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LIBRARY OF THE

UNIVERSITY OF ILLINOIS

AT URBANA-CHAMPAIGN

BIOLOGY

TROPICAL AMERICAN MYRTACEAE, II

NOTES ON GENERIC CONCEPTS

AND DESCRIPTIONS OF

PREVIOUSLY UNRECOGNIZED SPECIES

ROGERS McVAUGH

FIELDIANA: BOTANY

VOLUME 29, NUMBER 8

Published by

CHICAGO NATURAL HISTORY MUSEUM

MAY 31, 1963

BIOLOGY LIBRARY

101 BURRILL HALL

TROPICAL AMERICAN MYRTACEAE, II

NOTES ON GENERIC CONCEPTS

AND DESCRIPTIONS OF

PREVIOUSLY UNRECOGNIZED SPECIES

ROGERS McVAUGH

Curator of Vascular Plants, University Herbarium,

The University of Michigan

FIELDIANA: BOTANY

VOLUME 29, NUMBER 8

Published by

CHICAGO NATURAL HISTORY MUSEUM

MAY 31, 1963

3

CONTENTS

PAGE

Introduction 395

1. Calyptranthes Sw 397

The Genus in Continental North America 397

Miscellaneous Notes 412

2. Eugenia L 413

Synopsis of Mexican Species West of the Isthmus of Tehuantepec . . 413

Notes on Miscellaneous Species 464

3. Marlierea Camb 470

4. Myrcia DC 470

Miscellaneous Notes 470

5. Myrcianthes Berg 473

Synopsis of the Genus (Eastern South America Excluded) 473

Nomenclature 476

6. Certain Eugenioid Genera 498

Revision of Myrciaria Berg, in North America and Northern South

America 499

A New Species of Plinia L., from Costa Rica 505

The Genus Siphoneugena Berg 507

7. Pimenta Lindl 511

8. Psidiumi, 512

Delimitation of the Genus 512

Additions to the Genus 519

Species Described by Standley from British Honduras 522

Miscellaneous Notes 524

Index . . 530

393

Tropical American Myrtaceae, II1

The following notes have been prepared as a necessary prelimi-

nary to a formal treatment of the Myrtaceae of Guatemala. There

has been no comprehensive revision of North American Myrtaceae

since the work of Berg (1855-62), and it was found impossible to

revise the Guatemalan species without full consideration of those of

Mexico, the rest of Central America, and the West Indies.

The myrtaceous flora of Central America is a comparatively

small one. The number of genera represented in the flora of Guate-

mala, for example, is 8 (the same as in Costa Rica and Panama).

The number of species is about 95 as far as known, as against 35-40

in Costa Rica and about as many in Panama. The flora of Surinam

includes about 90 species. The myrtaceous flora of Peru includes

14 genera and about 175 species; that of Brazil, according to esti-

mates by Berg, includes more than 600 species in 33 genera.

The genera represented in Central America are all, with the pos-

sible exception of Pimento,, represented in South America by more

diversified populations and larger numbers of species. There seems

to be little or no direct relationship between the North American

and South American species, except for weedy species like Psidium

guajava and a few like Myrcianthes fragrans and Myrciaria flori-

bunda that are widespread in the West Indies and range to northern

South America and continental North America.

In general there seems to have been a strong phylogenetic con-

nection between the myrtaceous flora of the West Indies and that of

Central America, especially of the Atlantic lowlands that extend from

Honduras to Yucatan. In all the genera having lowland representa-

tives (that is, all except Ugni), some of the species are identical with

West Indian species or scarcely distinguishable from them. The

greatest amount of relative similarity between the two floras, as

pointed out above, is where they are geographically closest. Other

connections, however, are suggested between the flora of Jamaica

and that of Costa Rica (in Myrcianthes), and most curiously (and

1 An earlier paper with the same title dealt primarily with the Myrtaceae of

Peru (Fieldiana, Bot. 29: 143-228. 1956).

395

396 FIELDIANA: BOTANY, VOLUME 29

as yet unexplainedly) along an interrupted axis extending from Cuba

to Yucatan and thence to western Jalisco and the Tres Marias Islands.

There seems to be a certain amount of endemism in both western

and eastern Mexico and in the Central American highlands, and a

relatively great amount — perhaps a reflection of greater thorough-

ness of exploration — in Pete"n and British Honduras. Any sup-

posedly undescribed species from Central America should be carefully

compared with West Indian material, not only because of the pos-

sibility that the same species may occur in both regions, but also

because of the high probability that the West Indian plant has

already been named and described. This is true not only because

of the many species described by Linnaeus, Lamarck, Swartz and

others in the eighteenth and early nineteenth centuries, but also

because of the many Myrtaceae described by Urban and others in

more recent times. Between 1895 and 1931, for example, about 70

species of Calyptranthes were described from the West Indies; this is

almost exactly the same number as all the species of Calyptranthes

known to Berg a century ago.

Notable contributions to our knowledge of the flora of Central

America have been made in the last few decades by Paul C. Standley

and C. L. Lundell. More than half of all the known Guatemalan

species of Myrtaceae were first recognized and described by one or

the other of these workers.

In my own work of revision I have been aided by many friends

and colleagues who have lent herbarium materials and have assisted

in other ways. To the authorities at the Jardin Botanique de 1'Etat,

Brussels (BR); the Botanical Museum, Copenhagen (C); Chicago

Natural History Museum (F); the Harvard University Herbaria

(A, GH) ; the Royal Botanic Gardens, Kew (K) ; the Herbarium of

the Botanical Institute of the Academy of Sciences, Leningrad (LE) ;

the Herbario Nacional del Institute de Biologia, Mexico City

(MEXU); the University of California, Berkeley (UC); the United

States National Herbarium, Washington (US); and the Naturhis-

torisches Museum, Vienna (W), I am grateful for permission to

study types and other unique and valuable specimens.

Dr. C. L. Lundell has been most gracious in permitting me free

access to his large collections of Myrtaceae from Pete"n and elsewhere

in the Maya area, including the types and in most instances abundant

duplicate materials of species he has recently described. I am also

much indebted to Dr. Faustino Miranda for providing me with a

study set of his own collections from Chiapas and elsewhere in south-

eastern Mexico.

McVAUGH: TROPICAL AMERICAN MYRTACEAE 397

In the systematic notes that follow, the terminology is intended

to conform to that used in my earlier papers and also in the Flora

of Peru (Field Mus. Bot. 13, pt. 4: 569-818. 1958), and a paper on

Calyptranthes and Marlierea (Mem. N. Y. Bot. Gard. 10: 61-91.

1958). Some changes in concepts are noted below.

1. CALYPTRANTHES Sw.

THE GENUS IN CONTINENTAL NORTH AMERICA

There appear to be several centers of endemism in Calyptranthes.

Few species are widely distributed. At least one species, C. fascicu-

lata Berg, ranges from the Guianas to the Lesser Antilles, but no

primarily South American species is known to range northward into

North America or even into Colombia except Colombian Guayana.

About 16 species are known from the region between the Amazon

and the Orinoco (cf. Mem. N. Y. Bot. Gard. 10: 65-79. 1958), and

about 20 were treated recently in the Flora of Peru (Field Mus. Bot.

13, pt. 4: 591-616. 1958). The genus is almost absent from the

whole Andean region of South America and from the cis-Andean low-

lands bordering the Pacific. North of the Amazon and west of the

Andean foothills I know of but two local species, viz. C. killipii

Standl., from the coastal lowlands about Buenaventura, Colombia,

and C. meridensis Steyerm., from mountains in the State of MeYida,

Venezuela.

In view of the restricted distribution of a majority of the South

American species, and the apparent absence of the genus from much

of northwestern South America, it is perhaps not surprising to find

that most of the North American species are distinctive and com-

paratively distantly related. There is apparent, however, among the

North American species, a strong West Indian influence.

The genus Calyptranthes in the West Indies was reviewed by

Urban many years ago (Bot. Jahrb. 19: 591-603. 1895), at which

time he recognized about 30 species. About 70 additional West

Indian species, almost all from the Greater Antilles, have since been

described by Urban, Britton, and others. A new revision of all the

Antillean species is much needed, but indeed there seems to be more

diversity among the species of Calyptranthes in the Caribbean basin

than in any other area in tropical America.

Two well-known West Indian species, C. pollens and C. zuzygium,

occur in southern Florida as well; C. zuzygium has not been found

398 FIELDIANA: BOTANY, VOLUME 29

elsewhere on the North American continent, but C. pattens is repre-

sented in Mexico and Central America by several varieties. A species

of Cuba and Jamaica, C. chytraculia, is represented on the mainland

by the var. americana, which ranges from Mexico to northern Colom-

bia. Other species common to the West Indies and the continent

may become known as the flora is revised.

The following key includes all the species known to occur natur-

ally in continental North America. Descriptions of species are not

included unless they are unavailable elsewhere.

KEY TO THE SPECIES

Leaf-bases broadly cordate or subcordate, the blades nearly sessile or subclasping,

on stout short petioles 3 mm. long or less; blades usually acute or obtuse,

sometimes obscurely acuminate, never caudate; young leafy branchlets 2- or

4-angIed or -winged except in C. chiapensis with leaves 12-23 cm. long.

Young branchlets and inflorescence densely ferruginous-tomentose with con-

torted and mostly 1- or 2-branched but nearly erect hairs 0.4-1.3 mm. long.

Leaves 1.3-5 cm. wide, 4-12 cm. long, mostly lanceolate; branchlets bicari-

nate or winged.

Branchlets bialate, the wings terminating distally at each node between the

leaf-bases; buds 5.5-7 mm. long; panicles 9-flowered; San Luis Potosf.

C. hernandezii McVaugh.

Branchlets bicarinate, the keels terminating distally in the median line of

the leaf-bases; buds 4.5-5 mm. long; panicles 30-60-flowered; British

Honduras C. bartlettii Standl.

Leaves 6.5-7.5 cm. wide, 12-23 cm. long, oblong; branchlets terete or com-

pressed, not carinate; Chiapas C. chiapensis Lundell.

Young branchlets and inflorescence glabrous or thinly pubescent with appressed

flat dibrachiate hairs 0.1-0.2 mm. long; branchlets 2- or 4-angled or -winged.

Branchlets biconvex in cross section, bicarinate, the winglike angles termi-

nating distally in the midlines of the leaf-bases; Guatemala (Alta Vera-

paz, PetSn) C. contrerasii Lundell.

Branchlets 4-angled, 2 of the angles terminating distally in the midlines of

the leaf-bases, the other 2 winged, terminating distally between the

leaf-bases.

Plants quite glabrous; leaves ovate-oblong; Mexico (Mesa Chica, Vera-

cruz) C. karwinskyana Berg.

Plants thinly pubescent with minute appressed hairs; leaves lanceolate or

elliptic-lanceolate; Tabasco, Chiapas, British Honduras, Pete"n.

C. karlingii Standl.

Leaf-bases acute or cuneate, or sometimes abruptly rounded, never cordate; blades

usually definitely petiolate, often acuminate, sometimes caudate; branchlets

various.

Flowers 9 or fewer in each panicle, the two lateral branchlets and the terminal

one each bearing a single pedicellate flower or a cluster of 2-3 sessile flowers;

inflorescence 3-5 cm. long, the very slender or even capillary peduncle

usually much longer than the rest of the panicle; leaves long-acuminate,

3-7.5 cm. long.

Branchlets 4-angled or -winged, glabrous or essentially so; Mexico (Chiapas)

to Panama C. hylobates Amsh.

McVAUGH: TROPICAL AMERICAN MYRTACEAE 399

Branchlets terete, when young with abundant appressed or contorted rufous

dibrachiate hairs.

Branchlets tomentulose with coarse contorted hairs; leaves rounded at base;

petioles 2 mm. long; axis of inflorescence filiform, bearing 3 (-5) flowers;

Mexico C. tenuipes McVaugh.

Branchlets appressed-pubescent with straight longitudinally oriented hairs;

leaves acute at base; petioles 3-6 mm. long; peduncle up to about 1 mm.

wide, the panicle with 9 flowers or fewer; Guatemala.

C. paxillata McVaugh.

Flowers more numerous, usually 20-30 or more in each panicle, often sessile or

subsessile in small clusters near the tips; panicles as a whole usually 5 cm.

long or more, twice or more compound; leaves various, 10-16 cm. long if the

flowers are 9 or fewer (in C. perlaevigata with buds 7-8 mm. long).

Plants glabrous or sparingly pubescent; pubescence, if any, usually consisting

of minute flat appressed dibrachiate hairs 0.1-0.3 mm. long, these often

nearly confined to the base of the hypanthium, the nodes of the inflores-

cence, and the undeveloped vegetative buds.

Panicles 15-20 cm. long; peduncles compressed, 10-12 cm. long, 3-4.5 mm.

wide; leaves as far as known 11-12 cm. wide, up to 35 cm. long; in-

florescence and buds thinly reddish-pubescent; British Honduras-

Peten boundary C. megistophylla Standl.

Panicles mostly 10 cm. long or less (if infrequently to 15 cm. long, the pedun-

cles 3-5 cm. long, less than 2 mm. wide); leaves mostly 12-15 cm. long

or less, in one Panamanian species up to 28 cm.

Leaves 12-28 cm. long, caudate-acuminate; buds obovoid, 2-2.5 mm.

long, glandular-verrucose; plants sparingly pubescent at flowering

time, the panicles and the bases of the buds appressed-pubescent

with reddish hairs; Panama C. tumidonodia Schery.

Leaves shorter (or the plants quite glabrous) and merely obtuse or acute

or short-acuminate; northern Central America to Mexico.

Leaves prevailingly obovate to elliptic, obtuse or rounded at tip, 3.5-

6.5 cm. long; small connecting veins evident on the lower surface

in dried leaves as a raised reticulum; branchlets not winged;

pubescence appressed, rather sparse, grayish- or yellowish-white;

British Honduras C. cuneifolia Lundell.

Leaves prevailingly elliptic to lanceolate or ovate, acute or acuminate

(if small and obtuse the branchlets winged, the small veins ob-

scure, the pubescence, if any, reddish).

Leaves small and obtuse, mostly 2.5-4.5 cm. long, usually narrow,

often varying on the same plant from 2-5 (-7) times as long as

wide; branchlets persistently and prominently corky- winged;

pubescence sparse, evanescent, of rather loosely appressed yel-

low- or reddish-brown hairs; Honduras to British Honduras.

C. hondurensis Standl.

Leaves usually larger and acute or acuminate, usually 2-3 times as

long as wide; branchlets wingless or somewhat obscurely winged.

Flower-buds 3-4.5 mm. long, glabrous; whole plant essentially

glabrous; calyptra (2.5-) 3-4 mm. wide; persistent collar-like

base of the calyx 3-4 mm. wide in fruit.

Midvein narrowly impressed above; inflorescence pyramidal,

many-flowered, up to 15 cm. long, the lowest branches of

the panicle up to 6 cm. long, often as long as the peduncle;

flowers clustered near the tips; branchlets winged; state of

Mexico . . . . C. hintonii Lundell.

400 FIELDIANA: BOTANY, VOLUME 29

Midvein convex or flat above, sometimes concave, not narrowly

impressed; inflorescence mostly 5-7 cm. long or less, with

5-15 flowers, the lowest branches of the panicle up to 2 cm.

long and much shorter than the peduncle; flowers mostly

solitary.

Leaves 3-6 cm. long, obtuse to acuminate; branchlets winged;

calyptra membranaceous, 2.5-3.5 mm. wide; flowers con-

spicuously pedicellate and solitary; West Indies, Florida.

C. zuzygium (L.) Sw.

Leaves 10-12 cm. long, prominently acuminate; branchlets

compressed, not winged; calyptra conic, thickened, 4 mm.

wide; flowers nearly sessile in pairs at the 1-2 nodes of

the panicle; Chiapas C. perlaevigata Lundell.

Flower-buds [as far as known] (2?-) 2.5-3 mm. long; plants and

buds glabrous or pubescent; calyptra [as far as known] 1.5-

2.5 mm. wide; persistent collar-like base of the calyx 1.5-

3 mm. wide in fruit; midvein impressed or sulcate above.

Flower-buds and distal nodes of the panicle sparingly pubescent

with appressed reddish or pale hairs.

Leafy branchlets (at least some of them) wing-angled in most

plants; inflorescence rather slender, the principal second-

ary branches of the panicle less than 1 mm. wide; moun-

tains of Chiapas.

C. pollens var. mexicana (Lundell) McVaugh.

Leafy branchlets not wing-angled; inflorescence stout, the

principal secondary branches of the panicle much flat-

tened, their lowest internode often 6-10 mm. long,

1.5 mm. wide; southern Veracruz(?), British Honduras,

Yucatan [C. euryphylla Standl.] . . . .C. millspaughii Urb.

Plants glabrous except for a few hairs on the youngest vegeta-

tive parts; branchlets not winged.

El Salvador and British Honduras; petioles 7-10 mm. long;

veins and glands scarcely apparent in mature leaves.

C. calderonii Standl.

Veracruz; petioles 3.5-6 mm. long; veins sometimes incon-

spicuous, but evident beneath; glands apparent, the

leaves pellucid-punctate or impressed-punctate.

Leaves 3-4 times as long as wide, 1.5-2.5 cm. wide, coria-

ceous, impressed-punctate but scarcely pellucid-punc-

tate; lateral veins strongly ascending, leaving the

midvein at an angle of 45° or less; ventral surface of

the petiole channeled, the thick obtuse margins bor-

dering an ill-defined V-shaped or somewhat concave

trough; fruit 5-8 mm. in diameter, the fruiting disk

1.5-2 mm. wide C. schlechtendaliana Berg.

Leaves 1.6-3.3 times as long as wide, 1.5-4.5 cm. wide,

thin, evidently pellucid-punctate but rarely or only in

age impressed-punctate; lateral veins divergent, leav-

ing the midvein at an angle of much more than 45°;

ventral surface of the petiole broadly and smoothly

concave or channeled, the angles acute and continu-

ous; fruit 4-6 mm. in diameter, the fruiting disk 1.3-

1.5 mm. wide C, schiedeana Berg.

McVAUGH: TROPICAL AMERICAN MYRTACEAE 401

Plants rather generally pubescent on the young growth and the inflorescence,

often softly and loosely so, at least some of the hairs branched and spread-

ing or erect, often up to 0.5 mm. long, at least to some extent persistent

on the branchlets, the inflorescence and the fruits.

Leaves prevailingly obovate to elliptic, obtuse or rounded at tip, 3.5-

6.5 cm. long; branchlets not winged; small connecting veins evident

on the lower surface in dried leaves as a raised reticulum; pubescence

appressed, rather sparse, grayish- or yellowish-white; British Honduras.

C. cuneifolia Lundell.

Leaves prevailingly elliptic to lanceolate or ovate, acute or acuminate (if

small and obtuse the branchlets winged, the small veins obscure, the

pubescence reddish).

Leaves small and obtuse, mostly 2.5-4.5 cm. long, usually narrow, often

varying on the same plant from 2-5 (-7) times as long as wide;

branchlets persistently and prominently corky- winged; pubescence

sparse, evanescent, of rather loosely appressed yellow- or reddish-

brown hairs; Honduras to British Honduras . .C. hondurensis Standl.

Leaves usually larger and acute or acuminate, if narrow usually consist-

ently so; branchlets usually wingless.

Young leafy branchlets softly brown- to rufous-tomentose with hairs

of two types: straight and nearly erect, simple or short-branched

hairs up to 2 mm. long, these standing above the tomentum of

contorted and somewhat appressed hairs with two equal or

strongly unequal branches; hairs of the panicles of the contorted,

appressed type; buds 3 mm. long or less, obovoid, mostly short-

apiculate; branchlets not winged.

Leaves narrowly elliptic-lanceolate, 1-2.3 cm. wide, 4-6 times as

long as wide; Tabasco, British Honduras and northern Guate-

mala C. lindeniana Berg.

Leaves elliptic-ovate, 2.5-6 cm. wide, 2-3 times as long as wide.

Leaves mostly 4-6 cm. wide, 7-13 cm. long; pubescence pale red-

dish, reddish-brown or yellowish-brown; southern Mexico to

northern Colombia. .C. chytraculia var. americana McVaugh.

Leaves mostly 2-4 cm. wide, 5-10 cm. long; pubescence reddish

brown to dark red; Cuba and Jamaica.

C. chytraculia (L.) Sw., var. chytraculia.

Pubescence of leafy branchlets not as above, not noticeably different

from that of the panicle-branches.

Hairs coarse, 0.3-1.5 mm. long, erect and the branchlets tomentose,

or appressed but then the individual hairs 0.5-1 mm. long; buds

[as far as known] 4.5 mm. long or more; branchlets not wing-

angled.

Leaves broadly rounded at base; lateral veins impressed above;

petioles 8-11 mm. long; buds ovoid, short-apiculate, 4.5-

6 mm. long; tomentum dense, yellowish-brown, the very

numerous erect hairs simple or forked, up to 1.3-1.5 mm.

long; Guatemala C. macrantha Standl. & Steyerm.

Leaves acute at base; lateral veins not impressed above; petioles

4-7 mm. long; hairs various.

Branchlets bristly-tomentose, the very coarse hollow erect pale

yellowish-brown forked hairs up to 0.3-0.7 mm. high; buds

unknown, the calyptra shallowly dome-shaped, 2 mm. wide,

short-apiculate; Honduras C. zanquinensis A. Molina.

402 FIELDIANA: BOTANY, VOLUME 29

Branchlets clothed with appressed nearly sessile longitudinally

oriented golden-brown dibrachiate hairs 0.5-1 mm. long;

buds 9-14 mm. long, the trumpet-shaped calyptra 6-9 mm.

long; Panama C.johnstonii McVaugh.

Hairs minute, dibrachiate, often appressed, sometimes crisped,

mostly 0.1-0.3 mm. long, the pubescence of the young branch-

lets and inflorescence rusty-brown, yellowish brown or, especially

on the leaves, nearly colorless.

Peduncle 2.5-3 mm. wide just below the lowest branches; buds

4-4.5 mm. long; calyptra 2.5-3 mm. wide; hairs dark reddish

brown; Costa Rica, at 2600 m C. pittieri Standl.

Peduncle 1.5-2 mm. wide or less; buds 3 mm. long or less; calyptra

2.5 mm. wide or usually less; hairs coppery to yellow or gray-

white.

Branchlets wing-angled; leaves cuneate at base, often 4-7 cm.

long, usually glandular-punctate above; inflorescence copi-

ously soft-pubescent with reddish or sometimes pale hairs;

West Indies; southern Florida; Isthmus of Tehuantepec;

Yucatan C. pallens Griseb., var. pallens.

Branchlets terete or compressed, not wing-angled; leaves acute

at base, varying to somewhat rounded or occasionally to

cuneate; Mexico and Central America.

Inflorescence stout, the principal secondary branches of the

panicle much flattened, their lowest internodes often

6-8 mm. long, 1.5 mm. wide; pubescence of very small

flat closely appressed longitudinally oriented hairs; Vera-

cruz; British Honduras to Yucatan; [C. euryphylla

Standl.] C. millspaughii Urban.

Inflorescence more slender, the internodes of the secondary

panicle-branches terete or slightly compressed, 1 mm.

wide or less; pubescence, at least in part, of crisped or

soft hairs; mountains of Mexico and Guatemala.

Leaf-blades often 7-12 cm. long, eglandular above; petioles

8-10 mm. long; branchlets terete or compressed; pan-

icles often 50-60-flowered; inflorescence with often

densely aggregated coppery hairs about the base of

the hypanthium but otherwise rather sparingly pubes-

cent; mountains of Guatemala and southern Mexico.

C. pendula Berg.

Leaf-blades often 3-7 cm. long, often glandular-punctate

above but sometimes inconspicuously so; petioles 2.5-

6 mm. long; branchlets sometimes wing-angled.

Branchlets of the inflorescence copiously soft-pubescent

at least in the flowering stage, the hairs often pale

and the whole inflorescence appearing somewhat

grayish; leafy branchlets rarely if ever wing-angled;

panicles often 50-60-flowered; Costa Rica.

C. pallens var. williamsii (Standl.) McVaugh.

Branchlets of the inflorescence glabrous or sparingly

appressed-pubescent with minute coppery hairs;

leafy branchlets sometimes wing-angled; panicles

mostly 30-35-flowered; Mexico.

C. pallens var. mexicana (Lundell) McVaugh.

McVAUGH: TROPICAL AMERICAN MYRTACEAE 403

Calyptranthes chytraculia (L.) Sw. Prodr. 79. 1788. Myrtus

chytraculia L. Syst. Nat. ed. 10. 1056. 1759.

Known originally from the West Indies only, this species seems

to be represented in Central America by a slightly different taxon,

described below as a variety. The characteristic pubescence is so

nearly identical in all these plants, and so distinctive, that there can

be little doubt of close relationships among them. The mostly red-

dish-brown hairs are abundant, rather soft, those of the panicles

mostly much contorted, loosely appressed and with two equal or

strongly unequal branches; the young leafy branches bear similar

hairs, and bear also a number of fine, straight, nearly erect hairs up

to 2 mm. long, these simple or with a short branch near the base.

As far as I know, these two kinds of hairs, that give the young leafy

branches a unique shaggy appearance, are found only in C. chytra-

culia and in C. lindeniana Berg, which hardly differs except in its

very narrow leaves.

Some Central American specimens are virtually indistinguishable

from West Indian ones, but usually the leaves of Central American

plants are larger (4-6 cm. wide, 7-13 cm. long, as against 2-4 cm.

wide, 5-10 cm. long, in typical chytraculia). The hairs in West In-

dian specimens are often reddish-brown or dark red, whereas in

Central American plants the hairs are paler, varying from reddish-

brown to yellowish-brown or pale reddish. A specimen from north-

ern Colombia (region of Santa Marta, H. H. Smith 897}, agrees

perfectly with specimens from northern Central America. Miss

Amshoff, in the Flora of Panama (Ann. Missouri Bot. Card. 45:

169. 1958), referred a single Panamanian specimen to C. chytraculia

with the note that it "quite agrees with specimens from British

Honduras distributed as C. millspaughii Urb."

The plant of British Honduras and Central America has long

been confused with Calyptranthes millspaughii Urb. (Symb. Antill.

7: 294. 1912), which was based on Millspaugh 1537 from Cozumel

Island. This plant belongs to a quite different species. The pubes-

cence is scanty, of short flat appressed coppery hairs, not at all like

the shaggy abundant pubescence of chytraculia. The panicles are

4 cm. long or less, about 30-flowered, contrasting with the larger,

many-flowered panicles of chytraculia.

The taxonomy of C. chytraculia may be summarized as follows:

Calyptranthes chytraculia (L.) Sw., var. chytraculia.

Cuba and Jamaica; for synonymy and citation of specimens see

Urban, Bot. Jahrb. 19: 597-598. 1895.

404 FIELDIANA: BOTANY, VOLUME 29

Calyptranthes chytraculia (L.) Sw., var. americana McVaugh,

var. nov.

