Jalapa; Chimaltenango; Escuintla; Santa Rosa; Chiquimula.
Mexico; Honduras; El Salvador to Panama and Colombia.

A shrub or small tree, sometimes 8 meters high, the branchlets stout, glabrous;
leaf buds ciliate and minutely ferruginous-tomentose; leaves with marginate peti-
oles, usually short, sometimes up to 1.2 cm. long; leaf blades oblanceolate, obovate-
oblong, elliptic or obovate-elliptic, mostly 9-19 cm. long, 3.5-8 cm. wide, obtuse or
often rounded at apex, attenuate to the decurrent base, glabrous, subcoriaceous or
chartaceous, entire, drying pallid, the costa prominent beneath, the veins slender
and obscure; inflorescences terminal, punctate, minutely ferruginous-tomentulose
in bud, sparsely puberulent at anthesis, glabrescent, the racemes in sessile or sub-
sessile panicles, the panicles usually less than 15 cm. long; pedicels usually 4-6 mm.
long, sometimes shorter, sometimes up to 1 cm. long; flowers white, 6-9 mm. long
at anthesis; sepals ovate-oblong or elliptic, 1.5-2.8 mm. long, rounded at apex,
ciliate, the margin scarious, black-punctate medially with lineate or oblong glands,
papillose within at base, otherwise glabrous; petals dextrorsely imbricate, oblong
or oblong-elliptic, up to 8 mm. long, rounded and laterally notched at apex, united
at base into tube about 2 mm. long, glandular-papillose within at base, the petals
otherwise glabrous, black-punctate with conspicuous lineate glands; stamens 4-
6.5 mm. long, attached near base of tube; filaments glabrous, slender, sometimes
up to 4.8 mm. long; anthers dorsifixed 0.5-0.8 mm. above base, narrowly triangu-
lar, 2-3.2 mm. long, apiculate, epunctate; ovary glabrous; style slender, 5-6 mm.

152 FIELDIANA: BOTANY, VOLUME 24

long; ovules numerous, pluriseriate, immersed; fruit coarsely punctate, subglobose,
4-5 mm. in diameter when dry, turning purple-black at maturity.

I have not seen the type of A. revoluta, and this species is pre-
sumed to be the common one of Mexico and Central America. The
photograph of the type in the Willdenow Herbarium, F. M. Neg.
4881, is my basis for this conclusion.

A. scoparia Mez, known to me from a fragment of the type (F)
and the type photograph (F. M. Neg. 31992), does not appear to
differ from A. revoluta. Mez lists A. revoluta in his key (I.e., p. 60)
under the species with petals glabrous within, and separates A. sco-
paria by the presence of lepidote-papillose indument at base within.
The common species of Mexico and Central America has petals lepi-
dote-papillose within.

Described from Mexico, where it is found from Durango and Sina-
loa south to Oaxaca, it is curiously unknown from eastern Mexico,
Chiapas and the Yucatan Peninsula. It is abundantly represented
from all countries of Central America.

Ardisia Schippii Standl., Field Mus. Bot. 12: 412. 1936.
Wet forest, at low altitudes; Pete"n. British Honduras (type,
W.A. SchipplZGS).

Small tree, up to 11 meters high, 25 cm. in diameter; branchlets minutely glan-
dular-puberulent; leaves with marginate petioles 1.5-2.5 cm. long; leaf blades
elliptic or oblong-elliptic, 12.5-20 cm. long, 5.5-9.5 cm. wide, apex acutish or ob-
tuse-rounded and apiculate, acute or subobtuse at base and decurrent, chartaceous,
entire below and at apex, elsewhere finely denticulate or serrulate-denticulate, at
first minutely puberulent along costa beneath, glabrous otherwise, costa prominent
and elevated beneath, nearly plane above, primary lateral veins slender; inflores-
cences paniculate, the panicles 2-4-pinnate, large and broad, sometimes longer
than leaves, open, minutely glandular-puberulent; flowers numerous, 6-7 mm.
long, corymbose or in racemiform corymbs past anthesis; pedicels 3-7 mm. long;
sepals dextrorsely imbricate, oblong-ovate, 2-2.2 mm. long, scarious-marginate,
short ciliate with gland-tipped hairs, coarsely black-punctate; petals dextrorsely
imbricate, united about 1 mm. at base, oblong-lanceolate, 5.5-6 mm. long, black-
punctate medially, glandular pubescent within at base, otherwise glabrous; sta-
mens 3.75 mm. long; filaments slender, about 2 mm. long, densely glandular-
pubescent; anthers dorsifixed one-third above base, narrowly lanceolate, 2.7 mm.
long, attenuate apically and conspicuously caudate, with several small pellucid
glands dorsally above point of attachment, longitudinally dehiscent; ovary glo-
bose, glabrous; style slender, 5 mm. long; placenta subglobose; ovules pluriseriate,
numerous.

Ardisia sexpartita Lundell, Wrightia 3: 29. 1962. Huesa de
montana (Quezaltenango) ; polo cruz (San Marcos).

LUNDELL: FLORA OF GUATEMALA 153

Forested slopes, mostly of volcanoes, 3,000 meters or lower, mostly
at middle elevations; Quezaltenango (type from San Juan Patzulin,
Volcan Santa Maria, J. A. Steyermark 33608); San Marcos; Suchi-
tepe"quez; Huehuetenango. Mexico (Chiapas).

A tree, sometimes 16 meters high, the branches thick, glabrous; leaves with
stout marginate petioles 0.5-2 cm. long; leaf blades oblanceolate or oblanceolate-
elliptic, 13-38 cm. long, 4.5-11 cm. wide, base attenuate, decurrent, apex broadly
acuminate, glabrous, coriaceous or subcoriaceous, obscurely crenulate or subentire,
costa stout and elevated beneath, the veins slender; inflorescences terminal, gla-
brous, broadly paniculate, 2-3-pinnate, sometimes up to 17 cm. long, usually small,
the primary axis and branches stout; pedicels stout, variable in length, up to 22
mm. long, the lower longest; flowers 5- or 6-parted, 8-12 mm. long at anthesis,
subcorymbose or in racemiform corymbs; sepals quincuncial or imbricate, coria-
ceous, with scarious ciliolate or ciliate margin, ovate-orbicular, 4-5 mm. long,
rounded at apex, conspicuously black-punctate with lines and dots, minutely glan-
dular-lepidote within at base, otherwise glabrous; corolla 9-12 mm. long, black-
punctate, the medial lines slender, the petals oblong-elliptic, obtuse-rounded at
apex, united at base into cylindrical tube 3.5-5 mm. high, the tube glandular-
papillose within above middle; stamens attached about middle of tube, 7.5-8 mm.
long; filaments slender above, expanded and united at base, 3-4 mm. long, pubes-
cent with gland-tipped hairs; anthers thick, narrowly-triangular, dorsifixed about
1 mm. above base, 3-5 mm. long, acute or apiculate, epunctate; ovary glabrous;
style slender, 7.5-9 mm. long; ovules numerous, pluriseriate; fruits globose, about
8 mm. in diameter.

Although closely resembling A. verapazensis, the species may be
readily separated by its smaller, ciliolate or ciliate, suborbicular sepals
which do not have plate-like glands on the outer surface.

Ardisia Tuerckheimii Donn.-Sm. Bot. Gaz. 13: 74. 1888.
Wet, mountain forest, 800-1,500 meters; Alta Verapaz (type from
Pansamala, H. von Tuerckheim 1035) ; Baja Verapaz.

A shrub up to 3 m. high, the branchlets very slender, glabrous or sometimes
very minutely and sparsely puberulent; leaves with slender narrowly marginate
petioles 4-8 mm. long; leaf blades lanceolate-elliptic or lanceolate-oblong, 5-11.5
cm. long, 1.6-4 cm. wide, apex subabruptly caudate-acuminate, acute or obtusish
at base, chartaceous, glabrous, entire or obscurely crenulate, slightly paler beneath,
the midvein elevated beneath, narrowly sulcate above, the veins very slender,
black-punctate; inflorescences terminal, lax, slender with few branches, usually
glabrous; pedicels very long and slender, up to 2 cm. long; flowers white, subcorym-
bose-umbellate, 7-9 mm. long at anthesis; sepals free, quincuncial, oblong-lanceo-
late or ovate, 2-3 mm. long, rounded or acutish, ciliolate, densely punctate; petals
up to 8 mm. long, connate about 2 mm. at base, oblong-elliptic, sparsely glandular-
puberulent within around base of filaments, glabrous otherwise, conspicuously
punctate; stamens 6-6.5 mm. long; filaments stout, glandular-puberulent, 2-2.5
mm. long, inserted about middle of corolla tube; anthers linear, dorsifixed about

154

FIELDIANA: BOTANY, VOLUME 24

0.5 mm. above base, about 4 mm. long, apiculate; ovary ellipsoid, glabrous, style
slender, up to 5.5 mm. long; placenta ellipsoid; ovules 10 or more, pluriseriate,
immersed.

Ardisia verapazensis Donn.-Sm. Bot. Gaz. 46: 113. 1908.
In high forest, 1,000-1,600 meters; Alta Verapaz (type from Co-
ban, H. von Tuerckheim II. 2093).

A tree, sometimes 15 meters high, glabrous, the branches thick and stout, often
densely leafy at the tips; leaves with stout broadly marginate petioles about 1 cm.

B

FIG. 39. Ardisia Tuerckheimii. A, Flowering branchlet; X J^. B, Branchlet
with fruits; X K. C, Undersurface of leaf; X H. D, Calyx; X 5. E, Flower,
dissected; X 5. F, Stamen; X 5. Flowers illustrated from isotype, Tuerckheim
1035 (F).

FIG. 40. Ardisia verapazensis. A, Flowering branchlet; X ^. B, Calyx and
style; X 5. C, Flower, showing petals and stamens; X 5. D, Stamen; X 5. Illus-
trated from type, Tuerckheim II 2093 (US).

155

156 FIELDIANA: BOTANY, VOLUME 24

long; leaf blades oblanceolate-oblong or oblanceolate, mostly 18-25 cm. long, 6.5-
9 cm. wide, short-acuminate to subobtuse, attenuate to the decurrent base, charta-
ceous to subcoriaceous, entire, glabrous, the costa stout and elevated beneath,
nearly plane above, the veins slender, obscurely reticulate on both surfaces; inflores-
cences terminal, glabrous, large and many-flowered but shorter than the leaves, up
to 15 cm. long, 25 cm. wide, paniculate, 2-pinnate, the primary axis and branches
stout; pedicels stout and thick, 0.7-3 cm. long; flowers corymbose or in corymbi-
form racemes with lower pedicels elongated, the flowers 5-merous, about 1.1 cm.
long at an thesis; calyx coriaceous, the sepals nearly free, quincuncial, ovate-elliptic
or ovate-oblong, 5-6 mm. long, rounded at apex, the margin entire, scarious, me-
dially black-punctate in two lines, the dots scattered and few, dorsally with con-
spicuous small round plate-like glands, glandular-lepidote within at base; corolla
pink, firm, the petals connate into tube 4-5 mm. high, the tube constricted medi-
ally where stamens attached, the apical half of tube densely and minutely glandu-
lar-lepidote within, the lobes broadly ovate-elliptic, usually cucullate, rounded
apically, glabrous, medially linear-punctate in two lines, with few other black
glands; stamens 7-8 mm. long; filaments slender above, wider below, 4-5 mm. long,
sparingly glandular-pubescent below with short gland-tipped hairs; anthers dorsi-
fixed about 1 mm. above base, narrowly triangular, about 4 mm. long, epunctate,
apiculate; ovary ovoid, glabrous; style rather slender, 8.5-9 mm. long; ovules
numerous, pluriseriate, immersed.

GENTLEA Lundell
Reference: Lundell, Wrightia 3: 100-108. 1964.

Shrubs or trees with alternate leaves, petiolate; inflorescences terminal, panicu-
late, broader than long, short pedunculate or subsessile; bracts and bractlets thin,
caducous; flowers corymbose, perfect, usually 5-merous, sometimes 4- or 6-merous,
greenish or white; sepals inconspicuously imbricate, united at base or nearly free;
petals connate one-fourth to one-half into tube, the lobes imbricate in bud, spread-
ing and sometimes appearing valvate at an thesis; stamens exserted, exceeding the
petals; filaments long and slender; anthers very small, cordate, dorsifixed, epunc-
tate or inconspicuously glandular-punctate; ovary ovoid or subglobose; style slen-
der, sometimes subequaling corolla; stigma punctiform, minute; placenta few to
multiovulate, the ovules in two or more series; fruits subglobose.

There are six species, ranging from Mexico (Jalisco) south to
Venezuela and Peru. All except Gentlea venosissima (Ruiz & Pavon)
Lundell, the type species, are of local distribution.

Branchlets furfuraceous; leaves small, mostly obovate, 2-4 cm. long, coarsely
crenate-dentate above the middle or subentire G. Vatteri.

Branchlets glabrous, ferruginous-puberulent at first, or minutely and densely
rufous-tomentose.

Sepals incrassate, concave, scarcely 1 mm. long; branchlets minutely rufous-
tomentose G. venosissima.

Sepals thin, 1.5-3 mm. long; branchlets glabrous or ferruginous-puberulent at
first.

LUNDELL: FLORA OF GUATEMALA 157

Inflorescences glabrous; corolla glabrous on outer surface G. tacanensis.

Inflorescences glandular-puberulent; corolla glandular-lepidote on outer sur-
face G. micrantha.

Gentlea micrantha (Donn.-Sm.) Lundell, Wrightia 3: 107. 1964.
Ardisia micrantha Donn.-Sm. Bot. Gaz. 14: 27. 1889, not H.B.K.,
1818. Parathesis micranthera Donn.-Sm. Bot. Gaz. 18: 205. 1893,
not Ardisia micranthera Pitard, 1930. Ardisia staminosa Lundell,
Wrightia 3: 78. 1964.

Wet, mountain forest, 3,000 meters or less; Alta Verapaz (type
from Coban, H. von Tuerckheim 1365); Quiche"; Huehuetenango; El
Progreso; Zacapa. Honduras.

A shrub or tree of 6-9 meters, the branchlets stout, minutely ferruginous-
puberulent when young, sometimes apparently glabrous; leaves with conspicu-
ously marginate petioles usually less than 1 cm. long; leaf blades oblanceolate,
oblanceolate-elliptic or oblong-elliptic, 7.5-23 cm. long, 2.5-6.5 cm. wide, acumi-
nate or subabruptly acuminate, base acutish and decurrent, entire or subentire,
subcoriaceous, glabrous, the midvein elevated and conspicuous beneath, impressed
above, the veins slender and obscure; inflorescences terminal, broadly paniculate,
1-3-pinnate, 3-8 cm. long, glandular-puberulent, glabrescent; pedicels 3-12 mm.
long; flowers greenish-white, corymbose, up to 9 mm. long including exserted sta-
mens; sepals almost free, lanceolate-elliptic or ovate, 2-3 mm. long, thin, sparsely
orange-red punctate, ciliolate, glabrous otherwise; petals lanceolate-oblong or ob-
long-elliptic, 5-6 mm. long, united 1.5-2 mm. at base, glandular-lepidote or nearly
glabrous on outer surface, minutely glandular-puberulent within at base, epunctate
or with occasional small glands; stamens inserted near base of tube, 6.5-8.5 mm.
long; filaments slender, glabrous, up to 8 mm. long; anthers cordate or ovate,
0.6-1 mm. long, punctate dorsally with several minute glands, versatile; ovary
ovoid, sometimes sparingly glandular-puberulent; style 3-5 mm. long; placenta
globose, apiculate; ovules numerous, pluriseriate, immersed; fruits subglobose, up
to 8 mm. in diameter when ripe, coarsely punctate.

Gentlea tacanensis (Lundell) Lundell, Wrightia 3: 105. 1964.
Ardisia tacanensis Lundell, Contr. Univ. Mich. Herb. 4: 21. 1940.

Wet mixed forest, 1,300-3,000 meters; San Marcos (Volcan Ta-
cana and Volcan Tajumulco). Mexico (Chiapas, type from Volcan
Tacana, E. Matuda S-226).

A shrub, 1.5-4.5 meters high, the branchlets rather slender, glabrous; leaves
small, on short marginate petioles up to 7 mm. long; leaf blades narrowly oblan-
ceolate or oblong-oblanceolate, 6-13 cm. long, 1.5-3.5 cm. wide, apex long-acumi-
nate, attenuate to the decurrent base, inconspicuously crenate-serrate, chartaceous,
glabrous, the midvein elevated beneath, slightly impressed above, the primary
lateral veins slender and inconspicuous on both surfaces; inflorescences terminal,
paniculate, 1-branched, 5 cm. long or less, rather congested, few-many-flowered,
sessile or nearly so, glabrous; pedicels slender, 4-8 mm. long; flowers corymbose,
up to 7 mm. long including exserted stamens; sepals 5, nearly free, dextrorsely im-

158

FIELDIANA: BOTANY, VOLUME 24

bricate, glabrous, broadly ovate, up to 2 mm. long, minutely erose, glandular-
ciliolate at first, punctate with small round orange glands; corolla greenish-white,
glabrous on outer surface, the petals united almost to middle, the tube up to 2 mm.
long, glandular-puberulent within, the corolla lobes dextrorsely imbricate, broadly
ovate, up to 2.5 mm. long, rounded at apex, sparsely and minutely punctate; sta-
mens 5, free, opposite the petals and borne at base of tube, up to 6.5 mm. long;
filaments glabrous, 4.5-6 mm. long, rather stout; anthers exserted, versatile, ovate
and shallowly cordate, 0.8-1.3 mm. long, apiculate; ovary ovoid, glabrous; style
4-5 mm. long; placenta ovoid; ovules pluriseriate, numerous, immersed; fruits
globose, black at maturity, drying 4 mm. in diameter.