A var. chytraculia foliis maioribus 7-13 cm. longis, pilis pallidioribus pallide

rufis, vel rufo-flavidis usque ad fulvis, differt.

Near the Atlantic Coast of Central America, mostly at elevations

of 200 meters or less, often along rivers or lake shores; British Hon-

duras and Department of Pete"n, Guatemala, to northern Colombia.

Representative specimens examined :

BRITISH HONDURAS: Maskall, P. H. Gentle 1217 (MICH); Big

Fall, Belize River, Lundell 4343 (MICH) ; Cohune Ridge, El Cayo

Dist., Lundell 6431 (MICH); Temash River, W. A. Schipp 1295

(MICH).

GUATEMALA: Pete"n: Uaxactun, 4 Apr. 1931, H. H. Bartlett 12 W

(MICH, type; UC), 24 Apr. 1931, Bartlett 12723 (MICH).— Izabal:

Along Rio Frio, Steyermark 41632 (MICH).

NICARAGUA: Isla de Providencia, Proctor 3414 (US); Pearl La-

goon, Allen 6524 (F).

Calyptranthes hernandezii McVaugh, sp. nov.

Frutex 1-1.5 m. altus, ramulis bialatis, alis inter folia terminantibus; ramuli

paniculaeque tomentosae, pilis flaccidis flavidis saepe inaequaliter dibrachiatis

usque ad 1.3 mm. longis tectae; folia subsessilia, cordata, obtusa, 4-9 cm. longa,

2-3.7-plo longiora quam latiora; paniculae 5-6 cm. longae, 9-florae, floribus tri-

geminis sessilibus, alabastris 5.5-7 mm. longis apiculatis, calyptra 3-5 mm. longa,

3.5-4 mm. lata; stylus 10 mm. longus; stamina ca. 100.

A shrub 1-1.5 meters high, the branchlets bialate, the wings 0.5 mm. high,

terminating distally between the leaf-bases; branchlets and inflorescence loosely

tomentose with erect flaccid simple or unequally forked pale yellow-brown hairs

up to 1.3 mm. long; leaves coriaceous, glabrate, nearly sessile, lanceolate to oblong

or ovate on the same plant, 1.3-3 cm. wide, 4-9 cm. long, 2-3.7 times as long as

wide; blades narrowed to a broad or slender but always obtuse tip, cordate at base,

the stout petiole 1-2 mm. long, 2 mm. wide, ventrally concave; midvein broadly

and deeply concave above; lateral veins rather close and parallel, 10-20 including

some intermediate ones, inconspicuous, more evident beneath; marginal vein about

equaling the lateral ones and but slightly arched between them, 1-1.5 mm. from

the margin; blades green and sometimes impressed-punctate above, paler and with

convex resinous glands beneath; paired panicles 5-6 cm. long, 9-flowered, the

peduncle 3-4 cm. long, stout, compressed, up to 2.5 mm. wide; flowers sessile in

3's at the tip of a terminal branch 1.5 cm. long and at the tips of a pair of lateral

branches 8 mm. long, 2.5 mm. wide; bracts at the base of the flower-cluster boat-

shaped, scarious, deciduous at anthesis, enveloping the lateral bud, 6 mm. long;

bracteoles similar, deciduous, ovate, 3 mm. long; buds 5.5-7 mm. long, broadly

fusiform or the body subglobose and 4 mm. in diameter, the base tapering and

sessile, the tip bearing an apiculum 1-2.5 mm. long; calyptra domelike, 3-5 mm.

McVAUGH: TROPICAL AMERICAN MYRTACEAE 405

long including the apiculum; hypanthium after anthesis funnelform, 3.5-4 mm.

wide at the mouth; style about 10 mm. long; stamens about 100, 7-8 mm. long,

the anthers 0.5 mm. long; petals and fruit not seen.

MEXICO: San Luis Potosi: Near river edge, Taman, 15 km. south-

west of Tamazunchale, [elevation ca. 200 m.], 14 Apr. 1944, E. Her-

nandez X. 159 (LL, type).

Calyptranthes johnstonii McVaugh, sp. nov.

Arbor vel frutex, ramulis teretibus, ramulis foliisque juvenilibus et inflores-

centia copiose pilis flavidis, nitidis, dibrachiatis, 0.5-1 mm. longis, obsitis; folia

late elliptica, 6-11 cm. longa, basi acuta, petiolis 4-6 mm. longis, nervo medio

supra ad basin impresso; inflorescentiae geminatae, spicatae, 5-7 cm. longae,

pauciflorae; alabastra cornuta, 9-14 mm. longa, hypanthiis campanulatis, 3-5 mm.

longis; stylus 8-10 mm. longus; fructus globosus, glabrescens, ca. 1.5 cm. crassus.

Shrub or tree 2.5-10 m. high, the "bark on trunk and stems pale, moderately

smooth, not exfoliating" (Johnston); branchlets terete; inflorescence, and young

branchlets and leaves, copiously appressed-pubescent with lustrous golden-brown

dibrachiate hairs 0.5-1 mm. long; leaves coriaceous, broadly elliptic, about equally

narrowed to the prominently short-acuminate tip and the acute base, (2.5-)

3-5 cm. wide, 6-11 cm. long, the margins decurrent on the inner angles of the

channeled petiole 4-6 mm. long, 1-1.5 mm. thick; midvein impressed above near

base; lateral veins rather numerous and parallel, 12-15 pairs, not very conspicu-

ous (more so on the lower surface) ; marginal vein about equaling the lateral ones

and somewhat arched between them, 1-2 mm. from the margin; leaves dark green

and glandular-punctate above, sparingly pubescent and inconspicuously glandular

beneath; inflorescence paired at the terminal nodes of old branchlets, spicate,

5-7 cm. long; flowers few, probably 2-3 clustered at a node 2-4 cm. above the

base and 2-3 clustered at the tip of the spike; buds cornute, 9-14 mm. long, the

calyptra trumpet-shaped, 6-9 mm. long, 4 mm. wide at mouth, thence abruptly

contracted to a slender tapering tip 5-8 mm. long; hypanthium 3-5 mm. long,

campanulate, prolonged 2.5 mm. beyond the summit of the ovary; style 8-10 mm.

long; stamens about 50; petals none(?), or 1-2, narrow, 1-2 mm. long; fruit glo-

bose, glabrescent, about 1.5 cm. in diameter, surmounted by a collar about 4 mm.

wide, 1 mm. high; mature seeds not seen.

PANAMA: Canal Zone: South of Ft. Sherman, in wet places back

of the mangrove swamp, 1 Apr. 1956, 1. M. Johnston 1764 (MICH,

type, in flower), 1764 (MICH, isotype, with nearly mature fruits);

3 miles northwest of Gatun Locks, edge of marsh, 9 Aug. 1955, John-

ston 1571 (MICH); near Ft. Sherman, P. C. Standley 30933 (US).

Calyptranthes millspaughii Urb. Symb. Antill. 7: 294. 1912.

C. euryphylla Standl. Contr. U. S. Nat. Herb. 23: 1034. 1924.

Examination of the type of C. millspaughii, a flowering specimen

from Cozumel Island, Quintana Roo, Millspaugh 1537 (F), shows

this to represent a distinctive taxon, quite different from the copi-

UNIVERSITY OF

406 FIELDIANA: BOTANY, VOLUME 29

ously soft-hairy Central American plant that has in recent years

passed under the name of C. millspaughii. The more pubescent

plant is C. chytraculia var. americana McVaugh, well known along

the Atlantic coast from British Honduras to Colombia. The true

C. millspaughii seems to be known from but a single collection in

addition to the type; the second collection is also a flowering speci-

men, from British Honduras, Stann Creek District, Freshwater

Creek, J. Kelly no. 1 (F, US).

There is a strong superficial resemblance between C. millspaughii

and C. pendula Berg, and likewise between C. millspaughii and the

several varieties of C. pollens. From all of these C. millspaughii

may be distinguished by the broad, flat, short internodes of the pan-

icle branches. The hairs of the inflorescence of C. millspaughii are

distinctive also; they are copper-colored, longitudinally oriented, very

closely appressed and straight, the individual hairs readily observed.

In C. pendula and in the varieties of C. pollens the hairs are crisped

and often somewhat matted together.

The type and only known specimen of C. euryphylla Standl., col-

lected at Catemaco, Veracruz, in April 1894, Nelson 1+21 (US), bears

half -grown fruit; flowers and buds are wanting. The leaves and the

panicle-branches are sparingly pale-pubescent, the appressed hairs

very like those of C. millspaughii but almost colorless. Apparently

the paleness of the hairs is related to the maturity of the plant when

collected, for the hairs persisting on the fruit and on some of the dis-

tal nodes of the panicle are copper-colored and precisely like those

of C. millspaughii. The lower internodes of the lateral panicle-

branches in the type of C. euryphylla are 10 mm. long or less and

about 1.5 mm. wide, exactly comparable to those of C. millspaughii.

The foliage of C. euryphylla, as noted by Standley, is notable in that

the leaves are ovate and rather broadly rounded at base, and the

petioles are short; I suspect that this represents an individual dif-

ference. In other respects the leaves of euryphylla and millspaughii

are closely similar; both are impressed-punctate above, with im-

pressed midvein; the lateral veins are but indistinctly seen in the

rather opaque brownish lower surface of the rigid dried leaves.

Allowing for the differences between flowering and fruiting speci-

mens, there can be little doubt that C. euryphylla and C. millspaughii

are conspecific.

Calyptranthes pallens Griseb. Veg. Kar. 67. 1857. ?Eugenia

pollens Poir. in Lam. Encyc. Suppl. 3: 122. 1813 (fide Urban, Bot.

McVAUGH: TROPICAL AMERICAN MYRTACEAE 407

Jahrb. 19: 598. 1895). C. chytraculia 8 pauciflora Berg, Linnaea 27:

27. 1855.

A species of the Caribbean basin, the nomenclaturally typical

variety occurring in the West Indies, southern Florida, the Yucatan

Peninsula and the Isthmus of Tehuan tepee, Mexico; other varieties,

distinguished somewhat subjectively, occur in Mexico and Central

America.

A tree or tree-like shrub 5-8 m. high; leaves coriaceous, rather broadly elliptic,

or sometimes broadest a little above or below the middle, narrowed about equally

to both ends, 2-4 (-6) cm. wide, 3-7 (-10) cm. long, 2-3 (-3.5) times as long as

wide; margins of the blade decurrent at base on the inner angles of the channeled

petiole; midvein impressed above at least near base; lateral veins close and par-

allel, obscure on both surfaces, 10-20 pairs including some poorly denned inter-

mediate ones; marginal vein about equaling the lateral ones and slightly arched

between them, 0.5-1.5 mm. from the margin; leaves smooth and dark green above;

panicles usually bracteate and paired at the base of an otherwise leafy shoot,

sometimes axillary on ordinary leafy branches, the individual panicles pyramidal,

3-4 times compound, the flowers in small clusters near the tips, not appearing

closely sessile; peduncles 2-5 cm. long; buds 2.5-3 mm. long including the narrow

base, obovoid, rounded and often slightly apiculate; calyptra domelike; hypan-

thium after opening campanulate, 2 mm. long; style 4-5 mm. long; stamens about

50-60, up to 3.5 mm. long; petals tiny, adhering to the calyptra and falling with it;

ripe fruit dark red, globose or oblate, 5-8 mm. in diameter.

Young leafy branchlets 2-winged; leaves cuneate at base, usually glandular-

punctate above; inflorescence copiously soft-pubescent with reddish or some-

times pale hairs; panicles often 50-60-flowered; West Indies; southern Florida;

Isthmus of Tehuantepec var. pallens.

Young leafy branchlets not winged, or some plants with a few branchlets winged;

leaves acute at base, varying to somewhat rounded or occasionally to cuneate,

often glandular-punctate above but sometimes inconspicuously so; Mexico

and Central America.

Branchlets of the inflorescence copiously soft-pubescent at least in the flowering

stage, the hairs often pale and the whole inflorescence appearing somewhat

grayish; leafy branchlets rarely if ever winged; panicles often 50-60-flow-

ered; Costa Rica var. williamsii.

Branchlets of the inflorescence glabrous or sparingly appressed-pubescent with

minute coppery hairs; leafy branchlets sometimes winged; panicles mostly

30-35-flowered; Mexico var. mexicana.

Calyptranthes pallens Griseb., var. pallens.

Vegetative branches prominently 2-winged, the wings terminating distally

between the leaf-bases; inflorescence, branchlets, and to some extent the foliage,

rather persistently appressed-pubescent with pale rusty-brown, yellowish brown

or, especially on the leaves, nearly colorless dibrachiate hairs 0.1-0.3 mm. long;

leaves prominently acuminate at tip, broadly cuneate at base, usually glandular-

punctate above; petioles 6-8 mm. long; panicles 10-12 cm. long or less, usually

with 50-60 or more flowers, the lowest branches often as long as the peduncles,

the latter 2-4 cm. long; calyptra 1.3-1.7 (-2) mm. across.

408 FIELDIANA: BOTANY, VOLUME 29

West Indies and southern Florida. By Urban (Bot. Jahrb. 19:

598. 1895) reported from Cuba, Jamaica, Haiti, Cayman Islands,

St. Thomas, St. Croix and Guadeloupe. I have accepted Urban's

identification of this species but I have not seen the type. The

nomenclature of C. pollens is somewhat involved. The name is often

cited as C. pollens (Poir.) Griseb., with the implication that it was

based upon Eugenia pattens Poir. The latter is included by Urban

in the synonymy of C. pattens, but without the customary sign of

verification [!]. Grisebach, however, in his first publication of C. pal-

lens, made no mention of Eugenia pattens Poir., citing only C. chytra-

culia var. 8 pauciflora Berg. Berg's variety pauciflora was based

wholly upon a specimen collected in Guadeloupe by Bertero, and

studied by Berg in Sender's herbarium. Urban indicated with a [!]

that he had verified the identity of var. pauciflora, and that it was

indeed the same as C. pattens Griseb. The type of Eugenia pattens

Poir., however, was collected on the island of St. Thomas by Ledru;

it may well represent the same species, as supposed by Urban, but

it does not form part of the nomenclatural basis of Calyptranthes

pattens Griseb. The latter is readily identified from Grisebach's de-

scription in Fl. Brit. W. Ind. 233. 1860, as it is the only species with

2-edged branchlets and "cymes rusty-sericeous."

The following specimens are quite indistinguishable from West

Indian material of C. pattens, except that the petioles are no more

than 3-5 mm. long:

MEXICO: Oaxaca: Mena, C. D. Mell 7, 20 Jan. 1927 (US); Chi-

vela, Mell 2275, 1 Mar. 1934 (US) ; Rincon Antonio, C. R. Orcutt 3233

(F, US). — Yucatan: 3 km. de El Cuyo en el camino a Colonia Yuca-

tan, 0. G. Enriquez 479 (MICH).

Calyptranthes pallens Griseb., var. williamsii (Standl.)

McVaugh, comb. nov. C. williamsii Standl. Ceiba 1: 156. 1950.

?C. costa-ricensis Berg, Linnaea 27: 20. 1855.

Vegetative branchlets terete or slightly compressed; inflorescence, branchlets,

and to some extent the foliage rather persistently appressed-pubescent with pale

rusty-brown, yellowish brown or nearly colorless dibrachiate hairs 0.1-0.3 mm.

long; leaves acuminate at tip (sometimes obscurely so), the base usually not

cuneate but the margins narrowed and rounded to an acute base; blades incon-

spicuously if at all glandular, sometimes glandular-punctate above; petioles 4-

6 mm. long; panicles 6-8 cm. long, about 50-60-flowered, the lowest branches

shorter than the peduncles, the latter 3-5 cm. long; calyptra 2-2.5 mm. wide.

Mountain slopes, river banks and open pastures, at elevations of

800-2000 m., central Costa Rica, as far as known mostly in the prov-

inces of Cartago and San Jose", flowering March-April, fruiting in June.

McVAUGH: TROPICAL AMERICAN MYRTACEAE 409

A similarity between the West Indian pattens, and the species

described as C. costa-ricensis Berg, was suggested by Urban. Berg's

original description of "C. Costa-Ricensis" seems to apply satisfac-

torily to the plant described above, but I have not seen the type

("Costa Rica et Veragua, de Warszewicz, . . . v.s. in hb. Berol.")

and therefore prefer to base this new variety on a species that can

be satisfactorily typified.

Specimens examined:

COSTA RICA: Rio Maria Aguilar, J. D. Smith 6499 (F); Rio Ciru-

elas, massif du Barba, Tonduz 2211 (F) ; Sta. Maria de Dota, Standley

41716 (F); San Sebastian, Standley 32683 (F); San Jose", Stand-

ley 34816 (F), 47371 (F); Cerro de la Carpintera, Standley 34516

(F); Mt. Carpintera, H. E. Stork 314 P-P- (MICH); Alto de Ocho-

mogo, Williams & Molina 13841 (F; US, type), Williams 16001

("probably from same tree from which type came," MICH) ; Cerro

Carpintera, above Tres Rios, Williams 16141 P-P- (MICH); La Car-

pintera, Cerro del Aguacate, Williams 16579 (MICH); along Rio

Sucio near Corinto, Williams 16555 (MICH).

Calyptranthes pallens Griseb., var. mexicana (Lundell)

McVaugh, comb. nov. C. mexicana Lundell, Wrightia 2: 166.

1 May 1961.

Vegetative branches winged or wingless, the wings sometimes apparent only

on a few of the young branchlets of individual plants; inflorescence and branchlets

very sparingly pubescent, the appressed and usually coppery hairs mostly 0.1-

0.2 mm. long, and aggregated chiefly about the base of the hypanthium and the

distal nodes of the panicle; leaves acuminate at tip (sometimes obscurely so or

some leaves merely obtusely rounded), acute or sometimes cuneate at base, often

glandular-punctate above; petioles 2.5-5 mm. long; panicles 3-7.5 cm. long, 30-

35-flowered, the lowest branches shorter than the peduncles, the latter 2-4.5 cm.

long; calyptra 2-2.5 mm. wide.

Arroyos, brushy slopes, stream-sides, wet mountain forests,

southern and western Mexico, at elevations from ca. 500 m. to

2600 m.

MEXICO: Chiapas: Mt. Ovando, near Escuintla, E. Matuda 1838

(LL, type; MO); Tres Cruces, 2600 m., Matuda 5024 (F; LL, para-

type). — Jalisco: Pihuamo, M. E. Jones 18 (F, US); 10 miles south

of Autlan, Wilbur & Wilbur 1415 (MICH); San Sebastian, Hda.

del Ototal, Y. Mexia 1806 (MICH, US).— Nayarit: 8 miles west of

Tepic, McVaugh 18903 (MICH); Maria Madre Island, T. S. Maltby

110 (US).— -Sinaloa: Without collector or locality (US 1014144);

Ixtagua, Mun. San Ignacio, J. Gonzalez Ortega 776 (MEXU).

410 FIELDIANA: BOTANY, VOLUME 29

The following specimen, superficially differing from the preced-

ing ones in its long internodes and rather narrow and slenderly

acuminate leaves up to 10 cm. long and 3-4 times as long as wide, is

apparently merely a flowering stump-sprout or other vigorous shoot:

MEXICO: Sinaloa: San Ignacio, M. Narvaez M. & A. E. Salazar

846 (US).

Calyptranthes paxillata McVaugh, sp. nov.

Arbor parva, ramulis subteretibus vel compressis; ramuli inflorescentiaeque,

et folia juvenilia, pilis rufidulis appressis dibrachiatis subsessilibus longitudinaliter

directis 0.5-1 mm. longis obsiti; folia elliptica vel elliptico-lanceolata, caudato-

acuminata, 4-7 cm. longa, basi acuta vel cuneata, petiolis 3-6 mm. longis, nervo

medio supra versus basin canaliculate; inflorescentiae 3-4.5 cm. longae, ca. 9-florae,

e basi ramulorum foliosorum vel abortivorum geminati oriundi; pedunculi 3.5 cm.

longi, statu frutescenti 1 mm. crassi; alabastra obovoideo-fusiformia, 4.5-5 mm.

longa, apiculo obtuso 1-1.3 mm. longo paxillato; stylus 7-8 mm. longus.

A small tree with nearly terete or compressed branchlets; branchlets, inflores-

cence and young foliage covered with appressed, nearly sessile, dibrachiate and

longitudinally oriented coppery hairs 0.5-1 mm. long; leaves glabrate, elliptic or

elliptic-lanceolate, 1.5-3.5 cm. wide, 4-7 cm. long, 2.3-3 times as long as wide,

caudate-acuminate, the slender tapering tip 1-1.5 cm. long, obtuse; base of blade

acute or cuneate, the margins decurrent on the inner angles of the petiole 3-6 mm.

long, 0.8-1.2 mm. thick; midvein channeled above near base; lateral veins obscure,

close and parallel, about 15 on each side, including some intermediate ones; mar-

ginal vein about equaling the lateral ones, nearly straight between them, 1 mm.

from the margin; inflorescence of paired panicles from an abortive axillary axis or

from the lowermost (bracteate) node of a leafy shoot; panicles 3-4.5 cm. long,

twice compound, bearing 9 flowers or fewer; peduncle up to 3.5 cm. long, com-

pressed, 1 mm. wide below the lowest branches; buds obo void-fusiform, 4.5-5 mm.

long, 2-2.5 mm. in diameter, broadest above the middle, tapering to the base,

rather abruptly contracted to a blunt peglike apiculum 1-1.3 mm. long; hypan-

thium after anthesis funnelform, 2.5-3 mm. high, the rim prolonged 2 mm. beyond

the summit of the ovary; calyptra about 3 mm. across, 2.5 mm. long including the

apiculum; style 7-8 mm. long; stamens about 100, up to 4 mm. long; petals 1-2,

minute, remaining attached to the calyptra and falling with it; fruit unknown.

GUATEMALA: Alta Verapaz: Wet forest above Tactic, above the

bridge across Rio Frio, alt. 1400-1500 m., 30 Mar. 1941, P. C. Stand-

ley 90457 (US, type).

This species is distinctive by virtue of its copious appressed

pubescence of long hairs, the few-flowered panicles, long styles and

long-apiculate fusiform buds. This peculiar "peglike" apiculum

gives the name to the species; I do not find the word paxillatus in

any Latin dictionary at my disposal, but as the word "paxillate" may

be found in English dictionaries, it seems reasonable to provide a

Latin equivalent for it.

McVAUGH: TROPICAL AMERICAN MYRTACEAE 411

Calyptranthes schiedeana Berg, Linnaea 27: 28. 1855. Myr-

cia aromatica Schlecht. Linnaea 13: 415, pro max. part. 1839. Not

C. aromatica St. Hil.

This species and the next were based by Berg upon the two col-

lections assigned by Schlechtendal to Myrcia aromatica. The type

of C. schiedeana was from Hacienda de la Laguna, Veracruz, Schiede

(in herb. Schlecht.), and was presumably the collection upon which

M. aromatica was chiefly based. Several collections by C. A. Purpus

(e.g. nos. 2927, 10778), all from Veracruz, seem to represent the

same species as a presumed isotype shown in Field Mus. Neg. 23392.

Calyptranthes schlechtendaliana Berg, Linnaea 27: 29. 1855.

Myrcia aromatica Schlecht. Linnaea 13: 415, quoad spec, in sylvis

Papantlae coll. 1839.

The type of C. schlechtendaliana was Schiede no. 25 (in herb.

Schlecht.), collected at Papantla, Veracruz, and presumably the

same specimen mentioned by Schiede on page 416 as similar to his

Myrcia aromatica, described on the previous page. A photograph

purporting to be C. schlechtendaliana (Field Mus. Neg. 36864) seems

actually to represent a specimen of C. schiedeana.

One collection, Purpus 8810 (UC, US), from Veracruz without

definite locality, seems to represent C. schlechtendaliana. The dif-

ferences between it and C. schiedeana are listed in the key above.

As far as can be determined from the scanty material at hand, the

two species are distinct.

Calyptranthes tenuipes McVaugh, sp. nov.

Frutex, ramulis teretibus; ramuli teneri densiuscule pilis crassis, contortis,

nitidis, fuscis, dibrachiatis 0.5-0.8 mm. longis obsiti; folia ab initio subglabra,

lanceolato-ovata, caudato-acuminata, 3-5 cm. longa, basi rotundata, petiolis 2 mm.

longis, nervo medio supra impresso; inflorescentiae 3-4 cm. longae, ut videtur

3-5-florae, e basi ramulorum foliosorum geminati oriundi; pedunculi filiformes,

statu fructescenti 0.3-0.4 mm. crassi; flores non vidi; fructus globosus, 7-9 mm.

crassus, umbilicatus; seminis maturi cotyledones carnosi, contorto-plicati.

A shrub 2 m. high, glabrate, the branchlets terete; young branchlets, and to a

lesser extent the very young inflorescence, more or less thickly pubescent with

coarse contorted lustrous reddish-brown dibrachiate hairs 0.5-0.8 mm. long; leaves

glabrous from the first except for a few hairs on the lower surface near the mid-

vein, the blades lance-ovate, caudate-acuminate, 1-2.3 cm. wide, 3.5 cm. long, the

narrow acumen 1-1.8 cm. long, rounded and 1-2 mm. wide at its tip; base of blade

rounded, the margins passing abruptly onto the inner angles of the deeply sulcate

petiole 2 mm. long, 1 mm. thick; midvein impressed and sulcate above, the other

veins obscure, the lateral ones about 10 pairs, the marginal vein about 1 mm. from

412 FIELDIANA: BOTANY, VOLUME 29

the margin; blades dark green and nearly featureless above, paler beneath, the

glands obscure; inflorescences 3-4 cm. long, paired at the base of a leafy shoot;

peduncle 2-2.5 cm. long, filiform, 0.3-0.4 mm. thick in fruit; flowers unknown,

probably 3-5, on pedicels or lateral branches up to 3 mm. long; fruit globose,

7-9 mm. in diameter, surmounted by a collar about 2 mm. wide, less than 1 mm.

high; seed 1, globose, lustrous, up to 8 mm. in diameter; cotyledons contorted.

In the absence of flowers this species is assigned to the genus

Calyptranthes on the basis of the contorted cotyledons, the collar-

like remains of the calyx suggesting that the buds open by a calyptra,

and the dibrachiate hairs characteristic of Calyptranthes but not

found in other genera of the Myrcioideae.

MEXICO: Veracruz: In a deep barranca below Atzalan, elev. ca.

1650 m., 13 Mar. 1946, A. J. Sharp 46165 (MICH, type).

Calyptranthes zuzygium (L.) Sw. Prodr. 79. 1788. Myrtus

zuzygium L. Syst. Nat. ed. 10. 1056. 1759.

A West Indian species, known from Hispaniola, Jamaica and

Cuba; also in southern Florida. Reported from "Michoacan and

Tres Marias Islands" by Standley (Contr. U. S. Nat. Herb. 23: 1034.

1924) ; judging from specimens annotated by Standley, these reports

are based upon M. E. Jones 18, from Pihuamo, Jalisco (not Micho-

acan, as labeled by Jones), and T. S. Maltby 110, from Maria Madre

Island. I should refer these specimens to Calyptranthes pattens var.

mexicana, q.v.; C. zuzygium is so far unknown in Mexico.

MISCELLANEOUS NOTES

Calyptranthes tovarensis (Berg) Steyermark, Fieldiana, Bot.

28: 1010. 1957. Myrcia tovarensis Berg, Linnaea 27: 118. 1855.

This name resulted from a misinterpretation of Myrcia tovarensis,

and a misidentification of Tamayo 227, which is indeed not a Myrcia

but a species of Calyptranthes. The type of M. tovarensis, however

(Moritz 17 If?}, represents an undoubted and distinctive species of

Myrcia,. In addition to the type collection, which I have seen at

Chicago Natural History Museum, the following specimens from

Venezuela seem to be representative of M . tovarensis: Colonia Tovar

[the type locality], Fendler 42 (PH, NY); Caracas, Linden 13 (F).

Calyptranthes vexata McVaugh, nom. nov. C. apoda McVaugh,

Mem. N. Y. Bot. Card. 10: 70. 1958, not C. apoda Urban, Symb.

Antill. 9: 96. 1923. I am indebted to Dr. Richard S. Cowan for

calling this nomenclatural lapse to my attention.

McVAUGH: TROPICAL AMERICAN MYRTACEAE 413

2. EUGENIA L.

SYNOPSIS OF THE MEXICAN SPECIES WEST OF THE

ISTHMUS OF TEHUANTEPEC

In 1924, in Standley's Trees and Shrubs of Mexico, 24 species of

Eugenia were recorded; two additional species now known to belong

to Eugenia (Myrtus oaxacana and M. ledophylla) were also included

in this work but under the genus Myrtus. Now after almost four

decades of active botanical exploration in Mexico, approximately

60 species of Eugenia are known to occur as native plants in that

country. The following synopsis includes the approximately 40 spe-

cies known from west and north of the Isthmus of Tehuantepec.

The myrtaceous flora of Chiapas and Tabasco, and that of the Yuca-

tan peninsula, is quite a different one and as yet imperfectly known.

Essentially all the known species are treated in the Flora of Guate-

mala (Fieldiana, Bot. 24). From western Chiapas and Tabasco

very few collections are available; it may be supposed that some

new species will come to light from this region, and that some addi-

tional species will be found to extend into the region from the isth-

mus. Eventually it may be convenient and useful to treat all

Mexican species in one publication, but it is expedient in the pres-

ent state of knowledge to treat as a unit what may be called the

endemic Mexican flora, as distinct from that with Central American

or West Indian affinities.

It is difficult to make a generally useful key to any large group

of species of Myrtaceae, for several reasons. Vegetative similarities

throughout the family are so great that sterile specimens may be

quite unrecognizable even to one familiar with many species. Floral

morphology varies little from one species to another. The most use-

ful "recognition characters" (i.e., those possible to use in a key) are

those of the inflorescence and those of pubescence. The inflorescence

in Eugenia is almost always an unbranched axillary axis bearing one

or more opposite and decussate pairs of pedicellate flowers. In any

one species the length of this "raceme," the number of pairs of

flowers, and the characteristics of the bracts and pedicels, are con-

stant within rather narrow limits. Unfortunately for the taxono-

mist, almost all the species abandon on occasion their characteristic

modes of flowering. Apparently when stimulated to vigorous growth

almost any axillary branch may develop not into a short leafless and

completely fertile raceme, but into a leafy twig bearing normal pe-

dicellate flowers at its lower nodes. Often the lowermost one or two

nodes bear small deciduous bracts instead of ordinary green leaves.

414 FIELDIANA: BOTANY, VOLUME 29

When the raceme becomes a leafy branch in this way, the lowermost

flowers on casual inspection may seem to be borne at leafless nodes,

and the upper flowers solitary and pedicellate in the axils of ordinary

green leaves. Obviously a key based on the morphology of the in-

florescence must be interpreted with care.

Pubescence in Eugenia provides a series of characters most use-

ful to the taxonomist. Again, however, this is of limited use because

the pubescence, if present at all, usually disappears soon after the

plants flower, at least before the fruit matures. It would be quite

feasible to construct a key to flowering material, based largely on

characters of the pubescence; fruiting material of the same species

would be quite indeterminable by the use of the same key.

The following synopsis thus does not pretend to present a me-

chanically perfect key to all the species involved. The plan is the

same as that of the key in the Myrtaceae of Guatemala; those inter-

ested in the exceptions and qualifications of such a key may consult

that work.

KEY TO THE SPECIES

Species that are noticeably appressed-pubescent or tomentose (at least at flower-

ing time), and have at the same time relatively slender and elongate racemes

(the pairs of flowers well separated).

Bracteoles thin and conspicuous, mostly glabrous or nearly so and contrasting

with the heavily pubescent hypanthium, persistent in flower and fruit, 0.7-

1.5 mm. long, usually connate along the proximal margins, involucre-like;

pubescence copious, appressed, of pale sordid simple hairs.

Bracteoles acute or acuminate, broadly ovate-deltoid, 1-1. 5 mm. long; bracts

thin, persistent, the upper ones often much the longest and 1-2 mm.

long; raceme-axis usually more than 1 cm. long, with well-separated

nodes; lateral veins 10-20 on each side of the mid vein [group of Eugenia

biflora] E. karunnskyana.

Bracteoles obtusely pointed or rounded, less than 1 mm. long; bracts ovate-

deltoid, somewhat indurated at base, less than 1 mm. long; axis of the

raceme rarely as much as 1 cm. long, the nodes approximate; lateral

veins 4-6 (-8) on each side E. argyrea.

Bracteoles not conspicuous, usually pubescent like the hypanthium and the

pedicel, persistent (and then somewhat thickened at base), or deciduous

at an thesis, not connate and involucre-like even if persistent; pubescence

various, the hairs often reddish and often dibrachiate.

Leaves narrowly lanceolate or oblanceolate, 1 cm. wide or less, (3-) 5-8 times

as long as wide; hairs reddish, appressed, dibrachiate; region of Acapulco,

Guerrero E. avicenniae.

Leaves broader, mostly 2-4 (-7) cm. wide, 1.5-3 times as long as wide.

Herbage and inflorescence densely hirsute-tomentose with dark reddish-

brown erect hairs about 0.5 mm. long, these at flowering time quite

obscuring the surfaces; raceme 1-1.5 cm. long; disk 2.5 mm. wide;

blades 10-14 cm. long; petioles 10-15 mm. long; Veracruz.

E. praeterita.

McVAUGH: TROPICAL AMERICAN MYRTACEAE 415

Herbage and inflorescence paler and often sparingly hairy, the hairs cop-

pery red, yellowish red, yellowish white or silvery, often closely

appressed or loosely contorted and somewhat matted.

Flowers small; disk in flower mostly 1.3-2 mm. wide; stamens 50-75;

larger calyx-lobes 1.5-2.5 mm. wide; leaves broadly elliptic to ovate

or obovate, thinly pubescent with pale appressed hairs, 1.6-2 (-2.5)

times as long as wide; petioles 4-6 mm. long; raceme-axis usually

less than 5 mm. long; Pacific lowlands of Mexico, to Guatemala.

E. rekoi.

Flowers larger; disk in flower 2-3 (-5) mm. wide; stamens 100-200;

larger calyx-lobes up to 2.5-4 mm. wide and long; raceme-axis

1-2.5 (-8) cm. long, seldom shorter.

Bracteoles deciduous at anthesis or before, 2.5-5 mm. long; racemes

mostly 2-4 cm. long; pubescence often of loosely matted and

contorted hairs, yellowish white to pale reddish brown.

Leaf-margins cuneately decurrent on the petiole, the blades often

appearing acute at base; leaves soon glabrate, usually glabrous

beneath or nearly so, even at flowering time; hairs of the inflores-

cence sparse, appressed E. salamensis var. rensoniana.

Leaf-margins abruptly contracted to the petiole, the base of the

blade rounded or subauriculate, often asymmetric; leaves

densely and persistently pubescent or tomentulose beneath;

hairs of the inflorescence abundant, stalked, forming a loose

tomentum E. salamensis var. salamensis.

Bracteoles persistent through anthesis and usually much longer; axis

of the raceme usually less than 2 cm. long; pubescence closely

appressed, silvery to pale reddish or yellowish.

Leaves permanently silvery-scurfy beneath, obovate or obovate-

elliptic, rounded at apex; mid vein flat or convex above; axis of

raceme usually less than 1 cm. long E. hypargyrea.

Leaves at maturity glabrous or with fine scattered pale hairs beneath

(when young sometimes with a silvery coating), mostly elliptic

with acuminate tip; mid vein impressed or channeled above;

axis of the raceme usually 1-2 cm. long.

Leaves broadly elliptic, 1.6-2 times as long as wide, 2.5-5.5 cm.

wide, abruptly short-acuminate; style 7-8 mm. long; larger

calyx-lobes 3.5 mm. long and wide; fruit with 8 prominent

wing-angles, ellipsoid, 1 cm. in diameter, 2 cm. long; Jalisco

and Nayarit E. pleurocarpa.

Leaves narrowly elliptic, (2-) 2.3-3 times as long as wide, 1.5-

2.5 cm. wide, gradually acuminate; style 6-7 mm. long;

larger calyx-lobes 2.5 mm. long and wide; fruit smooth, not

winged, oblong or obovoid, 1 cm. long or a little more; state

of Mexico and Oaxaca; to Central America. .E. guatemalensis.

Species either glabrous or nearly so at flowering time, or the hairs small and up-

standing, or if the plants heavily pubescent or tomentose then the racemes

short (usually less than 1 cm. long) and the nodes approximate.

Racemes slender and elongate, 1-2.5 (-4) cm. long; plants glabrous or nearly

so at maturity.

Plants glabrous; midvein (when dry) elevated on the upper leaf-surface;

Colipa, Veracruz, known from the type only E. colipensis.

Branchlets and petioles minutely hispidulous, the tender growth bearing tiny

appressed coppery dibrachiate hairs also, a few of the latter persisting

416 FIELDIANA: BOTANY, VOLUME 29

near the base of the lower leaf-surface; mid vein impressed above, spar-

ingly hispidulous in the furrow; a widespread species. . . .E. oerstedeana.

Racemes short and crowded, if more than 1 cm. long then with numerous approx-

imate nodes; flowers often fasciculate or glomerate, sometimes solitary at

the lowermost nodes of new leafy branchlets and on the same plant in

abbreviated racemes.

Cauliflorous species, the flowers mostly long-pedicellate in clusters at usually

leafless nodes on old wood, the axis of the raceme almost none, or 1-3 mm.

long; coarse plants with relatively large leaves, flowers and fruits (leaves

7-20 cm. long; disk 2.5-6 mm. wide; pedicels (3-) 8-30 mm. long; fruit

1.5-2.5 cm. long.

Leaves cordate-auriculate at base.

Pedicels very stout, 3 mm. long; petioles 8-10 mm. long; buds 10-11 mm.

long, the hypanthium canescent; disk 6-7 mm. wide; San Luis Potosf.

E. xilitlensis.

Pedicels slender, 8-10 mm. long; petioles 5-7 mm. long; buds 5-6.5 mm.

long, the hypanthium glabrous or the base somewhat pubescent;

disk 3.5 mm. wide; Veracruz E. trunciflora.

Leaves acute to cuneate at base, or somewhat rounded to a cuneately de-

current base.

Midvein broad and flat or convex above (sometimes with narrow median

channel near base); lateral veins not impressed; plants essentially

glabrous; disk 3.5-4 mm. wide; larger calyx-lobes 2.5 mm. long,

3-3.5 mm. wide; buds 4.5-6 mm. long; Oaxaca to Guatemala.

E. choapamensis.

Midvein deeply and narrowly impressed above; lateral veins depressed

below the upper surface but the veins and veinlets convex, in a reticu-

late pattern; herbage and inflorescence sparingly puberulent with

tiny erect hairs; disk 3.5 mm. wide; larger calyx-lobes 6 mm. long,

5 mm. wide; buds 8 mm. long; Chiapas and Tabasco. .E. teapensis.

Species not cauliflorous, the inflorescences usually in the axils of new leaves,

if occasionally at leafless nodes then the raceme-axis prolonged or the

pedicels very short, or both; flowers relatively small, the calyx-lobes at

most 3.5-4 mm. long and wide, usually much less; disk at most 3.5 mm.

wide, usually much less.

Plants abundantly and conspicuously hairy at flowering time, the inflores-

cence and usually the branchlets hairy, setose, appressed-pubescent or

tomentose, the hairs often 2-branched (the branches equal or very

unequal); small erect hairs if present few, or obscured by the other

pubescence Key A, page 416.

Plants glabrous, puberulent or minutely hispidulous, the hairs if present

0.1-0.3 mm. long, often incurved (bracts and flower-parts sometimes

longer-ciliate, or the branchlets with slightly longer hairs, but the

leaves then essentially or quite glabrous) ; dibrachiate hairs none except

in one species Key B, page 418.

KEY A

Appressed dibrachiate hairs numerous on the branchlets and pedicels (one branch

of the hairs often much longer than the other); bracteoles and calyx-lobes

usually pubescent to about the same extent as the hypanthium.

Flowers glabrous or nearly so; plants as a whole soon glabrous, the youngest

branchlets and foliage bearing small evanescent partly dibrachiate hairs;

flowers usually in slender elongate racemes E. oerstedeana.

McVAUGH: TROPICAL AMERICAN MYRTACEAE 417

Flowers and pedicels appressed-pubescent, usually densely so; hairs more or

less persistent, the plants in age eventually glabrate.

Leaves narrowly lanceolate or oblanceolate, 1 cm. wide or less, (3-) 5-8 times

as long as wide; region of Acapulco, Guerrero E. avicenniae.

Leaves broader, 1.5-3 times as long as wide, usually much more than 1 cm.

wide.

Species with small, relatively broad, short-petiolate leaves [blades 5-7 cm.

long or often less, 1.5-2.5 (-3) times as long as wide; petioles 1-6 mm.

long].

Leaves more narrowly elliptic, less than 2 cm. wide, 2-3 times as long as

wide; petioles 1-2 mm. long; larger calyx-lobes up to 3 mm. long

and wide; Sinaloa only E. inconspicua.

Leaves broader, 1.5-4 cm. wide, 1.5-2.5 times as long as wide.

Leaves blunt or obscurely acuminate; petiole 3 mm. long; flowers

larger (disk 2.5-3 mm. wide; style 5-6 mm. long or more; larger

calyx-lobes 2.5-3 mm. wide); Morelos; Mexico; Veracruz.

E. yautepecana.

Leaves prominently or sometimes obscurely acuminate (on short

branches sometimes blunt); petioles 4-6 mm. long; flowers smaller

(disk 1.3-2 mm. wide; style 4-4.5 mm. long; larger calyx-lobes

1.5-2 mm. wide); Pacific lowlands, Sinaloa to Oaxaca; Guatemala.

E. rekoi.

Species with larger leaves and flowers and usually longer racemes (blades

5-9 cm. long; petioles mostly 5-10 mm. long; disk 2-3 mm. wide or

more; calyx-lobes 2.5-4 mm. long and wide; raceme mostly 1-2 cm.

long) ; ordinarily to be sought in the key above (E. hypargyrea, E. gua-

temalensis, E. pleurocarpa) See key above, page 415.

Hairs of the branchlets and pedicels not dibrachiate, either simple and antrorsely

appressed or spreading-ascending, or sometimes loosely contorted or erect;

hypanthium often heavily pubescent and then often contrasting with the

more nearly glabrous bracteoles and calyx-lobes.

Flowers glomerate, sessile or nearly so, the pedicels and the axes of the racemes

usually concealed by the clustered flowers; plants often somewhat hirsutu-

lous, at least in the inflorescence.

Leaves elliptic-oblong, rounded at apex and usually at base also, 1.6-2 times

as long as wide, 3.5-6 (-8?) cm. long, short-petiolate; herbage rather

generally short-hirsutulous; disk in fruit 1.6-1.7 mm. wide; bracteoles

persistent, not connate, broadly rounded, 1-1.5 mm. wide, 1 mm. long;

Sinaloa E. sinaloae.

Leaves acute or acuminate, sometimes bluntly acuminate.

Herbage at maturity nearly glabrous (or the mid veins pilose), the inflores-

cence hirsute with coarse, straight or contorted, coppery hairs up to

0.5-0.8 mm. long; calyx-lobes and the connate bracteoles nearly gla-

brous except the margins; leaves 3-7 cm. long, the petioles 5-8 mm.

long; Sierra Madre Oriental, Veracruz to Tamaulipas.

E. symphoricarpos.

Herbage and inflorescence abundantly and similarly pubescent or hirsutu-

lous.

Petioles 3-4 mm. long; herbage and inflorescence tawny-hirsutulous with

erect hairs 0.5-1 mm. long; bracteoles distinct, ovate, acute, 0.5-

1 mm. long; flowers small, the disk 1-1.5 mm. wide; eastern Mexico

to Panama E. origanoides.

418 FIELDIANA: BOTANY, VOLUME 29

Petioles 5-12 mm. long; bracteoles 1-2 mm. long, connate at base and

forming a broad 2-lobed involucre investing and often concealing the

hypanthium; flowers somewhat larger.

Young herbage and inflorescence closely appressed-pubescent with pale

sordid hairs; leaves abruptly acuminate; mid vein deeply impressed

above; calyx-lobes pubescent on both sides; disk 2.5 mm. wide;

Oaxaca E, oaxacana.

Inflorescence appressed-pubescent, the branchlets, petioles, leaf-mar-

gins and midvein (beneath) short-hispidulous with erect hairs;

leaves bluntly deltoid-acuminate; midvein shallowly concave in

the nearly flat blade; calyx-lobes glabrous within except at tips;

disk 1.5 mm. wide; western Michoacan E. alnifolia.

Flowers evidently pedicellate, the raceme-axis usually evident also but some-

times very short.

Flowers solitary in the axils of foliage-leaves, slender-pedicellate; pubescence

gray to white; Veracruz E. ledophylla.

Flowers in short or elongate bracteate racemes.

Bracteoles thin and conspicuous, glabrous or nearly so and contrasting with

the canescent hypanthium; flowers evidently slender-pedicellate.

Bracteoles acute or acuminate, 1-1.5 mm. long; pubescence gray to white;

flowers mostly in racemes 2-4 cm. long [the group of Eugenia biflora}.

E. karwinskyana.

Bracteoles obtuse or rounded, mostly less than 1 mm. long; flowers mostly

in racemes 2-6 mm. long, with several approximate nodes; young

herbage coppery-pubescent, the hairs of the inflorescence coppery or

yellowish E. argyrea.

Bracteoles usually not contrasting with the hypanthium, either glabrous or

pubescent, if glabrous the hypanthium also glabrous or nearly so, or

the flowers sessile; flowers in congested racemes mostly less than 1 cm.

long.

Branchlets and petioles setose-hispid with pale sharp erect hairs 0.3-

0.5 mm. long (the inflorescence sparingly so); hypanthium glabrous,

after flowering prolonged into a narrow neck 0.5 mm. high, 1 mm.

in diameter; longer calyx-lobes deltoid-ovate, 1.5-2.3 mm. long.

E. principium.

Pubescence of herbage and inflorescence appressed, or in addition with

numerous very short erect hairs 0.1-0.3 mm. long.

Calyx-lobes 1.5-2 mm. long, membranous, reflexed from the base soon

after an thesis; raceme-axis 2-4 mm. long, the pedicels filiform,

8-12 mm. long; bracteoles distinct, scarious; midvein slightly en-

graved above in the nearly plane blade; plants nearly glabrous,

the inflorescence and young herbage sparingly appressed-pubes-

cent; coastal Michoacan E. turneri.

Calyx-lobes 1-1.5 mm. long or less, neither evidently membranous nor

strongly reflexed; pedicels mostly shorter; bracteoles usually some-

what united by the proximal margins, involucre-like; midvein

concave or sulcate. [Here are to be found exceptionally pubescent

individuals of E. axillaris or E. acapulcensis.]

See key below, page 421.

KEY B

Plants nearly glabrous, the very young herbage with small coppery appressed

dibrachiate hairs, a few such hairs persisting on the lower leaf-surface and

McVAUGH: TROPICAL AMERICAN MYRTACEAE 419

the mid vein near base; flowers at least partly in racemes 1-2.5 (-4) cm. long;

bracteoles broad, membranous, persistent, up to 1.5 mm. long and wide;

widespread in Mexico and Central America E. oerstedeana.

Dibrachiate hairs none; flowers never in long slender racemes.

Flowers large and nearly sessile (disk 3-4 mm. wide; larger calyx-lobes 3.5-

4 mm. wide; style 7-9 mm. long; pedicels 0.5 mm. thick, 1.5 mm. long or

less); plants glabrous; Veracruz E. mexicana.

Flowers much smaller, or the pedicels slender and 6-10 mm. long.

Pedicels evidently slender and elongate, 6-16 mm. long; bracteoles distinct,

scarious; calyx-lobes foliaceous or scarious, more or less reflexed in an-

thesis; staminal ring bristly; flowers fasciculate, 1-3 pairs from a very

short axis.

Flowers large (disk 2.5-3.5 mm. wide; larger calyx-lobes up to 3.5 mm.

wide); Veracruz to Chiapas; primarily West Indian E. rhombea.

Flowers small (disk 1.5 mm. wide; larger calyx-lobes up to 2 mm. wide);

western Michoacan E. turneri.

Pedicels usually 5 mm. long or less (the lower ones in E. acapulcensis often

5-9 mm. long); calyx-lobes mostly somewhat thickened near base, erect

and imbricate.

Leaves 1.4-4 cm. long, oblong-linear, 6-8 times as long as wide, 2-4 mm.

wide; plants glabrous; Oaxaca E. standleyi.

Leaves broader and usually much longer, if only 2-4 cm. long then mostly

2-3 (-4) times as long as wide.

Leaves small, 1-2.5 cm. long, 3-10 mm. wide, obovate to ovate or elliptic,

rounded or obtusely pointed at apex; midvein slightly concave above,

at least near base; herbage and inflorescence minutely hispidulous-

puberulent; Tamaulipas to Oaxaca E. liebmannii.

Leaves usually larger, generally of an elliptic or a lanceolate type, nar-

rowed toward the acuminate or sometimes blunt apex.

Leaves narrowly lanceolate or linear-lanceolate, 1 cm. wide or less,

6-8 times as long as wide; Oaxaca, Tabasco E. lindeniana.

Leaves mostly 2-3 (-4) times as long as wide.

Leaves flat in drying, the midvein not depressed beneath the general

level of the upper surface of the blade, the vein itself flat or

slightly elevated or convex, sometimes shallowly channeled

especially near the base; leaves often opaque and featureless

above, the veins scarcely or not at all apparent.

Pedicels 3-5 mm. long; fruit about 5 mm. in diameter; hypanthium

usually finely puberulent; base of the style glabrous; eastern

Mexico to Honduras, little known from western Mexico.

E. capuli.

Pedicels 1-2.5 (-4) mm. long; fruit about 6-8 mm. in diameter;

hypanthium glabrous; disk sparingly hairy about the base of

the style; Jalisco, Colima, western Michoacan.

E. michoacanensis.

Leaves in drying curving down toward the midline, the midvein

thus depressed below the general level of the upper surface of

the blade, the resulting furrow concave or sometimes very nar-

row and (in section) V-shaped.

Midvein deeply and narrowly impressed above; leaves acutely

narrowed at both base and apex, dark green and lustrous

above, with inconspicuous veins; pedicels mostly 3-5 mm.

420 FIELDIANA: BOTANY, VOLUME 29

long, slender; bracteoles connate, involucrate; disk 1.5 mm.

wide (2-2.5 mm. wide in fruit); fruit 10-13 mm. in diameter;

Sierra Madre Oriental E. xalapensis.

Midvein usually depressed in a broad concave channel, neither

deeply nor very narrowly impressed; pedicels often shorter

and stouter; combined characters never as above.

Larger calyx-lobes 2-2.5 mm. wide; disk 2-3 mm. wide; pedicels

1-3 mm. long; fruit globose, 1-1.6 cm. in diameter; moun-

tains of south-central Mexico, mostly at elevations of 1000-

2000 meters.

Leaves glabrous, lustrous and nearly featureless above, the

mid vein glabrous, rather narrowly impressed; disk 2.5

mm. wide in flower, 3 mm. in fruit; Jalisco; state of

Mexico E. culminicola.

Petioles and midveins of the leaves persistently puberulent,

the midvein concave on the upper surface, often appear-

ing flat because of the persistent hairs; disk 1.5-2 mm.

wide in flower, 2.5 mm. wide in fruit.

Herbage and inflorescence hispidulous and somewhat vis-

cid-puberulent; fruit unknown; Morelos. .E. mirandae.

Herbage puberulent when young, the inflorescence some-

times nearly glabrous; fruit globose, 1-1.6 cm. in diam-

eter; Puebla; Hidalgo; San Luis Potosf. . .E. pueblana.

Larger calyx-lobes 1-2 mm. wide; disk 1-2 mm. wide; pedicels

various; fruit globose, oblate or oblong, usually less than

1 cm. in diameter.

Pedicels stout and short [0.4-0.6 mm. thick in flower, 1-2

(-3.5) mm. long]; midvein concave or channeled above,

the sides of the channel hispidulous with short golden or

pale hairs; bracteoles 0.5 mm. long, truncate or obtuse,

distinct and persistent but not involucre-like; fruit ellip-

soid, 7 mm. in diameter, 9-10 mm. long; state of Mexico

to Nayarit E. crenularis.

Pedicels, if only 1-2 mm. long, more slender; midvein flat or

somewhat elevated or concave, not hispidulous as above;

bracteoles usually somewhat united by their proximal

margins, involucrate; fruit globose or oblate; lowlands

of eastern Mexico, one species extending along the Pacific

coast to Sinaloa.

Leaves drying yellow-green or yellow; midvein convex on

the upper surface of the leaf, or toward the base of the

blade flat or even sulcate; hypanthium densely and

very minutely puberulent with pale appressed hairs

smaller than those of the pedicels, closely invested by

and partly covered by the bracteoles; calyx-lobes finely

pubescent on both sides; fruit transversely oval, 7-

8 mm. wide in the longer axis, 5-6 mm. long.

E. farameoides.

Leaves drying brownish green, green, brownish red or red-

dish; midvein flat or sulcate above; hypanthium usu-

ally glabrous, sometimes finely hispidulous like the

pedicels, mostly or wholly exserted beyond the bracte-

oles; calyx-lobes mostly quite glabrous; fruit globose.