B

FIG. 41. Gentlea tacanensis. A, Branchlet with inflorescence; X J^. B, Flower,
opened out; X 5. C, Calyx and pistil; X 5. D, Stamens; X 5. Illustrated from
isotype, Matuda S-226 (F).

Gentlea Vatteri (Standl. & Steyerm.) Lundell, Wrightia 3: 102.
1964. Ardisia Vatteri Standl. & Steyerm. Field Mus. Bot. 23: 220.
1947.

Wet, mountain forest, 1,500-3,000 meters; El Progreso; Zacapa;
Chiquimula; Huehuetenango (type from Cerro Huitz, Sierra de los
Cuchumatanes, between Mimanhuitz and Yulhuitz, J. A. Steyermark
48558). El Salvador.

A shrub of 1-3 meters, densely branched, the branchlets densely brown-
furfur aceous, glabrate in age, densely leafy; leaves very small, the marginate peti-
oles 3-5 mm. long; leaf blades oblanceolate, obovate-elliptic or elliptic, 2-4 cm.
long, 0.8-1.8 cm. wide, apex obtuse or acute, base cuneate, coarsely crenate-dentate
above the middle or sometimes subentire, glabrous above, sparsely and minutely
brownish-lepidote beneath or almost glabrous, the costa slender and prominent
beneath, subimpressed above, the lateral nerves inconspicuous beneath, obscure

LUNDELL: FLORA OF GUATEMALA

159

above; inflorescences terminal, small, few-flowered, corymbiform, simply branched,
scarcely more than 2.5 cm. long, leafy-bracted at the base, the upper bracts oblong,
obtuse, 4 mm. long or shorter; pedicels slender, erect, 3-9 mm. long, glandular-
puberulent; flowers about 6 mm. long, including exserted stamens; sepals 5, almost
free, oblong-lanceolate, 1.8 mm. long, acutish or obtuse, punctate, glandular-cilio-
late; corolla greenish-white, 3.2-4 mm. long, the petals united one-third at base,
minutely glandular-puberulent within the tube, the 5 lobes dextrorsely imbricate,
oblong-lanceolate, obtuse at apex, obscurely punctate and minutely papillose, the
margin glandular-ciliolate; stamens 5, free, up to 6 mm. long, borne about middle
of corolla tube; filaments slender, glabrous, 4-5.5 mm. long; anthers minute, ovate-
cordate, about 1 mm. long, black-punctate dorsally with several minute glands;
ovary ovoid, glabrous; style slender, 4.5-5.5 mm. long; placenta small, ovoid,
apiculate; ovules 7, large, pluriseriate, immersed; fruit globose, drying 4 mm. in
diameter.

FIG. 42. Gentlea Vatteri. A, Portion of stem and flowering branchlets; X 1.
B, Branchlet with fruits; X 1. C, Calyx, opened out, and pistil; X 5. D, Corolla,
dissected; X 5. E, Stamen, showing anther dehiscent by introrse slits; X 10,
F, Stamen, showing dorsal side of anther; X 10. Flowers illustrated from type,
Steyermark 48558 (F).

160 FIELDIANA: BOTANY, VOLUME 24

Gentlea venosissima (Ruiz & Pavon) Lundell, Wrightia 3: 103.
1964. Caballeria venosissima Ruiz & Pavon, Syst. 281. 1798. Myr-
sine venosissima (Ruiz & Pavon) Spreng. Syst. 1: 664. 1825. Ardisia
breviflora A.DC. Prodr. 8: 122. 1844. Ardisia Robinsonii Mez, Pflan-
zenreich IV. 236: 77. 1902. Ardisia meiantha Donn.-Sm. Bot. Gaz.
44: 115. 1907. Stylogyne phaenostemona Donn.-Sm. Bot. Gaz. 46:
113. 1908. Ardisia venosissima (Ruiz & Pavon) Macbride, Field
Mus. Bot. 13: pt. 5, 186. 1959. Ardisia phaenostemona (Donn.-Sm.)
Lundell, Wrightia 3: 78. 1964.

Wet, mountain forest, 1,350 meters or more; Alta Verapaz (type
of Stylogyne phaenostemona Donn.-Sm. from Coban, H. von Tuerck-
heim II. 1814); Quiche". Costa Rica; Venezuela; Peru.

A tree, the young branchlets slender, very minutely rufous- tomentose at first;
leaves with marginate petioles 3-6 mm. long, at first minutely ferruginous-tomen-
tose beneath; leaf blades lanceolate or lanceolate-elliptic, 7.5-10 cm. long, 2.5-4.5
cm. wide, apex obtusely acuminate, base acutish and decurrent, the margin entire,
coriaceous, glabrous, the midvein elevated beneath, plane above, the primary veins
very slender, obscurely areolate on both surfaces; inflorescences terminal, panicu-
late, 2- or 3-pinnate, sessile or nearly so, 4-6.5 cm. long, minutely ferruginous-
tomentose, many-flowered, compact; pedicels slender, 2-5 mm. long; flowers small,
subcorymbose, about 4 mm. long including exserted stamens; sepals united about
one-third at base, quincuncial, broadly ovate, about 1 mm. long, with scarious
minutely erose-ciliolate margin, densely punctate with small orange-red glands;
corolla 2 mm. long, the petals united to the middle, ovate-oval, rounded at the
apex, ciliolate, punctate with small orange-red glands; stamens inserted at the
middle of corolla tube, about 3 mm. long, exserted; filaments slender, glabrous, up
to 2.75 mm. long; anthers minute, ovate-cordate, about 0.6 mm. long, apiculate,
versatile, not punctate dorsally; ovary ovoid, glabrous; style slender, 2-2.4 mm.
long; placenta oval, apiculate; ovules 6-8, pluriseriate, immersed.

PARATHESIS Hooker f.

Trees or shrubs, young branchlets commonly ferruginous-tomentose with stel-
late or dendroid hairs, sometimes tawny, often glabrescent; leaves petiolate, usually
pubescent on lower surface, the hairs stellate or dendroid, often appressed and
bizonal, the margins entire, crenulate or dentate; inflorescences paniculate, axil-
lary or terminal; flowers perfect, usually 5-parted, sometimes 4-parted, rarely 6-
parted, mostly pink or white, umbellate, corymbose or subcorymbose-racemose;
sepals small, open in bud, connate at base, commonly tomentulose and papillose;
corolla rotate, usually tomentulose outside, papillose-tomentose inside at least api-
cally and along edges, the petals united at base, valvate, narrow, acutish; stamens
usually 5, sometimes 4 or 6, inserted near the base of corolla tube, the filaments
well developed, slender to stout; anthers subsagittate, ovate-lanceolate or oblongish
and slender, acute or mucronate or obtuse at the apex, dehiscent by introrse slits
or apical introrse pores, dorsifixed above the base, erect or versatile; ovary ovoid or

LUNDELL: FLORA OF GUATEMALA 161

subglobose, the style long and slender, the stigma punctiform; ovules few to numer-
ous, usually 1-seriate, sometimes partially 2-seriate, or rarely pluriseriate, enclosed
or exposed apically; fruit 1-seeded, the endocarp crustaceous; embryo cylindric,
transverse.

About 70 species, in tropical America, chiefly in mountainous re-
gions of Central America with Guatemala the center of greatest diver-
sity. Parathesis serrulata (Swartz) Mez is the type of the genus.

Inflorescences usually axillary, sometimes terminal in P. subcoriacea; leaf bud pres-
ent at apex of the branches bearing strictly axillary inflorescences; anthers
dorsifixed, strictly erect, usually orange- or black-punctate dorsally, often with
minute glands, sometimes epunctate and concolorous.

Pubescence of branchlets and undersurface of leaves usually coarse, hairs either
stellate with erect and spreading rays or dendroid, then either stipitate and
bearing apical or lateral branches, or the trunk of hairs branched to base.

Margin of leaves dentate with sharp regular teeth; hairs of branchlets and
undersurface of leaves simple or long stipitate and bifid or stellate api-
cally P. vestita.

Margin of leaves entire to crenulate.

Hairs of branchlets and undersurface of leaves stellate with erect and
spreading rays, usually short stipitate P. pleurobotryosa.

Hairs of branchlets and undersurface of leaves dendroid, the trunk of hairs
with lateral branches usually to base.

Leaves loosely floccose-tomentose beneath; petals up to 5 mm. long; fila-
ments 1.5-2 mm. long P. reflexa.

Leaves densely tomentose beneath, not with loose floccose hairs; petals
up to 7 mm. long; filaments 2.5-4 mm. long P. calophylla.

Pubescence of branchlets and undersurface of leaves consisting of fine sessile stel-
late hairs with appressed rays (the hairs sometimes dendroid but small in
P. leptopa).

Anthers usually large, broad and thick, ovate-lanceolate (sometimes abnormal
and small in P. leptopa).

Anthers with dorsal area conspicuously black-punctate, usually with large
glands; filaments stout, much shorter than anthers.

Leaves linear-lanceolate, up to 30 cm. long, 5 cm. wide, apex attenuate-
acuminate P. stenophylla.

Leaves oblanceolate, oblanceolate-elliptic or oblong, up to 19 cm. long,
6.5 cm. wide, subabruptly acuminate P. vulgata.

Anthers with dorsal area orange-punctate, or punctate with a few small
blackish glands; filaments sometimes subequaling anthers; leaves not
long linear-lanceolate.

Flowers pale orange-punctate; ovary glabrous except for sparse hairs api-
cally and at base of style P. aurantiaca.

Flowers red-punctate, the glands drying red-black; ovary finely tomen-
tose almost to base, base of style short villous with branched hairs.

P. leptopa.

Anthers slender, usually lanceolate and small, sometimes less than 1.5 mm.
long, epunctate and concolorous dorsally, or orange-punctate, or with
scattered minute reddish or black glands; filaments slender, often sub-
equaling anthers, sometimes longer than anthers.

Branchlets conspicuously scaly, the epidermis peeling transversely and ir-
regularly P. tartarea.

162 FIELDIANA: BOTANY, VOLUME 24

Branchlets not scaly.

Ovary and basal half of style tomentose, the hairs dense and thickly
matted (see P. leptopa, also).

Panicles branched to base, the branches very slender; pedicels filiform,
5-9 mm. long; leaves less than 10 cm. long, 5 cm. wide.

P. tomentosa.

Panicles pedunculate, the branches sharply angled, thick, the terminal
ones often reduced; pedicels stout, 3-6 mm. long; leaves large,
up to 25 cm. long, 9 cm. wide.

Flowers 3-4 mm. long at anthesis; sepals 0.5-0.75 mm. long; petals
densely papillose-tomentose over entire inner surface; anthers
up to 1.75 mm. long, concolorous or with one to several small
black glands dorsally; leaves oblong-lanceolate or oblanceolate,
the acumen acute P. Skutchii.

Flowers 5 mm. long at anthesis; sepals 1-1.5 mm. long; petals with
glabrous area above base on inner surface, papillose at base;
anthers up to 2.5 mm. long, conspicuously punctate dorsally
and in lobes with small red-black glands; leaves obovate-elliptic,
the acumen obtuse P. latifolia.

Ovary glabrous, puberulent or tomentulose apically and at base of style,
or finely tomentulose over the entire surface, the hairs never dense
and thickly matted.

Leaves subcoriaceous, drying pallid; inflorescences stout, congested,
with short terminal branches; pedicles of flowers up to 3 mm. long;
ovary glabrous P. subcoriacea.

Leaves thin, not drying pallid; inflorescences slender, with elongated
branches; pedicels mostly 5-17 mm. long, sometimes only 3 mm.
long.

Inflorescences simple or 1-branched, small, slender, 3-6 cm. long;
apex of leaf caudate-acuminate; ovary tomentulose. .P. subulata.

Inflorescences usually branched two or more times, sometimes 1-
branched, mostly over 10 cm. long, usually much larger; apex
of leaf obtuse or obtusely acuminate; ovary glabrous.

P. macronema.

Inflorescences usually terminal, sometimes axillary in P. cubana and P. subcori-
acea; leaf bud absent at apex of the branches when inflorescences strictly
terminal; anthers dorsifixed, versatile or erect, black-punctate dorsally.
Anthers versatile, dorsifixed, attached medially; filaments longer than anthers.

Flowers 4- or 5-parted; anthers with several minute black glands dorsally;
inflorescences strictly papillose; ovary sparsely papillose at apex.

P. papillosa.

Flowers 5- rarely 6-parted; anthers conspicuously black-punctate dorsally;
inflorescences minutely rufous-tomentose and papillose; ovary tomentose,

the hairs erect P. chrysophylla.

Anthers erect, dorsifixed, never versatile; filaments shorter than anthers, com-
monly one-half as long.

Pubescence of young branchlets and undersurface of leaves coarse and of dis-
tinct reddish hairs, the hairs dendroid and sessile or stipitate, or the hairs
stellate with erect and speading rays, the stellate hairs either sessile, sub-
sessile or stipitate; pubescence beneath leaves uniform or sometimes ob-
scurely bizonal.

Sepals and pedicels papillose-puberulent; the tomentum of branchlets ap-
pearing to be waxy; anthers slender, linear-oblong, often adnate and

forming a column P. columnaris.

Sepals and pedicels short hirsute with stellate or branched hairs, these often
stiff and bristly; the tomentum of branchlets not appearing waxy; an-
thers ovate or lanceolate.

LUNDELL: FLORA OF GUATEMALA 163

Flowers 5-7 mm. long; sepals usually large, commonly 2-3 mm. long,
rarely smaller, stamens 3-4 mm. long.

Branchlets conspicuously villous, dendroid hairs long-stipitate, shaggy;
hairs of costa on lower leaf surface long-stipitate, coarse and den-
droid P. sessilifolia.

Branchlets not villous, dendroid hairs short-stipitate or sessile and
branched to base; hairs of costa and lower surface of leaf stellate,
stipitate, subsessile or sessile, sometimes appressed.

Pedicels 4-12 mm. long, sometimes up to 17 mm. long in fruit, often
reflexed; hairs of lower leaf surface stipitate P. chiapensis.

Pedicels 3-5 mm. long; hairs of lower leaf surface sessile P. rufa.

Flowers less than 5 mm. long; sepals small, 1-1.5 mm. long; stamens
2-2.75 mm. long P. belizensis.

Pubescence of young branchlets and undersurface of leaves fine, mostly stel-
late and usually closely appressed, sometimes subappressed and rather
loose, the lower surface often glabrescent; pubescence beneath leaves
usually bizonal, areolate.
Ovary tomentose, at least apically, ridged, the pubescence persistent.

Young branchlets tawny, minutely appressed-pubescent with fine stellate
tawny hairs, glabrescent early; anthers 1.5-2.5 mm. long.

P. Donnell-Smithii.

Young branchlets dull red, densely ferruginous-tomentose with fine stel-
late appressed red hairs; anthers 2.7-3 mm. long, sparsely barbate
apically P. guatemalensis.

Ovary glabrous, or with scattered hairs apically, the hairs usually erect and
stiff, rarely sparsely short pilose or villous apically.

Apex of leaves acutish or obtuse; petals densely papillose- tomentose over
entire inner surface; ovules 12-16, partially biseriate P. cubana.

Apex of leaves acuminate or subabruptly acuminate; petals glabrous at
base on inner surface; ovules 4-12, strictly uniseriate.

Flowers 5- rarely 4-parted, less than 4 mm. long at anthesis; stamens
about 2.5 mm. long P. lanceolata.

Flowers 5-parted, 4-6.5 mm. long at anthesis; stamens 2.5-4.5 mm.

long.

Leaf blades entire or nearly so, chartaceous; pubescence compara-
tively coarse, subappressed P. oblanceolata.

Leaf blades conspicuously dentate, membranaceous; pubescence
minute, very fine and closely appressed P. membranacea.

Parathesis aurantiaca Lundell, Wrightia 3: 79, fig. ^9. 1963.

A cloud forest endemic of El Salvador, known only from the type
locality; Chalatenango, Los Esesmiles, east of La Palma, 2,430 me-
ters, Paul H. Allen & Mario Lewy van Severen 7310 (type).

Shrub or tree up to 7 m. high, branchlets closely appressed tomentose with
very fine hairs, glabrescent; leaves with petioles 1-2.5 cm. long; leaf blades lanceo-
late, lanceolate-elliptic or oblanceolate, 7-20 cm. long, 2.5-6 cm. wide, membra-
naceous, crenulate-serrulate, apex slightly contracted and acuminate, base acumi-
nate, decurrent, at first pubescent beneath with fine small pale appressed stellate
hairs, glabrescent early, pellucid-punctate, midvein slender, elevated beneath,
nearly plane above, primary lateral veins slender; inflorescences axillary, long-
pedunculate, 5-17 cm. long, lax, 1- or 2-branched, minutely tomentulose with ap-
pressed hairs; pedicels slender, 7-11 mm. long, puberulent; flowers umbellate or

164 FIELDIANA: BOTANY, VOLUME 24

corymbose, 5- or 6-parted, orange-punctate, up to 7 mm. long at anthesis, the
calyx puberulent, corolla minutely tomentulose; sepals triangular, up to 1.5 mm.
long, obtusish; petals linear-lanceolate, up to 8 mm. long, orange-punctate in lines,
papillose-tomentose inside except at base, with ring of minute glands at base; sta-
mens 3-4 mm. long; filaments stout, glabrous, 1.5-2 mm. long, eglandular; anthers
erect, ovate-lanceolate, thick, dorsifixed two-fifths above base, lanceolate, 2.2-2.7
mm. long, sometimes with a few short red hairs dorsally at apex, grooved and rather
sparingly orange-punctate dorsally, the glands extending into lobes; ovary ovoid,
glabrous except for a few scattered hairs apically and at base of style; style 5-6
mm. long; placenta subglobose, apiculate; ovules 7-10, ellipsoid, exposed apically,
uniseriate.

To be expected in Guatemala, P. aurantiaca is notable for having
all flower parts punctate with very pale orange glands.

Parathesis belizensis Lundell, Wrightia 3: 61. 1963.

Wet forest, 500 meters or less; Izabal. British Honduras (Stann
Creek District, type from Middlesex, Percy H. Gentle 2779) .