McVAUGH: TROPICAL AMERICAN MYRTACEAE 421

Leaves ovate or rhombic-ovate to elliptic, 3-6.5 cm. long,

opaque, the veins scarcely evident above in dried

leaves; petioles ventrally flat or concave, dilated into

the base of the blade; pedicels in flower 3 mm. long

or less; buds 1.5-2.7 mm. long; disk 1 mm. wide;

style 3-4.5 mm. long; fruit 7-8 mm. in diameter;

plants usually glabrous; West Indies; Yucatan Pen-

insula; rare or doubtful from Veracruz to Oaxaca.

E. axillaris.

Leaves elliptic or obovate, or ovate, 6-12 cm. long, veiny,

the lateral and marginal veins pale above in dried

leaves; petioles deeply channeled, usually passing

abruptly into the blade; midvein often flat and

wrinkled near base, often somewhat depressed below

the surface of the blade, and most of it in a furrow

between two low parallel convex ridges; pedicels (at

least the lower ones) 5-13 mm. long; buds 2.5-4 mm.

long; disk 1.3-2 mm. wide; style (5-) 7-8 mm. long;

fruit 1 cm. or more in diameter; plants often hispid-

ulous; Central America; Yucatan Peninsula; Chi-

apas; west of the Isthmus of Tehuantepec mostly

near the Pacific Coast, north to Sinaloa.

E. acapulcensis.

Eugenia acapulcensis Steud. Norn. ed. 2. 1: 601. 1840. ?E.

carthagenensis Jacq. Sel. Amer. 152, t. 178, f. 53. 1763. ?E. cartha-

genensis /3 baruensis Jacq. I.e. 153. ?E. baruensis (Jacq.) Jacq. Coll.

3: 183. 1790. Myrtus maritima HBK. Nov. Gen. & Sp. 6: 146 [folio

ed. 116]. 1823. ?E. guadalupensis DC. Prodr. 3: 275. 1828. E. mari-

tima (HBK.) DC. I.e. 282, not E. maritima DC. I.e. 271. ?E. bonplan-

diana Berg, Linnaea 27: 228. 1856. E. mosquitensis Berg, Linnaea

31: 257. 1862. E. deltoidea Standl. Contr. U. S. Nat. Herb. 23: 1045.

1924. E.antiquae~Ri\ey,Kew Bull. 1927:121. 1927. E. escuintlensis

Lundell, Phytologia 2: 4. 1941. E. ovatifolia Lundell, Bull. Torrey

Club 69: 396. 1 May 1942. E. bartlettiana Lundell, Contr. Univ.

Mich. Herb. 7: 30 May 1942. E. bracteolosa Lundell, I.e. 31. E.

campechiana Lundell, I.e. 32. E. sibunensis Lundell, I.e. 34. E. comi-

tanensis Lundell, Wrightia 3: 12. 1961.

As the above long list of synonyms suggests, the taxonomy of

this species-complex is little understood and apparently involved.

The members of the complex seem to be most abundant and diverse

in northern Central America and adjacent Mexico, but some of them

occur in the West Indies and in northern South America. What

appears to be a single and rather homogeneous species along the

Pacific Coast of Mexico from Guerrero westward, and also in eastern

Central America, becomes in Guatemala and the Yucatan Penin-

sula rather more variable. In the Yucatan Peninsula it comes into

contact with the morphologically similar E. axillaris (Sw.) Willd.,

422 FIELDIANA: BOTANY, VOLUME 29

the latter primarily a West Indian species, and it is of course possible

that some of the puzzling variability has resulted from past or con-

tinuing hybridization. The distinction between the mainland spe-

cies (which may for the present be called E. acapulcensis) and E.

axillaris is not always easy and is often subjective. The distinctions

between the two species are set forth above in the key. My inter-

pretation of axillaris is essentially that of Urban in his revision of

the West Indian Myrtaceae (Bot. Jahrb. 19: 639-641. 1895). Urban

interpreted a Swartzian specimen as representative of the plant that

has commonly been passing as E. axillaris in Florida and the West

Indies. It is a glabrous plant with relatively thick opaque ovate

leaves, the midvein sulcate on the upper surface of the leaf, the

flowers small and in very short crowded axillary racemes. On the

continent of North America it seems to be of limited distribution;

it is frequent in southern Florida, and it occurs on the Yucatan

Peninsula and probably at a number of localities in the Atlantic

lowlands of Guatemala and Honduras. Elsewhere in Central Amer-

ica it is replaced by E. acapulcensis.1

Urban seems to have included in his concept of Eugenia axillaris

not only this most widely distributed West Indian type, but also

some variants with larger, thinner, more veiny and sometimes ellip-

tic or obovate leaves, viz. those that on the mainland would be

referred to E. acapulcensis. Throughout the West Indies these

larger-leaved plants seem to be in the minority, but they are repre-

sented in herbaria by numerous specimens from peripheral areas,

e.g. Guadeloupe, central and western Cuba, and the Isle of Pines.

On the mainland, as suggested above, acapukensis is more wide-

spread and abundant, whereas axillaris is of very limited distribu-

tion. The exact relation between the two is as yet not understood,

and further taxonomic studies, especially in the West Indies, are

much needed.

On the North American continent E. acapulcensis occupies a con-

siderable range, and is morphologically rather homogeneous over

most of this range. The nomenclaturally typical population ranges

along the Pacific coast of Mexico at least from Guerrero to Sinaloa,

and is known from a large series of specimens. The following char-

1 In the past few decades many hundreds of herbarium specimens from Mexico

and Central America have been determined as Eugenia axillaris without much

regard for the actual identity of the plants. The name became a handy tag to

apply to almost any Eugenia haying small flowers in crowded axillary racemes.

Many of these misidentified specimens are now to be referred to E. acapulcensis,

but any one of a score of other species may occasionally turn up among the speci-

mens labeled axillaris.

McVAUGH: TROPICAL AMERICAN MYRTACEAE 423

acterization is based primarily on these, but also applies to most

Central American specimens from Honduras to Panama:

Branchlets, petioles and inflorescence usually minutely hispidulous (some-

times quite glabrous), the raceme sometimes thickly pubescent in addition with

small golden hairs; leaves usually elliptic, varying to obovate or rarely to ovate,

abruptly short-acuminate with a broad blunt tip, less often slenderly acuminate,

or some or all leaves obtuse; blades acute or cuneate at base, or sometimes rounded

in the broader leaves on the same plants; axis of the raceme mostly 3-6 mm. long.

In the lowlands from Guatemala to Tabasco may be observed

certain tendencies toward variation from this general pattern. Gla-

brous individuals are relatively more frequent; leaves are seldom

obtuse, often slenderly acuminate, more often ovate and rounded

at base; relatively long and many-flowered racemes (up to about

15 mm. long) occur more frequently; occasional individuals with

small flowers seem otherwise identical with the larger-flowered mem-

bers of the same populations. Some of these variations may be

normal for the species, some may represent unusual individuals,

some may have resulted from hybridization. Some small popula-

tions in this geographical area seem reasonably distinct, e.g. Eugenia

flavoviridis Lundell; others (including those listed above in synon-

ymy) are questionably distinct, but some of them may represent

local populations. Of the species recently described from the Guate-

mala-Yucatan area, E. bartlettiana (known from the type only) has

rather loosely few-flowered racemes but is otherwise not unusual for

E. acapulcensis; E. bracteolosa is glabrous, with rather long racemes

and narrowly pointed leaves; E. campechiana is not distinctive;

E. escuintlensis, known from the type only, was a glabrous plant

having the flowers partly in racemes and partly solitary at the bases

of new vigorous shoots; E. ovatifolia was based primarily on individ-

uals with short petioles and pedicels, but the leaves, in spite of the

specific epithet, are not significantly broader than the average for

the species; E. sibunensis is glabrous, with slenderly acuminate

leaves. I have studied the types of these species in the Herbarium

of the University of Michigan.

Of the earlier synonyms of Eugenia acapulcensis (listed above),

E. deltoidea Standl. is known only from the type (Nelson 2292, US!).

This is a glabrous specimen but otherwise very like the type of

Myrtus maritima HBK. (E. acapulcensis) ; cf . Field Mus. Neg. 36894.

The type localities of the two species are only a short distance apart.

Two specimens from farther east on the Pacific slope of Mexico,

Seler 1793 (US!) and Seler 2029 (US!), both from Juchitan, Oaxaca,

were determined at Berlin by Loesener as Eugenia bonplandiana

424 FIELDIANA: BOTANY, VOLUME 29

Berg. This name is included in the above synonymy of E. acapul-

censis because of the assumption that Loesener probably had access

to the type of E. bonplandiana when he made the determinations of

Seler's specimens. Berg based E. bonplandiana on a specimen col-

lected by Humboldt and Bonpland in "Columbia" ("Eugenia lauri-

folia Willd. herb. no. 9486"); if Loesener's comparison was a critical

one, it may well be that the type of E. bonplandiana was a member

of the acapulcensis complex, collected perhaps somewhere in north-

ern Colombia along the lower Rio Magdalena, or possibly even at

Cartagena, where Humboldt and Bonpland spent three weeks.

Also from the vicinity of Cartagena came the specimens described

by Jacquin under the names of Eugenia carthagenensis and its vari-

ety baruensis. In general terms E. carthagenensis was described as

having the leaves ovate, obtusely acuminate, glabrous, short-petio-

late, 4 inches long; the common peduncles [i.e. presumably the axis

of the raceme] short and many-flowered; the flowers 4 lines in diam-

eter; the fruit globose, the size of a pea, red, finally black. The

figure published with this description unfortunately shows but a

single detached leaf, which could perhaps represent our plant.

Urban (Bot. Jahrb. 19: 639. 1895) regarded E. carthagenensis

ft baruensis as probably a synonym of E. axillaris, but if he was

right in his understanding of Jacquin's concepts I suspect that the

specimens from Cartagena were not true axillaris but acapulcensis.

It has not been possible at this writing for me to learn the identity

of the taxa described by Jacquin.

An earlier name that may well prove to be synonymous with

E. acapulcensis is E. guadalupensis DC. A photograph of the type,

Field Mus. Neg. 33492, shows a broad-leaved plant extraordinarily

like those common along the American mainland. The type should

be compared with Central American material.

Two species described from eastern Central America, E. mosqui-

tensis Berg and E. antiquae Riley, seem to represent almost identical

populations. The type of E. mosquitensis (Wullschlagel in 1855,

from "Pearl Key Lagoon, Mosquito" [Nicaragua]), which I have

seen through the courtesy of the authorities at Brussels, consists of

a few twigs in young bud. The branchlets and pedicels are hispid-

ulous, as commonly in specimens from Panama; the leaves are

broadly elliptic, shortly and bluntly acuminate, rounded to a cuneate

base. In my judgment this is the common plant of Panama, the

same as that named E. antiquae. I have not seen the type of the

latter, but judging from the original description it is the same plant.

McVAUGH: TROPICAL AMERICAN MYRTACEAE 425

This was referred by Amshoff in the Flora of Panama (Ann. Missouri

Bot. Gard. 45: 189. 1958) to E. acapulcensis.

One species referred by Amshoff (I.e.) and previously by Standley

(Contr. U. S. Nat. Herb. 23: 1046. 1924) to the synonymy of E. aca-

pulcensis, proves to be very different. This is Eugenia colipensis

Berg, Linnaea 29: 243. 1858. The type (from Colipa, Veracruz,

Karwinsky 241, LE!) is an over-mature fruiting specimen, evidently

a species close to or identical with E. oerstedeana Berg, but differing

from that species in having the leaves impressed-punctate above,

and the midvein in the dried leaves elevated above in a narrow ridge.

In other respects, including the racemes 3-5 cm. long, the fruit about

1 cm. in diameter, the thin persistent bracteoles, the venation of the

leaves, the short pubescence of the branchlets and the minute flat

dibrachiate hairs at the base of the lower leaf-surface, it exactly

simulates a specimen of the oerstedeana-florida alliance. Apparently

nothing like it has since been collected.

One species not listed above, Eugenia itzana Lundell, Bull. Torrey

Club 69: 395. 1 May 1942, surely belongs with the acapulcensis com-

plex, but its status is in doubt. The type (Yucatan, Chichen Itza,

Lundell 7589, MICH!) is an immature flowering specimen that super-

ficially suggests a small-leaved example of acapulcensis. One para-

type, Steer e 1373, is scarcely distinguishable from E. axillaris. All

four specimens cited in the protologue of E. itzana are small-flowered

for acapulcensis, resembling in this respect E. axillaris. Hardly an

independent species, E. itzana seems to fall between acapulcensis

and axillaris.

The very recently published Eugenia comitanensis, from near

Comitan, Chiapas, E. Matuda 5753 (LL, type!), represents in my

opinion a glabrous and rather narrow-leaved phase of E. acapulcen-

sis. The leaf-margins are somewhat revolute, as they often are in

E. acapulcensis, but quite entire. Superficially they appear to be

"obscurely crenulate," as noted in the protologue.

Eugenia alnifolia McVaugh, sp. nov.

Frutex vel arbor parva, ramulis petiolisque, foliorum marginibus et nervo medio

(subtus) breve hispidulis; folia elliptico-oblonga, 6-10 cm. longa, 1.7-2.2-plolongi-

ora quam latiora, deltoideo-acuminata, petiolis 5-7 mm. longis, nervo medio supra

concavo; flores numerosi glomerati sessiles, bracteolis connatis extus pubescenti-

bus, hypanthio velutino: calycis lobi extus sericei, 1.5-1.8 mm. lati; discus 1.5 mm.

latus; stylus 7 mm. longus; ovarium biloculare, ovulis quoque loculo ca. 5; fructus

ignotus.

A shrub or small tree, the branchlets and petioles, and the leaf-margins and

lower surfaces of the midveins short-hispidulous with hairs 0.1-0.2 (-0.5) mm.

426 FIELDIANA: BOTANY, VOLUME 29

long; inflorescence densely appressed-pubescent; leaves elliptic-oblong, 3-5 cm.

wide, 6-10 cm. long, 1.7-2.2 times as long as wide, on shoots relatively longer

and sometimes obovate, but on flowering branchlets broad at the middle and

about equally rounded to the bluntly deltoid-acuminate tip and the acute base,

the margins at the very base cuneately but shortly decurrent on the channeled

petiole 5-7 mm. long, 1-1.5 mm. thick; midvein shallowly concave above in the

nearly flat blade, somewhat persistently hirsutulous at base, on the lower surface

elevated nearly its whole diameter and strongly hirsutulous, the other veins and

the leaf surface essentially glabrous; lateral veins 7-10 on each side in addition

to some intermediate ones, inconspicuous above, elevated and rather prominent

beneath, not forming a well-defined marginal vein but diminishing distally and

each arching over irregularly, 2-5 mm. from the margin, to join the next succeed-

ing one; leaves dark green and probably lustrous above, paler and dull and finely

dark-dotted beneath; inflorescences consisting of dense many-flowered axillary

glomerules 1-1.5 cm. in diameter, at leafy or leafless nodes on old wood, the

racemes very short, often 2-3 at one node, each bearing 2-4 closely approximate

pairs of sessile flowers; bracts persistent, very short; bracteoles membranous,

appressed-pubescent without, orbicular-reniform, 1-2 mm. wide and almost as

long, narrowed at base and somewhat connate, forming a broad 2-lobed involucre

nearly concealing the hypanthium; buds 3.5 mm. long, obovoid, the broadly

cup-shaped or hemispheric hypanthium velutinous, 1 mm. wide and almost as

high; calyx-lobes in unequal pairs, broadly rounded, closely appressed-silky with-

out, glabrous within except near the tips, the outer ones 1 mm. long, 1.5 mm. wide,

the inner pair 1.3 mm. long, 1.8 mm. wide; disk about 1.5 mm. wide, hairy around

the base of the style; style 7 mm. long or more, glabrous; stamens about 35, about

as long as the style, the anthers 0.7-0.8 mm. long; petals white, elliptic-obovate,

3 mm. long, 2.5 mm. wide; ovary bilocular, the ovules about 5 in a compact group

in each locule. "Cafecillo" (Hda. Coahuayula, Emrick).

Michoacan: Distr. Coalcoman, Aquila, in woods, 15 Jan. 1942

(fl.), G. B. Hinton 16292 (K; MICH, type; NY; UC; US); Hacienda

Coahuayula, Nov. 1906 (fl.), G. M. Emrick 74 (F); Hda. California,

Mun. Apatzingan, 1200 m., Leavenworth & Hoogstraal 1^7 (F).

A species somewhat resembling E. origanoides but differing obvi-

ously in the distribution of the foliar pubescence, in the larger flowers,

and in the differently pubescent inflorescence.

Eugenia argyrea Lundell, Wrightia 3: 10. 1961.

This very recently described species was known to its author

from the type only. It proves to be widely distributed and appar-

ently abundant in Central America; it is in fact represented in her-

baria by more specimens than almost any other Central American

species. In view of the distinctive appearance of the flowering speci-

mens, and its wide distribution, it is indeed odd that neither Berg

nor any of the more recent special students of the Myrtaceae seems

to have noticed it. Among the small-flowered species of Eugenia

bearing the flowers in abbreviated racemes the plant is easily recog-

McVAUGH: TROPICAL AMERICAN MYRTACEAE 427

nized by the contrast between the glabrous or nearly glabrous brac-

teoles and calyx-lobes, and the tawny-pubescent hypanthium. The

characteristic coppery-silky pubescence of the young growth is dis-

tinctive but does not long persist.

A formal description of this species as I understand it can be

found in the Flora of Guatemala (Fieldiana, Bot. 24) and need not

be repeated here. The geographical distribution of E. argyrea is

indicated by the following list of specimens examined :

MEXICO: Veracruz: Santa Lucrecia, C. D. Mell 580 (F, US).—

Oaxaca: Chivela, Mell 32 (US), Mell s.n., 10 Dec. 1928 (US) ; 17 km.

northwest of La Ventosa, R. M. King 605 (MICH). — Chiapas: Ca-

mino Berriozabal a S. Fernando, F. Miranda 5829 (MICH); Oco-

zocuautla, Miranda 7454 (MICH); Siltepec, E. Matuda 1609 (F,

MICH), Matuda s.n., 8 Aug. 1937 (K, MICH); Cascada, near Sil-

tepec, Matuda 5074 (LL, type).

GUATEMALA: Alta Verapaz: Between Semococh and Chajamayic,

J. A. Steyermark 45725 (F); pantano 2.5 miles west of Cubilgiiitz,

Steyermark 44309 (F). — Izabal: Puerto Barrios, Standley 25118 (US).

HONDURAS: Atlantida: Standley 53775 (F), 56628 (F, US).—

Yoro: J. B. Edwards P-820 (F, US).— Colon: Von Hagen 1334 (F).—

Comayagua: Standley 56511 (F, US), J. Valerio R. 2892 (F), Yuncker

et al. 5742 (F).

NICARAGUA: Chontales: La Libertad, Standley 9028 (F).

COSTA RICA: Rodeo, H. Pittier 1642 (US), 821*1 (US); vicinity

of San Ramon, A. M. Brenes 5372, 6751, 14977, 20357, 20358 (all F);

San Ramon, Austin Smith P2332 (UC).

PANAMA: Campana, P. H. Allen 1324 (F, US) ; Canal Zone, Pittier

2567 (F).

COLOMBIA: Region of Santa Marta, H. H. Smith 907 (MICH),

Smith 1927 (MICH, US).

Eugenia avicenniae Standl. Contr. U. S. Nat. Herb. 23: 1043.

1924.

Because of the very narrow leaves this plant seems distinctive.

It is presumably related to the other west-Mexican species with red-

dish and somewhat appressed, dibrachiate hairs, viz. E. rekoi and

E. salamensis. The species is still unknown except for a few col-

lections from the vicinity of Acapulco, Guerrero, the type locality.

428 FIELDIANA: BOTANY, VOLUME 29

Eugenia axillaris (Sw.) Willd. Sp. PI. 2: 970. 1800. Myrtus

axillaris Sw. Prodr. 78. 1788.

This species appears to intergrade to some extent with E. aca-

pulcensis Steud.; see the discussion under that species. In Mexico

E. axillaris is known certainly from the Yucatan Peninsula, but rec-

ords from the Mexican mainland should be regarded with suspicion.

Eugenia biflora (L.) DC. Prodr. 3: 276. 1828. Myrtus biflora

L. Syst. Nat. ed. 10: 1056. 1759. E. fieldingii Berg, Linnaea 29:

242. 1858.

As pointed out in the Flora of Peru (pt. 4, no. 2: 684-685), this

is one of the most variable species of Eugenia. Widely distributed

in South America and the West Indies, it is well known in Panama

and perhaps in Costa Rica, but in northern Central America it is

scarcely known except in the lands bordering on the Gulf of Hon-

duras. The latter region is already known for its strong phytogeo-

graphic connections with the West Indies.

The commonest form in Central America is one much resembling

the West Indian var. virgultosa (Sw.) Krug & Urban, and also much

like the prevailing form throughout much of South America, viz. a

plant with lanceolate leaves 3-4 times as long as wide, long-acumi-

nate and bristle- tipped, impressed-punctate on the upper surface;

the style is 5-6 mm. long, the bracts of the raceme narrow and

mostly 3-4.5 mm. long, the bracteoles often 1.5 mm. long, the whole

inflorescence densely silky, and the flowers only about 6-8 to a ra-

ceme. This plant is represented in herbaria by a number of collec-

tions from Belize and Stann Creek Districts, British Honduras, and

from the Department of Izabal, Guatemala.

Apparently more widely distributed on the North American

mainland than the above variety of E. biflora are several so-called

species that are certainly closely related to one another and to E. bi-

flora. A comprehensive revision of the whole species complex may

indicate that the "species" of Mexico and Central America are not

distinct from the inclusive biflora. All are alike in having abundant

pale appressed silky or silky-strigose pubescence of simple hairs, the

inflorescence an elongate raceme with compressed internodes, usu-

ally 3-6 pairs of flowers, the bracts persistent and the distal ones

increasingly longer, the bracteoles somewhat coherent by their basal

margins, closely investing the hypanthium, scarious and often thinly

strigose and so contrasting with the canescent hypanthium, the calyx-

McVAUGH: TROPICAL AMERICAN MYRTACEAE 429

lobes broad, and silky or silky-strigose within. The distinctions be-

tween the species appear to depend chiefly on leaf-shape, and on

the presence and abundance of glandular depressions on the upper

leaf-surface. Distinctions between E. karwinskyana and E. yuca-

tanensis seem to be particularly weak; the few apparent differences

in flower-size and in pubescence have been studied in relatively few

specimens, and are not very suggestive.

The following key contrasts E. biflora (sensu stricto) with the

closely related "species" of the mainland:

Leaves lanceolate, 3-4 times as long as wide, bristle-tipped, impressed-punctate

above, the glandular depressions numerous and conspicuous E. biflora.

Leaves elliptic to ovate or broadly lanceolate, not bristle-tipped (acuminate, but

the tip obtuse or sometimes subacute), not at all or sparingly and obscurely

impressed-punctate above.

Leaves mostly 3-4 times as long as wide; eastern Mexico to Guatemala.

E. karwinskyana.

Leaves mostly 1.7-2.5 times as long as wide; Yucatan Peninsula.

E. yucatanensis.

Confined as far as known to the Yucatan Peninsula, E. yucatan-

ensis Standl. seems to be frequent in the lowlands from the State

of Yucatan to Pete"n. It is scarcely distinguishable from some Ja-

maican material of E. biflora in which the leaves are rather broadly

elliptic, obtusely acuminate, 1.7-2.5 times as long as wide, not at all

or scarcely impressed-punctate above. The plants are somewhat

less prominently silky than those of E. biflora, and the flowers appar-

ently a little smaller.

A third "species," ranging from San Luis Potosi, Mexico, to

northern Guatemala, is Eugenia karwinskyana Berg, Linnaea 29:

244. 1858 (E. tabascensis Lundell, Phytologia 1: 482. 1941). This

plant has ovate or elliptic leaves narrower than those of E. yucatan-

ensis, obtusely acuminate, 2.6-4 times as long as wide, sometimes

impressed-punctate above. The plants are more prominently pubes-

cent than those of E. yucatanensis, sometimes indeed approaching

E. biflora in this respect. The flowers are usually 8-10, or sometimes

as many as 18, in a raceme.

The type of E. karwinskyana Berg, which I have seen through

the courtesy of the authorities at Leningrad, is a flowering specimen

in which the inflorescence is notably canescent; the type of E. tabas-

censis Lundell is less strongly pubescent and has somewhat narrower

leaves but is surely conspecific. The following are to be referred to

E. karwinskyana as now understood :

430 FIELDIANA: BOTANY, VOLUME 29

MEXICO: San Luis Potosi: Tamazunchale, Lundell 7113 (MICH).

—Hidalgo: Km. 339 between Chapulhuacan and Tamazunchale,

Lundell 12400 (MICH); Huejutla, Karwinsky 242 (LE, type).—

Tabasco: Boca Cerro, Tenosique, Matuda 3561 (MICH, type of E.

tabascensis) .

GUATEMALA: Alta Verapaz: Region of Chelae, northeast of Car-

cha, Standley 70388 (F).

Another member of the species complex that includes E. biflora

is apparently the following:

Eugenia ledophylla (Standl.) McVaugh, comb. nov. Myrtus

ledophylla Standl. Contr. U. S. Nat. Herb. 23: 1038. 1924.

The type, Purpus 7804 (US !) is a silky-pubescent plant in which

the leaves are small for biflora, up to about 1.5 cm. wide, 5 cm. long,

lanceolate or lance-ovate, cuspidate, impressed-punctate above. The

pubescence throughout is like that of biflora, as are the bracteoles

subtending the buds. The type bears nearly mature fruit; another

specimen from Zacuapan, Veracruz, presumably a topotype, Purpus

8053 (UC), is in bud. The flowers in both collections are borne on

slender pedicels 1-1.5 cm. long, solitary in the axils of foliage leaves

on new tender branchlets. It is conceivable that these specimens

came from a plant or plants abnormally stimulated so as to cause

excessive development of the axillary branches that normally be-

come short, leafless, bracteate racemes in the species of Eugenia

related to E. biflora. Whatever may prove to be the relationship

of this plant, it is quite out of place in Myrtus.

A synonym of E. biflora is Eugenia fieldingii Berg ["in Mexico

(Fielding)"], the type of which proves to be an average specimen of

E. biflora. The racemes are young but otherwise characteristic, the

leaves are silky and impressed-punctate above and often cuspidate.

The source of the specimen, however, can hardly be Mexico. The

Fielding cited as collector was presumably Henry Barren Fielding

(1805-1851; cf. Britten & Boulger, Biog. Ind. Brit. Bot. ed. 2. 107.

1931), who amassed a large private herbarium but who, as far as I

can learn, did not himself collect in tropical America.

In 1858 Berg published three new taxa based on American col-

lections attributed to "Fielding." Only one of the specimens, the

type of Myrtus acerosa (Linnaea 29: 253) was cited by number

(Fielding 1448)- Berg saw these specimens "in herb. hort. bot. Pe-

trop." Through the kindness of the authorities at Leningrad I have

been able to study all three specimens.

McVAUGH: TROPICAL AMERICAN MYRTACEAE 431

The types of E. fieldingii and Myrtus acerosa, and that of Myrcia

saxicola /3 grandifolia (Linnaea 29: 219), all seem to have been col-

lected by the same person. Each bears a small collector's field-label,

slit to pass over the end of the twig; each bears on this label in a

large smooth hand in faded black ink a short notation evidently by

the collector, and in addition a serial number in another hand (evi-

dently added in different ink); also the words "Hb. Fischer" in a

small angular hand, and the word "Fielding" in what appears to

be still another hand. Berg has added his annotation in one instance

to this label, and in the other two on a separate label. The pertinent

data are as follows:

Name Serial No. Collector's Notations

Myrtus acerosa 1448 636 Myrtaceae Bajasan

Eugenia fieldingii 1449 Indana fruit eatable

Myrcia saxicola /3 grandifolia 1958 205 Casapi Myrtacea

From the data on No. 1449 the collector apparently was English-

speaking. From the handwriting in which the field-labels are written

I think there can be little doubt that these are collections of Andrew

Mathews, whose principal Peruvian collections came to Kew in the

1830's. His Myrtaceae were arranged in systematic sequence, ac-

cording to the then-current practice, and assigned serial numbers

replacing the original field-numbers. The serial numbers assigned

the Myrtaceae included at least 1447-1454, and in another series

1957 and 1958. No. 1449 was cited under Eugenia biflora in the

Flora of Peru (p. 685), where the locality was given as Tarapoto.

I assume that "Indana" (if I have interpreted correctly the writing

on the label) is a locality near Tarapoto, and that the type of E.

fieldingii is from eastern Peru. It is certainly not Mexican.

No. 1958 was cited under Myrcia splendens (Sw.) DC. in the

Flora of Peru (p. 661), where the locality was given as Cassapi. Pre-

sumably the type of M. saxicola /3 grandifolia is a duplicate of this

collection. The specimen belongs to one of the compact-leaved types

of M. splendens.

The type of Myrtus acerosa [Myrteola acerosa (Berg) Burret], was

cited in the Flora of Peru (p. 806) as Fielding 1448. I have not seen

a sheet of this number definitely attributed to Mathews, but he

traveled in this part of Peru and there seems no reason to doubt

that he collected this particular specimen. The few buds on the

type are immature; some are seemingly 4-merous, not 5-merous as

432 FIELDIANA: BOTANY, VOLUME 29

Berg described them, and others are 5-merous. Vegetatively the

plant much resembles Ecuadorean specimens of Myrteola micro-

phylla var. microphylla, but as noted in the Flora of Peru the branch-

lets in M. acerosa are spreading-setose and the very narrow leaves

are revolute-sagittate when dry; in M. microphylla the branchlets

are canescent and the leaves revolute-linear or -lanceolate.

Fielding's herbarium, the principal part of which went to Oxford

University, is said to have contained 2000 sheets of Mathews' Peru-

vian plants, viz. the set reserved by the collector for himself (cf.

Hook. Journ. Bot. & Kew Misc. 6: 283. 1854). Apparently some of

these specimens, probably sent out as duplicates, found their way to

St. Petersburg where Berg saw them and assumed that Fielding was

the original collector.

Eugenia capuli (Schlecht. & Cham.) Berg, Linnaea 27: 238.

1856. Myrtus capuli Schlecht. & Cham. Linnaea 5: 561. 1830.

Eugenia schiedeana Schlecht. Linnaea 13: 418. 1839. E. contrerasii

Lundell, Wrightia 2: 206. 1961.

In his revision of West Indian Myrtaceae, Urban referred the

Mexican E. capuli to the synonymy of E. monticola (Sw.) DC., var.

latifolia (DC.) Krug & Urb. (Bot. Jahrb. 19: 636. 1895). In this

course he has not been followed, as far as I can learn, by any bota-

nist working with the floras of continental North America. Probably,

however, E. capuli and E. monticola actually are conspecific; some

Central American specimens are quite indistinguishable from indi-

vidual West Indian ones. On the other hand most Mexican and

Central American plants differ from typical monticola in having the

leaves wider and longer (but neither so long nor so wide as described

for var. latifolia!); the petioles longer; the blades relatively longer

on the average because of the frequent occurrence of forms with

caudate-acuminate tips; the pedicels longer; but the flowers in gen-

eral smaller and the fruits smaller. Aside from these quantitative

differences I cannot distinguish between West Indian and Central

American specimens. Pubescence in both populations is variable;

plants from the mainland are usually but not always less conspicu-

ously puberulent than West Indian specimens, which may indeed

be very obscurely puberulent or quite glabrous. After examination

of a large series of specimens from both eastern and western Mexico,

from the Yucatan Peninsula and from the adjacent lowlands in

Central America, I cannot find any convincing basis for recognizing

more than a single species in the population.

McVAUGH: TROPICAL AMERICAN MYRTACEAE 433

From the standpoint of practical taxonomy it seems preferable

to recognize E. capuli as an independent species at least until the

relationships of the several West Indian populations of E. monticola

are better understood. There is much as yet unexplained variation

among these insular populations, whereas E. capuli is abundant and

relatively homogeneous over a considerable lowland area from Ta-

maulipas to northern Central America.

The species complex that includes E. monticola and E. capuli

has also several representatives in northern South America and in

eastern Central America. A revision of the entire complex is much

to be desired.

The principal species that are involved are as follows:

Eugenia micrantha (HBK.) DC. Prodr. 3: 282. 1828. Myrtus

micrantha HBK. Nov. Gen. & Sp. 6: 144 [folio ed. 115]. 1823. Type

locality, Honda, Colombia.

Distinguished from West Indian monticola chiefly by the longer

racemes, the axis of which may be 1 cm. long or more in micrantha,

usually 3-6 mm. long or less in monticola. The pubescence in mi-

crantha is short and inconspicuous. The known range extends from

about the vicinity of Caracas, Venezuela, to the valley of the Rio

Magdalena in Colombia.

Eugenia casearioides (HBK.) DC. Prodr. 3: 275. 1828. Myrtus

casearioides HBK. Nov. Gen. & Sp. 6: 145 [folio ed. 115]. 1823. Type

locality, "in monte Cocollar Cumanensium," i.e. on the Orinoco.

Distinctive chiefly in having the branchlets rather long-pubes-

cent with spreading hairs up to 0.5-0.7 mm. long. The pedicels are

short and the flowers glomerate, i.e. the axis of the raceme is very

short. The species is known chiefly from northern Venezuela, and

from one locality in Colombia (Norte de Santander).

Eugenia pittieri Standl. Field Mus. Bot. 8: 145. 1930. Type

locality, Panama.

Specimens from Costa Rica and Panama, identified as E. pittieri,

are essentially indistinguishable from E. monticola of Tobago, e.g.

Broadway s.n., 19 Sep. 1910 (US). The branchlets are setose with

coarse, antrorsely curved, sordid hairs 0.3-0.5 mm. long, these occur-

ring also on the mid veins and margins of the leaves; the hypanthium

434 FIELDIANA: BOTANY, VOLUME 29

and the pedicels are nearly glabrous. The type, Pittier 44-84 (F), is

in young bud but otherwise not different from other specimens

(F and US).

Eugenia chepensis Standl. Field Mus. Bot. 8:144. 1930. Type

locality, Panama.

The type of this species, Pittier 4716 (F) was the only specimen

cited in the protologue, where the pedicels were described as "2-

3.5 mm. longis," and a little later as "comparatively long." Actually

the pedicels are not long when compared with those of E. capuli or

even with those of E. monticola; Standley was comparing them with

those of E. pittieri, whose flowers he described as sessile. Amshoff,

in the Flora of Panama (Ann. Missouri Bot. Card. 45: 185. 1958)

describes the pedicels of chepensis as 4-10 mm. long. It is doubtful

that chepensis is distinct from pittieri, or that either one is distinct

from monticola.

The monticola complex seems to be represented on the Isthmus

of Tehuantepec rather sparingly, but interestingly enough two addi-

tional species have been described from this area, both on the basis

of their unusually narrow leaves. One of these is Eugenia lindeniana

Berg, Linnaea 29: 240. 1858. The type collection, from Teapa,

Tabasco, Linden 619 (LE, type; MICH), has narrowly lanceolate

leaves 11 mm. wide or less, 6-8 times as long as wide; except for

this one feature the specimen seems to be an average moderately

hispidulous flowering specimen of E. capuli. Another specimen that

presumably should be referred to E. lindeniana is:

Tabasco: Boca Cerro, Tenosique, E. Matuda 3592 (MICH).

The recently published Eugenia tenuissima Lundell (Wrightia 3:

19. 1961) was based entirely on Martinez Calderon 546, from Chilte-

pec, District of Tuxtepec, Oaxaca. I have seen isotypes (MEXU

and UC). The specimens bear nearly mature fruit; a great deal of

the pubescence has been eroded away, but seems to be characteristic

of E. capuli as I understand that species. The leaves are unusually

narrow for E. capuli, 3.2-4.5 times as long as wide, and up to no

more than 1.4 cm. wide. In other respects the specimens are quite

typical of E. capuli; the type was compared in the protologue of

E. tenuissima with E. lindeniana, which does indeed have even nar-

rower leaves than tenuissima. The leaves of the type collection of

lindeniana, however, although very narrow, are not acutely pointed

McVAUGH: TROPICAL AMERICAN MYRTACEAE 435

but are obtuse exactly as in tenuissima. It is not likely that these

two narrow-leaved extremes are specifically different from E. capuli.

A second specimen, very like the type of E. tenuissima, is:

Oaxaca: Upper Coatzacoalcos River, C. D. Mell 601 (F, US).

In recent months Dr. C. L. Lundell has performed a signal service

by calling attention to what he supposes to be distinct populations

among the Eugenias related to E. capuli, specifically those species

with puberulent herbage, small elliptic or lanceolate leaves having

the midvein flat or convex (not sulcate or impressed) above, the

flowers small in short crowded racemes and the fruit small and glo-

bose. The fruits in E. capuli are typically small and slender-pedicel-

late, 4-6 mm. in diameter, the fruiting pedicels 0.3 mm. thick, and

the axis of the raceme rarely more than 4-5 mm. long.

In Eugenia tikalana Lundell (Wrightia 2: 210. 1961) the fruits

are mostly 9-12 mm. in diameter, the fruiting pedicels 0.4-0.6 mm.

thick, and the axis of the longer racemes on most plants ranges from

6-10 (or up to 25) mm. long. Through the courtesy of Dr. Lundell

I have seen a fine series of collections of E. tikalana from Pete"n.

In E. michoacanensis Lundell (Wrightia 3: 16. 1961) the plants

are superficially very similar to those of E. capuli and E. tikalana,

but the fruits are intermediate in size (6-8 mm. in diameter). As

apparently plants with large, intermediate and small fruits all occur

in the same general areas in southwestern Mexico, and as the plants

are otherwise so similar, it seems likely that these species are not

genetically isolated. These large-fruited forms need further study

but apparently represent one or more populations distinct from

E. capuli.

The small-fruited E. contrerasii Lundell, described in the same

publication as E. tikalana, seems amply distinct from that species

in Pete"n, where both are common. In the protologue of E. contre-

rasii, however, it was compared not with E. tikalana but in a very

general way with E. capuli. I do not find any way in which E. con-

trerasii differs from E. capuli as that species occurs in eastern Mexico.

Eugenia choapamensis Standl. Field Mus. Bot. 22: 93. 1940.

The type, Mexia 9254 (F!), was from near San Juanito, Distr.

Choapam, Oaxaca. The species is known from several additional

collections from Oaxaca to Guatemala. It is treated at length in

the Flora of Guatemala.

436 FIELDIANA: BOTANY, VOLUME 29

Oaxaca: Pluma Hidalgo, C. Leyva in 1947 (US); Distr. Tuxtepec,

Chiltepec, 20 m., G. Martinez Calderon 525 (US).— Tabasco: El

Azufre, Teapa, 25 Mar. 1957 (fr.), F. Miranda 8J>88 (MICH).

Eugenia colipensis Berg, Linnaea 29: 243. 1858.

Glabrous or essentially so in age, the racemes minutely and sparingly hispid-

ulous; leaves elliptic, acute to acuminate at both ends, 3-4 cm. wide, 6-10 cm.

long, the petioles 5-8 mm. long; blades impressed-punctate above; midvein ele-

vated as a narrow ridge on the upper surface; racemes 2-4 cm. long, each with

4-7 pairs of flowers on straight pedicels 4-6 mm. long; bracteoles persistent, not

connate, broadly ovate-deltoid, 1 mm. long; fruit about 1 cm. in diameter, sub-

globose, the disk 1.5-2 mm. wide, the calyx-lobes rounded, erect, the larger ones

1.5-1.7 mm. wide.

Still known from the type only, this apparently distinctive spe-

cies should be sought in the Atlantic lowlands in Veracruz. See the

discussion above under E. acapulcensis.

Eugenia crenularis Lundell, Wrightia 3: 12. 1961. E. hintonii

Lundell, Wrightia 3: 14. 1961.

A shrub or small tree 3-4 m. high, the branchlets and petioles uniformly but

rather sparsely hispidulous with sharp erect hairs 0.1-0.3 mm. long; axis of inflores-

cence, and pedicels, similarly but often very sparsely pubescent or essentially

glabrous, the bracteoles coarsely ciliate, the bracts usually strigose especially dis-

tally with coarse appressed reddish hairs 0.2-0.4 mm. long; leaves thin, often

crenulate, or apparently so because of differentially revolute margins, the blades

lanceolate to elliptic or narrowly ovate, 1-3 cm. wide, (2-) 3-7.5 cm. long, (2-)

2.3-3 (-3.7) times as long as wide, often much narrowed toward both ends from

the middle or below, the tip blunt-pointed or obscurely or sometimes narrowly

acuminate, the base usually less drawn out than the apex, acute or rounded, the

margins at the very base cuneately but usually shortly decurrent on the ventrally

hispidulous and channeled petiole 3-6 mm. long; midvein concave or channeled

above, the sides of the channel more or less densely and persistently hispidulous

with short golden or pale hairs; lateral veins 8-10 on each side in addition to some

poorly differentiated intermediate ones, somewhat convex on both surfaces and

forming with the small veins a reticulum when dry, not forming a well-defined

marginal vein but connected by a series of irregularly angled arches 0.7-2.5 mm.

from the margins; blades lustrous above, paler beneath; glands usually evident

as scattered convexities above, as small dark or resinous dots beneath; inflores-

cence an abbreviated axillary raceme, or 2 racemes superposed, the stout axis

1-4.5 (-7) mm. long, bearing 1-3 (-5) opposite and decussate pairs of flowers on

rather stout pedicels 1-2 (-3.5) mm. long, 0.4-0.6 mm. thick in flower; terminal

flower often present, sessile in the bracts (simulating bracteoles) at the terminal

node; bracts persistent, ovate or slightly elongate, 0.5-0.8 mm. long; bracteoles

similar to the bracts, about 0.5 mm. long and wide, truncate or obtuse, persistent,

distinct; buds rather closely sessile in the bracteoles, short-pyriform, 2-2.5 mm.

long; hypanthium (usually) glabrous or sparingly puberulent, campanulate. 1 mm.

McVAUGH: TROPICAL AMERICAN MYRTACEAE 437

long; calyx-lobes short and broad, thin and membranous, soon irregularly reflexed,

the outer ones 0.5-1 mm. long, 1-1.5 mm. wide, the inner 0.8-1.3 mm. long, 1.5-

2 mm. wide; disk 1.3-1.4 mm. wide, glabrous; style 5-6 mm. long; stamens 50-60,

up to 5 mm. long, the anthers 0.5-0.6 mm. long; petals white, obovate, ciliate,

3.5 mm. long; ovary bilocular, the ovules about 10 in a compact group in each

locule; fruit ellipsoid, black, about 7 mm. in diameter, 9-10 mm. long, obtuse at

both ends, or slightly prolonged distally to the erect or appressed calyx-lobes.

State of Mexico: Dist. Temascaltepec: Temascaltepec, 18 Mar.

(fl.), Hinton 405 (K); 1 Feb. (imm. fr.), Hinton 3255 (K, US);

5 May (fl.), Hinton 7679 (US); 2 June (fl.), Hinton 7695 (US, iso-

type); San Jose", 21 Feb. (fr.), Hinton 291 (K); 7 Mar. (bud), Hin-

ton 359 (K). — Guerrero: Dist. Mina: Manchon, 28 Sep. (imm. fr.),

Hinton 9603 (MICH, isotype of E. hintonii); 21 Apr. (fl.), Hinton

10076 (US); Tierras Blancas, 6 May (fl.), Hinton 10146 (US).

The following specimens appear to belong to the same species,

as indicated by the quality and distribution of the pubescence, leaf-

shape, texture, venation and crenulation, presence of the terminal

flower, general flower-morphology, and fruit shape. They differ,

however, in having the racemes longer (often 4-6 mm. long with

up to 4 pairs of flowers), the pedicels 1.5-3 mm. long and apparently

fleshy (shrinking in drying), the disk up to 1.5 mm. wide, and the

hypanthium distinctly and rather closely puberulent.

State of Mexico: Dist. Temascaltepec: San Lucas, 7 July (fl. &

imm. fr.), Hinton 4294 (K, MICH, US).— Jalisco: Sierra del Halo,

23 June (fl.), McVaugh 15029 (MICH); 14 Aug. (past fl.), McVaugh

16187 (MICH).— Nayarit: 9.5 miles west of Tepic, 11 Sep. (past fl.),

McVaugh 18949 (MICH).

Eugenia culminicola McVaugh, sp. nov.

Frutex subglaber, ramulis petiolisque breve hispidulis, demum glabris; folia

ovato-lanceolata, 2.7-4 cm. longa, 2.3-2.5-plo longiora quam latiora, obtusa vel

vix acuminata, nervo medio supra glabro, impresso; laminae nitidae, nervis later-

alibus inconspicuis; racemi perbreves, floribus subsessilibus, bracteolis distinctis,

hypanthio glabro infundibuliforme; calycis lobi rotundati, majores 0.8 mm. longi,

1.7 mm. lati; discus floris 2.5 mm. latus, fructus 3 mm.; stylus 5-6 mm. longus;

ovarium biloculare, ovulis quoque loculo 6-8.

Shrub 1.5 meters high, the branchlets short-hispidulous, the petioles at first

sparingly so; leaves ovate-lanceolate, 1-1.8 cm. wide, 2.7-4 cm. long, 2.3-2.5 times

as long as wide, blunt or bluntly acuminate at tip, acute at base, the margins more

or less cuneately decurrent on the channeled petiole 3-4 mm. long; midvein nar-

rowly impressed above; lateral veins 4-6 on each side, scarcely apparent in dried

leaves; blades lustrous and nearly featureless above, probably paler and dull be-

neath, and there obscurely gland-dotted; inflorescence a much-abbreviated axillary

raceme or 2 racemes superposed, the axis up to 2.5 mm. long, bearing up to 6

438 FIELDIANA: BOTANY, VOLUME 29

closely approximate opposite decussate pairs of flowers on pedicels 1 mm. long or

less (up to 2.5 mm. long in fruit); bracts indurate-persistent, ovate-deltoid, divari-

cate, 0.8 mm. long; bracteoles suborbicular-deltoid, ciliate, persistent, distinct,

0.5 mm. long and 0.7 mm. wide; buds 3.5-4 mm. long, turbinate, the glabrous

funnelform hypanthium 1.5-2 mm. long; calyx-lobes rounded, unequal, the smaller

ones 0.4 mm. long, 1.2 mm. wide, the larger pair 0.8 mm. long and 1.7 mm. wide;

disk about 2.5 mm. wide in flower (3 mm. in fruit); style 5-6 mm. long; stamens

about 75; ovary bilocular, the ovules 6-8 in a compact group in each locule; fruit

purple-black, ellipsoid or obovoid, substipitate, 8-10 mm. in diameter, 10-12 mm.

long, crowned by the calyx-lobes up to 1.5 mm. long and 2.5 mm. wide.

State of Mexico: Dist. Temascaltepec, Cumbre-Pena Blanca,

Alnus woods, 1 Mar. 1936 ["3-1-36"] (bud), G. B. Hinton 8957 p.p.

(TEX); in fruit, Hinton 8957 (US, TEX p.p.); in flower, with few

detached fruits, Hinton 8957 (GH, type).

The fruiting specimens associated with the budding plants of

Hinton 8957 may conceivably belong to another species and are

probably from another plant; the leaves are more veiny and the

pedicels are longer (up to 8 mm.).

A common and often very abundant treelike shrub in several

localities in high barrancas in Jalisco, in forests of pine or fir or with

various deciduous trees, is apparently the same species but is known

only in fruit. Fruiting plants are quite glabrous; the dark green

and lustrous leaves are up to 9.5 cm. long, on petioles 5-8 mm. long;

the fruits are borne in clusters, 1-3 pairs on an axis 2-4 mm. long, the

pedicels 1 mm. thick or more, 1-3 mm. long, the fruits themselves

subglobose, turning red and then black, 8-15 mm. in diameter; the

disk is about 3 mm. wide, the calyx-lobes persistent but inconspic-

uous on the fruit, broad and low, about 1 mm. long, 2 mm. wide.

Jalisco: Sierra de Cuale, southwest of Talpa de Allende, barrancas

in fir zone, 1800-2250 m., 19-21 Nov. 1952, McVaugh 14301 (MICH);

Sierra de Manantlan, 15-20 mi. southeast of Autlan, cloud-forests

ca. 2000 m., 6 Nov. 1952, McVaugh 13944 (MICH); ca. 40 km. west

of Ayutla, fir forest, elev. 2100-2200 m., 29 Nov. 1960, McVaugh

21559 (MICH).

This species apparently has much in common with E. pueblana

Lundell, which apparently occupies a similar ecological niche in the

mountains to the east and northeast of E. culminicola. Neither spe-

cies is adequately understood. The flowers of E. pueblana are

smaller in several respects than those of E. culminicola, and the

plants are more generally and persistently puberulent. The mid-

vein in E. culminicola is somewhat more deeply impressed and is

consistently glabrous.

McVAUGH: TROPICAL AMERICAN MYRTACEAE 439

Eugenia farameoides A. Rich. Ess. Fl. Cub. 588. 1845.

This plant seems not to have been reported in the literature ex-

cept from Cuba, but it is well represented in herbaria by collections

from the Yucatan Peninsula, where it has been passing under the

name of Eugenia flavifolia Standl. Full description and synonymy

are given in the Flora of Guatemala. The plant is also known from

Veracruz and from the lowlands of Oaxaca. Dried specimens are

easily recognized because of their yellow-green color suggestive of

some species of Symplocos.

Veracruz: Between Acayucan and Minatitlan, 22 Sep. 1957 (fl.),

F. Miranda 85^6 (MICH). — Oaxaca: Cerro Campana, 400 m., 12

July 1926 (fl.), E. Makrinius 619 (US).

Eugenia guatemalensis Donn. Sm. Bot. Gaz. 23: 245. 1897.

Description and full synonymy are given in the Flora of Guate-

mala. This species is very similar on the one hand to E. rekoi of the

Pacific slope of Mexico, and on the other to the species with wing-

angled fruit and similarly pale rufous-pubescent branchlets and ra-

cemes, E. pleurocarpa of western Mexico and E. octopleura of Cen-

tral America and the West Indies. Additional material, especially

fruiting material that can be matched with flowering specimens of

known origin, is much needed to clarify the relationships between

these species. A few specimens from Mexico are referred without

much doubt to E. guatemalensis:

State of Mexico: Dist. Temascaltepec, 11 Nov. 1933 (bud),

Hinton 3368 (A). — Oaxaca: Mountains, Joyacatlan, ca. 1800 m.,

18 June 1894 (fl.), L. C. Smith 29 (GH).— Chiapas?: "Chiapas, etc.,"

Sep. (fr.), Ghiesbreght 893 (GH).

Eugenia hypargyrea Standl. Contr. U. S. Nat. Herb. 23: 1044.

1924.

This plant belongs to the complex of species that includes E. gua-

temalensis and E. rekoi, but seemingly is distinct by virtue of its

very fine silvery rather than reddish indumentum, and by its rela-

tively large obtuse obovate leaves. It is known from but few col-

lections; the type was from Zacuapan, Veracruz, Purpus 6171 (US!)

and a recent collection is from Chiapas. Description and full synon-

ymy are given in the Flora of Guatemala.

440 FIELDIANA: BOTANY, VOLUME 29

Chiapas: Bajada del Puerto de la Sepultura, carretera al N. de

Arriaga, 600 m., 23 Aug. 1951 (fl.), F. Miranda 7262 (MICH).

Eugenia inconspicua Standl. Contr. U. S. Nat. Herb. 23: 1043.

1924.

"Compact upright shrub 8 feet high" [Palmer], the foliage nearly glabrous

[from the first?], the branchlets and inflorescence [and the youngest leaves?]

densely appressed-pubescent with pale dibrachiate hairs mostly 0.2-0.4 mm. long,

the whole plant ultimately glabrate; leaves elliptic to elliptic-obovate, 1-1.8 cm.

wide, 3-4.5 cm. long, 2-3 times as long as wide, narrowed about equally to an

obtusely pointed or obscurely and bluntly acuminate tip, and to an acute base,

the margins acutely or cuneately decurrent on the channeled petiole 1-2 mm.

long; mid vein concave above at least near base, or shallowly channeled between

two obscure parallel ridges; lateral veins 5-6 on each side, ascending, pale and

readily visible on both surfaces of dried leaves, scarcely forming a marginal vein

but arching over 0.5-1 mm. from the margin in a series of irregular angled arches

to join the next succeeding ones; blades darker and very minutely (or obscurely)

pale-verruculose above, paler beneath, with scattered small dark resinous glands

apparent on both surfaces; inflorescence a much-abbreviated axillary raceme, the

axis up to 2 mm. long, bearing 1-2 opposite decussate pairs of flowers on slender

pedicels, these in fruit up to 4.5 mm. long, 0.6 mm. thick; bracts ovate, sub-

persistent, 1 mm. long or less; bracteoles oblong, divaricate, distinct, persistent, up

to 1.3 mm. long; calyx-lobes broadly rounded, concave, sparingly beset with large

resinous convex glands 0.2 mm. in diameter, appressed-pubescent on both surfaces,

but persistently and more densely on the inner; larger lobes up to 3 mm. long

and wide; disk in fruit 2.5-3 mm. wide; immature fruit obovoid, 6 mm. in diameter,

8 mm. long, topped by the erect concave calyx-lobes.

Sinaloa: Culiacan, Palmer 1786 in 1891 (US, type!).

Still unknown except in the vicinity of Culiacan, this species is

strikingly close to E. yautepecana, with which it was compared by

Lundell in the protologue of the latter. The above description of

E. inconspicua, and the one of E. yautepecana (q.v., infra), were pre-

pared with awareness of the similarities between the species. It is

probable that yautepecana and inconspicua are no more than sub-

specifically different, but flowering specimens of the latter, and in-

deed additional material of both species, are needed to resolve this

question.