A small tree, branchlets rather stout, tomentose with rather coarse dark red
short stipitate dendroid and stellate hairs; leaves with stout marginate petioles up
to 1 cm. long; leaf blades oblong-elliptic or lanceolate-elliptic, 10-20 cm. long, 4-
6.5 cm. wide, apex subabruptly acuminate, base narrow and acute, decurrent,
thinly chartaceous, rather obscurely denticulate, glabrescent early above, uni-
formly stellate pubescent over entire surface beneath with red short stipitate or
subsessile hairs, the costa prominent beneath, slightly impressed above, the pri-
mary lateral veins slender and elevated beneath, plane above; inflorescences shorter
than leaves, paniculate, 12 cm. long, red-tomentose with subsessile rather coarse
multibranched dendroid hairs; pedicels short, 2-4 mm. long, papillose and spar-
ingly stellate-hirtellous; flowers corymbose, crowded, bristly tomentose with short
red stellate hairs and papillose, up to 4.5 mm. long at anthesis; sepals narrowly
triangular, up to 1.5 mm. long, acuminate, black-punctate; petals linear-lanceolate,
4.5 mm. long, black-punctate in lines, papillose-tomentose within except at base;
stamens less than 3 mm. long; filaments about 1.5 mm. long; anthers erect, dorsi-
fixed one-third above base, elliptic-lanceolate, about 2.2 mm. long, acutish, with a
few short reddish hairs apically, prominently black-punctate, with glands extend-
ing down into lobes; ovary ovoid, tapering into style, with scattered short erect
hairs apically; style about 3 mm. long; ovules 6, uniseriate, enclosed; fruits globose,
borne on stout pedicels about 5 mm. long.

This species closely resembles P. sessilifolia Donn.-Sm., but dif-
fers at once in not having the conspicuous long shaggy villous hairs
which are so characteristic of the latter.

Parathesis calophylla Donn.-Sm. Bot. Gaz. 18: 205. 1893. Uva;
cinco negritos (Huehuetenango) .

Moist or wet, mountain forest, 2,000-3,500 meters; Quiche* (type
from San Miguel Uspantan, Heyde & Lux 2909); Huehuetenango;
El Progreso. Mexico (Chiapas).

LUNDELL: FLORA OF GUATEMALA 165

Tree, up to 15 m. high; branchlets stout, densely and persistently ferruginous-
tomentose, the hairs multibranched to base; leaves with rather stout to slender
petioles up to 3.5 cm. long; leaf blades lanceolate or elliptic-lanceolate, up to 18 cm.
long, 7 cm. wide, apex acuminate or subabruptly acuminate, base acute or acumi-
nate, subcoriaceous, entire or obscurely crenulate, glabrous above except along
midvein and petiole, shiny, persistently ferruginous-tomentose beneath with fine
matted multibranched hairs, costa stout and elevated beneath, impressed above,
the primary lateral veins slender, slightly impressed above, the surface sometimes
slightly bullate; inflorescences axillary, pyramidal, the terminal leaves usually re-
duced, ferruginous-tomentose with multibranched hairs, paniculate, 1- or 2-branched,
6-15 cm. long, multiflowered; pedicels 3-6 mm. long; flowers umbellate or subcorym-
bose, tomentose, 6-7 mm. long at anthesis; sepals narrowly triangular, 2-2.5 mm.
long, acuminate, orange-punctate in lines; petals linear-lanceolate, up to 7 mm. long,
papillose-tomentulose on outer surface and on inner surface except at base, punc-
tate in lines; stamens 3.75-5.5 mm. long; filaments slender, 2.5-4 mm. long, gla-
brous, punctate; anthers narrowly lanceolate, 2-3 mm. long, apiculate, dorsifixed
one-fourth above base, with several small glands dorsally in a narrow area above
point of attachment; ovary ovoid, glabrous at base, villous above middle and at
base of style; style punctate in lines, 5-6.5 mm. long, rather stout; placenta sub-
globose, small, apiculate; ovules 7 or 8, erect, uniseriate, tardily exposed apically;
fruits depressed-globose, black and shiny when ripe, drying up to 9 mm. in diameter.

Parathesis chiapensis Fernald, Proc. Amer. Acad. 36: 497.
1901. Uva; pish-match (Huehuetenango).

Wet mountain forest, often along rivers, 2,000 meters. or less;
Huehuetenango. Mexico (Chiapas, type from mountain forest be-
tween San Martin and Ococingo, C. & E. Seler 2226).

Arborescent shrub or small tree up to 10 m. high, branchlets rather stout,
densely red tomentose with rather fine stipitate or subsessile dendroid hairs; leaves
with petioles 5-18 mm. long, the petioles canaliculate, tomentose; leaf blades ob-
lanceolate, elliptic or lanceolate, 6-20 cm. long, 2.5-7 cm. wide, apex subabruptly
acuminate, base narrowed and decurrent on the petiole, chartaceous, at first stel-
late pubescent above, glabrescent, densely and uniformly pubescent on under-
surface with fine red, stipitate, stellate hairs, the rays of hairs comparatively short
in typical collections, margin closely denticulate, sometimes inconspicuously so or
entire at base, the costa elevated beneath, nearly plane above, the lateral veins
conspicuous on undersurface, slightly impressed above; inflorescences terminal,
paniculate, sometimes small and congested, shorter than leaves, sometimes broadly
pyramidal and up to 15 cm. long, puberulent and hirtellous or hirtellous-tomentose
with sessile or short-stipitate red, stellate hairs; flowers corymbose or subracemose-
corymbose; pedicels rather slender or sometimes clavate, 4-12 mm. long, sometimes
up to 17 mm. long, often reflexed; flowers at anthesis 5-7 mm. long, the calyx and
corolla densely black-punctate, the corolla in lines; sepals lanceolate, (1.2) 1.8-
3 mm. long, acuminate; petals narrowly lanceolate, 5-8 mm. long, papillose-
tomentose within apically and along margins; stamens 3-4 mm. long; filaments
glabrous, stout, 1.4-1.8 mm. long; anthers erect, ovate-lanceolate, (1.5) 2.2-2.8
mm. long, apiculate, dorsifixed below middle, with conspicuous black-punctate

166 FIELDIANA: BOTANY, VOLUME 24

area above attachment and extending down into lobes; ovary glabrous, or with a
few scattered erect hairs at apex and base of style; style slender, up to 5.5 mm.
long; placenta subglobose, apiculate; ovules 9, uniseriate, enclosed.

The species ranges north to the State of Mexico where it was col-
lected at Nanchititla (Matuda 30865). The flowers are sometimes
abnormal and small, possibly due to either disease or insect infestation.

Parathesis chrysophylla Lundell, Wrightia 3: 82, fig. 50. 1963.
Camaco (Guatemala); uva (British Honduras); cuya (Honduras).

Wet forest, 600 meters or less; type from Santa Ines, Guatemala,
C. Galusser 3. British Honduras; Honduras.

Tree, branchlets stout, closely tomentose with appressed matted hairs; leaves
large, with long marginate petioles 1-4 cm. long; leaf blades lanceolate, oblong-
elliptic or oblanceolate, usually 12.5-25 cm. long, sometimes up to 45 cm. long,
4.5-9 cm. wide, sometimes up to 14 cm. wide, apex acuminate, or subabruptly
short acuminate, base subacuminate and decurrent, entire but undulate, thinly
chartaceous, finely pubescent beneath with golden-brown fine closely appressed
stellate hairs, bizonal, the marginal zone glabrescent early, the midvein sulcate
above, prominently elevated beneath, the lateral veins very slender; inflorescences
terminal, pyramidal, paniculate, up to 30 cm. long, minutely rufous-tomentose,
the tomentum consisting of a densely papillose lower layer and an upper layer of
branched hairs, the glands conspicuous; pedicels up to 7 mm. long; flowers corym-
bose, finely rufous-tomentose and papillose, up to 6 mm. long at an thesis; sepals
triangular, spreading, 1-1.4 mm. long, acute or acuminate, often almost as wide
at base as long, punctate; petals lanceolate, about 6 mm. long, black-punctate in
lines, papillose-tomentose along edges, otherwise glabrous within; stamens up to
4.5 mm. long; filaments slender, black-punctate, up to 3.5 mm. long; anthers small,
versatile, dorsifixed medially, lanceolate-oblong, about 2 mm. long, conspicuously
black-punctate dorsally, the glands occasional in lobes; ovary tomentose, the hairs
erect; style slender, up to 5 mm. long, glabrous; placenta depressed-globose; ovules
small, 11-14, partly biseriate, partially exposed.

The tree is reported to reach a height of 60 feet and a diameter of
16 inches near Progreso in Honduras (fide Hottle). Leaves of vigor-
ous shoots reach a length of 18 inches and a width of about 6 inches,
the largest in the genus. The numerous partially biseriate ovules
suggest a linkage with Ardisia.

P. acuminata Lundell (Wrightia 3: 125. 1965), described from
El Salvador, is closely related to P. chrysophylla. It should occur
in Guatemala.

Parathesis columnaris Lundell, Wrightia 3: 62. 1963. Capulin
(Suchitepequez) ; cerezo silvestre (Retalhuleu) ; cinco negritos (Chiapas) ;
cerezin, cerecita (El Salvador).

FIG. 43. Parathesis columnaris. A, Branch with inflorescence; X K- B, Flower;
X 5. C, Calyx, spread out; X 5. D, Pistil; X 5. E, Leaf, undersurface showing
bizonal pubescence; X }/%. Illustrated from type, Steyermark 33205 (F).

167

168 FIELDIANA: BOTANY, VOLUME 24

Wet forest or thickets, usually in lowlands, 1,500 meters or less;
Amatitlan; Escuintla; Quezaltenango (type from lower slopes of Vol-
can Santa Maria, Finca Pirineos, J. A. Steyermark 33205) ; Retalhu-
leu; San Marcos; Santa Rosa; Suchitepequez. Mexico (Chiapas);
El Salvador.

Shrub or small tree, twigs sordidly tomentose with red stalked multibranched
dendroid hairs, often branched to base, glandular, appearing waxy; leaves with
slender canaliculate petioles up to 2 cm. long, glabrous above, tomentose beneath;
leaf blades oblanceolate, 10-18 cm. long, 2.5-5 cm. wide, apex subabruptly acumi-
nate, base attenuate and acuminate, decurrent on petiole, thin, chartaceous, pellu-
cid-punctate, entire or irregularly and obscurely crenulate, glabrous above, bizonal
beneath and pubescent with fine red appressed stellate hairs, densely so at first
along costa and in costal zone, glabrescent, the lateral veins fine but evident on
undersurface, obscure above; inflorescences terminal, paniculate, up to 20 cm. long,
equaling or exceeding leaves, red tomentose with short branched hairs, sparingly
so on branches; flowers corymbose, densely papillose; pedicels very slender, 4-9
mm. long, papillose; flower buds slender, up to 7 mm. long at anthesis; sepals
papillose-puberulent, narrowly triangular, 1-1.4 mm. long, acuminate, black-punc-
tate in lines; petals linear-lanceolate, united at base, up to 7 mm. long, attenuate-
acuminate, black-punctate in lines, short papillose-tomentose inside over entire
surface; stamens 3-4.5 mm. long; filaments slender, 1.3-1.7 mm. long, punctate;
anthers erect, linear-lanceolate, 2.3-3.5 mm. long, dorsifixed about one-fourth
above base, acutish, black-punctate in narrow band almost to apex and with glands
extending into lobes; ovary with a few scattered apical hairs, sometimes glabrous,
the style slender; placenta broadly ovoid, rounded; ovules 7 or 8, uniseriate, enclosed.

Parathesis cubana (A.DC.) Molinet & Gomez Maza, Dice. Bot.
11. 1889. Ardisia cubana A.DC. Trans. Linn. Soc. 17: 124. 1834;
Prodr. 8: 120. 1844. P. corymbosa Hemsl. Biol. Centr.-Amer. Bot. 2:
291. 1882 (Mexico, type from Sierra San Pedro Nolasco, Talea, & c.,
C. Juergensen 611). Tinus cubana (A.DC.) 0. Ktze, Rev. Gen. 2:
974. 1891. P. obovata Standl. Field Mus. Bot. 4: 250. 1929 (British
Honduras, type from Tower Hill, J. S. Karling 29). Ixpanol, ixpan-
panjul (Pete"n).

Wet thickets, savannas, pinelands, and swamps, common in second
growth, 200 meters or lower; Pete"n; Alta Verapaz; Izabal. Mexico
(Veracruz, Oaxaca, Tabasco, Campeche); British Honduras; Cuba.

Shrub or small tree up to 5 m. high, branchlets slender, at first ferruginous-
tomentulose with fine sessile appressed stellate hairs; leaves with slender petioles
0.5-2.2 cm. long; leaf blades elliptic, obovate-elliptic or obovate, 3.5-16 cm. long,
2-7 cm. wide, apex obtuse to acutish, base acute, chartaceous or sometimes thinner,
slightly paler beneath, at first pubescent on undersurface with fine appressed ses-
sile stellate ferruginous hairs, glabrescent early, entire, the midvein elevated be-
neath, the primary lateral veins slender; inflorescences usually terminal and leafy
below, rarely terminal and axillary on some branches, pyramidal, paniculate, 1- or

FIG. 44. Parathesis cubana. A, Flowering branch; X >£. B, Typical obo-
vateleaf;X^- C, Calyx, spread out; X 5. D, Flower; X 5. E, Pistil, pubescent
as illustrated or glabrous; X 5. F, Fruits; X 3.

169

170 FIELDIANA: BOTANY, VOLUME 24

2-branched, up to 20 cm. long, ferruginous-tomentulose with fine sessile stellate
hairs; pedicels slender, 2-7, sometimes 10 mm. long; flowers corymbose, 5- or
sometimes 6-merous, up to 7 mm. long at anthesis, the corolla pinkish, densely
and minutely papillose-tomentulose on outer surface; sepals ovate- triangular, 0.7-
1 mm. long, acutish, pellucid-punctate with orange glands; petals narrow, lanceo-
late-linear, 5-7 mm. long, densely papillose-tomentose over entire inner surface,
orange-red punctate in lines; stamens 3-5 mm. long; filaments thick, 1-2 mm. long,
orange-red punctate in lines; anthers erect, lanceolate-linear, 2-3.4 mm. long, at-
tached almost basally to filaments, the lobes small, sometimes vestigial, punctate
dorsally with orange-red glands which sometimes dry blackish, dehiscent with
apical pores; ovary ovoid, glabrous, or short villous apically; style glabrous, or
short villous at base, 3.5-5.5 mm. long; placenta depressed-ovoid, apiculate; ovules
small, numerous, 12-16, partially biseriate, tardily exposed apically; fruits de-
pressed-globose, up to 1 cm. diameter.

In cleared wet lowlands, this is one of the first invaders to ap-
pear. If such a shrub can be called weedy, the species belongs in
that category.

The anthers of P. cubana are atypical, but similar to those in
many species of Ardisia, and this characteristic together with the
numerous partially biseriate ovules indicate the affinity of Parathesis
to Ardisia through this species and several others in the genus.

Parathesis Donnell-Smithii Mez, Pflanzenreich IV. 236: 176.
1902. P. platyphylla Lundell, Wrightia 1 : 160. 1946 (type from Vaca,
El Cayo District, British Honduras, Percy H. Gentle 2615) . Palo de
uva (Huehuetenango).

Wet forest, sometimes in second growth, 500 meters or less; Alta
Verapaz (type from Cubilquitz, H. von Tuerckheim 7919) ; Huehuete-
nango; Chiquimula; Suchitepe"quez; Quiche". Mexico (Chiapas, Oax-
aca) ; British Honduras.

Arborescent shrub, branchlets stout, turgid and brittle when fresh, drying sub-
angular, very minutely appressed pubescent with stellate tawny hairs, glabrescent
and blackish-gray early; leaves with marginate petioles up to 2.5 cm. long; leaf
blades oblanceolate or elliptic, 10-25 cm. long, 3.5-9 cm. wide, apex subabruptly
long acuminate or acuminate, base acuminate, membranaceous or chartaceous,
bullate, obscurely crenulate and sometimes coarsely undulate-dentate, appearing
glabrous but at first with very fine closely appressed stellate hairs on undersurface,
completely glabrous very early, costa elevated beneath, plane or slightly impressed
above, the primary lateral veins slender, plane or slightly impressed above, incon-
spicuously reticulate; inflorescences fleshy and brittle, terminal, large, many-flow-
ered, tripinnately paniculate, oblongish or pyramidal, usually exceeding the leaves,
up to 30 cm. long, rather sparsely but minutely appressed pubescent with minute
stellate indument; floriferous rachis corymbose-racemose, accrescent, the racemes
at length up to 3.5 cm. long; pedicels pink, up to 8 mm. long; flowers 4-6 mm. long
at anthesis, the smaller flowers terminal, minutely ferruginous-tomentose; sepals

LUNDELL: FLORA OF GUATEMALA 171

0.8-1.7 mm. long, narrowly triangular, acuminate, black-punctate; petals pink,
linear-lanceolate, 4-6 mm. long, short papillose-tomentose within except at base,
black-punctate in lines; stamens 2-3.8 mm. long; filaments stout, 1-1.5 mm. long;
anthers erect, lanceolate-oblong, 1.5-2.5 mm. long, acutish, dorsifixed one-third
above base, with conspicuous acutely triangular elevated black-punctate dorsal
area, the glands large; ovary ovoid, ridged, finely tomentose apically; style 3-4.5
mm. long, pubescent at base; placenta minute; ovules 4-6, uniseriate, enclosed;
fruits depressed-globose, costate, punctate with small glands, acid at maturity,
red-black, drying 8 mm. in diameter.

The minute fine stellate tawny pubescence and corymbose-race-
mose flowers of the late flowering stage serve to distinguish P. Don-
nell-Smithii. Some plants have branchlets and leaves essentially
glabrous, even the young growth.