Eugenia karwinskyana Berg, Linnaea 29: 244. 1858.

Description and full synonymy are given in the Flora of Guate-

mala. For a discussion of the taxonomic position of E. karwinskyana,

see above under E. biflora.

Eugenia ledophylla (Standl.) McVaugh.

See above under E. biflora.

McVAUGH: TROPICAL AMERICAN MYRTACEAE 441

Eugenia liebmannii Standl. Contr. U. S. Nat. Herb. 23: 1044.

1924.

A shrub 5 meters high, persistently but minutely hispidulous-puberulent, the

branchlets, petioles and inflorescence thickly beset with sharp erect hairs 0.1-

0.2 mm. long; leaves soon glabrate, at first sparingly strigose like the very young-

est branchlets with coarse coppery hairs; leaves elliptic to obovate or ovate, 3-

10 mm. wide, 10-25 mm. long, 1.6-3 (-4) times as long as wide, rounded or obtusely

pointed at apex, gradually rounded to a narrow base, the pale thick revolute mar-

gins rather abruptly decurrent on the channeled petiole 1-1.5 mm. long, 0.5 mm.

thick; blades sometimes apparently acute at base because of the inrolled margins;

midvein slightly concave above at least near base; lateral veins inconspicuous,

sometimes scarcely apparent above, 3-4 pairs, the lowest 1 (or 2) strongly ascend-

ing, nearly parallel to the margin and 1-1.5 mm. within it, the distal veins diverg-

ing at a wide angle from the midvein and joining the basal (marginal) one at or

above the middle of the blade; blades smooth above or with a few convex glands,

dull and paler beneath and there rather abundantly dark-dotted; inflorescence an

abbreviated axillary raceme, or 2 racemes superposed, the axis 2 mm. long or less,

bearing 1-4 opposite and decussate pairs of flowers on pedicels 1.5-3 mm. long;

flowers sometimes solitary and opposite at the lowermost (bracteate) nodes of

new branchlets; bracts dark, scarious, 0.5 mm. long; bracteoles ovate, obtuse,

distinct but persistent, 0.5-0.6 mm. long; buds 2-2.5 mm. long, the short-campan-

ulate hispidulous hypanthium 0.5-0.8 mm. long, abruptly expanded into the globe

of the petals; calyx-lobes broadly rounded, nearly flat, fimbriate-ciliate but other-

wise nearly glabrous, about 1 mm. long and wide; disk 1-1.5 mm. wide; style 4-

4.5 mm. long; stamens about 40, up to 4 mm. long, the anthers 0.5 mm. long and

about as wide; petals white, prominently gland-dotted, orbicular-obovate, 3 mm.

long; ovary bilocular, the ovules 10-11 in a compact group near the center of the

dissepiment in each locule; fruit glandular- verru cose, globose or transversely oval,

4-5 mm. in diameter or up to 6 mm. wide, 4 mm. broad and high. — U. of Mich.

Neg. 1415.

Tamaulipas: 9 mi. south of C. Mante, 31 Aug. 1948 (fr.), Kenoyer

& Crum 3678 (MICH); area of Sierra de Tamaulipas, between "La

Chona" and Rio Sta. Olaya, elev. 50 m., 26 Sep. 1956 (fr.), F. Mar-

tinez M. & G. Borja L. F-2150 (TEX).— Veracruz: Near Ebano

along Rio Panuco, H. LeSueur 55-4 (TEX). — San Luis Potosi: Villa

Juarez, lowland thickets, July 1937 (fl.), Lundell & Lundell 7270

(MICH).— Oaxaca: Villa Alta, Aug. 1842 (fr.), Liebmann 3969 (US,

type).

Plants of this species are superficially similar to very small-leaved

examples of E, buxifolia; the two species may indeed be closely re-

lated. Their geographical ranges, however, are quite distinct. Mor-

phologically the two may be distinguished by the differences in

leaf-size, by the considerably more veiny leaves of E. buxifolia and

the more abundant and larger hairs; and by the flowers, which are

consistently smaller in E. buxifolia.

442 FIELDIANA: BOTANY, VOLUME 29

Sterile specimens of Eugenia liebmannii may be distinguished

from those of Myrtus ehrenbergii Berg, a similarly small-leaved plant

of eastern Mexico, by the petioles (glabrous in M. ehrenbergii even

on the young antrorsely hispidulous branchlets), by the midvein

(distinctly convex on the upper surface in M. ehrenbergii) and by

the yellowish resinous convex glands on both leaf-surfaces in M.

ehrenbergii. The latter is a plant of desert mountains in San Luis

Potosi, at least as indicated by modern specimens:

San Luis Potosi: San Luis Potosi to Tampico, Palmer 112^ in

1878-79 (NY); Cerro de las Minas, Guadalcazar, 25 Oct. 1891 (fl.),

P. Maury 7557 (MEXU); San Jose" Pass, 10 Oct. (fr.), Pringle 352^

(A, GH); Minas de San Rafael, May 1911, Purpus 5211 (GH, MO,

UC), Purpus 8588 in 1910 (UC).

Eugenia lindeniana Berg, Linnaea 29: 240. 1858.

See above under Eugenia capuli.

Eugenia mexicana Steud. Norn. Bot. ed. 2. 1: 603. 1840.

E. macrocarpa Schlecht. Linnaea 5: 560. 1830; op. cit. 13: 417. 1839,

not E. macrocarpa Roxb., 1814.

Probably a tree, 5-6 meters high, glabrous except for reddish hairs on the

vegetative buds; leaves elliptic-ovate, 2.5-4 (-5.5) cm. wide, 6-7.5 (-9) cm. long,

1.7-2.2 times as long as wide, the blades about equally rounded to a blunt decurved-

deltoid-acuminate tip and a broadly angular or rounded base, the margins at the

very base cuneate-decurrent on a channeled petiole 5-10 mm. long, 1-1.3 mm.

thick; midvein impressed above as a V-shaped channel; lateral veins 8-12 on each

side including some intermediate ones, inconspicuous, straight and divaricate,

mostly passing directly into a weak marginal vein 1-2 (-4) mm. from the margin;

blades dark green and lustrous above, paler and perhaps glaucous beneath and

there finely dark-dotted; inflorescence a stout axillary raceme, the axis 3-6 mm.

long (in Pringle 8192 up to 10-16 mm. long), 1-1.7 mm. thick, straw-color in dry-

ing, the internodes strongly compressed and angled; flowers in 1-5 opposite and

decussate pairs in the racemes, or opposite at the lowermost (bracteate) nodes of

new leafy branchlets; pedicels almost none, or stout, glandular-verrucose, up to

1.5 mm. long, 0.5 mm. thick; bracts broad and scarious, membranous, (or their

bases indurate-persistent), sparingly ciliate, glandular-verrucose, loosely enfolding

the base of the pedicel, about 1.5 mm. long, 1.5-3 mm. wide; bracteoles glandular-

verrucose, short-ciliate, broadly rounded and connate, forming a deep involucre

3 mm. long and 2 mm. wide that envelops the base of the hypanthium; buds 5-

6 mm. long, obovoid, the broadly campanulate hypanthium more or less hidden

by the bracteoles; calyx-lobes broadly rounded, membranous, unequal, the outer

pair 1.5 mm. long and 3 mm. wide, the inner 2-2.5 mm. long and 3.5-4 mm. wide;

disk 3-4 mm. wide; style 7-9 mm. long; stamens about 150, up to 6-8 mm. long,

the anthers 0.5-0.7 mm. long; petals obovate or ovate, 5-6 mm. long, up to 5 mm.

wide at base; ovary bilocular, the ovules about 6-10 in a compact group in each

locule; fruit unknown.

McVAUGH: TROPICAL AMERICAN MYRTACEAE 443

Veracruz: Jalapa, woods, 4000 ft., 19 May 1899 (fl.), Pringle 8192

(MEXU, MICH, UC); in sylvis jalapensibus, "Schiede & Deppe"

(G) ; Jalapa, Schiede 544 (NY, ex herb. Meisner., "comm. cl. Schlecht-

endal 1840")-

As indicated in the description above, the specimens collected

by Pringle are somewhat more luxuriant, with larger leaves, longer

racemes and pedicels, and slightly larger flowers than the plants

collected by Schiede. All the specimens are otherwise very similar

and certainly belong to the same species.

Schlechtendal in 1839 reported the number of ovules as 3-5 in

each locule of the ovary; this was based on the flowering specimens

collected by Schiede and distributed to several herbaria including

those cited above. My own examination of these same specimens

indicates that the number of ovules varies from 6 to 8; in Pringle's

specimens the number is about 10. The type of Eugenia macrocarpa

Schlecht. was a fruiting specimen also collected near Jalapa but

some years earlier; its identity is unknown, but there is no reason

to suppose it was a different species.

Eugenia michoacanensis Lundell, Wrightia 3: 16. 1961.

Shrub or small tree up to 10-12 meters high, 12-15 cm. in diameter, the

branchlets very minutely puberulent with erect sharp hairs; ventral surface of

petiole, and of the mid vein near base, pubescent; axis of the raceme, and the

bracts and bracteoles, sometimes strigose with lustrous reddish hairs 0.2-0.4 mm.

long; leaves thin, elliptic-lanceolate, lanceolate or ovate, 1-3 cm. wide, 3-7.5 cm.

long, 2.3-3 times as long as wide, usually prolonged distally to a blunt, gradually

acuminate tip, the acumen broad and short or narrowly triangular and up to more

than 1 cm. long; base acute (the margins often about at right angles) or somewhat

rounded, the margins at the very base cuneately decurrent on the channeled peti-

ole 3-6 mm. long; midvein when dry flat or broadly convex on the plane surface

of the blade, pubescent at least near base; lateral veins 6-8 on each side, incon-

spicuous, more apparent beneath, ascending and nearly straight; marginal vein

about as strong as the laterals and but slightly arched between them, 1-1.5 (-3)

mm. from the margin; blades dull green and darker above, paler beneath; glands

often visible as small convexities above, as dark dots beneath; inflorescence an

abbreviated axillary raceme or 2-3 racemes superposed, the axis 3-6 (-10) mm.

long, bearing 5-7 opposite and decussate pairs of flowers on slender, minutely

puberulent pedicels 1-2.5 (-4) mm. long; bracts persistent, broad, scarious or in-

durate, 0.5-0.8 mm. long; bracteoles ovate or broader, persistent, contiguous or

nearly so but scarcely connate, 0.5 mm. long; buds 2-2.5 mm. long, short-pyriform;

hypanthium short-campanulate, glabrous, 0.7 mm. long; calyx-lobes broad and

low or obtusely triangular, membranous, glabrous, up to about 0.7 mm. long,

1-1.3 mm. wide; disk 1-1.2 mm. wide in flower, sparingly hairy about the base

444 FIELDIANA: BOTANY, VOLUME 29

of the style; style 4-4.5 mm. long, hairy below the middle; stamens about 50,

up to 4.5 mm. long, the anthers 0.5 mm. long; petals white, obovate, 2.5-3 mm.

long; ovary bilocular, the ovules 5 in a compact group in each locule; fruit de-

pressed-globose, ripening red, probably finally black, 6-8 mm. in diameter, the

calyx-lobes appressed together over the disk and outlining a circular figure 1.5-

1.7 mm. wide.

Michoacan: Dist. Coalcoman: San Pedro, 600 m., 19 June (fl.),

Hinton 13812 (US, isotype); Huizontla, 12 Dec. (imm. fr.), Hinton

16212 (MICH, US); Aquila, Hinton 162U (MICH).— Colima: Foot-

hills of the Nevado de Colima south of Hda. San Antonio, 11 Aug.

(fl.), McVaugh 16101 (MICH); Rio Cihuatlan 13 mi. north of San-

tiago, 27 July (fl.), McVaugh 15797 (MICH).— Jalisco: Puente San

Pedro 5 mi. southwest of Tecalitlan, 2 Dec. (imm. fr.), McVaugh &

Koelz 1307, 1308 (MICH) ; 9 mi. north of west end of Bahia Navidad,

12-13 Dec. (imm. fr.), McVaugh & Koelz 1725 (MICH); same tree,

1 Oct. (fl.), McVaugh 19717 (MICH) ; Santiago-Durazno road, 11 mi.

north of Rio Cihuatlan, 1 Aug. 1957, McVaugh 15973 (MICH).—

Guerrero: Nanorida(?), 1500 m., 16 June 1899 (fl.), Langlasse 1062

(US); Dist. Adama, Achotla, 570 m., 25 Nov. 1937 (fr.), Y. Mexia

8898 (US) ; Acahuizotla, 10 May 1903 (fr.), E. W. Nelson 7041 (US).

This species is very close to and perhaps not distinct from E. ca-

puli, which differs in its smaller fruit, longer pedicels, and usually

puberulent hypanthium, but glabrous style-base; see discussion above

under E. capuli.

Eugenia mirandae D. Ramirez Cantu, An. Inst. Biol. Mex. 14:

487. 1944.

Tree probably 5-10 meters high, viscid and viscid-pubescent, the branchlets,

petioles and inflorescence densely short-hispidulous with viscid hairs 0.1-0.2 mm.

long, the surfaces between the hairs apparently also viscid; leaf -margins undulate-

crenate, somewhat revolute, slightly viscid-pubescent; leaves thin, elliptic or lan-

ceolate, 1.5-2.5 cm. wide, (4-) 5-7 cm. long, 2.5-3.5 times as long as wide, usually

narrowed about equally to the gradually and bluntly acuminate tip and to the acute

base, the tip sometimes more prolonged and the base then somewhat rounded;

margins at base cuneately decurrent on the channeled but nearly terete petiole

5-7 mm. long, 0.7-0.9 mm. thick; midvein impressed above in a concave channel

in the otherwise plane leaf, the channel hairy; lateral veins 6-8 on each side, in-

conspicuous but in dried leaves convex on both surfaces, in a reticulum including

the smaller veinlets; marginal vein not well defined, about as strong as the lateral

ones and formed by the strongly arched and asymmetrical continuations of these,

1.5-3 mm. from the margin; blades more or less concolorous, more lustrous above,

at flowering time with numerous convex glands on both surfaces; inflorescence an

abbreviated axillary raceme (or 2 racemes superposed), the axis up to 5 mm. long,

bearing 1-3 opposite and decussate pairs of flowers on pedicels 1.5-3 mm. long, the

terminal flower sometimes present between bracts simulating bracteoles; bracts

McVAUGH: TROPICAL AMERICAN MYRTACEAE 445

broad and short, hairy near the tips, 0.5 mm. long; bracteoles scarious, persistent,

distinct, broadly ovate-deltoid, 0.5-0.7 mm. long and about as wide; buds 3-4 mm.

long, pyriform, the hypanthium minutely puberulent, viscid, campanulate, some-

what attenuate at base; calyx-lobes broadly rounded, glabrous, up to 1.8-2 mm.

wide, 0.8-1.3 mm. long; disk 1.7-2 mm. wide, hairy around the base of the style;

style 5-6.5 mm. long, hairy in the proximal two-thirds; stamens about 75, about

as long as the style, the anthers 0.5 mm. long; petals white, ovate, 3.5 mm. long;

ovary bilocular, the ovules about 10 in a compact group in each locule; fruit prob-

ably black, oblong-obovoid, 8 mm. in diameter, 12 mm. long.

Morelos: Sierra de Tepoxtlan, 7500 feet, May 1900 (fl.), C. G.

Pringle 8333 (MEXU, MICH, UC).

The above description is based entirely on Pringle 8333. Dr.

Faustino Miranda informs me that these specimens are essentially

topotypes of E. mirandae. I find no very convincing differences

between Pringle 8333 and specimens of Eugenia pueblana Lundell,

except that the Pringle specimens are somewhat viscid almost

throughout. As this is a most unusual condition in this group of

Myrtaceae, it is possible that these plants are abnormally coated

with insect exudations or other substance.

Very recently, through the kindness of Srta. De"bora Ramirez

Cantu, I have studied type-material of Eugenia mirandae. The

type [Morelos, "bajando del Parque a Texpoxtlan," 4 May 1943

(fl.), D. Ramirez C. s.n. (MEXU)] is a nearly glabrous flowering

specimen. I cannot distinguish it from Pringle 8333 except that it

completely lacks the viscid covering described above, and the mid-

veins are glabrous rather than pubescent. In this latter feature it

differs from E. pueblana and resembles the newly described E. cul-

minicola. From E. culminicola it differs in having smaller flowers

(disk 1.5-2 mm. wide) and in the slightly longer pedicels. In the

one (terminal) flower dissected the type of mirandae has about 10

ovules in each locule as in Pringle 8333, whereas in pueblana the num-

ber of ovules is usually 4-5, and in culminicola 6-8. None of the three

species is well known, and it is possible that these several popula-

tions in the highlands of Morelos, Puebla, Hidalgo and Mexico may

ultimately prove to be conspecific.

Eugenia oaxacana Berg, Linnaea 30: 683. 1861.

Shrub or tree, the branchlets, petioles and inflorescence, and probably the

young foliage, closely (antrorsely) appressed-pubescent with pale sordid hairs

about 0.3 mm. long, the leaves soon glabrate, a few hairs persistent on the lower

surface; leaves thin, coriaceous, elliptic-oblong, 3.5-6.5 cm. wide, 8.5-15 cm. long,

about twice as long as wide, about equally rounded to the abruptly acuminate tip

446 FIELDIANA: BOTANY, VOLUME 29

and to the acute or cuneate base, the acumen obtuse, about 1 cm. long, the mar-

gins long-decurrent on the petioles 8-12 mm. long, 1.3-1.5 mm. thick; midvein

deeply impressed above, elevated nearly its whole diameter beneath; lateral veins

rather close and parallel, 10-12 on each side in addition to some intermediate

ones, rather inconspicuous but in dried leaves convex on both surfaces, rather

more prominent beneath; marginal vein about as strong as the lateral ones and

forming a series of angled arches between them, 2-3 mm. from the margin; blades

lustrous and yellowish green above, paler and dull and drying brown beneath,

minutely dark-dotted beneath; flowers in axillary glomerules, the inflorescence

a much-congested axillary raceme (or 2 racemes superposed), the axis 5 mm.

long or less, up to 1.5 mm. thick, bearing 1-4 pairs of sessile flowers; bracts

ovate, indurate, 1 mm. long; bracteoles persistent, very broadly rounded, gla-

brous within, about 2 mm. wide and 1.5 mm. long, slightly connate, forming a

laterally notched involucre closely investing the hypanthium; buds and open

flowers not seen; hypanthium in post-anthesis densely pale-sericeous, subglobose,

1.5-2 mm. long and high, about as long as the prominent erect concave calyx-

lobes, which are broadly rounded, strongly imbricated, pubescent both sides, the

inner ones longer, the outer pair 1.5 mm. long, 2-2.5 mm. wide; disk 2.5 mm. wide;

style about 5 mm. long; ovary bilocular, the ovules 7-8 in a compact group in

each locule.

Oaxaca: Chinantla, Nov. 1842, Liebmann 3963 (C); "Dep. Oa-

jaca," sin. dat., Liebmann 3962 (C); Tepinapa, B. P. Reko 1*198 in

1920 (US).

I have not located the type of this species. Berg gave the type

locality merely as "in Mexico ad Oaxacam," and of the type he said

merely "v. specimen unicum valde mancum in herb. Buchinger."

From his detailed description I presume the above specimens belong

to the same species.

Eugenia oerstedeana Berg, Linnaea 27: 285. 1856. E. vincen-

tina Krug & Urb. ex Urb. Bot. Jahrb. 19: 621. 1895. E. conzattii

Standl. Contr. U. S. Nat. Herb. 23: 1041. 1924. E. cocquericotensis

Lundell, Bull. Torrey Club 64: 554. 1937. E. petenensis Lundell,

Bull. Torrey Club 69: 397. 1942.

This Central American species seems to be a close relative of

Eugenia florida DC., a widespread species of the lowlands of north-

ern South America. The two are virtually indistinguishable except

that in E. florida the inflorescence is usually a raceme bearing 4-5

(or up to 10) pairs of flowers on pedicels 3-4 (or occasionally to 7)

mm. long. The terminal flowers usually abort. Occasional speci-

mens have some or most of the flowers solitary in the axils of small

bracts at the lowermost nodes of leafy branchlets. In E. oerstedeana

the racemes are developed in relatively few plants (fewer than half

the specimens examined) ; terminal flowers are often present; solitary

McVAUGH: TROPICAL AMERICAN MYRTACEAE 447

bracteate flowers are usually present, either in addition to the flowers

in racemes, or to the exclusion of these; the pedicels are 6-10 (or up

to 25) mm. long.

Typical florida apparently has not been reported from Panama,

but the following specimen is to be referred there:

Chiriqui: Progreso, Cooper & Slater 158 (F).

The Central American and Mexican populations I interpret as a

single series that has been derived from E. florida or something very

like it, by the gradual restriction of the flowers to the lower axils of

leafy branches, the reduction in frequency of non-leafy racemes and

an increase in the frequency with which a terminal flower develops,

the elongation of the pedicels and perhaps also a reduction in the

number of ovules (in E. florida the usual number seems to vary from

4 to 7; in E. oerstedeana the number may be 4-6, but is often only

2 or 3) . A further specialization seems to have occurred in the some-

what localized Mexican population that was described as E. conzattii;

here solitary flowers and short racemes may be found on the same

plant with 3-flowered "dichasia." I interpret the dichasium as a

development from a solitary terminal flower, lateral pedicels having

arisen from the axils of the floral bracteoles. The distinctive appear-

ance of this 3-flowered inflorescence which Standley called "cymose"

led him to associate this species with Eugenia [Myrcianthes] fragrans,

to which it has little relationship. The following specimens from

eastern Mexico show this tendency to the production of "dichasia":

Oaxaca: Galeotti 2882, 2887; Mell 602; Conzatti et al. 3113 (type

of E. conzattii) (all US).— San Luis Potosi: Palmer 148 in 1904 (US);

R. M. King 4405 (TEX).

The type of E. oerstedeana Berg, which I have seen through the

courtesy of the authorities at Copenhagen, is an over-mature flower-

ing specimen; the leaves have the typical venation of the florida-

oerstedeana complex, and also bear on the lower surface near the

base the minute sessile reddish flat dibrachiate hairs that mark these

otherwise nearly glabrous species. A few of the flowers are axillary,

but most of them are in racemes 1-2 cm. long, bearing 4 or 5 pairs of

flowers on filiform pedicels 7-9 mm. long; the flowering disk is 1.5-

1.7 mm. wide, and the ovules are 3-4 in each locule of the ovary.

The type of E. petenensis Lundell (Lundell 3746, MICH) is a

flowering specimen bearing relatively few flowers and most of these

solitary in the axils of bracts or ordinary leaves; the ovules appear

to be 2 in each locule.

448 FIELDIANA: BOTANY, VOLUME 29

A large series of specimens is available for study, exhibiting many

conditions intermediate between those in which the inflorescences are

all racemose and those in which the flowers are all solitary. I can-

not believe that more than one species is involved. I should rather

question the propriety of recognizing E. oerstedeana as a species dis-

tinct from E. florida. Some Central American specimens are scarcely

distinguishable from E. florida, e.g. the following, in which the flowers

are almost all in racemes, the pedicels are 3-5 mm. long or less, and

the ovules 4-6:

BRITISH HONDURAS: El Cayo District, Vaca, P. H. Gentle 2533

(MICH) ; Little Cocquericot, Belize River, C. L. Lundell 4090 (MICH,

type of E. cocquericotensis).

HONDURAS: Comayagua: Malcotal, Minas de Oro, J. B. Edwards

P-195 (F).

The type of E. vincentina [St. Vincent, "in sylvis ad leeward side

rara 200-500 m. alt., m. April, flor.," H. H. Smith & G. W. Smith

1521 (NY!)] is a flowering specimen with new thin leaves on young

shoots, racemes 3-4 cm. long, filiform pedicels 8-15 mm. long, ter-

minal flowers often present. The leaves have the special features

of the oerstedeana-complex, namely the sunken midvein which is

hispidulous above in the furrow; small sessile dibrachiate hairs on

the lower leaf -surf ace near the base of the petiole; and the slightly

"engraved" arching veins on the upper leaf-surface. According to

Krug and Urban the ovules are 3 in each locule. The plant is indis-

tinguishable from many Central American specimens of E. oerstedeana.

Eugenia origanoides Berg, Linnaea 29: 229. 1858.

Here has been treated a variety of plants, all with short-veluti-

nous foliage and branchlets, short petioles, and very small flowers in

sessile glomerules. Such plants, varying in leaf-size and shape,

in compactness of the inflorescence, and in density of pubescence,

range from northern Veracruz to Colombia. The type, Karwinsky

238, from Papantla, Veracruz, which I have seen through the cour-

tesy of the authorities at Leningrad, is a strongly hirsute specimen

with rather broadly ovate leaves 3.5-6 cm. wide, 6.5-11 cm. long,

rounded or even subcordate at the base; most other specimens as

far as I know have the leaves averaging 2-3 cm. wide, and acute or

merely rounded at base. The flowers in the type are not unusual

except that the hypanthium is more than ordinarily hirsute. In the

type the ovules are 5-6 in each locule of the ovary; in other speci-

mens the number may be as small as 4, or as many as 9-10.

McVAUGH: TROPICAL AMERICAN MYRTACEAE 449

Perhaps more than one species is included under what is now

called E. origanoides. Variation seems to be considerable. Few

specimens in the fruiting condition are available. Dr. C. L. Lundell

has recently described, in the course of his intensive studies on the

flora of the Maya area, a species that should be compared with

E. origanoides when a thorough revision of that species is under-

taken. The type of this new species, E. rubella Lundell (Wrightia 3:

18. Dec. 1961) was originally thought by Dr. Lundell to represent

E. origanoides, but to be atypical in having pedicellate flowers. The

specimen certainly resembles other plants of E. origanoides except

that the buds are borne on pedicels up to 4 mm. long. Two addi-

tional collections from the same locality in Oaxaca [District of Tux-

tepee, Chiltepec, in llanos, G. Martinez Calderdn 157 (MEXU, UC,

US), 218 (UC, US,)] are to be referred to Eugenia rubella if that spe-

cies be a valid one, but some features of these specimens cast doubt

upon that validity. The racemes in the Oaxaca specimens are at

first crowded, but in most instances become 8-14 mm. long, 2-3

superposed in an axil, closely simulating the racemes of Eugenia

acapulcensis. The pedicels are not 4 mm. long as in the type of

E. rubella, but 1-2 (-2.5) mm. long. The pubescence of the branch-

lets and the racemes (in the type of rubella and also in the Oaxaca

plants) seems exactly that of E. origanoides, as does the impressed

midvein of the leaves. It is possible that E. rubella is a valid species,

but it may be of hybrid origin or merely the result of some abnor-

mality (e.g., the developing ovaries in Martinez 218 are partially

transformed into galls).

Full description and synonymy of E. origanoides are given in the

Flora of Guatemala. Not to be confused with E. origanoides is the

Guatemalan E. chinajensis Standl. & Steyerm., a very distinctive

species marked by its coarsely setose foliage, branchlets and inflores-

cence, and by the broad-based subcordate and subsessile leaves.