Para thesis guatemalensis Lundell, Wrightia 3: 65. 1963.
In pine-oak forest, 1,350 meters; Alta Verapaz (type from Coban,
H. von Tuerckheim 1202). Honduras.

Shrub, branchlets dull red, rather stout, at first densely ferruginous- tomentose
with fine appressed hairs, glabrescent at length and appearing lepidote; leaves with
marginate petioles up to 1.3 cm. long; leaf blades lanceolate or lanceolate-elliptic,
9-16 cm. long, 3-6.5 cm. wide, apex subabruptly acuminate, base cuneate and de-
current, firm and subcoriaceous, repand and denticulate or subentire, midvein
prominent beneath, the primary lateral veins conspicuous on undersurface, slightly
impressed above, upper surface somewhat rugulose, and at first stellate pubescent,
glabrescent very early except along costa, bizonal, the ferruginous pubescence of
fine short-rayed appressed stellate hairs, glabrescent, the marginal zone early; in-
florescences terminal, tripinnately paniculate, up to 12.5 cm. long, narrow, only
slightly wider at base, finely tomentose at first with ferruginous stellate hairs and
papillose-puberulent, glabrescent; pedicels 4-7 mm. long; flowers minutely ferrugi-
nous-tomentose, subcorymbose, up to 5 mm. long at anthesis; sepals lanceolate,
about 1.5 mm. long, acuminate, black-punctate; petals linear-lanceolate, 5 mm.
long, black-punctate with lines, villous within except at base; stamens 3.5-4 mm.
long; filaments about 1.4 mm. long; anthers erect, lanceolate, 2.7-3 mm. long,
acutish and bearing a few reddish hairs apically, dorsifixed one-third above base,
with conspicuous acutely triangular black-punctate area dorsally, the glands ex-
tending into lobes; ovary ovoid, attenuate apically into the style, conspicuously
hirtellous-tomentose along ridges; style slender, about 4 mm. long, with a few hairs
basally; ovules 5 or 6, uniseriate, enclosed; fruits subglobose, black, up to 8 mm.
diameter (dry).

Parathesis lanceolata Brandeg. Univ. Calf. Publ. Bot. 6: 188.
1915.

Damp barrancas; Mexico (Chiapas, type from Finca Irlanda, C. A.
Purpus 7371) ; doubtless extending into mountains of San Marcos.

Shrub or tree, twigs rather slender, minutely ferruginous-tomentose; leaves
with slender petioles up to 1 cm. long, ferruginous-tomentose beneath, canalicu-

172 FIELDIANA: BOTANY, VOLUME 24

late; leaf blades lanceolate or oblanceolate, 8-15 cm. long, 2-4.5 cm. wide, apex
and base acuminate, the apex subcaudate, the base decurrent on the petiole, thin,
paler beneath, entire or essentially so, pubescent beneath with small fine stellate
appressed reddish hairs, bizonal, glabrescent, pellucid-punctate, costa elevated be-
neath, the veins obscure; inflorescences terminal, laxly paniculate, the branches
very slender, pyramidal, up to 18 cm. long, exceeding the leaves, minutely ferrugi-
nous-tomentose; flowers small, corymbose or subracemose-corymbose, the florif-
erous rachis accrescent; pedicels slender, 2.5-4 (6) mm. long; flower buds ovoid,
very minutely ferruginous-tomentose, about 4 mm. long before anthesis, the calyx
and corolla black-punctate in lines; sepals narrowly triangular, 0.8-1 mm. long;
petals narrowly lanceolate, up to 4 mm. long, united at base, short papillose-
tomentose within except at base; stamens about 2.5 mm. long; filaments glabrous,
black-punctate, up to 1.4 mm. long; anthers erect, lanceolate-oblong, 1.5-1.8 mm.
long, dorsifixed one-third above base, with a conspicuous orange- or black-punctate
area dorsally, with glands sometimes extending into lobes, apex obtuse and minutely
apiculate; ovary glabrous, but style with a few scattered hairs at base, the style
up to 3.5 mm. long; placenta minute, subglobose; ovules 4 or 5, uniseriate, enclosed.

Para thesis latifolia Lundell, Wrightia 3: 83, fig. 51. 1963.

Cloud forest, at 1,500 meters; Mexico (Chiapas, type from Colo-
ma Tizcao, on the Guatemalan border, L. Irby Davis & Edgar Kincaid
55-50).

Shrub, branchlets stout, the indument of terminal buds and apical internodes
smooth and tightly appressed, tan in color; leaves with short thick petioles up to
1.5 cm. long, at first sparsely papillose above at base and along petiole, otherwise
glabrous; leaf blades obovate-elliptic, 15-25 cm. long, 7.5-9 cm. wide, apex abruptly
short acuminate, the acumen obtuse, base cuneate, decurrent, chartaceous, irregu-
larly crenate, costa plane above, prominent beneath, primary lateral veins very
slender; inflorescences axillary, paniculate, up to 10 cm. long including peduncle
3 cm. long, angled, 1- or 2-branched, compact, the secondary branches reduced,
sometimes only 1 mm. long, the indument minute, very closely appressed; pedicels
papillose-puberulent, 4-6 mm. long; flowers subcorymbose, 5 mm. long at anthesis,
finely papillose-tomentulose; sepals triangular, 1-1.5 mm. long, acute, red-punc-
tate; petals lanceolate, about 6 mm. long, papillose-tomentose on inner surface
except for glabrous area above base, papillose at base, punctate in lines; stamens
3-3.4 mm. long; filaments stout, glabrous, 1.5-1.7 mm. long; anthers erect, lanceo-
late, up to 2.5 mm. long, acute, punctate dorsally and in lobes with small red-black
glands, dorsifixed one- third above base; ovary ovoid, ribbed, finely tomentose over
entire surface and almost to middle of style, the apical hairs short villous and
longer; style 4 mm. long; placenta depressed-globose, apiculate, ovules 8 or 9, sub-
globose, uniseriate, enclosed.

Parathesis leptopa Lundell, Contr. Univ. Mich. Herb. 7: 41.
1942.

Mountain forest, 3,000 meters or less; Alta Verapaz; Chimalte-
nango; Santa Rosa. Mexico (Chiapas, type from Saxchanal, E. Ma-

LUNDELL: FLORA OF GUATEMALA 173

Shrub or small tree; branchlets rather stout, angled, ferruginous-tomentose
with sessile stellate appressed hairs often with some short dendroid hairs present;
leaves with petioles 1-3 cm. long; leaf blades lanceolate-elliptic or oblanceolate,
10-22 cm. long, 3-7 cm. wide, apex subabruptly acuminate, base acuminate and
decurrent, membranaceous to chartaceous, entire to obscurely crenulate, at first
sparsely papillose-puberulent above at base and along petiole, the young leaves
ferruginous-tomentose beneath with fine stellate subappressed stellate hairs, sparse
subappressed indument persisting to maturity, especially along midvein, glabres-
cent, costa slightly impressed above, elevated beneath, the primary lateral veins
9-26 on each side, slender, slightly impressed above; inflorescences axillary, some-
times appearing to be terminal with greatly reduced leaves in axils, panicles
long-pedunculate, 1- or 2-branched, many-flowered, 5-20 cm. long, ferruginous-
tomentulose, the small sessile hairs stellate, subappressed but often with erect or
spreading rays, sometimes papillose-puberulent; pedicels 3-7 mm. long; flowers 5-
or 6-parted, umbellate, densely papillose-puberulent, up to 7 mm. long at anthesis;
sepals triangular, 1.3-1.75 mm. long, acuminate, orange-red punctate; petals linear-
lanceolate, 5-7 mm. long, orange-red punctate in lines, papillose-tomentose on in-
ner surface; stamens 3-4.5 mm. long; filaments 1.75-3 mm. long, equaling or
slightly longer than anthers, punctate, united at base into a shallow ring; anthers
erect or sometimes appearing versatile, ovate-lanceolate or lanceolate, 1.75-2.7
mm. long, dorsifixed one-third above base, dorsally punctate with few small red-
dish-black glands, sometimes appearing eglandular; ovary finely tomentose almost
to base, the indument minute and appressed below, short villous above and at base
of style mostly with branched hairs, the ovary tapering into style, often slightly
ribbed; style about 5 mm. long; placenta subglobose, apiculate; ovules 5-9, rather
large, uniseriate, enclosed.

Parathesis macronema Bullock, Hooker's Ic. Plant. 34, t. 3397.
1939. Ardisia chiapensis Brandeg. Univ. Calif. Publ. Bot. 10: 413.
1924 (Chiapas, type from Hacienda Monserrate, C. A. Purpus 9273),
not Parathesis chiapensis Fernald, 1901.

Along streams, 1,000 meters or less. Mexico (Guerrero, type from
Banco, District of Montes de Oca, Geo. B. Hinton 10803; Chiapas).

Shrub or small tree, up to 4 m. tall, branchlets rather slender, at first ferrugi-
nous-tomentose with closely appressed hairs, glabrescent; leaves very variable in
size and shape, with slender petioles 5-20 mm. long; leaf blades oblong-elliptic,
lanceolate or oblanceolate, 5-15 cm. long, 1.2-5 cm. wide, apex obtuse to obtusely
acuminate, base cuneate, subentire to finely crenulate-serrulate, membranaceous
to subchartaceous, at first rather sparsely pubescent on lower surface with small
appressed stellate hairs, glabrous at maturity, costa elevated beneath, the primary
lateral veins very slender; inflorescences axillary, slender, lax, paniculate, 5-18 cm.
long, 1- or 2-branched, bracteate, with long slender tomentulose peduncles, the
branches sparsely puberulent; pedicels filiform, 5-17 mm. long, accrescent; flowers
corymbose or racemose-corymbose, minutely granular-puberulent, 4.5-6 mm. long
at anthesis; sepals ligulate, 0.6-1 mm. long, orange-punctate; petals triangular-
linear, 4.5-6.5 mm. long, densely papillose-tomentose over entire inner surface,
orange-punctate in lines; stamens 3.75-4.5 mm. long; filaments rather slender,
2.75-3.5 mm. long, papillose-puberulent below middle; anthers erect, linear-lanceo-

174

FIELDIANA: BOTANY, VOLUME 24

late, 1-1.3 mm. long, base sagittate, apex acuminate to cuspidate, usually with
several orange-colored glands dorsally, sometimes eglandular; ovary ovoid, gla-
brous; style slender, 3.5-5 mm. long, rather sparsely puberulent basally; placenta
subglobose, apiculate; ovules (6) 7 or 8 (10), uniseriate, enclosed or tardily exposed
apically; fruits globose or depressed-globose, up to 7 mm. in diameter.

The species is doubtfully distinct from Parathesis Rekoi Standl.
of Oaxaca. It should be found in the mountains of San Marcos.

• y

D

FIG. 45. Parathesis membranacea. A, Branch with inflorescence; X ^.
B, Calyx, spread out; X 5. C, Petal, showing inner surface; and stamen, dorsal
view; X 5. D, Pistil; X 5. Illustrated from isotype, Bartlett 13140 (MICH).

LUNDELL: FLORA OF GUATAMALA 175

Para thesis membranacea Lundell, Wrightia 1: 58. 1945.
Lowland forest; British Honduras (type from Tea Kettle, El Cayo
District, H. H. Bartlett 13140).

Branchlets rather stout, persistently brown tomentose, the hairs minute; leaves
with petioles 1-2 cm. long, canaliculate; leaf blades membranaceous, elliptic or
obovate-elliptic, 8.5-15 cm. long, 4.5-7.5 cm. wide, apex abruptly acuminate, base
decurrent on the petiole, stellate-pubescent on the undersurface at first, the fine
hairs appressed, essentially glabrous with age, glabrous on upper surface, black-
punctate, finely reticulate- veined, primary veins 12-15 on each side, conspicuous
on undersurface, plane or slightly impressed on upper surface, margin conspicu-
ously dentate almost to base; inflorescences leafy, terminal, narrowly paniculate,
finely tomentose, 8-10 cm. long; flowers corymbose, finely tomentose, borne on
pedicels up to 4 mm. long; buds 4 mm. long at anthesis; sepals narrowly triangular,
1.3-1.5 mm. long, acute; petals about 5.5 mm. long, narrowly lanceolate, attenuate
to the apex, punctate; filaments stout, about 1.3 mm. long; anthers ovate-lanceo-
late, 2.2 mm. long, apiculate, conspicuously black-punctate; ovary with a few hairs
at apex, glabrous otherwise; style 3.5 mm. long, glabrous.

P. oblanceolata closely resembles this species.

Para thesis oblanceolata Lundell, Wrightia 3: 67. 1963.

In wet forest, and along streams, mostly in lowlands, 300 meters
or less; Pete"n; Alta Verapaz (type collected along Rio Santa Isabel,
between mouth of Rio Sebol and El Porvenir, J. A. Steyermark 45853).

Shrub, branchlets slender and short to rather stout and elongated, ferruginous-
tomentose with fine sessile stellate hairs; leaves with rather slender petioles 5-15
mm. long; leaf blades oblanceolate, 7.5-20 cm. long, 2-5.5 cm. wide, apex caudate-
acuminate or acuminate, base acuminate and decurrent, entire, or inconspicuously
crenulate, chartaceous, punctate with small black glands, at first uniformly pubes-
cent beneath with fine sessile stellate ferruginous hairs, the rays spreading and sub-
appressed, glabrescent, midvein prominent beneath, slightly impressed above,
primary lateral veins elevated beneath, obscurely impressed above; inflorescences
terminal, tripinnately paniculate, the branches slender, sparsely pubescent, the
stellate spreading hairs with slender rays, and papillose-puberulent; pedicels rather
slender, 3-5 (7) mm. long; flowers corymbose, sparingly ferruginous-tomentose,
the buds very slender, about 4.5 mm. long at anthesis; sepals narrowly triangular,
1-1.4 mm. long, acuminate, black-punctate; petals linear-lanceolate, up to 5 mm.
long, papillose-tomentose within along margins and apically, black-punctate in
lines; stamens about 3 mm. long; filaments stout, epunctate, 1.5-1.8 mm. long;
anthers erect, narrowly lanceolate, about 2.2 mm. long, acutish, attached one-third
above base, with conspicuous black-punctate dorsal area extending into lobes;
ovary tapering into style, pubescent apically, usually with scattered short rather
stiff hairs; style about 4.5 mm. long, with a few hairs at base; ovules 6 or 7, uni-
seriate.

A fruiting specimen, Paul C. Standley 88357 (F) from Dept. Re-
talhuleu, may be referable here.

FIG. 46. Parathesis oblanceolata. A, Flowering branch with typical caudate-
acuminate leaves; X Yz- B, Inflorescence; X Yz- C, Calyx, spread out; X 5.
D, Petal, showing inner surface; and stamen, dorsal view; X 5. E, Pistil; X 5.
Illustrated from type, Steyermark 45853 (F), and isotype (LL).

176

B

FIG. 47. Parathesis papillosa. A, Branch with inflorescence; X %. B, Flower;
4-parted; anthers versatile; X 5. C, Pedicel and calyx; X 5. D, Pistil; X 5.
Illustrated from isotype, Tuerckheim 921 (US).

177

178 FIELDIANA: BOTANY, VOLUME 24

Para thesis papillosa Lundell, Wrightia 2: 69. 1963.

At 1,500 meters or less; Alta Verapaz (type from Pansamala,
H. von Tuerckheim 92) ; Huehuetenango.

Shrub or tree, branchlets rather stout, minutely lepidote; leaves large, long
petiolate, the petioles stout, up to 3 cm. long; leaf blades oblanceolate or oblanceo-
late-oblong, 12-25 cm. long, 3.5-6.5 cm. wide, apex subabruptly acuminate, base
acute, decurrent on petiole, entire, obscurely pellucid-punctate, subcoriaceous,
minutely pubescent with fine sessile, stellate, appressed hairs on undersurface,
bizonal, glabrescent, costa elevated beneath, the lateral veins fine, evident but not
conspicuous; inflorescences terminal, large, paniculate, up to 30 cm. long, 25 cm.
wide at base, densely papillose; flowers 4- or 5-parted, densely papillose, umbellate
or subcorymbose, the buds at an thesis pyriform, about 4 mm. long; pedicels 3-5
mm. long; calyx small, densely black-punctate; sepals ovate-triangular, about
1 mm. long, acuminate; petals lanceolate-linear, about 4 mm. long, united at base,
black-punctate in lines, papillose- tomentose within apically and along margins;
stamens up to 2.5 mm. long; anthers small, versatile, dorsifixed medially, lanceo-
late-oblong, 1.2-1.4 mm. long, minutely and sparingly black-punctate above point
of attachment of filaments, rarely eglandular; filaments slender, 2 mm. long, usu-
ally orange-punctate; ovary minutely papillose at apex, the style glabrous; pla-
centa obovoid; ovules 9-13, uniseriate or partly biseriate, partially exposed.

Para thesis pleurobotryosa Donn.-Sm. Bot. Gaz. 16: 195. 1891.

At 1,650 meters; Baja Verapaz (type from Santa Rosa, H. von
Tuerckheim 1442).

Branchlets thick, densely villous with coarse ferruginous stellate stipitate hairs;
leaves with marginate petioles 1-1.5 cm. long; leaf blades lanceolate or lanceolate-
elliptic, 7-13.5 cm. long, 2.5-4.5 cm. wide, apex acuminate, base acutish and de-
current, rather obscurely crenulate, subchartaceous, at first sparsely hirtellous on
upper surface, persistently pubescent on lower surface with red stipitate short-
rayed stellate hairs, the costa villous and elevated beneath, nearly plane above,
the primary lateral veins 12-15 on each side, rather conspicuous beneath, slightly
impressed above; inflorescences axillary, villous-tomentose with coarse red hairs,
the hairs of secondary branches shorter and less dense, panicles 1- or 2-branched,
5-15 cm. long, long-pedunculate; pedicels slender, 5-12 mm. long, villous-hirtel-
lous; flowers umbellate, papillose-tomentose, about 5 mm. long at anthesis; sepals
lanceolate-triangular, 1.2-1.8 mm. long, acuminate or cuspidate, punctate, papil-
lose-hirtellous; petals narrowly lanceolate, 5.5-6.5 mm. long, punctate in lines,
papillose-tomentose on inner surface along edges and above middle, glabrous be-
low; stamens about 3 mm. long; filaments thick, glabrous, 1.2-1.4 mm. long;
anthers ovate, 1.6-2 mm. long, thick, dorsifixed about one-third above base, apicu-
late, sparingly black-punctate dorsally, the area grooved and narrowly triangular,
sometimes apically barbate dorsally, the basal lobes rounded; ovary ovoid, gla-
brous below, short villous-hirtellous apically; style slender, 4.5-6 mm. long, glabrous;
placenta subglobose, apiculate; ovules 5-7, uniseriate, enclosed.

Para thesis reflexa Brandeg. Univ. Calif. Publ. Bot. 6: 189. 1915.
Lichui (Chimaltenango) .

LUNDELL: FLORA OF GUATEMALA 179

Wet forests of barrancas and mountain sides, 3,000 meters or
lower; Santa Rosa; Chimaltenango; San Marcos. Mexico (Chiapas,
type from barranca near Finca Irlanda, C. A. Purpus 7334).

Shrub or small tree, branchlets rather stout, loosely ferruginous-tomentose
with dendroid hairs; leaves with petioles 1-2.5 cm. long; leaf blades narrowly ellip-
tic, lanceolate or oblanceolate, 10-23 cm. long, 3-7.5 cm. wide, apex subabruptly
acuminate, base acuminate and decurrent, membranaceous to chartaceous, paler
beneath, entire to crenulate, at first pubescent on upper surface at base and along
petiole, at first floccose-tomentose beneath with dendroid ferruginous hairs, these
sometimes somewhat appressed marginally, glabrescent except along the prominent
midvein, the primary lateral veins slender, slightly impressed above; inflorescences
axillary, the terminal leaves sometimes reduced and bract-like, 2- or 3-paniculate,
long-pedunculate, minutely floccose-tomentulose, with ferruginous dendroid hairs,
pyramidal, 8-25 cm. long; pedicels slender, 5-12 mm. long; flowers 5- or rarely 6-
merous, umbellate, minutely papillose-tomentulose, up to 5 mm. long at anthesis;
sepals narrowly triangular, up to 1.5 mm. long, acuminate, densely punctate, papil-
lose-puberulent; petals lanceolate, 5 mm. long, papillose-tomentose over entire in-
ner surface, black-punctate in lines; stamens 2.75-3 mm. long; filaments glabrous,
sometimes black-punctate, 1.5-2 mm. long; anthers erect, slender, lanceolate, about
2 mm. long, acutish, dorsifixed one-third above base, inconspicuously punctate
dorsally with minute orange-black glands, these usually dispersed, occasionally in
lobes; ovary ovoid, tomentose over entire surface to base, the hairs above and at
base of style multibranched or simple; style about 4.5 mm. long; placenta subglo-
bose, apiculate; ovules small, 8-10, uniseriate, enclosed; fruits subglobose, up to
1 cm. in diameter when ripe.

Parathesis rufa Lundell, Wrightia 3: 73. 1963.

Wet forests and riverbanks, sea level to 1,200 meters; British
Honduras (type from Rio Grande, Botan Creek, Toledo District,
Percy H. Gentle 4649). Mexico (Chiapas).

Arborescent shrub or small tree up to 17 cm. in diameter, branchlets stout, to-
mentose, with fine rather loose red multibranched dendroid hairs; leaves with stout
marginate petioles up to 2 cm. long, the petioles tomentose beneath, stellate pubes-
cent above at first with appressed hairs; leaf blades oblong-elliptic, oblanceolate or
obovate, 12-27 cm. long, 5-9 cm. wide, apex subabruptly acuminate or caudate-
acuminate, base acute and decurrent, thin at first, becoming subcoriaceous and
rugose at maturity, obscurely bizonal, at first pubescent above with stellate hairs,
especially along midvein, densely pubescent on undersurface with small fine sessile
stellate hairs, the hairs spreading or subappressed in costal zone, usually appressed
in marginal zone, glabrescent, the margin usually conspicuously crenulate, costa
and lateral veins prominent beneath, slightly impressed above; inflorescences red,
terminal, pyramidal, tripinnately paniculate, up to 22 cm. long, papillose-puberu-
lent and tomentulose with red loose or multibranched dendroid hairs; pedicels
3-5 mm. long; flowers subcorymbose, about 5 mm. long at anthesis, papillose-
puberulent and finely stellate tomentulose; sepals narrowly triangular, 1.75-2.2
mm. long, subulate; petals narrowly lanceolate, about 5.5 mm. long, papillose-
tomentose within except at base, punctate in lines; stamens about 3 mm. long;

FIG. 48. Parathesis rufa. A, Branch with inflorescence; X %. B, Flower;
X 5. C, Calyx, opened out; X 5. D, Pistil; X 5. Illustrated from type, Gentle
4649 (LL).

180

FIG. 49. Parathesis sessilifolia. A, Flowering branch; X J^. B, Flower;
X 5. C, Calyx, spread out; X 5. D, Stamen, dorsal view; X 5. E, Pistil; X 5.
F, Hair from stem, greatly enlarged. Illustrated from type, Tuerckheim 1443

(US).

181

182 FIELDIANA: BOTANY, VOLUME 24

filaments stout, glabrous, about 1.3 mm. long; anthers erect, dorsifixed one-third
above base, lanceolate-oblong, about 2.4 mm. long, with conspicuous triangular
black-punctate area above attachment, the large glands extending into lobes; an-
thers conspicuously acute-apiculate; ovary ovoid, glabrous, or sparsely hirtellous
and papillose apically and at base of style; style 4.5 mm. long; placenta columnar
or obovoid; ovules 6 or 7, elliptic-oblong, uniseriate, enclosed; fruits black, de-
pressed-globose, up to 8 mm. in diameter when dry.

Parathesis sessilifolia Donn.-Sm. Bot. Gaz. 16: 195. 1891.

Wet forests, sea level to 1,500 meters; Pete"n; Alta Verapaz (type
from Coban, H. von Tuerckheim 1443); Izabal. Mexico (Tabasco);
British Honduras.

Arborescent shrub or tree up to 17.5 cm. in diameter, 12 m. high, branchlets
rather stout, densely villous-tomentose with long stipitate, dark red, shaggy den-
droid hairs, with lower layer of stellate hairs, papillose; leaves with thick petioles up
to 1.5 cm. long, sometimes appearing subsessile due to decurrent and marginate leaf
blades, the petioles villous-tomentose, glabrescent above; leaf blades oblanceolate,
elliptic or obovate-elliptic, 10-25 cm. long, 3.5-7.5 cm. wide, apex acuminate, the
acumen often caudate and falcate, base cuneate and decurrent on petiole, charta-
ceous, nigro-punctate, glabrous above, uniformly pubescent over undersurface with
red stipitate dendroid hairs, the margin subcrenulate, costa prominent beneath,
nearly plane above, the lateral veins slender but conspicuous on undersurface,
slightly impressed above; inflorescences terminal, paniculate, usually large and
broadly pyramidal, up to 22 cm. long, sometimes small and shorter than leaves,
red villous with shaggy stipitate stellate hairs and papillose-puberulent; pedicels
slender, 3-7 mm. long; flowers corymbose or subracemose-corymbose, 5-7 mm.
long at an thesis; the calyx and corolla bristly tomentose with stellate hairs and
papillose-puberulent; sepals narrowly triangular-linear, 1.5-2 mm. long, attenuate-
acuminate, black-punctate; petals black-punctate in lines, united at base, linear-
lanceolate, up to 7 mm. long, acutish, short papillose-tomentose along margins and
inside above middle, otherwise glabrous; stamens up to 3.5 mm. long; filaments
glabrous, up to 1.8 mm. long; anthers erect, dorsifixed slightly below middle, lan-
ceolate-oblong, 2.2-2.8 mm. long, acutish, with a narrowly triangular conspicuous
black-punctate area above point of attachment and extending into lobes; ovary at
apex and base of style with a few scattered erect hairs; style up to 5 mm. long;
placenta with 5 or 6 ovules; ovules uniseriate, enclosed; fruits globose, about 7 mm.
in diameter, densely black-punctate.

Parathesis Skutchii Lundell, Contr. Univ. Mich. Herb. 6: 54.
1941.

Cloud forest, 2,500 meters or less; Quiche" (type from Nebaj, A. F.
Skutch 1681); Alta Verapaz; Huehuetenango.

Shrub or tree, up to 10 m. high, branchlets angled, stout, the indument of ter-
minal buds and apical internodes smooth and closely appressed, tan in color, the
branchlets at length minutely appressed-lepidote; leaves with petioles 1.5-2.5 cm.
long; leaf blades papillose above at base and along petiole, oblong-lanceolate or

FIG. 50. Parathesis Skutchii. A, Flowering branch; X 1A- B, Flower, show-
ing epunctate anthers, and petals papillose-tomentose over entire inner surface;
X 5. C, Calyx, spread out; X 5. D, Pistil; X 5. Illustrated from type, Skutch
1681 (F).

183

184 FIELDIANA: BOTANY, VOLUME 24

oblanceolate, 12-25 cm. long, 2.5-7 cm. wide, apex acuminate, base cuneate and
decurrent, thinly chartaceous, finely crenulate to subentire, thinly stellate pubes-
cent beneath or glabrous, costa slightly impressed above, prominent beneath, the
primary lateral veins up to 25 on each side, slender; inflorescences axillary, panicu-
late, up to 20 cm. long, angled, short pedunculate, divaricately 2- or 3-branched,
compact, the secondary branches reduced, the indument minute, very closely ap-
pressed; pedicels minutely papillose, appearing pruinose, 3-5 mm. long; flowers
umbellate to short racemose-corymbose, minutely papillose-puberulent, 3-4 mm.
long at an thesis, the buds conspicuously angled; sepals minute, triangular, 0.5-
0.75 mm. long, obtuse, punctate; petals lanceolate, 3-4 mm. long, densely papillose-
tomentose over entire inner surface, punctate in lines; stamens 2-2.5 mm. long;
filaments stout, up to 1.4 mm. long, epunctate; anthers erect, ovate-lanceolate, up
to 1.75 mm. long, dorsifixed about one-third above base, acute, concolorous or with
1 to several small black glands dorsally; ovary depressed-ovoid, minutely tomen-
tose over entire surface, apically the hairs slightly longer, basal third of style
minutely tomentose; style short, about 2.5 mm. long, punctate in lines; placenta
subglobose or strongly depressed-globose, small, minutely apiculate; ovules 7-11,
rather small, crowded, uniseriate, enclosed.

Parathesis stenophylla Lundell, Wrightia 3: 86, fig. 53. 1963.
Wet forest; Quiche" (known only from the type, Jose Ignacio Agui-
Zar793).

Branchlets rather slender, ferruginous-tomentose with fine closely appressed
stellate hairs, the rusty tomentum persistent; leaves with petioles up to 2 cm. long;
leaf blades linear-lanceolate, 10-30 cm. long, 2.5-5 cm. wide, apex acuminate, base
acuminate and decurrent, chartaceous, crenulate-denticulate, at first ferruginous-
pubescent on lower surface with closely appressed fine stellate hairs, glabrescent,
papillose above at base and along petiole, the costa prominent beneath, plane
above, the primary lateral veins slender; inflorescences axillary, with the apical
leaves reduced, long-pedunculate, lax and few-flowered, 1- or 2-branched, up to
23 cm. long, finely tomentulose with rufous stellate hairs; pedicels 4-8 mm. long;
flowers corymbose, 5 mm. long at anthesis, finely tomentulose; sepals narrowly
triangular, about 1.5 mm. long, acuminate, black-punctate; petals linear-lanceo-
late, 5-6 mm. long, black-punctate in lines, papillose-tomentose on inner surface
except for area above base; stamens 3-3.4 mm. long; filaments glabrous, up to 2
mm. long, punctate; anthers erect, lanceolate, up to 2.7 mm. long, dorsifixed one-
third above base, black-punctate dorsally, the glands extending into lobes; ovary
ovoid, black-punctate, glabrous below, short villous-pubescent apically and at base
of style, glabrescent; style about 5 mm. long; placenta obovoid, acicular-apiculate;
ovules 8, obovoid or ellipsoid, uniseriate, enclosed.

Parathesis subcoriacea Lundell, Wrightia 2: 73. 1963.

In forest, 1,950 meters; Mexico (Chiapas, type from Escuintla,

Small tree, 3-4 m. high, the twigs rather stout, minutely lepidote, drying
brownish; leaves long petiolate, the petioles up to 2 cm. long, obscurely and mi-
nutely lepidote beneath, canaliculate; leaf blades lanceolate, 8-15 cm. long, 2.5-

B

FIG. 51. Parathesis subcoriacea. A, Flowering branch; X ]/z- B, Flower;
X 5. C, Calyx, spread out; X 5. D, Pistil; X 5. Illustrated from type, Matuda
5985 (LL).

185

186 FIELDIANA: BOTANY, VOLUME 24

5 cm. wide, apex acuminate, the acumen obtusish, base cuneate and decurrent on
petiole, subcoriaceous, paler beneath, obscurely pellucid-punctate with orange
glands, essentially entire, the young leaves with minute closely appressed stellate
hairs, glabrescent early, mature leaves entirely glabrous, costa elevated beneath,
the lateral veins fine but evident on lower surface, obscure above; inflorescences
axillary or terminal, subequaling the leaves, obscurely and sparingly lepidote-
papillose, congested; flowers minutely papillose, in few-flowered umbels, the um-
bels congested; pedicels slender, 2.5-3 mm. long, minutely papillose; flower buds
at anthesis ovoid, 3 mm. long, the calyx and corolla black-punctate in lines; sepals
minute, narrowly triangular, 0.8-1 mm. long, acuminate; petals narrowly triangu-
lar, 3 mm. long, short papillose-tomentose within except at base; stamens 2 mm.
long, filaments slender, glabrous, epunctate, about 1 mm. long; anthers erect,
epunctate, dorsifixed basally, lanceolate, acutish, 1.2-1.4 mm. long; ovary gla-
brous; style about 3 mm. long, glabrous; placenta depressed-ovoid, apiculate,
ovules 10-12, very small, uniseriate.

Parathesis subulata Lundell, Wrightia 3: 74. 1963.

In forest, 1,500-2,000 meters; Huehuetenango (type from Cerro
Victoria near Barillas, Sierra de los Cuchumatanes, J. A. Steyermark
49714).

Shrub, up to 3 m. high, branchlets slender, at first sparsely appressed tomen-
tulose, glabrescent; leaves with slender petioles 5-15 mm. long; leaf blades lanceo-
late-oblong, 5-9.5 cm. long, 1.2-2.3 cm. wide, apex caudate-acuminate, base
acuminate, glabrous, membranaceous, slightly paler beneath, obscurely and mi-
nutely crenulate, pellucid-punctate, costa slender and elevated beneath, plane
above, primary lateral veins slender, 12-15 on each side; inflorescences axillary,
few-flowered, paniculate, 3-6 cm. long, usually 1-branched, glabrous or essentially
so, the peduncles slender but firmly erect; bractlets foliaceous, up to 7 mm. long;
pedicels slender, 3-6 mm. long; flowers corymbose; sepals subulate, about 0.75 mm.
long, sparsely and minutely puberulent, orange-punctate; corolla and stamens
unknown; ovary ovoid, minutely appressed tomentulose; style slender, 3-4 mm.
long, puberulent basally; placenta depressed-globose; ovules 7 or 8, uniseriate,
exposed apically; young fruits globose.

Parathesis tartarea Lundell, Wrightia 3 : 86, fig. 5|. 1963. Sapo-
tilla (San Marcos).

Wet mountain forest of upper slopes, 3,900 meters or less; Quiche";
San Marcos (type from Volcan Tacana, J. A. Steyermark 36023);
Solola; Suchitepe"quez. Mexico (Chiapas).

Tree, up to 16 m. high, branchlets thick, scaly with epidermis peeling trans-
versely and irregularly, the terminal buds and apical internodes densely ferrugi-
nous-tomentose with minute appressed sessile stellate hairs; leaves with rather
stout marginate petioles 2-5 cm. long; leaf blades lanceolate to oblong-elliptic,
10-20 cm. long, 2.5-7 cm. wide, apex acuminate or subabruptly acuminate, base
rounded and acutish or acute, coriaceous, paler beneath, denticulate, the teeth
usually conspicuous but small, rarely obscure, glabrescent above, persistently and
minutely pubescent on lower surface with very fine small closely appressed stellate

LUNDELL: FLORA OF GUATEMALA 187

hairs, the midvein prominent beneath, slightly impressed above, the primary lat-
eral veins very slender; inflorescences axillary, paniculate, 5-15 cm. long, 1- or
2-branched, the peduncle and branches thick and angled, minutely ferruginous-
tomentose with minute appressed hairs; pedicels rather stout, drying wrinkled,
5-10 mm. long; flowers corymbose, minutely ferruginous- tomentose, 5-6 mm. long
at anthesis; sepals thick, triangular, 1.2-1.6 mm. long, acute, inconspicuously and
sparsely orange-punctate; petals thick, linear-lanceolate, about 6 mm. long, papil-
lose-tomentose over entire inner surface, inconspicuously orange-punctate in lines;
stamens up to 3.75 mm. long; filaments thick, united at base, about 1.4 mm. long,
orange-punctate; anthers erect, lanceolate, up to 2.75 mm. long, inconspicuously
punctate with small pale orange colored glands; ovary ovoid, glabrous except for
minute tomentum apically and at base of style, tapering into the thick style; style
5-6 mm. long; placenta obovoid, apiculate; ovules 9, uniseriate, enclosed; fruits
subglobose.