The flowers are unknown, but are probably larger than those of

E. origanoides.

Eugenia pleurocarpa Standl. Field Mus. Bot. 4: 243. 1929.

Still unknown except from western Jalisco and southern Nayarit,

this species is apparently related to Eugenia octopleura Krug &

Urban, which I take to be the same as E. doubledayi Standl. and

E. koepperi Standl.; this is discussed more fully elsewhere in this

paper. A full description of E. octopleura may be found in the Flora

of Guatemala. The prominently wing-angled fruits seem to be found

450 FIELDIANA: BOTANY, VOLUME 29

in these two species alone among the North American ones; in the

absence of fruit, sterile specimens of either species may be confused

with E. guatemalensis or E. rekoi. The following material of E. pleuro-

carpa has been seen:

Nayarit: Hills 6-7 miles south of Compostela, 900-1000 m.,

11-12 July 1957 (fl.), McVaugh 15322 (MICH), 5 Sep. 1960 (imm.

fr.), McVaugh 18743 (MICH); mountains above La Cucaracha, sea-

ward slopes 12-13 miles south of Las Varas, 300-400 m., 20 Sep. 1960

(imm. fr.), McVaugh 19194 (MICH).— Jalisco: Quimixto, 14 km.

southwest of Puerto Vallarta, 28 Mar. 1959 (fr.), A. Carter & F. Chi-

saki 1238 (MICH); Quimixto, 70 m., 29 Nov. 1926 (fr.), Y. Mexia

1176 (F, type; MICH) ; vicinity of San Sebastian, 1500 m., Mexia 1446

Eugenia praeterita McVaugh, sp. nov.

Frutex vel arbor tomentuloso-hirsuta, pilis erectis crebris atrorufis ca. 0.5 mm.

longis; folia coriacea, supra glabrescentia nitidaque, subtus tomentulosa, elliptica,

apice acuminata, basi cuneata, 10-14 cm. longa, ca. 2.2-2. 5-plo longiora quam

latiora, marginibus revolutis, petiolis 10-15 mm. longis, nervo medio impresso;

racemus 1-1.5 cm. longus, florum 2-5 pares gerens; pedicelli crassi recti 2-3.5 mm.

longi; bracteolae divaricatae, distinctae, late ovatae, usque ad 1 mm. longae;

alabastra 3 mm. longa; hypanthium campanulatum vel urceolatum, post anthesin

3 mm. longum; calycis lobi late ovato-deltoidei, obtusi, intus glabri vel subglabri,

usque ad 2 mm. longi latique; discus 2.5 mm. latus; stylus 5 mm. longus; ovarium

biloculare, ovulis quoque loculo 8-10.

A shrub or tree, the branchlets, petioles, lower leaf-surfaces and inflorescence

densely short-hirsute-tomentulose with dark reddish-brown erect hairs about

0.5 mm. long; leaves coriaceous, soon glabrous and lustrous above, somewhat

persistently tomentulose beneath, elliptic, about equally narrowed to the acumi-

nate tip and the acute or cuneate base, 4-6 cm. wide, 10-14 cm. long, about 2.2-

2.5 times as long as wide, the revolute margins strongly decurrent on the broadly

channeled rugulose petiole 10-15 mm. long, 1.5-2 mm. thick; midvein sharply

impressed above; lateral veins 8-10 on each side, evident above, more conspicuous

beneath, stronger than the intermediate ones if any; marginal vein about equal-

ing the lateral ones and arched between them, 1.5-2.5 mm. from the margin;

blades probably darker green above, paler and minutely dark-dotted beneath;

inflorescence a stout raceme (or 2 racemes superposed), the axis 1-1.5 cm. long,

somewhat angular, 1-1.5 mm. thick, bearing 2-5 opposite and decussate pairs of

flowers on stout straight pedicels 2-3.5 mm. long, about 1 mm. thick including

the hairs; bracts suborbicular, indurated but subdeciduous, 1 mm. long; bracteoles

divaricate, distinct, broadly ovate, apparently indurated and somewhat persist-

ent, 1 mm. long or less; buds about 3 mm. long, ellipsoid, the calyx somewhat

closed over the petals until anthesjs; hypanthium just after anthesis 3 mm. long,

campanulate or urceolate; calyx-lobes broadly ovate-deltoid, blunt, incurved,

glabrous or nearly so within, the larger ones 2 mm. long and wide; disk 2.5 mm.

wide; style about 5 mm. long; petals 3-4 mm. long, hairy without; ovary bilocular,

the ovules 8-10 in a compact group in each locale.

McVAUGH: TROPICAL AMERICAN MYRTACEAE 451

Veracruz?: Amatlan, July 1842, Liebmann s.n. (C), Liebmann

3959 (C, type; US).

Extraordinarily different from any other species of Mexico or

Central America by virtue of the dense covering of dark reddish-

brown tomentum.

Eugenia principium McVaugh, sp. nov.

Frutex vel arbor parva, ramulis petiolisque setoso-hispidis, bracteis bracteo-

lisque ciliatis, racemis sparse hispidis; folia ovata vel elliptico-ovata, 2.2-4.5 cm.

longa, (1.7-) 2-2.4-plo longiora quam latiora, apice obtusa vel obtuse acuminata,

basi rotundata, petiolis 2 mm. longis, marginibus incrassatis, nervo medio im-

presso, basin versus hirsutulo; racemus 2-4 mm. longus, florum 2-4 pares gerens,

pedicellis (1-) 2.5-7 mm. longis; bracteolae in sicco atrofuscae, persistentes, non

connatae; alabastra 4-4.5 mm. longa; hypanthium campanulatum vel urceolatum,

post anthesin globosum vel obovoideum, supra ovarium constrictum; calycis lobi

erecti, rosei, inaequales, plani, vix imbricati, majores basi 1.2-1.7 mm. lati, 1.5-

2.3 mm. longi; discus in hypanthio prolongato vix conspicuus; stylus 5 mm. longus

vel longjor; ovarium biloculare, ovulis quoque loculo 4-9; fructus ignotus.

A shrub or small tree up to 3-10 meters high, the branchlets and petioles

setose-hispid with pale sharp erect lustrous hairs 0.3-0.5 mm. long; bracts and

bracteoles coarsely ciliate; raceme-axis and pedicels sparingly beset with hairs like

those of the branchlets; leaves coriaceous, ovate or elliptic-ovate, 1-2.5 cm. wide,

2.2-4.5 cm. long, (1.7-) 2-2.4 times as long as wide, narrowed from the middle or

below to a blunt or bluntly acuminate tip, rounded at base, the margins passing

abruptly onto the thick inner angles of the broadly channeled petiole 2 mm. long;

margins coarsely ciliate, scarcely revolute but cartilaginous-thickened, the nearly

terete vein-like thickening 0.2 mm. in diameter, evident from beneath; midvein

somewhat impressed above, hirsutulous toward the base; lateral veins 5-7 on each

side, inconspicuous, hardly apparent in mature leaves; marginal vein about equal-

ing the lateral ones, 1-1.5 mm. from the margin; leaves smooth and opaque above,

pale and almost glaucous beneath; the glands obscure on both surfaces; inflores-

cence a stout much abbreviated raceme (usually without an additional raceme

superposed), the axis up to 2-4 mm. long, bearing 2-4 approximate, opposite and

decussate pairs of flowers on pedicels (1-) 2.5-7 mm. long; bracts drying chestnut

color, indurate-persistent, convex on the backs, ovate-deltoid to oblong, 1-1.5 mm.

long; bracteoles drying chestnut, persistent but distinct, broadly ovate-deltoid,

divaricate and flat in age, 1 mm. long or a little more; buds about 4-4.5 mm. long,

the globe of the petals abruptly expanded above the short glabrous hypanthium

0.5-1 mm. high; hypanthium campanulate or urceolate, sometimes attenuate to

the base, in post-anthesis becoming globose or obovoid and surmounted by a

prominent narrow neck 0.5 mm. high and 1 mm. in diameter below the prominent

erect "rose-pink" calyx-lobes; lobes in unequal pairs, deltoid-ovate or elliptic and

rounded, flat, scarcely imbricate even in the bud, the larger ones 1.2-1.7 mm.

wide at base, 1.5-2.3 mm. long, the smaller pair deltoid, 1-1.2 mm. long and wide;

disk 1.7-2 mm. wide, poorly developed or at least not evident in the prolonged

neck of the hypanthium, slightly hairy about the base of the style; style 5 mm.

long or more; stamens about 50, about as long as the style, the anthers 0.5-0.6 mm.

452 FIELDIANA: BOTANY, VOLUME 29

long; petals white, elliptic-ovate, ciliate, 4-5 mm. long; ovary bilocular, the ovules

4-6 in a compact group in each locule (in the type), or 7-9 in each locule (in the

Panama specimens) ; fruit not seen.

MEXICO: Colima: Palm [Orbignya] forest near Playa de Santiago,

in dense shade, sandy soil near sea level, 21 July 1957 (past fl.),

McVaugh 15581 (MICH, type).

PANAMA: C.Z.: West of Limon Bay, Gatun Locks and Gatun

Lake, thin soil over limestone, 2 Apr. 1956 (fl.), /. M. Johnston 1773

(MICH).— Darien: Forests around Pinogana, 16-21 Apr. 1914 (fl.),

H. Pittier 6532 (US).

It seems strange that this species has never before been recog-

nized, as it has a combination of unusual morphological features in

the hypanthium and calyx-lobes, and it seems to be of wide geograph-

ical range. Except for the difference in ovule-number, which may

represent an individual peculiarity, the Panama specimens are very

like the type. The specific epithet refers to the great palms — since

unfortunately destroyed by the hurricane of 1959 — among which

the type was collected.

Eugenia pueblana Lundell, Wrightia 2: 167. 1961.

A tree or large shrub 2-6 meters high, up to 10-12 cm. in diameter, the branch-

lets when young more or less antrorsely puberulent with short reddish hairs, the

petioles and midveins of the leaves puberulent with short coarse hairs, the inflores-

cence often nearly glabrous; leaves lance-ovate or elliptic, 1-2.5 cm. wide, 2.5-

5.5 cm. long, 2-3 times as long as wide, gradually acuminate (the slender tip blunt),

acute to rounded at base, the margins cuneately decurrent on the slightly splayed

summit of the nearly terete but shallowly channeled petiole 3-6 mm. long; mid-

vein concave above, appearing flat because of the persistent stiff hairs; lateral

veins about 4-6 on each side, inconspicuous, more evident beneath; marginal vein

ill-defined, 1-2 mm. from the margin; blades probably darker above, drying brown-

ish and dull beneath, finely verruculose above, scarcely glandular beneath; inflores-

cence a much-abbreviated axillary raceme (sometimes at leafless nodes on old

wood), the axis almost none, the flowers in 1-2 (-4) approximate pairs, on pedicels

0.5-1.3 mm. long, 0.4 mm. thick; bracts minute, hairy; bracteoles persistent,

minute, deltoid-orbicular, ciliate, 0.3 mm. long; buds 3.5 mm. long, pyriform, the

globe of the petals 2.5 mm. in diameter, much surpassing the short-stipitate

minutely puberulent hypanthium 1-1.5 mm. long; calyx-lobes rounded, the larger

ones 0.5-0.7 mm. long, 1.5 mm. wide; disk 1.5 mm. wide (2.5 mm. wide in fruit);

style 5 (-6?) mm. long; stamens about 50, about as long as the style, the anthers

0.5 mm. long; petals suborbicular, 3 mm. long; ovary bilocular, the ovules 4-5 in

a compact group in each locule; fruit globose, red, probably at length black, 1.3-

1.6 cm. in diameter, the low calyx-lobes usually persistent.

MEXICO: Puebla: Wet mixed forest west of Huauchinango, 1 Nov.

1943 (fr.), C. L. Lundell 12618 (isotype, MICH).— Hidalgo: Zacual-

tipan, M.Martinez, in June 1940 (GH) ; forested ravines 6 miles from

McVAUGH: TROPICAL AMERICAN MYRTACEAE 453

Zacualtipan, road to Tlahuelompa, 2000 m., 14 Oct. 1949 (bud),

H. E. Moore 5295 (MICH); Capatla below Alumbres near Zacualti-

pan, ca. 1800 m., 21 Mar. 1946 (fr.), A. J. Sharp 46214 (MICH).

The following specimens apparently represent the same species,

but the leaves are up to 3.5 cm. wide, 7 cm. long, the axis of the

raceme up to 3.5 mm. long, the inflorescence and the hypanthium

strongly puberulent:

MEXICO: San Luis Potosi: Tamasopo Canyon, 25 June 1890 (fl.),

Pringle 3534 (A); Las Canoas, 25 June 1890 (fl.), Pringle 3534 (GH).

The following specimens differ from typical E. pueblana in hav-

ing the hypanthium more strongly puberulent, and the midvein on

the upper surface of the leaf glabrous:

MEXICO: State of Mexico: Dist. Temascal tepee, Calera, wet bar-

ranca, ca. 770 m., 19 Apr. 1933 (fl.), Hinton 3797 (K).

As noted above in the discussion of Eugenia mirandae, that spe-

cies is perhaps not distinct from E. pueblana.

Eugenia rekoi Standl. Contr. U. S. Nat. Herb. 23: 1044. 1924.

A relatively well-collected species, ranging from Sinaloa to Oaxaca

and occurring also in Guatemala. It seems to be most abundant in

tropical deciduous forests, on hills near the Pacific Ocean. The fol-

lowing specimens have been seen:

MEXICO: Sinaloa: Los Caballos, Sierra Tacuichamona, ca. 300 m.,

16 Feb. 1940 (fr.), H. S. Gentry 5643 (MICH); Hda. Cafia, San Ig-

nacio, 460 m., J. G. Ortega 858 (F, US; doubtfully this species).—

Jalisco: East of La Manzanilla on Bahia Tenacatita, 150-200 m.,

11 Nov. 1960 (fl.), McVaugh 20966 (MICH).— Colima: Manzanillo,

Dec. 1890 (fr.), E. Palmer 1020 (F, US); west side of Cuyutlan la-

goon, 28 Nov. 1925 (fr.), R. S. Ferris 6107 (F, US).— Guerrero: Aca-

pulco, Oct. 1894-Mar. 1895 (fr.), Palmer 357 (F, MICH, US), 21

Aug. 1935 (fl.), L. H. MacDaniels 144 (F).— Oaxaca: Cafetal Apango

(Cerro Huatulco), 400 m., 18 Aug. 1917 (fl.), B. P. Reko 3356 (US,

type); Distr. Tamiltepec, Playa de Minizo, 7 Dec. 1921 (fr.), Con-

zatti4416 (US).

Eugenia rhombea (Berg) Krug & Urb. ex Urb. Bot. Jahrb. 19:

644. 1895. E. foetida y rhombea Berg, Linnaea 27: 212. 1856. ?E.

leptopa Lundell, Bull. Torrey Club 69: 396. 1942.

Berg's Eugenia foetida 7 rhombea was based on two specimens,

one from Florida (Cabanis) and the other from "St. Domingo"

454 FIELDIANA: BOTANY, VOLUME 29

(Bertero). In the original publication (Linnaea 27: 212. 1856) the

name appears to be based directly upon E. foetida /3 maleolens (Pers.)

DC., and so superfluous when published, but this is evidently the

result of a typographical error on Berg's part; on the preceding page

(p. 211) Berg had published his var. /3 maleolens, again based directly

upon E. maleolens Pers. and with a reference to DeCandolle. Pre-

sumably Berg meant to indicate (on p. 212) that his rhombea had

made up a part of the more inclusive maleolens, as the latter was

understood by DeCandolle.

Apparently Eugenia leptopa was a mixture. As pointed out in

the protologue, the paratypes differ in some respects from the type,

which is a fruiting specimen with narrow leaves and with somewhat

immature obovoid fruit, unlike the oblate fruit of E. rhombea. Noth-

ing like the type of E. leptopa has come to my attention, but I should

refer the paratypes to E. rhombea.

For description of E. rhombea see the Flora of Guatemala. This

species is little known in Mexico except in the Yucatan Peninsula,

but the following specimen has been seen :

Chiapas: Cerros de Don Ventura, northwest of Tuxtla Gutierrez,

1000 m., 30 Dec. 1948 (fr.), F. Miranda 5101 (MICH).

The very similar Eugenia fiscalensis Donn. Sm. (Bot. Gaz. 54:

235. 1912) is thought to differ from E. rhombea in having slightly

smaller flowers and leaves, and in having the calyx-lobes and mar-

gins of the staminal disk more reflexed in anthesis. Until more ma-

terial of E. fiscalensis can be obtained, its status cannot be deter-

mined. The following incomplete specimen seems to represent a

species much like E. fiscalensis:

Oaxaca: 11 km. northwest of La Ventosa, Trans-Isthmian high-

way, 50 m., 14 July 1958 (past fl.), R. M. King 538 (MICH).

As suggested by their association in the key above, E. rhombea

and the newly described E. turneri have many superficial similarities.

The latter is known only from the type, a flowering specimen refer-

able neither to E. rhombea nor to E. fiscalensis. The leaves in E. tur-

neri are gradually rounded to the base, elliptic-lanceolate, 2.3-3

times as long as wide, the marginal vein relatively straight and less

than 1 mm. from the margin; the axis of the raceme is usually 2-4

mm. long and, like the young branchlets, appressed-pubescent. In

E. rhombea and E. fiscalensis the leaves are more abruptly rounded

at base, ovate, 1.5-2 times as long as wide, the marginal veins more

strongly arched between the laterals and 1-3 mm. from the margins;

McVAUGH: TROPICAL AMERICAN MYRTACEAE 455

the axis of the raceme is scarcely developed (or in extreme instances

up to 2 mm. long), and the whole plant is glabrous. It is probable

that additional and perhaps more fundamental differences will be

apparent when E. turneri is better known.

Eugenia salamensis Bonn. Sm. Bot. Gaz. 27: 333. 1899. Psid-

ium rensonianum Standl. Journ. Wash. Acad. 14: 241. 4 June 1924.

Eugenia oaxacana Standl. Contr. U. S. Nat. Herb. 23: 1043. 1 Dec.

1924, not E. oaxacana Berg, 1861. E. tomentulosa Standl. I.e. E. pur-

pusii Standl. Contr. U. S. Nat. Herb. 23: 1678. 15 Nov. 1926. E.

mexiae Standl. Field Mus. Bot. 4: 243. 1929. E. hiraeifolia Standl.

Field Mus. Bot. 17: 202. 1937.

Apparently comprising a single species is a series of almost iden-

tical populations ranging from Panama to northwestern Mexico.

The oldest name for the inclusive species seems to be that of Smith;

it seems odd that no member of this rather widely distributed Cen-

tral American group ever came to the attention of Berg.

Trees 3-5 m. high, or large shrubs, at least when young rather loosely pale-

or rufous-tomentose with somewhat contorted stalked or partly sessile dibrachiate

hairs; leaves rather large, elliptic-oblong or -obovate, sometimes broadly elliptic

or obovate, 6-16 (-20) cm. long, blunt or rounded or sometimes short-acuminate

at tip; inflorescence a stout tomentose raceme 2-4 cm. long, with several pairs of

rather large flowers on pedicels often 5-10 mm. long; buds 4-5.5 mm. long; disk

(3-) 4-5 mm. wide; calyx-lobes broadly rounded, tomentose on both sides; bracte-

oles linear or oblong, 2-5 mm. long, deciduous at or before anthesis; fruit ellipsoid-

oblong, 1.5-2 cm. long.

Certain rather ill-defined regional populations can be recognized,

as follows:

Leaves soon glabrate, usually glabrous beneath or nearly so even at flowering time;

hairs of the inflorescence appressed, sparse; leaf-margins cuneately decurrent

on the petiole var. rensoniana.

Leaves persistently and densely pubescent or tomentulose beneath; hairs of the

inflorescence abundant, stalked, forming a loose tomentum; leaf -margins

abruptly contracted to the petiole, the base of the blade sometimes sub-

auriculate, often asymmetric.

Leaves broadly obovate or broadly elliptic, 1.4-2 times as long as wide; lateral

veins with few intermediates, widely spaced, 6-8 on each side; Costa Rica

and Panama var. hiraeifolia.

Leaves elliptic to elliptic-obovate, 2-3 times as long as wide; lateral veins

8-12 on each side, the number indefinite because of the presence of inter-

mediate ones; Guatemala and Mexico var. salamensis.

Eugenia salamensis Bonn. Sm., var. salamensis. E. tomen-

tulosa Standl. Contr. U. S. Nat. Herb. 23: 1043. 1924. E. mexiae

Standl. Field Mus. Bot. 4: 243. 1929.

456 FIELDIANA: BOTANY, VOLUME 29

Leaves elliptic or obovate, blunt or rounded or short-acuminate at tip, nar-

rowed and then abruptly rounded to the base, scarcely decurrent, often subauricu-

late, 3-7 cm. wide, 6-12 (-16) cm. long, 2-3 times as long as wide; petioles 4-10

mm. long; midvein depressed and usually tomentulose, often with narrow elevated

central ridge when dry; pedicels 2-6 (-8) mm. long; bracteoles 2.5-5 mm. long;

style 7-9 mm. long.

Specimens examined :

MEXICO: Sinaloa: Without locality, Ortega 299 (US); San Ignacio,

Conzatti 299 (MEXU); Rosario to Colomas, Rose 3174 (US); San

Ignacio, Hda. Cana, Ortega 858 (K). — Nayarit: Acaponeta, Rose 1476

(US, type of E. tomentulosa) , M. E. Jones 22903 (UC); Acaponeta,

Tiger Mine, Jones 22902 (UC) ; 19 miles northwest of Tepic, McVaugh

& Koelz 725 (MICH). — Jalisco: South of Puerto Vallarta, Mexia

1129 (F, type of E. mexiae; MICH; UC), Carter & Chisaki 1204,

1205, 1209 (all MICH).— Colima: 7 miles north of Santiago, McVaugh

15915 (MICH).— Michoacan: Dist. Coalcoman, Hinton 16124, 16266,

16274 (all K, MICH & UC).— Guerrero: Dist. Montes de Oca, San

Antonio, Hinton 10578 (MICH, UC, US).

GUATEMALA: Salama: Llano Grande, Bergwald, Seler 2445 (US,

type).— Guatemala: Without locality, Aguilar 137 in 1939 (F).—

Quich<§ : Without locality, Aguilar 893, 1176 (both F).— Jalapa: Be-

tween Jalapa and Paraiso, Standley 77345 (F).

Eugenia salamensis Bonn. Sm., var. rensoniana (Standl.)

McVaugh, comb. nov. Psidium rensonianum Standl. Journ. Wash.

Acad. 14: 241. 1924. E. oaxacana Standl. Contr. U. S. Nat. Herb.

23: 1043. 1924, not E. oaxacana Berg, 1861. E. purpusii Standl.

Contr. U. S. Nat. Herb. 23: 1678. 1926.

Leaves on short shoots obovate with 5-8 lateral veins, obtuse or rounded,

on longer shoots elliptic, obtuse or acuminate, about twice as long as wide with

8 or more lateral veins; blades often acute at base and the margins evidently

and usually cuneately decurrent; petioles 5 mm. long; pedicels 3-10 mm. long;

bracteoles 2 mm. long; disk 3.5 mm. wide.

Specimens examined :

MEXICO: Oaxaca: San Geronimo [Ixtepec], July 1914, Purpus

7139 (F; UC; US, type of E. oaxacana Standl.).

EL SALVADOR: S. Calderon 838 (US, type of P. rensonianum),

1930 (US), Renson 339 (F, US).

This taxon is poorly known. It is characterized by little except

the thin pubescence of the inflorescence at anthesis, and by the gla-

brous or nearly glabrous leaves.

McVAUGH: TROPICAL AMERICAN MYRTACEAE 457

Eugenia salamensis Dorm. Sm., var. hiraeifolia (Standl.)

McVaugh, comb. nov. E. hiraeifolia Standl. Field Mus. Bot. 17:

202. 1937. Psidium rensonianum sensu Standl. I.e. 18: 779. 1937.

Leaves broadly obovate, broadly elliptic or oblong-obovate, rounded or ob-

scurely acuminate at tip (obtusely deltoid-acuminate on shoots), narrowed toward

the base from the middle or above but abruptly contracted to the petiole and often

unequally subauriculate, 4-6 (-11) cm. wide, 9-12 (-20) cm. long, 1.4-2 times as

long as wide; petioles 4-7 mm. long; midvein rather deeply but broadly depressed,

usually with elevated median ridge when dry; pedicels 3-8 mm. long; bracteoles

2-3.5 mm. long; style 5-6 mm. long.

Specimens examined:

COSTA RICA: Entre Canas y Tilaran, Guanacaste, Brenes 126 88 A

(NY), 12688 (F, NY); San Pedro de San Ramon, Brenes 21873 (F);

San Jose", Werckle (Herb. Nat. C. R. 16437) (F, NY); El Rodeo,

Villa Colon, M. Valeria 94-4 (F) ; Camino de Conception a Desmonte,

J. A. Echeverria 4-180 (F, UC); Pres du pont de Rio Grande, Chemin

de fer au Pacifique, Pittier (Herb. Nat. C. R. 16405) (US).

PANAMA: Monte Oscuro near Panama City, J. Zetek 3550 (F,

type of E. hiraeifolia).

The type of Eugenia hiraeifolia is a well-matured vigorous leafy

branchlet bearing a single raceme and one fruit; the pubescence has

almost all fallen, a little persisting in protected places on the leaves

and the fruit; the leaves toward the end of the shoot are unusually

elongated and bluntly acuminate, but I believe the specimen belongs

to the same species as the Costa Rican plants cited above. Miss

Amshoff has already pointed out (Act. Bot. Ne"erl. 5: 278-279. 1956)

the similarity between these Costa Rican specimens and the Pana-

manian E. hiraeifolia.

Plants of this variety are surely conspecific with those of var.

salamensis but are readily distinguished, at least in the specimens at

hand, by the larger and broader, more obtuse leaves with fewer and

more distinct veins.

Eugenia sinaloae Standl. Contr. U. S. Nat. Herb. 23:1042. 1924.

This species is known to me from the type only: Sinaloa: Guada-

lupe, Rose et al. 14793 (US). Very little can be added to the original

description, except to say that the pubescence is very like that in

Eugenia origanoides, namely of abundant pale erect hairs, readily

observed with the unaided eye. The flowers are unknown.

458 FIELDIANA: BOTANY, VOLUME 29

Eugenia standleyi McVaugh, nom. nov. Myrtus oaxacana

Standl. Contr. U. S. Nat. Herb. 23: 1038. 1924. Not Eugenia oaxa-

cana Berg, 1861, norE". oaxacana Standl. 1924.

The short-racemose inflorescence and the bibracteolate tetramer-

ous flowers establish this plant without question as a species of Eu-

genia. It is apparently still known from the type only:

Oaxaca: Between Juchitan and Chivela, Nelson 2631 (US).