Para thesis tomentosa Lundell, Wrightia 3 : 74. 1963. Chimiche.
Endemic; known only from the type; Los Andes to Entre Rios,
Izabal, S. J. Record G.44.

Small tree, branchlets slender, minutely appressed tomentulose at first, gla-
brescent; leaves with slender petioles 5-14 mm. long; leaf blades obovate or obo-
vate-elliptic, 5-9.5 cm. long, 3.5-4.5 cm. wide, apex rounded and abruptly short
acuminate, base cuneate, membranaceous, very sparsely and minutely stellate-
lepidote on lower surface at first, essentially glabrous, pellucid-punctate, finely
crenulate above, costa elevated beneath, plane above, primary lateral veins slen-
der, 8 or 9 on each side, prominent beneath; inflorescences terminal, paniculate,
up to 5 cm. long, 1- or 2-branched to base, the branches very slender, few-flowered,
sparsely puberulent; pedicels filiform, 5-9 mm. long, sparsely and minutely puberu-
lent; flowers corymbose; sepals narrowly triangular, 1-1.2 mm. long, subulate,
pellucid-punctate, puberulent; petals linear-lanceolate, 3.5 mm. long, sparingly
and minutely puberulent on outer surface, densely papillose-tomentose over entire
inner surface, orange-punctate; ovary ovoid, tapering into style, densely tomentose
over entire surface; style about 2.75 mm. long, tomentose to middle; placenta de-
pressed-globose, short apiculate; ovules 8 or 9, uniseriate, apparently enclosed.

Para thesis vestita Lundell, Wrightia 3: 75. 1963. Hueso bianco,
huesito bianco (Quezaltenango) .

Wet forest, mostly upper mountain slopes, 1,200-3,800 meters;
Chimaltenango; Quezaltenango (type from Finca Pirineos, P. C.
Standley 68219); San Marcos; Solola; Suchitepe"quez.

Shrub or tree, up to 10 m. high; branchlets rather stout, densely villous-tomen-
tose with long red mostly simple hairs; leaves with petioles 5-15 mm. long; leaf
blades oblanceolate, oblanceolate-elliptic or obovate-elliptic, 10-30 cm. long, 3-10
cm. wide, apex subabruptly acuminate, base acutish to acuminate, decurrent,
membranaceous, conspicuously dentate with numerous small sharp regular teeth
or sometimes bidentate with larger irregular teeth, persistently hirtellous on upper
surface, long villous along midvein beneath, the lower surface rather thinly and

FIG. 52. Parathesis tomentosa. A, Flowering branchlets; X %. B, Calyx
spread out; X 5. C, Petal, inner surface; X 5. D, Pistil, showing densely tomen-
tose ovary and base of style; X 5. Illustrated from type, Record G.44 (US).

188

LUNDELL: FLORA OF GUATEMALA 189

persistently villous with simple hairs, or long stipitate hairs, these bifid or stellate
tipped, the small terminal rays of hairs small and inconspicuous, the midvein prom-
inent beneath, impressed above, the primary lateral veins slender on undersurface,
plane to slightly impressed above; inflorescences axillary, long-pedunculate, py-
ramidal and lax, 10-25 cm. long, villous to short villous, dull red; pedicels slender,
hirtellous-villous, 6-12 mm. long; flowers white, corymbose or racemose-corymbose,
up to 7 mm. long at anthesis, papillose-tomentose; sepals lanceolate-triangular or
triangular, 1.4-2 mm. long, acuminate or cuspidate, punctate; petals linear-lanceo-
late, 6-7 mm. long, papillose-tomentose on inner surface except at base, punctate
in lines; stamens up to 3.5 mm. long; filaments stout, glabrous, 1.4-1.5 mm. long,
punctate; anthers erect, dorsifixed about one-fourth above base, ovate or ovate-
lanceolate, 2-2.5 mm. long, thick, obtusish, with grooved black-punctate dorsal
area; ovary ovoid, short villous-tomentose almost to base; style slender, 5.5-6.5
mm. long, short villous basally; placenta subglobose, apiculate; ovules 6 or 7, erect,
uniseriate, enclosed; fruits black at maturity, depressed-globose, drying 7-9 mm.
in diameter.

Para thesis vulgata Lundell, Wrightia 3: 88. 1963. Mora pava
(Jalapa), Cerecil de montana, sirasil de montana (Chiquimula) .

Wet or mixed forest, mostly middle and upper mountain slopes,
often in cloud forest, 1,200-2,500 meters; Baja Verapaz; Chiquimula;
Jalapa; Jutiapa; Zacapa. Honduras (type from Mt. Uyuca, S. F.
Glassman 1616).

Shrub or small tree up to 8 m. high, branchlets rather stout, closely appressed
tomentose with fine sessile stellate hairs; leaves with petioles mostly less than 1 cm.
long, sometimes up to 1.5 cm. long; leaf blades oblanceolate, oblanceolate-elliptic
or oblong, 7-19 cm. long, 3-6.5 cm. wide, apex subabruptly acuminate, base acumi-
nate and decurrent, membranaceous, paler beneath, subentire or crenulate, at first
pubescent on lower surface with fine closely appressed stellate hairs, glabrescent
early, appearing glabrous, densely black-punctate, costa nearly plane above, ele-
vated beneath, the primary lateral veins slender; inflorescences axillary, long pe-
dunculate, paniculate, the panicles 1- or 2-branched, 6-18 cm. long, minutely
papillose-puberulent with some longer hairs intermixed; pedicels usually 5-8, rarely
10 mm. long; flowers corymbose, densely papillose-puberulent, 5-6 mm. long at
anthesis; sepals ovate-triangular, 1.4-2 mm. long, acuminate, red-black punctate;
petals narrowly lanceolate, 5-7 mm. long, punctate with red-black lines, papillose-
tomentose on inner surface except at base, with a papillose ring at base; stamens
3-3.5 cm. long; filaments stout, up to 1.5 mm. long, punctate; anthers thick, ovate-
lanceolate, 2-2.5 mm. long, apiculate, with conspicuous black-punctate area dor-
sally, the glands often extending almost to apex, dorsifixed one-third above base;
ovary ovoid, short villous apically to villous-tomentose, short villous at base of
style; style 5-7 mm. long, punctate in lines; placenta subglobose, apiculate; ovules
6-9, erect, uniseriate, enclosed; ripe fruits wine-red, depressed-globose, about 1 cm.
in diameter.

The collections from Guatemala are unsatisfactory, either sterile,
in flower bud or fruit, but they appear to be referable to P. vulgata.
Evidently the species is common in cloud forest.

190

FIELDIANA: BOTANY, VOLUME 24
RAPANEA Aublet

Shrubs or trees, glabrous or pubescent; leaves petiolate, entire or nearly so;
flowers small, dioecious, 4-5-merous, lateral or axillary, appearing fasciculate by
reduction of inflorescence, bracteolate; sepals small, usually connate at base, im-
bricate or valvate, ovate or triangular, often ciliolate, usually punctate or puncticu-
late; petals connate below, spreading or recurved, usually lineate, often papillose
on the margins; stamens inserted in throat of corolla, the filaments obsolete; an-

FIG. 53. Rapanea myricoides. A, Flowering branchlet; X 1A,. B, Pistillate
flower, opened out showing abortive anthers; X 10. C, Incrassate sepals, and
ovary of pistillate flower, showing large, sessile, morchelliform stigma; X 10.

LUNDELL: FLORA OF GUATEMALA 191

thers sessile, dehiscent by introrse slits, abortive in pistillate flowers; ovary globose
or ellipsoid; stigma sessile in pistillate flowers, large, subcapitate, morchelliform
or lobed; ovules few, uniseriate; fruit 1-seeded, dry or fleshy, the endocarp crusta-
ceous to ligneous; seed globose, smooth, intruded at the base, the endosperm corne-
ous, sometimes slightly ruminate; embryo elongate, transverse, usually curved.

About 140 species, in the tropics of both hemispheres. Other spe-
cies are found in southern Central America.

Young branchlets and petioles sparsely villous with reddish hairs; leaves narrowly
lanceolate R. myricoides.

Young branchlets glabrous; leaves mostly obovate-elliptic or elliptic.

Leaves rounded at apex; stigma of pistillate flowers subcapitate and shallowly
lobed, less than 1 mm. long; in lowland forests R. guianensis.

Leaves narrowed above and broadly obtuse at apex; stigma of pistillate flowers
deeply sinuate-lobed, up to 2.5 mm. long; in wet forest of high mountains.

R. Juergensenii.

Rapanea myricoides (Schlecht.) Lundell, Wrightia 3: 109. 1964.
Myrsine myricoides Schlecht. Linnaea 8: 525. 1833. Myrsine guate-
malensis Gandoger, Bull. Soc. Bot. Fr. 65: 57. 1918 (type from Coban,
Alta Verapaz, H. von Tuerckheim 1001). Array&n (Alta Verapaz);
paln& (Huehuetenango) .

Moist to wet or dry thickets or mixed forest, often in pine-oak
or Liquidambar forest, sometimes in swamps, often in second
growth, 1,000-2,700 meters; Alta Verapaz; Bajo Verapaz; Chimalte-
nango; Chiquimula; Guatemala; Jalapa; Jutiapa; Huehuetenango;
El Progreso; Quezaltenango; Quiche"; Sacatepe"quez; San Marcos;
Santa Rosa; Solola; Zacapa. Mexico; El Salvador and Honduras
to Panama.

Usually a shrub of 2-3 meters, sometimes a small tree up to 10 meters, branch-
lets sparsely villous to densely villous-tomentose with reddish hairs, rarely glabrous,
often densely leafy; leaves usually short-petiolate, the petioles sometimes up to
12 mm. long, rarely longer; leaf blades lanceolate or oblong-lanceolate, mostly
6-13 cm. long, 1-2.5 cm. wide, acute, acuminate or obtuse, attenuate to the base,
chartaceous or subcoriaceous, paler and usually silver-green beneath, usually vil-
lous at first, especially along the midvein on both surfaces, glabrescent, entire; in-
florescences 3-9-flowered, glomerate, arising in the leaf axils or from defoliate
nodes; pedicels 1-1.5 mm. long; flowers dioecious, 2-3.5 mm. long, glabrous or
nearly so; calyx lobes triangular-ovate, less than 1 mm. long, acute or subacute,
more or less black-punctate; petals connate one-third at base, subacute to rounded
at apex, black-punctate; anthers sessile, those of pistillate flowers abortive, less
than 1 mm. long; anthers in staminate flowers shorter than petals, 1.5-1.8 mm.
long, attached at apex of corolla tube; ovary abortive in staminate flowers; ovary
subglobose in pistillate flowers, with large conical morchelliform stigma; ovules 3,
uniseriate, imbedded in fleshy placenta; fruit blackish at maturity, glabrous, 2.5-
3.5 mm. in diameter.

192 FIELDIANA: BOTANY, VOLUME 24

This shrub, long known from our area as R. ferruginea (Ruiz &
Pavon) Mez, a species of South America, much resembles Myrica in
habit and general appearance, and frequently is associated with it.
Abundant in some regions, especially in the mountains of Alta Vera-
paz, it is a characteristic element of the vegetation.

Rapanea guianensis Aubl. PI. Guian. 121. t. 1*6. 1775.

Usually in or near mangrove swamps, common in tintal, 200 me-
ters or less; Izabal; Pete"n. Southern Florida; southern Mexico;
British Honduras south to South America; West Indies.

A glabrous shrub or small tree, 9 meters high or less, the trunk 15 cm. or less
in diameter; leaves with short petioles up to 7 mm. long; leaf blades oblong-obovate,
oblong-elliptic, or oblanceolate, 4-10 cm. long, 2.5-4 cm. wide, apex rounded or
obtuse-rounded, cuneately narrowed to the base, lustrous above, somewhat paler
beneath, entire, coriaceous; inflorescences shorter than petioles, 3-7-flowered, the
flowers glomerate mostly on very short lateral spur-like branches; pedicels 1.5 mm.
long or less; flowers dioecious, glabrous, 2-2.5 mm. long, greenish, punctate; sepals
connate at base, ovate or ovate-lanceolate, less than 1 mm. long, puncticulate;
petals short-connate, elliptic, up to 2.5 mm. long, puncticulate; anthers included,
sessile, attached at apex of corolla tube, up to 1.5 mm. long in staminate flowers,
apiculate, epunctate, abortive in pistillate flowers; ovary subglobose; stigma in
pistillate flowers thick, and subcapitate, irregularly short-lobed; ovules 2 or 3;
immature fruit pellucid-punctate, black and globose at maturity, about 4 mm. in
diameter, smooth.

Rapanea Juergensenii Mez, Pflanzenreich IV. 236: 388. 1902.

Wet mixed mountain forest, 1,300-3,000 meters; El Progreso;
Guatemala; Zacapa; Chiquimula; Jalapa; Solola; Huehuetenango;
Quezaltenango; San Marcos. Western Mexico; Honduras.

A large shrub or a tree of 6-12 meters, glabrous throughout, the branches
thick; leaves with marginate petioles up to 1 cm. long; leaf blades narrowly ellip-
tic, oblanceolate or oblong-elliptic, 6-12 cm. long, mostly 3-4 cm. wide, subacute
to obtuse at apex, cuneately narrowed to base, subcoriaceous, paler beneath; in-
florescences 5-9-flowered, shorter than petioles; flowers dioecious, glabrous, green-
ish, 2-2.75 mm. long, puncticulate; sepals connate at base, ovate, about 1 mm.
long, thin; petals short-connate, elliptic, about 2.75 mm. long in pistillate flowers;
anthers included; ovary subglobose; stigma large, capitate, sinuate-lobed, up to
2 mm. long; immature fruit punctate with large pellucid glands.

STYLOGYNE A. DeCandolle

Shrubs or trees, glabrous or nearly so; leaves alternate, petiolate; inflorescences
paniculate, terminal or axillary; flowers dioecious (in Guatemala), small, umbellate
or subcorymbose, pedicellate, usually white, usually 5-parted, rarely 4-parted;
sepals dextrorsely contorted in bud, free or short-connate at base, punctate or line-

LUNDELL: FLORA OF GUATEMALA 193

ate; petals short-connate at base, dextrorsely contorted in bud, commonly lineate;
stamens usually shorter than petals, the filaments slender; anthers elongate, sub-
sagittate at base, dorsifixed or basifixed, usually dehiscent by introrse slits; ovary
in pistillate flowers ovoid, with slender style subequaling or exceeding the stamens;
ovary in staminate flowers abortive, with short style less than 1 mm. long; placenta
with 3-5 uniseriate ovules; fruit drupaceous, 1-seeded, the endocarp crustaceous or
osseous; seed globose or depressed, the endosperm corneous, excavate, not rumi-
nate; embryo transverse, elongate.

About 50 species in tropical America. Several additional ones are
represented in Central America, all of which appear to be dioecious.

Inflorescences axillary, 3.5 cm. long or less S. laevis.

Inflorescences terminal, usually 5-10 cm. long S. guatemalensis.

Stylogyne guatemalensis Blake, Contr. U. S. Nat. Herb. 24:
16. 1922 (type from Quebradas, Izabal, H. Pittier 8624). S. perpunc-
tata Lundell, Bull. Torrey Club 69: 398. 1942 (British Honduras, the
type from Silk Grass Reserve, Stann Creek District, Percy H. Gentle
2990). Pigeon berries (British Honduras).

Moist or wet forest, 300 meters or lower; Izabal. Mexico (Oaxaca,
Chiapas); British Honduras; Honduras.

A shrub or small tree, entirely glabrous; branchlets rather stout, terete; leaves
with canaliculate petioles mostly 1-1.5 cm. long, sometimes up to 3 cm. long; leaf
blades elliptic or obovate-elliptic, 6-15 cm. long, 3-8 cm. wide, apex rounded and
rather abruptly short-acuminate or acutish, base cuneate or acute and decurrent
on the petiole, entire, membranaceous to subcoriaceous, coarsely punctate, finely
and closely veined, the costa plane or nearly so above, elevated beneath; inflores-
cences terminal, pinnately paniculate, sessile or nearly so, usually less than 6 cm.
high and wide, sometimes up to 10 cm. high, 12 cm. wide; flowers dioecious, 5-
parted, small, umbellate or subcorymbose; pedicels mostly 3-6 mm. long, some-
times 2-11 mm. long; pistillate flowers: about 5 mm. long at anthesis, rather con-
spicuously punctate; sepals oval-oblong to lanceolate-oblong, 1.4-2 mm. long,
rounded, dorsally punctate with elevated glands, rather thick, the margin thin and
paler; petals oblong-elliptic, 4.5-4.8 mm. long, connate about 1.5 mm. at base,
obtuse, rounded or obliquely emarginate, lineate; stamens about 2.2 mm. long;
filaments about 1 mm. long; anthers 1.2-1.7 mm. long, basifixed, epunctate; ovary
ovoid, 1.2 mm. long; style slender, about 2.5 mm. long; ovules 3 or 4, uniseriate,
erect; staminate flowers: sepals ovate-elliptic, 1.3-1.75 mm. long, punctate; petals
elliptic, up to 5.5 mm. long, connate 1.5 mm. at base, lineate; stamens 3.5 mm. long;
filaments 2 mm. long; anthers 1.8-2 mm. long, dorsifixed above base; pistil abortive,
with style about 0.75 mm. long; ripe fruits subglobose, about 6 mm. in diameter
when dry.

The vernacular name "pigeon berries" is applied in British Hon-
duras to various plants with small edible berries.

194

FIELDIANA: BOTANY, VOLUME 24

More than one species may be represented in the Guatemalan
collections referred here. S. perpunctata represents the plants which
have large obovate thin leaves and petioles often 2 to 3 cm. long.
In the type of S. guatemalensis, a pistillate specimen, the leaves are
subcoriaceous, small and elliptic. Most collections have been iden-
tified as S. laevis (Oerst.) Mez, which is often difficult to separate.

Stylogyne laevis (Oerst.) Mez, Pflanzenreich IV. 236: 268. 1902.
Ardisia laevis Oerst. Vid. Medd. Kjoebenhavn 125. 1861. Stylogyne
ramiflora (Oerst.) Mez, Pflanzenreich IV. 236: 272. 1902. Ardisia
ramiflom Oerst. Vid. Medd. Kjoebenhavn 132. 1861.

FIG. 54. Stylogyne laevis. A, Flowering branch, pistillate plant; X %.
B, Staminate flower, opened out showing stamens, and abortive pistil with short
style; X 5. C, Pistillate flower, dissected; X 5.

LUNDELL: FLORA OF GUATEMALA 195

Mixed lowland forest, 300 meters or less; Escuintla; Retalhuleu;
Suchetepe"quez. Mexico (Chiapas) ; Nicaragua to Panama.

A glabrous shrub, or small tree, commonly 1-3 meters high; leaves with stout
marginate petioles usually less than 1 cm. long, sometimes up to 2 cm. long; leaf
blades oblong-elliptic to elliptic or oblanceolate-oblong, up to 22 cm. long, 8.5 cm.
wide, apex rounded to obtuse and abruptly short-acuminate, base acute and de-
current, chartaceous, entire, finely veined on both surfaces, the costa shallowly
sulcate above; inflorescences axillary, sessile, small, pinnately paniculate, mostly
3.5 cm. long or less, the branches carmine, glabrous; flowers dioecious, small, 5-
parted, white, umbellate or subcorymbose; pedicels 2-6.5 mm. long, up to 1 cm.
long in fruit; bracts small, ovate, deciduous; pistillate flowers: glabrous, about
4.5 mm. long at anthesis, orange-punctate with dots and lines; sepals membrana-
ceous, short-connate at base, oblong, ovate-elliptic or ovate, rounded at the apex,
2 mm. long or less, orange-punctate; petals contorted in bud, connate about 1.5
mm. at base, elliptic, up to 5 mm. long, reflexed at anthesis, orange-lineate; sta-
mens 3 mm. long; filaments slender, 1.8-2 mm. long; anthers basifixed, 1.5-1.8 mm.
long; ovary glabrous, the style slender, 2.5-3.2 mm. long; ovules 2-4 erect, uniseri-
ate; staminate flowers: about 4 mm. long at anthesis, orange-punctate; sepals ob-
long, 1.2 mm. long or less; petals 4 mm. long, connate about 1.4 mm. at base,
oblong-elliptic; stamens 3 mm. long; filaments about 2 mm. long; anthers slender,
about 1.3 mm. long, basifixed; ovary abortive, the style about 0.6 mm. long; fruit
globose, 5 mm. in diameter when dry, black, juicy.

Excellent Guatemalan specimens of this dioecious species were
collected by W. A. Kellerman at Retalhuleu. His number 6627 (F)
is from a staminate plant, while 6701 (F) has both pistillate flowers
and fruits. A study of the types of S. laevis (Oerst.) Mez and S. rami-
flora (Oerst.) Mez, both from Costa Rica, shows that these two are
based on staminate and pistillate collections, respectively, of the
same species.

Of Ardisia guatemalensis Mez, which was based on Friedrichsthal
857, I have seen only the type photograph. This is evidently a spe-
cies of Stylogyne, and probably is referable to S. laevis.

SYN ARDISIA (Mez) Lundell

Shrub or tree, rarely epiphytic; leaves alternate, large, petiolate; inflorescences
large, terminal, densely pubescent with gland-tipped hairs; flowers perfect, 5-
merous, corymbose, borne on long pedicels; sepals dextrorsely imbricate, almost
free, spreading early, punctate; corolla campanulate or suburceolate, the petals
dextrorsely convolute in bud, connate up to three-fourths their length; stamens
included, attached at base of corolla tube; filaments slender, longer than anthers,
united at base into shallow ring; anthers small, ovate, dorsifixed near base, cau-
date-acuminate, epunctate, longitudinally dehiscent; ovary subglobose, punctate,
glabrous; style slender, included; ovules 8 or more, pluriseriate; fruits depressed-
globose.

196

FIELDIANA: BOTANY, VOLUME 24

A monotypic genus, known only from Mexico and northern Cen-
tral America.

Synardisia venosa (Mast.) Lundell, Wrightia 3: 90. 1963. Ardi-
sia venosa Mast, in Donn.-Sm. Bot. Gaz. 18: 205. 1893; Mez, Pflan-
zenreich IV. 236: 77, t. 10. 1902. Pish-match (Huehuetenango) ; flor
de lima (Quezaltenango) .

Wet cloud forest and cool barrancas, 1,300-3,200 meters; Chimal-
tenango; Solola; Huehuetenango; Quezaltenango; San Marcos; Za-
capa; Jalapa; El Progreso; Santa Rosa; Zacatepequez (type from
Volcan Acatenango, J. Donnell Smith 2485). Mexico (Jalisco, Gue-
rrero, Mexico, Chiapas) ; El Salvador; Honduras.

A shrub or tree, sometimes 20 meters high, rarely epiphytic, the branchlets
thick, glabrous; leaves glabrous with short broadly marginate petioles up to 2 cm.
long; leaf blades large, oblanceolate or oblanceolate-oblong, mostly 15-25 cm. long,
5-9 cm. wide, sometimes larger, apex acute or broadly acuminate with obtuse
acumen, attenuate to the decurrent base, entire, punctate, the costa elevated be-

FIG. 55. Synardisia venosa. A, Flowering branch; X ^. B, Flower; X 2^.
C, Calyx and pistil; X 2>£. D, Corolla, dissected; X 2>£. E, Stamen, dorsal
view; X 5.

LUNDELL: FLORA OF GUATEMALA 197

neath, the veins slender, reticulate on both surfaces; inflorescences terminal, many-
flowered, subpyramidal, 3-4-pinnate-paniculate, often exceeding the leaves, up to
45 cm. long, the branches densely glandular-pubescent with gland-tipped hairs;
basal bracts leafy, oblanceolate, up to 10 cm. long, glabrous; pedicels slender, up
to 2 cm. long, usually shorter, densely pubescent with gland-tipped hairs; flowers
corymbose, 6-12 mm. long; sepals almost free, dextrorsely imbricate in young buds,
spreading early, very thin and slender, narrowly lanceolate, 2.5-5 mm. long, cau-
date-acuminate, punctate, pubescent with gland-tipped hairs; corolla campanulate
or suburceolate, thin, semi-transparent, pink, 5-11 mm. long, the petals dextrorsely
convolute in bud, connate up to three-fourths their length, subovate, obtuse-
rounded apically, sparsely punctate, glandular-puberulent at base within, other-
wise glabrous; stamens small, included, about two-thirds length of tube, attached
at base of corolla, up to 3.5 mm. long; filaments very slender, 1-2.5 mm. long,
glabrous, broadened at base and united into a shallow ring; anthers small, ovate,
1-1.8 mm. long, dorsifixed near base, caudate-acuminate or acuminate, epunctate,
longitudinally dehiscent; ovary subglobose, punctate, glabrous; style very slender,
included, 2.5-4 mm. long; ovules 8 or more, pluriseriate; fruit depressed-globose,
up to 1 cm. in diameter, wine-purple when ripe.

Among the Myrsinaceae of Mexico and Central America, S. ve-
nosa may be recognized by its ample, terminal, densely glandular-
pubescent inflorescences, petals connate up to three-fourths forming
a campanulate or suburceolate tube, and small included stamens and
style. The flowers, quite variable in size, are reported to be very
fragrant.

Collectors report that both the young shoots and the berries are
edible.

YUNCKERIA Lundell
Reference: Lundell, Wrightia 3: 111-114. 1964.

Shrubs or small trees, glabrous or nearly so; branches terete; leaves mostly
large and stoutly petiolate, punctate, entire, finely veined; flowers perfect, in ter-
minal panicles, usually 5-merous, rarely 4-merous, umbellate or corymbose; pedi-
cels long, thickened above; calyx lobed to or below middle, tapering at base into
pedicel; sepals dextrorsely imbricate, rounded, thick and punctate medially; petals
dextrorsely imbricate, connate into basal tube, linear-oblong, widest at or near
middle, punctate medially; stamens subequaling petals, attached near base of
corolla tube; filaments free, shorter than anthers; anthers large, linear-lanceolate,
attenuate to apex, erect, rigid, reddish, epunctate, dehiscent through apical pores;
ovary ovoid, glabrous; style slender, subequaling petals, the stigma minute, puncti-
form; placenta obovoid, with 6-8 large, erect, uniseriate ovules; fruits drupaceous,
black at maturity; seed globose, with basal depression extending into center of
endosperm; embryo transverse, elongate.

Three species in southern Mexico and Central America of which
two are known only from Chiapas, and one from British Honduras,
Honduras and Nicaragua. Yunckeria, named for the late T. G.

198 FIELDIANA: BOTANY, VOLUME 24

Yuncker, is a distinctive genus, referable to the Tribe Myrsineae.
With erect uniseriate ovules, it has possible affinity to Tapeinosperma
Hook. f. of Australia, New Caledonia and the South Pacific islands.

Leaves elliptic, apex short-acuminate y. Purpusii.

Leaves obovate or oblanceolate, apex obtuse or rounded and apiculate.

Leaves pallid, subcoriaceous; pedicels sparsely papillate; sepals about 1.5 mm.
long y . ovandensis .

Leaves reddish-brown, conspicuously so on undersurface, thin; pedicels glabrous;
sepals 2-2.5 mm. long y. amplifolia.

Yunckeria amplifolia (Standl.) Lundell, Wrightia 3: 113. 1964.
Ardisia amplifolia Standl., Field Mus. Bot. 4: 249. 1929. Uva de
montana, uva de monte (Honduras) .

Wet forest and river banks, 200 meters or lower. British Hon-
duras; Honduras; Nicaragua (type from region of Braggman's Bluff,
F. C. Englesing23ty.

Shrub or small tree up to 5 m. tall, entirely glabrous, branchlets stout; leaves
with marginate petioles 1-1.5 cm. long; leaf blades oblong-obovate or obovate,
18-30 cm. long, 8-13 cm. wide, apex obtuse, base cuneate and decurrent, entire,
subchartaceous, the lower surface drying paler and reddish-brown, densely punc-
tate but sometimes obscurely so, the costa prominent and elevated beneath, the
lateral veins slender; inflorescences bright red, terminal, bipinnate, paniculate,
4-15 cm. long, the apical umbels of central axis often subsessile; pedicels slender,
up to 1 cm. long, accrescent and up to 1.5 cm. long in fruit; flowers usually 5-,
rarely 4-parted, subcorymbose-umbellate, the buds up to 1 cm. long, slender, gla-
brous; sepals dextrorsely imbricate, broadly ovate or elliptic, 2-2.5 mm. long,
rounded at apex, entire, rather thick, punctate; petals (in buds) linear-lanceolate,
dextrorsely imbricate, about 2.5 mm. wide, connate about 1.5 mm. at base, con-
spicuously punctate; stamens (in buds) 7.5 mm. long; filaments slender, 2 mm.
long; anthers erect, dorsifixed about 0.75 mm. above base, linear-lanceolate, 5.5-
6 mm. long, attenuate to the apex, dehiscent through apical pores, epunctate; ovary
small, ovoid, glabrous; style very slender, up to 9 mm. long; placenta obovoid;
ovules 6-8, uniseriate, erect; fruits globose at maturity, about 1 cm. in diameter,
black, apiculate; endocarp crustaceous; seed globose, with basal depression extend-
ing into center of endosperm; embryo transverse, elongate.

The species is easily recognized among Central American Myrsi-
naceae by its reddish-brown twigs and leaves.

Yunckeria ovandensis (Lundell) Lundell, Wrightia 3: 112. 1964.
Ardisia ovandensis Lundell, Contr. Univ. Mich. Herb. 4: 21. 1940.

Wet forest, 2,370 meters or less. Mexico (Chiapas, type from
Mt. Ovando, E. Matuda 2549).

D

FIG. 56. Yunckeria amplifolia. A, Flowering branchlet; X K- B, Branchlet
with fruits; X 1A. C, Calyx and style; X 2J^. D, Flower bud, showing imbricate
petals; X 2J^. E, Placenta, showing erect ovules; X 10. F, Placenta cross section;
X 10. Flowers illustrated from type, Englesing 230 (F).

199

200 FIELDIANA: BOTANY, VOLUME 24

A small tree, up to 9 meters high, 25 cm. in diameter, the branchlets stout,
glabrous; leaves with stout marginate petioles 1-1.3 cm. long; leaf blades obovate
or oblanceolate, 12.5-22 cm. long, 5.5-8.3 cm. wide, apex rounded and obtusely
apiculate, base cuneate and decurrent, entire, punctate, subcoriaceous, costa prom-
inent and elevated beneath, the primary lateral veins very slender; inflorescences
terminal, 2-3-pinnate, paniculate, up to 12.5 cm. long, glabrous, many-flowered;
pedicels slender, up to 1 cm. long at base of corymb, sparsely and minutely papil-
late at first; flowers in racemiform corymbs, about 1 cm. long at an thesis; calyx
lobed to about middle, tapering below into pedicel, minutely and sparsely papillate
below in bud; sepals dextrorsely imbricate, broadly rounded, 1.5 mm. long, punc-
tate, the margin scarious, entire; petals oblanceolate-oblong, 8-9 mm. long, 2.5 mm.
wide, connate about 2 mm. at base, glabrous, dextrorsely imbricate, conspicuously
punctate; stamens 8 mm. long; filaments slender, 3-4 mm. long; anthers erect,
linear-lanceolate, 4-5.5 mm. long, dorsifixed about 0.75 mm. above base, not punc-
tate, dehiscent through apical pores; ovary small, elongate, ovoid-oblong; style
slender, 9 mm. long; placenta obovoid, small; ovules 6 or 7, uniseriate, erect.

Yunckeria Purpusii (Brandegee) Lundell, Wrightia 3: 112. 1964.
Ardisia Purpusii Brandegee, Univ. Calif. Publ. Bot. 6: 189. 1915.

Known only from the type, collected in damp barrancas near
Finca Irlanda, Chiapas, Mexico, C. A. Purpus 7119.

Branchlets stout, terete, the plant entirely glabrous; leaves large with stout
marginate petioles 1-2 cm. long; leaf blades elliptic, 17-27 cm. long, 7.5-10 cm.
wide, apex short-acuminate, base cuneate or acute and decurrent, entire, thin, sub-
chartaceous, costa shallowly sulcate above, elevated beneath, the veins slender and
inconspicuous; inflorescences terminal, tripinnately paniculate, about 8 cm. long;
pedicels slender, 8-14 mm. long, thickened above; flowers perfect, corymbose,
about 9 mm. long at anthesis; sepals dextrorsely imbricate, ovate, 2 mm. long,
obtuse or rounded, rather thick, punctate with a few small black glands, drying
with few scattered plate-like glands on dorsal surface; petals dextrorsely imbricate,
connate about 3 mm. at base, linear-oblong, 8-9 mm. long, obscurely lineate medi-
ally; stamens about 7 mm. long, attached about 1 mm. above base of corolla;
filaments about 3 mm. long; anthers dorsifixed about 0.8 mm. above base, linear-
lanceolate, 4-5 mm. long, attenuate to the apex, dehiscent through apical pores,
epunctate, rigid; ovary ovoid; style slender, 6.5 mm. long; placenta obovoid; ovules
6 or 7, erect, uniseriate.

PRIMULACEAE. Primrose Family

Reference: F. Pax & R. Knuth, Primulaceae, Pflanzenreich IV.
237: 1-386. 1905.

Annual or perennial herbs, or rarely suffrutescent; leaves alternate, opposite,
or verticillate, sometimes all basal, usually simple, sometimes dentate or lobate;
flowers axillary and solitary, usually racemose, paniculate, spicate, or umbellate

STANDLEY AND WILLIAMS: FLORA OF GUATEMALA 201

at the apex of a scape, small or rather large and showy, regular, often dimorphous
or heterostylous, commonly 5-parted; calyx more or less persistent, often somewhat
foliaceous, gamosepalous; corolla gamopetalous, the tube short or elongate, the
lobes imbricate in bud, entire or emarginate; stamens epipetalous, opposite the lobes;
ovary superior, or half inferior in Samolus, 1-celled, the placenta central; ovules
numerous; fruit capsular or a pyxis in Anagallis and Centunculus, variously dehis-
cent, few-many-seeded; seeds angulate or angulate-compressed ; embryo small,
immersed in the carnose or corneous endosperm.

Genera about 28, some of them with many species, the plants
mostly in temperate and arctic regions. Very few are found in trop-
ical regions, and only the following genera and species are known in
Central America.

The common Cyclamen, Cyclamen persicum Mill., native of the
eastern Mediterranean regions, is grown in pots in Guatemala, par-
ticularly in Quezaltenango and San Marcos, but is rare. The plants
sometimes are sold at what would be in the United States fantasti-
cally high prices.

One or more species of Primula are sometimes grown in Guate-
mala in pots for ornament, but they are rare in cultivation in Central
America.

Flowers in terminal leafless racemes; leaves large, mostly 10-20 cm. long. .Samolus.
Flowers axillary; leaves often very small.

Capsule longitudinally dehiscent; leaves mostly 3-6 cm. long Lysimachia.

Capsule circumscissile; leaves small, less than 2 cm. long, often very small.

Corolla longer than the calyx; stamens inserted on the base of the corolla.

Anagallis.

Corolla shorter than the calyx; stamens inserted in the throat of the corolla.

Centunculus.

ANAGALLIS L.

Small herbs, simple or branched, with leafy stems, repent or erect, the stems
terete or tetragonous; leaves opposite or alternate, small, sessile or short-petiolate,
entire; flowers axillary, pedicellate, red, blue, or white; calyx 5-parted almost to
the base, the segments lanceolate or subulate, spreading; corolla hypogynous, parted
almost to the base, rotate or rotate-campanulate, the lobes obovate or linear, entire
or erose-denticulate, contorted; stamens 5, inserted on the base of the corolla,
united at the base to form a short tube, usually pilose, the anthers ellipsoid, obtuse;
ovary globose, the style filiform, the stigma obtuse; ovules numerous, immersed
in the globose placenta; capsule (pyxis) globose, circumscissile, many-seeded; seeds
small, flat dorsally, conic-umbilicate ventrally, the embryo transverse.

About 24 species, widely distributed in both hemispheres. Only
the following are known from tropical America.

Leaves opposite, mostly 8-15 mm. long A. arvensis.

Leaves alternate, mostly 5-6 mm. long A. pumila.

202

FIELDIANA: BOTANY, VOLUME 24

Anagallis arvensis L. Sp. PI. 148. 1753.

Native of Europe and other parts of the Old World; introduced
and naturalized in many temperate and tropical regions of America;
well established in Guatemala in many places, especially in Chimal-
tenango, mostly as a weed in cultivated or waste ground, 1,500-2,500
meters; Guatemala; Sacatepe"quez; Chimaltenango; Quezaltenango.

Plants annual, procumbent, usually much branched, the stems mostly 30 cm.
long or less, 4-angulate, subulate; leaves opposite or ternate, usually broadly ovate,
commonly 8-15 mm. long, very obtuse to subacute, pale green; pedicels axillary,
often twice as long as the leaves, slender, recurved in fruit; calyx lobes 5 mm. long,
lance-linear; corolla salmon-red or blue, rotate, about equalling the calyx lobes,
parted almost to the base, the lobes broadly obovate, obtuse; stamens half as long
as the corolla lobes.

FIG. 57. Ana gallis arvensis. A, Terminal portion of a branch, X 1. B, Flower,
X 6. C, Portion of corolla showing attachment of stamen, X 6. D, Capsule,
nearly mature, the line of rupture of dehiscence shown, X 5.

STANDLEY AND WILLIAMS: FLORA OF GUATEMALA 203

The Guatemalan material is referable to var. Phoenicia (Scop.)
Gren. & Godr. The plant is extensively naturalized in Mexico and
the United States. It has been found in Honduras also, where the
flowers are usually blue — while those in Guatemalan material are
usually salmon red.

Anagallis pumila Swartz, Prodr. Veg. Ind. Occ. 1: 40. 1788.
Centunculus pentandrus R. Br. Prodr. 427. 1810.

Wet meadows or moist banks, sometimes in pine forest or bogs,
1,900 meters or less; Pete"n; Alta Verapaz; Baja Verapaz; Jalapa;
Jutiapa; Santa Rosa; Guatemala; Chimaltenango; Quiche"; Huehue-
tenango. Florida; Mexico; British Honduras to Panama; West In-
dies; South America; Asia; Africa; and Australia.

Plants annual, often much branched, erect or procumbent, the stems mostly
10 cm. long or shorter, glabrous throughout; leaves sessile or nearly so, alterate,
ovate or lance-ovate, mostly 5-6 mm. long, acute, the lower leaves usually oppo-
site; pedicels axillary, spreading or reflexed, equaling or twice as long as the leaves;
calyx lobes scarcely 2 mm. long, linear-lanceolate; corolla white, funnelform, scarcely
equaling the calyx lobes, deeply 4-5-parted, the lobes ovate-lanceolate, obtuse, the
tube short; stamens much shorter than the corolla lobes.

Most of the rather numerous Guatemalan collections belong to
var. longipedunculata Chod. & Hassl., in which the pedicels equal or
exceed the leaves. One collection from Quezaltenango (Volcan de
Zunil, 2,500-3,800 meters, Steyermark 31+611+} is referable to var.
ovalis (Ruiz & Pav6n) R. Knuth, in which the flowers are sessile or
nearly so. A. pumila is a small and very inconspicuous plant, usually
hidden by the grasses and other larger herbs among which it grows.

CENTUNCULUS L.

Plants very small, annual, glabrous, branched or simple, the stems leafy; leaves
alternate or the lowest opposite, sessile or on very short petioles, entire; flowers
minute, axillary, sessile or pedicellate; calyx 4-5-parted almost to the base, the
lobes lanceolate, acute, longer than the corolla; corolla hypogynous, deeply 4-5-
parted, the lobes erect-spreading, entire, the tube subglobose; stamens 4-5, inserted
in the throat of the corolla, not exceeding the corolla lobes, the anthers ellipsoid,
obtuse at each end; style filiform, the stigma obtuse; ovary globose, the ovules
numerous, inserted on a globose placenta; capsule (pyxis) globose, circumscissile,
many-seeded; seeds flat dorsally, convex ventrally, the embryo transverse.

The genus consists of a single species.

204

FIELDIANA: BOTANY, VOLUME 24

Centunculus minumus L. Sp. PI. 116. 1753.

Wet, marshy meadows or wet open banks, 2,200-3,500 meters;
Chimaltenango (Cerro de Tecpam); Totonicapan ; San Marcos (vol-
canoes of Tajumulco and Tacana); Huehuetenango (Sierra de los
Cuchumatanes). United States; Mexico; Honduras; South America.
Old World.

Stems prostrate, mostly 5 cm. long or shorter, very slender, densely leafy;
leaves ovate, mucronate, sessile, generally 3-5 mm. long; flowers sessile or on very
short petioles, white; calyx parted almost to the base, 2.5 mm. long, the lobes subu-
late-linear, acuminate; corolla urceolate-rotate, shorter than the calyx, 4-parted
to the middle, the lobes lanceolate, acute, the throat slightly constricted; capsule
globose, membranaceous; seeds numerous, angulate, minute.

Pax and Knuth speak of this plant as introduced in America, but
there seems to be no good reason for doubting that it is native, like
Anagallis pumila.

JN\S

FIG. 58. Centunculus minimus. A, Habit of plant, X 1. B, Branch of plant,
X 5. C, Flower, X 10. D, Capsule nearly mature, X 10. E, Corolla dissected
to show anthers, X 15.

STANDLEY AND WILLIAMS: FLORA OF GUATEMALA

LYSIMACHIA L.

205

Reference: Ray, James Davis. The Genus Lysimachia in the New
World. 111. Biol. Monog. 24: 1-160, tt. 1-20. 1956.

Erect or procumbent, usually perennial herbs; leaves alternate, opposite, or ver-
ticillate, membranaceous, entire; flowers rather large, axillary, racemose, or corym-
bose-paniculate, sometimes umbellate, white or yellow; calyx 5-6-parted; corolla
hypogynous, subrotate or campanulate, 5-6-parted, the lobes contorted, entire or
dentate; stamens 5-6, inserted on the tube of the corolla, sometimes with inter-
posed staminodia, the anthers oblong, obtuse; ovary globose or ovoid, the style
filiform, the stigma obtuse; ovules few or numerous, inserted on a globose placenta;
capsule ovoid or globose, usually 5-valvate at the apex, few-many-seeded; seeds
oblong, orbicular, or angulate, sometimes winged, the embryo transverse.

More than 100 species, in both hemispheres, mostly in temperate
regions. Only the following is known from Central America.

Lysimachia steyermarkii Standl. Field Mus. Bot. 22: 369.
1940.

FIG. 59. Lysimachia steyermarkii. A, Branch, X Y^ B, Flower, X 6. C, Sta-
men showing attachment to portion of corolla, X ± 7. D, Underside of leaf to
show pubescence along nerves, X ± 7. E, Portion of multicellular hair, ± 18.

206

FIELDIANA: BOTANY, VOLUME 24

Moist shaded banks at base of cliffs, sometimes along streams in
Abies forest, 2,500-3,800 meters; endemic; Huehuetenango; Quezal-
tenango (type from Volcan de Zunil, Steyermark 34772).

An ascending or decumbent herb, primary stems very thick, rooting at the
nodes, the ultimate branches erect, 16-19 cm. long, simple, glabrous or sparsely
villous above with long spreading hairs; leaves alternate, on petioles 4-15 mm.
long, broadly ovate or elliptic, 3-6 cm. long, 1.5-2.5 cm. broad, acute or subacumi-
nate, narrowed to the base, sometimes contracted and decurrent, green above,
sparsely villous or glabrate, paler beneath, sparsely villous or almost glabrous,
densely punctate; flowers fasciculate in the upper leaf axils, the slender pedicels
1.5-2 cm. long, sparsely villous; sepals in fruit 5 mm. long, oblong-lanceolate, acute
or acuminate, densely punctate, 1-costate; capsule equaling the sepals, glabrous.

B

J.M.S. *.!

FIG. 60. Samolus ebracteatus. A, Habit of relatively small plant, X %.
B, Corolla from above, X 20. C, Portion of corolla showing attachment of stamen
and stipitate glands at throat, X 10. D, Flower, X 5. E, Stipitate gland from
pedicel of flower, X ± 50.

STANDLEY AND WILLIAMS: FLORA OF GUATEMALA 207
SAMOLUS L.

Annual or perennial herbs; leaves alternate, often succulent, usually scattered
along the stems, entire; flowers small, white, racemose or corymbose, the inflores-
cences terminal, the pedicels bracteate at the base; calyx tube adnate below to the
ovary, the limb 5-fid, persistent; corolla perigynous, subcampanulate, the limb
5-parted; stamens inserted on the tube or throat of the corolla, alternating with
the same number of staminodia, these subulate or ligulate; connective of the sta-
mens sometimes elongate; ovary globose, the style short, the stigma obtuse or
capitellate; ovules numerous; capsule ovoid or globose, 5-valvate at the apex; seeds
very numerous.

About 9 species, occurring in both hemispheres. Only one is
known from Central America.

Samolus ebracteatus HBK. Nov. Gen. & Sp. 2: 181, t. 129.
1817.

Moist, or wet, open places, 300 meters or lower; Huehuetenango
(Cie"naga de Lagartero, below Miramar, Steyermark 51518). South-
western United States; Mexico; British Honduras.

Plants stout, perennial, glabrous throughout, usually 70 cm. high or less, the
stems mostly simple; leaves fleshy, spatulate or rounded-spatulate, most of them
10-20 cm. long, rounded or very obtuse at the apex, somewhat abruptly narrowed
below the middle into a long, broadly winged petiole, this dilated and clasping at
the base, green above, pale beneath; racemes long and many-flowered, the pedicels
widely spaced, 1-2.5 cm. long, very slender, glabrous or nearly so, not bracteate
at the base; calyx green, 2 mm. long, lobate to the middle, puberulent, the lobes
ovate-deltoid, acute; corolla white, little longer than the calyx, the lobes rounded-
cuneate, obtuse or truncate, denticulate at the apex; staminodia none; capsule
ovoid-globose, about equaling the calyx.

PLUMBAGINALES

The order contains a single family, the Plumbaginaceae. The
family is presumed to be related to the Primulaceae and perhaps to
some Caryophyllaceae. It is distinguished by having five styles, or
stigmas, a one-celled ovary with a single pendulous ovule. The char-
acters of the order are those of the single family which it contains.

PLUMBAGINACEAE, Plumbago Family

Perennial herbs or suffrutescent plants with short or sometimes vine-like stems;
leaves alternate, sometimes rosulate, estipulate; inflorescence spicate to usually
racemose or paniculate, sometimes reduced to a single flower (in Armeria capitate) ;
flowers perfect, regular, small to large and showy, essile or short-pedicellate; calyx
5-lobed, gamosepalous, the tube 5-15-costate, sometimes membranaceous or hya-

208 FIELDIANA: BOTANY, VOLUME 24

line; corolla sympetalous or the petals often nearly separate, the lobes imbricate
in bud; stamens 5, opposite the corolla segments, the filaments adnate to base of
the segments or united for most of their length with the corolla tube; anthers ovate
or oblong, dorsifixed, the two cells parallel and contiguous, dehiscent by longi-
tudinal slits; ovary superior, 1-celled, with a single anatropous ovule; styles 5,
distinct or connate at the base, the stigmas terminal and capitate or sometimes
decurrent on the styles; fruit an utricle, indehiscent or circumscissile, often in-
cluded in the calyx; seed filling the cell; endosperm farinaceous, from copious to
scant or none; embryo straight, cotyledons linear or oblong.

A predominately Old World family of about eight genera and
some 300 species. Three genera are native in America, but only
Plumbago is indigenous in Central America.

Calyx glandular, tubular, the teeth erect; style simple below, branched above.

Plumbago.
Calyx not glandular; the teeth spreading; styles distinct Limonium.

LIMONIUM Adanson

Perennial or annual herbs, usually erect and branched, sometimes acaulescent;
leaves cauline or basal, narrow or broad, entire, or often sinuate or pinnatifid;
stems often winged; flowers rather small, in bracteate spikelets; calyx usually fun-
nelform, the tube commonly 10-costate or 5-costate, the limb scarious, plicate,
spreading in anthesis, the 5 costae excurrent as teeth or bristles; petals united only
near the base, usually obovate or obcordate; styles distinct, the stigmas oblong,
capitate, or minute; utricle included in the calyx, oblong, fusiform, or linear, inde-
hiscent or variously dehiscent at the apex.

About 150 species, mostly in the northern hemisphere of the Old
World, a few species native in temperate North America. None is
native in tropical America.

Limonium sinuatum (L.) Mill. Gard. Diet. ed. 8. no. 6. 1768.
Statice sinuata L. Sp. PI. 276. 1753. Estatice.

Native of the Mediterranean region; cultivated occasionally for
ornament in Guatemalan gardens.

Plants biennial or annual, erect, 30-60 cm. high, sparsely branched, sparsely
hispid-scabrous with broad-based hairs; lower leaves sinuate-pinnatifid, the lobes
broad, mostly rounded at the apex; branches broadly 3-winged; spikelets 3-5-
flowered, in short secund spikes at the ends of the branches; calyx corolla-like, 1 cm.
broad, broadly funnelform, blue or purple, glabrous, truncate, crenulate; corolla
small, yellowish white.

This is an occasional garden plant in the Guatemalan highlands
and in other places in Central America. The flowers are sometimes
used like "everlasting" flowers and remain for a long time in homes,
often on altars. Flower color, actually the calyx, is blue or purple

STANDLEY AND WILLIAMS: FLORA OF GUATEMALA 209

but forms (Bailey in Man. Cult. PI. 787. 1949 says hybrids) with
white and yellow "flowers" are known in Guatemala. Standley
thought L. bonduellii (Lest.) Kuntze was cultivated in Guatemala
but the form he saw may have been a yellow "flowered" L. sinuatum.

PLUMBAGO L. Plumbago

Plants perennial, herbaceous or suffrutescent, the stems usually weak and often
reclining or somewhat scandent; leaves alternate, sometimes clasping, entire, mem-
branaceous; flowers rather large, purple, blue, or white, in bracteate spikes; calyx
tubular, 5-costate, glandular, with 4-5 erect teeth; petals 4-5, their claws united
into a slender tube, the limb spreading, the corolla salverform; stamens 5, distinct,
the filaments dilated at the base, the anthers linear; styles filiform, partly united,
stigmatose on the inner side; fruit capsular.

About 10 species, in the Mediterranean region, western Asia, and
tropical America. Two native species occur in Mexico, but only one
of them extends to Central America.

Petioles conspicuously dilated and auriculate at the base; cultivated plants.

P. capensis.
Petioles not auriculate-dilated at the base; native plants P. scandens.

Plumbago capensis Thunb. Prodr. PL Cap. 33. 1794-1800.
Plumbago.

Native of South Africa; grown for ornament in gardens and parks
of Guatemala, from sea level to 2,200 meters or higher.

A slender sub-shrub, 1-2 meters long, the stems weak, often reclining or sub-
scandent; leaves oblong to spatulate, 3-7 cm. long, narrowed to the obtuse apex,
attenuate to the base, auriculate-dilated at the base of the petiole, sparsely lepidote
or glabrate; flowers large and showy, sky-blue, in short dense spikes; calyx tube
12 mm. long, covered over most of its surface with long-stipitate glands; corolla
tube as much as 4 cm. long, the limb 2.5 cm. broad.

A rather handsome plant, well known in cultivation in the United
States, where it usually is grown in hothouses. Known in El Salvador
as "umbela" and "lumbela"; "embeleso," "jazmin azul" (Yucatan).

Plumbago scandens L. Sp. PI. ed. 2. 215. 1762. Hierba del
pajaro; zapatilla.

Moist or dry thickets, sometimes in hedges or waste ground, 1,000-
1,800 meters; Zacapa; Guatemala; Chimaltenango; Quiche". Southern
Florida; Mexico to Costa Rica. West Indies; South America.

Plants suffrutescent or herbaceous almost throughout, a meter high or less,
erect or reclining, minutely lepidote or almost glabrous; leaves pale, ovate to ob-

210

FIELDIANA: BOTANY, VOLUME 24

long-lanceolate, 3-10 cm. long, acute or acuminate, acute and petiolate at the base,
the petioles 1 cm. long or shorter, not dilated at the base; spikes slender and inter-
rupted, several-many-flowered, 5-12 cm. long, the bracts lanceolate, acuminate,
persistent; calyx 1 cm. long, covered with long-stipitate glands, very viscid to the
touch; corolla white, the tube filiform, 2 cm. long, the lobes of the limb obovate,
mucronate, 5-7 mm. long.

Known in El Salvador as "guacochile" ; "chabac," "chapac"
(Maya); "hierba de alacran" (Yucatan).

If the crushed leaves of this and other species are applied to the
skin they cause almost instantaneous rubef action, and in a very short
time blisters. Taken internally, the plant is poisonous. In tropical
America the leaves sometimes are applied externally as a remedy for
itch and other cutaneous diseases, and an infusion of the leaves some-
times is administered as an emetic or purgative, although such use
must be rather dangerous.

FIG. 61 Plumbago scandens. A, Habit, X %. B, Flower, X 3. C, Corolla,
opened to show attachment of anthers, X 3. D, Mature capsule within calyx,
X 2^2. E, Single ovule showing attachment of petiole to stem, X ± 2. G, Stipi-
tate gland from calyx, X 10.

Publication 1002