Eugenia symphoricarpos McVaugh, sp. nov.

Arbor vel frutex, ramulis hispidulis; racemis foliorumque costis plusminusve

pilis crassiusculis aeruginosis rectis vel contortis usque ad 0.8 mm. longis obsitis;

folia elliptica vel ovata, 3-7 cm. longa, 1.5-2.8-plo longiora quam latiora, apice

acuminata, marginibus in petiolum 5-8 mm. longum cuneatim decurrentibus,

nervo medio supra depresso, laminis supra atroviridis lucidis, subtus pallidioribus;

flores numerosi, sessiles, glomerulis axillaribus dense congesti, bracteis oblongo-

lanceolatis vel ovatis obtusis 1.5-4 mm. longis, bracteolis conspicuis, vix connatis

sed imbricatis, sub hypanthium tomentulosum involucrum efformantibus; calycis

lobi concavi, majores 2 mm. longi latique; discus 1.5 mm. latus; stylus 7 mm.

longus; stamina ca. 60; ovarium biloculare, ovulis quoque loculo (3-) 5-8; fructus

ad modum Symphoricarpi glomerati, subglobosi, diametro 5 mm.

Probably a small tree, somewhat pubescent when young, finally glabrate;

branchlets hispidulous or antrorsely pubescent with dark reddish hairs; inflores-

cence, and both sides of the midveins of the foliage leaves, more or less beset

with rather coarse, straight or contorted coppery hairs up to 0.5-0.8 mm. long;

leaves elliptic or ovate, 1.5-4 cm. wide, 3-7 cm. long, 1.5-2.8 times as long as

wide, broadly and bluntly or rather slenderly acuminate at tip, rounded to the

cuneately narrowed base, the margins cuneately decurrent on the slender chan-

neled petiole 5-8 mm. long, 0.7-1 mm. thick; midvein depressed above in a

V-shaped channel, elevated at least half its diameter beneath, more or less per-

sistently pilose with contorted hairs; lateral veins 6-8 on each side including some

intermediate ones, convex on both surfaces in dry leaves but not very conspicu-

ous, straight and ascending; marginal vein about equaling the lateral ones and

arched between them, 0.5-1.5 mm. from the margin; blades dark green and lus-

trous above, when dry much paler and dull beneath, dark-dotted beneath; flowers

numerous in dense axillary glomerules up to 1-1.5 cm. in diameter, the raceme

axis (often 2 or more superposed) up to 3-6 mm. long, hirsute, bearing up to 3-6

approximate opposite decussate pairs of sessile flowers; buds 2.5 mm. long, obo-

void; bracts oblong-lanceolate to ovate, blunt-tipped, glabrous or coarsely ciliate,

about 1 mm. wide at base, 1.5-4 mm. long; bracteoles conspicuous, ovate, ca. 1.3

mm. long and more than 1 mm. wide, their margins overlapping but scarcely

connate, the whole forming a deep involucre enveloping and nearly concealing the

broadly campanulate, densely rufous-tomentulose hypanthium 0.7-1 mm. long;

calyx-lobes concave, thin, glandular-verrucose, suborbicular or a little longer or

shorter, the outer ones up to 1.5 mm. long and 2 mm. wide, the inner 2 mm. long

and wide; disk 1.5 mm. wide, hairy around the base of the style; style 7 mm.

long; stamens about 60, about as long as the style, the anthers 0.5 mm. long:

McVAUGH: TROPICAL AMERICAN MYRTACEAE 459

petals white, 3 mm. long; ovary bilocular, the ovules (3-) 5-8 in a compact group

in each locule; fruits in dense clusters, probably black, subglobose or obovoid,

up to 5 mm. in diameter, tipped by the prominent erect thin calyx-lobes.

At middle elevations, 800-1500 meters, in the Sierra Madre

Oriental, flowering April-May, fruiting May-September.

Tamaulipas: Sierra de Tamaulipas, Martinez M. & Borja L.

F-1923 (TEX), F-2076 (TEX).— Veracruz: Orizaba, C. Mohr in 1857

(US) ; Cerro del Carrisal, Orizaba, P. Tow in Apr. 1891 (MEXU) ;

Zacuapan, C. A. Purpus 2058 (GH, UC), 7523 (A, GH, NY; UC,

type) ; Paso del Correa, Liebmann 3970 (C) ; Barranca near Rancho

Viejo, Apr. 1934, Purpus 1625^ (A). — Locality unspecified: Mexico,

Sumichrast 1955 (K).

Eugenia teapensis McVaugh, sp. nov.

Frutex vel arbor minuscule sparseque puberula, foliis majusculis ellipticis

6-9 cm. latis, 14-20 cm. longis, obtuse acuminatis, venis lateralibus marginali-

busque et costa supra impressis, lateralibus prominentibus, intermediis vix evo-

lutis; racemi crassi usque ad 3 mm. longi 4-6-flori, pedicellis tenuibus 10-11 mm.

longis, bracteolis ovatis parvis persistentibus, floribus majusculis; alabastra 8 mm.

longa; calycis lobi concavi, inaequales, majores 6 mm. longi obovati; discus 3.5 mm.

latus; stylus 7-10 mm. longus; stamina plurima (ca. 200), antheris 1-1.2 mm.

longis; ovarium biloculare, ovulis quoque loculo ca. 12.

A tree or shrub, nearly glabrous, the foliage, branchlets and inflorescence

sparingly beset with minuscule erect hairlike processes; bracts, bracteoles and

calyx-lobes ciliate; leaves elliptic, rather large and prominently veined, 6-9 cm.

wide, 14-20 cm. long, 2-2.5 times as long as wide, about equally narrowed to the

bluntly acuminate tip and the acute to rounded base, the margins somewhat de-

current on the corky-rimose petiole 6-8 mm. long, 2-2.5 mm. in diameter; midvein

deeply and narrowly impressed above in a V-shaped cleft, elevated nearly its

whole diameter beneath; lateral veins 6-10 on each side, well separated and with-

out any conspicuous intermediate ones, prominent beneath, depressed below the

upper surface of the blade but the individual veins and veinlets convex in a re-

ticulate pattern; marginal vein about as strong as the lateral ones, similarly im-

pressed above, forming strong somewhat asymmetrical arches 5-10 mm. from the

margin; blades probably darker green and lustrous above and with some convex

glands, the lower surface sparingly dark dotted; inflorescence an abbreviated

raceme, the axis stout, dorsiventrally compressed, up to 1.5-3 mm. long, bearing

2-3 opposite decussate pairs of flowers on slender pedicels 10-11 mm. long; bracts

small; bracteoles ovate, persistent, distinct, 1 mm. long; buds 8 mm. long, obovoid,

the conic hypanthium 2.5 mm. long; calyx-lobes deeply concave, in unequal pairs,

the smaller pair orbicular, 4-5 mm. long and wide, the larger pair obovate, 5 mm.

wide, 6 mm. long; disk 3.5 mm. wide; style 7-10 mm. long; stamens about 200,

up to 9 mm. long, the anthers 1-1.2 mm. long; petals white, obovate, 10 mm. long;

ovary bilocular, the ovules about 12 in a compact group in each locule; fruit

unknown.

460 FIELDIANA: BOTANY, VOLUME 29

Tabasco: Forets sombres de Teapa, April (fl.), Linden 623 (K,

type). — Chiapas: Near El Ocote, ca. 30 km. northwest of Ocozo-

cuautla, 27-28 July 1958 (fl.), E. Hern&ndez X. (Hb. Miranda 8870)

(MICH).

Eugenia trunciflora (Cham. & Schlecht.) Berg, Linnaea 27:

223. 1856. Myrtus (?) trunciflora Schlecht. & Cham. Linnaea 5:

561. 1830.

The original collections of this species came from "sylvis umbro-

sis inter Mesachica et Mapilque" and from Papantla, both localities

in Veracruz, Schiede 547. I have not seen these specimens, but from

the original description and that given by Berg I suppose the follow-

ing belong to the same species; the leaves are mostly broadest at or

above the middle; the flowers are a little more than half as large as

those of E. xilitlensis (q.v.) and are borne on slender pedicels 8-

10 mm. long (the original description says "Pedunculi 3-9 lineas

longi"):

Veracruz: Zacuapan, C. A. Purpus 2434 (UC, US), 7663 (US).

Eugenia turneri McVaugh, sp. nov.

Arbor 6-7-metralis, subglabra, ramulis juvenilibus racemisque pubescentibus;

folia lanceolata vel elliptico-lanceolata, 3.5-5.5 cm. longa, 2.3-3-plo longiora quam

latiora, apice anguste sed obtuse acuminata, basi gradatim rotundata, petiolis

4.5-5 mm. longis, nervo medio supra inciso; racemi abbreviati 2-4 mm. longi, vel

flores ramulorum novorum nodis infimis solitarii bracteatique; pedicelli filiformes

8-12 mm. longi; bracteolae scariosae, distinctae, vix deciduae; alabastra 3.5 mm.

longa, supra hypanthium glabrum subglobosa; calycis lobi membranacei, post

anthesin reflexi, majores 2 mm. longi; discus 1.5 mm. latus, annulus staminalis

subhirsutulus; stylus 4 mm. longus; ovarium biloculare, ovulis quoque loculo 2

(-1); fructus ignotus.

Tree 6-7 meters high, nearly glabrous, the vegetative buds, axis of the raceme,

and the very youngest growth appressed-pubescent with pale hairs 0.1-0.3 mm.

long; leaves lanceolate or elliptic-lanceolate, 1.5-2 cm. wide, 3.5-5.5 cm. long,

2.3-3 times as long as wide, the tip slenderly but bluntly acuminate, the base

gradually rounded, the margins at base passing abruptly to the rather deeply

channeled petiole 4.5-5 mm. long, 0.6 mm. thick; midvein slightly engraved above

in the nearly plane surface of the blade; lateral veins 6-8 on each side including

some intermediate ones, slightly elevated when dry but very inconspicuous on

both sides; marginal vein about equaling the lateral ones and somewhat arched

between them, less than 1 mm. from the margin; blades nearly concolorous and

yellow-green when dry, very smooth but apparently not lustrous above, a little

paler beneath and roughened by many small inconspicuous glands; inflorescence

an abbreviated axillary raceme (sometimes at leafless nodes or terminal!), or the

flowers solitary and opposite at the lowermost (bracteate) nodes of new leafy

branchlets; raceme-axis 2-4 mm. long, bearing 2-3 pairs of flowers on filiform

McVAUGH: TROPICAL AMERICAN MYRTACEAE 461

pedicels 8-12 mm. long; bracts scarious, pubescent, 0.5 mm. long; bracteoles lance-

ovate, 0.5-0.7 mm. long, scarious, distinct, ciliate, subpersistent; buds 3.5 mm.

long, subglobose above the short-campanulate glabrous hypanthium 1 mm. long;

calyx-lobes membranous, ciliate, unequal, the outer pair broadly rounded or ellip-

tic-ovate, 1.2 mm. wide, 1.5 mm. long, the inner pair suborbicular-oval, 1.7 mm.

wide and 2 mm. long, all the lobes reflexed from near the base soon after anthesis;

disk about 1.5 mm. wide, the staminal ring bristly; style 4 mm. long; stamens

about 50, about as long as the style, the anthers about 0.7 mm. long and wide with

broad basal lobes; petals white, ovate, ciliate, 3.5-4 mm. long; ovary bilocular,

the ovules 2 and collateral, or only 1, in each locule; fruit not seen.

For a note on the relationships of this species, see above under

E. rhombea.

Michoacan: 1 mile east of La Placita, near the Pacific Coast, dry

stream bottom, 4 July 1950 (fl.), B. L. Turner 2073 (MICH, type).

Eugenia xalapensis (HBK.) DC. Prodr. 3: 276. 1828. Myrtus

xalapensis HBK. Nov. Gen. & Sp. 6: 145. 1823.

Shrub or small tree 1-4 meters high, soon glabrate, the growing branchlets

and the youngest petioles minutely hispidulous-puberulent, the mature leaves

and the pedicels and flowers quite glabrous; leaves ovate or ovate-elliptic, 1.5-

3.5 cm. wide, 3-7 cm. long, 1.7-2.7 times as long as wide, the tip prominently

acuminate (the very tip blunt), the base acute, the margins cuneately decurrent

on the inner angles of the channeled petiole 4-8 mm. long; midvein deeply and

narrowly impressed above; lateral veins 5-6 on each side in addition to some inter-

mediate ones, inconspicuous above, apparent beneath; marginal vein about as

strong as the lateral ones and little arched between them, 0.5-1.5 mm. from the

margins; blades dark green and lustrous above, much paler beneath; glands appar-

ent beneath as small dark or resinous dots, sometimes as convexities above; in-

florescence an abbreviated axillary raceme, or some flowers opposite at the lower-

most (bracteate) nodes of new leafy branchlets; raceme-axis up to 1 cm. long with

4 pairs of flowers, usually 2-4 mm. long with 1-3 opposite and decussate pairs of

flowers on pedicels 3-5 mm. long; bracts indurate-persistent, blunt, oblong or

ovate, 1-1.5 mm. long; bracteoles connate, oblong-ovate, obtuse, forming a deeply

2-lobed involucre with divaricate lobes 0.8 mm. long and wide; buds 3-3.5 mm.

long, pyriform, the hypanthium turbinate or campanulate, often somewhat atten-

uate at base, 1-1.5 mm. long; calyx-lobes suborbicular, thin-margined, the larger

ones 1.3-2 mm. long and wide; disk 1.5 mm. wide (2-2.5 mm. wide in fruit); style

(not seen expanded) probably about 5 mm. long; stamens 60-70, up to 4 mm.

long, the anthers 0.5 mm. long; petals white, about 3 mm. wide, concave, suborbic-

ular; ovary bilocular, the ovules about 10 in a compact group in each locule; fruit

black, globose, 10-13 mm. in diameter, the calyx-lobes scarcely persistent.

Veracruz: Xalapa, 770 hex., Bonpland (P, type, not seen; Field

Mus. Neg. 36911), Schiede 543 (NY), Rose & Hay 6173 (NY), L. H.

MacDaniels 802 (F), 849 (F), C. R. Orcutt 2855 (F), C. L. Smith 1507

(F), 1898 (F).— Hidalgo: Bluffs near Tutotepec, 5300 ft., 28 Aug.

1945 (fr.), A. J. Sharp 45866 (MICH); Dist. Molango, northwest

462 FIELDIANA: BOTANY, VOLUME 29

of Lake Atexca, deciduous woods, 1500 m., 9 Nov. 1946 (fr.), H. E.

Moore 1953 (UC). — San Luis Potosi: 20 mi. east of C. del Maiz, oak

forest, 1200 m., 9 May 1949 (fl.), McVaugh 101+85 (MICH).— Ta-

maulipas: 5 mi. northwest of Gomez Farias, 3600 ft., 15 May 1949

(fl.), B. E. Harrell 179 (MICH).— State:? Joya de Salas trail above

El Cielo, 1 Sep. 1952 (fr.), Sharp et al 52218 (MICH); San Luis

Potosi to Tampico, Dec. 1878-Feb. 1879, Palmer 1045 (GH, NY).

This rather local species was correctly interpreted by Schlechten-

dal and Chamisso (Linnaea 5: 560. 1830) and by Standley (Contr.

U. S. Nat. Herb. 23: 1040, in key; 1045. 1924), but the name E. xala-

pensis has been erroneously applied to many specimens from the

lowlands of eastern Mexico and adjacent Guatemala. Sterile speci-

mens of E. xalapensis may ordinarily be distinguished from those

of E. symphoricarpos (which occurs in the same general area) by the

glabrous branchlets; in E. xalapensis only the very youngest grow-

ing branchlets are pubescent.

Eugenia xilitlensis McVaugh, sp. nov.

Arbor 7-8-metralis, racemis abbreviates hirsutulis; folia lanceolato-elliptica,

16-20 cm. longa, ca. 3-plo longiora quam latiora, acuminata, basi cordato-

auriculata, auriculis rotundatis 1-1.5 mm. longis; petioli 8-10 mm. longi; nervus

medius supra concavus; lamina subtus venis elevatis glandulisque numerosis

maximis instructa; flores 1-2, subsessiles, e nodis defoliatis oriundes, pedicellis

hirsutis 3 mm. longis; alabastra 10-11 mm. longa, hypanthio canescenti, brac-

teolis deciduis 3.5 mm. longis; calycis lobi maiores 7 mm. lati, 9 mm. longi; discus

6-7 mm. latus; stylus 15 mm. longus; stamina ca. 300, antheris 1.2-1.5 mm.

longis; ovula quoque loculo ca. 15-20.

A tree up to 7-8 meters high, the young branchlets and foliage sparingly

hirsutulous, the inflorescence thickly hirsutulous with pale hairs; leaves lance-

elliptic, 5.5-6.5 cm. wide, 16-20 cm. long, about 3 times as long as wide, gradually

and bluntly acuminate; blades rounded from the middle or below to the cordate-

auriculate base, the auricles rounded, 1-1.5 mm. long, 3-4 mm. wide; petioles

8-10 mm. long, 2 mm. thick; midvein concave above; lateral veins 10-15 on each

side including some intermediate ones; marginal vein about as strong as the lateral

ones and forming angular arches between them, 2-5 mm. from the margin;

leaves green and smooth above, sparingly gland-dotted, the veins appearing as

fine pale parallel lines; lower surface paler, more prominently dotted with glands

0.2-0.4 mm. in diameter, the veins elevated and more evident; flowers 1-2,

nearly sessile at leafless nodes, the pedicels hairy, 3 mm. long, 1-1.5 mm. thick;

buds 10-11 mm. long, pyriform, the cup-shaped hypanthium canescent, 3-4

mm. long; bracteoles ovate, membranous, deciduous, 2 mm. wide, 3.5 mm.

long; calyx-lobes rounded, in unequal pairs, prominently glandular, thin, pu-

bescent along the edges, the larger ones 7 mm. wide, 9 mm. long, the smaller

4.5-5 mm. wide, 6 mm. long; sinuses splitting a little between the lobes after

anthesis (perhaps as a result of pressing in the specimen seen); disk 6-7 mm.

wide; style 15 mm. long; stamens about 300, about as long as the style, the anthers

McVAUGH: TROPICAL AMERICAN MYRTACEAE 463

1.2-1.5 mm. long; petals white, ovate, narrowed to the tip, ciliate-fringed, up

to 1.5 cm. long or more; ovary bilocular, the ovules 15-20 in a compact group

in each locule; fruit unknown.

San Luis Potosi : In forest west of Xilitla, Cerro Prieto, 3800 feet,

22 Apr. 1946 (fl.), A. J. Sharp 46284 (MICH, type).

This species occurs in the same general region as Eugenia trunci-

flora, which it somewhat resembles. For a key to the North Ameri-

can species of Eugenia with cordate-auriculate leaves, see below

under Notes on Miscellaneous Species.

Eugenia yautepecana Lundell, Wrightia 2: 107. 1960.

A shrub 3-4 meters high, the foliage nearly glabrous from the first, the

branchlets and inflorescence (and the youngest leaves) densely appressed-pubescent

with pale reddish or almost colorless dibrachiate hairs mostly 0.2-0.4 mm. long,

the whole plant ultimately glabrate; leaves elliptic or occasionally ovate, 1.5-

3.5 cm. wide, 3-6 cm. long, 2-2.4 times as long as wide (or leaves near the bases

of branchlets 1.4-1.6 times as long as wide); blades often about equally narrowed

to a blunt or obscurely acuminate tip and a rounded or acute base, the margins

usually passing abruptly to the channeled petiole 3 mm. long (bases of the broader

leaves rarely subcordate, or in the narrower ones rather long-decurrent) ; mid-

vein broadly and shallowly concave above at least near base, or shallowly chan-

neled between low parallel ridges; lateral veins 5-7 on each side, ascending, pale

and readily visible on both surfaces of dried leaves, scarcely forming a marginal

vein but recurving in a series of irregular angled arches 2-3 mm. from the margin

to join the next succeeding ones; blades dark green and lustrous and very minutely

pale-verruculose above, paler beneath, with numerous convex resinous glands

usually apparent on both surfaces; inflorescence an abbreviated axillary raceme

or two racemes superposed, the axis up to 3-6 mm. long, bearing 1-3 opposite

and decussate pairs of flowers on pedicels 3-5 (-7) mm. long, up to 0.6-0.8 mm.

thick in fruit; bracts ovate, 1 mm. long, the lower ones broader; bracteoles ovate,

divaricate, distinct, persistent, 1-1.4 mm. long; buds 4 mm. long; calyx-lobes

broadly rounded, concave, sparingly beset with large resinous convex glands

0.2-0.3 mm. in diameter, appressed-pubescent on both surfaces but persistently

and more densely on the inner; inner lobes broadly rounded, 2.5-3 mm. wide,

2.5 mm. long, the outer often ovate, 1.5-2 mm. wide and long; disk 2.5-3 mm.

wide, somewhat hairy about the base of the style; style 5-6 mm. long or more;

stamens about 75, about as long as the style, the anthers 0.5-0.7 mm. long;

ovary bilocular, the ovules about 10 in a compact group in each locule; fruit

ellipsoid-oblong, black(?), 8-10 mm. in diameter, 10-15 mm. long, crowned

by the erect concave calyx-lobes.

Veracruz: Barranca de Panoaya, Aug. 1919 (fl.), C. A. Purpus

8432 (UC, US), July 1920, Purpus 8505 (UC), Purpus 8561 (UC).—

Mexico: Temascaltepec, Acatitlan, 27 Dec. 1934 (fr.), G. B. Hinton

7172 (K, MICH, UC, US) .— Morelos : Cuernavaca, pedregal, 23 Sep.

1896 (imm. fr.), Pringle 7234 (MICH); Yautepec-Cuernavaca road,

lava field, 2 Oct. 1943 (imm. fr.), Lundell & Lundell 12502 (MICH,

US, isotypes).

464 FIELDIANA: BOTANY, VOLUME 29

NOTES ON MISCELLANEOUS SPECIES

Eugenia basilaris McVaugh, sp. nov.

Frutex vel arbor parva 2-4 m. alta, ramulis rufescentibus longitudinaliter

rimosis, decor ticantibus; ramuli hispido-hirsuti, pilis acutis fuscis 0.5-1 mm.

longis obsiti; folia elliptica vel ovata, basi plusminusve cordata; petioli crassi

3 mm. longi; nervi laterales utrinque 5-8, arcuatim ascendentes, nervum mar-

ginalem vix efformantes; flores glomerati subsessiles, pedicellis 1 mm. longis

subglabris; alabastra 3-4 mm. longa pyriformia, hypanthio hirsute anguste urce-

olato 1.2-1.5 mm. longo; calycis lobi subglabri mox reflexi; discus 1.5 mm. latus,

centre concavo, marginibus reflexis; fructus subglobosus diametro 1.7-2 cm.,

apice conico, disco peltato 2-2.5 mm. lato coronato; ovarium biloculare, loculis

basilaribus, ovulis quoque loculo 5-7.

Shrub or small tree 2-4 m. high, the younger branchlets wrinkling in sharp

ridges when dry, the thin light reddish brown outer bark in mature branchlets

cracking longitudinally and flaking off; branchlets, petioles, and young leaves

at least on the lower surface hispid-hirsute with erect or flexuous stout sharp

dark reddish-brown hairs 0.5-1 mm. long; leaves elliptic or elliptic-ovate, 3-5

cm. wide, 6-11 cm. long, 2-2.5 (-3) times as long as wide, broadly and bluntly

or sometimes slenderly acuminate, rounded toward the base from the middle

or below, the base itself broadly or narrowly rounded, cordate or cordate-

auriculate, with a definite shallow often asymmetric sinus between the rounded

auricles; petiole about 3 mm. long, 1.5 mm. thick; mid vein impressed above,

elevated its whole thickness beneath; lateral veins 5-8 on each side, seen as

fine lines above and often somewhat depressed when dry, prominent beneath,

scarcely forming a marginal vein, but diminishing distally and arching over at

a point 3-5 mm. from the margin to join the next succeeding vein; leaves often

blackening in drying, darker above, not prominently glandular; flowers in axillary

glomerules, the inflorescence a crowded axillary raceme or two racemes super-

posed, the axis up to 3-3.5 mm. long, 4-angled, bearing up to 4 pairs of flowers

on stout nearly glabrous pedicels 1 mm. long (in fruit up to 3 mm. long, 1.5 mm.

thick); bracts oblong, hirsute outside, 1.5 mm. long; bracteoles like the bracts,

subpersistent, not connate, 1 mm. long; buds 3-4 mm. long, pyriform, the globe

of the petals broader than the hirsute narrowly urceolate hypanthium 1.2-1.5

mm. long; calyx-lobes ciliate but nearly glabrous, soon reflexed, the larger ones

1.5-1.7 mm. wide, 2-3 mm. long; disk 1.5 mm. wide, the center concave, the

staminal ring hairy, the margins soon much reflexed; style about 6 mm. long;

fruit subglobose, glabrous, glandular- verrucose, 1.7-2 cm. in diameter, pro-

longed into a broadly conical apex surmounted by the peltate disk 2-2.5 mm.

wide, the calyx-lobes radiate, rounded, glabrous within; ovary bilocular, the

locules expanded near the base of the elongate hypanthium; ovules 5-7 in each

locule.

Specimens examined:

Costa Rica: San Ramon, bois des collines, 1200-1400 m., 5 May

1913, A. Tonduz (Herb. Nat. C. R. 17662) (F, MICH, US); San

Pedro de San Ramon, bois, collines, 850-1075 m., A. M. Brenes 4282

(F, type), 4267, 4323, 4538, 5022 (all F).

McVAUGH: TROPICAL AMERICAN MYRTACEAE 465

This most distinctive species has been confused with E. truncata

Berg, to which it bears little resemblance except that the leaves are

about the same size. The flowers in E. truncata are borne on slender

pedicels 10-25 mm. long.

Eugenia truncata Berg, Linnaea 27: 157. 1856. E. oreinoma

Berg, I.e. 158. ?E. guanacastensis Standl. Field Mus. Bot. 8: 144.

1930.

According to Berg E. truncata was distinguished from E. oreinoma

chiefly by the narrower and very short-petiolate leaves truncate at

base. Study of the abundant Costa Rican material now available

suggests that but one species is involved. The type of E. truncata,

which I have seen through the courtesy of the authorities at Copen-

hagen, is a strongly pubescent specimen consisting mostly of young

vegetative branchlets 3-5 cm. long; the leaves are all truncate and

small for the species (cf. Field Mus. Neg. 20982). An isotype of

E. oreinoma (seen at F) bears relatively large broad leaves (5.6-7 cm.

wide, 8.5-13 cm. long) which are merely rounded at the base. In

other respects the two specimens are similar.

The glabrous or nearly glabrous plant described as E. guanacas-

tensis is evidently very closely related to E. truncata and may be a

synonym of that species, as Standley ultimately decided. The young

shoots are sometimes hirsutulous exactly as in E. truncata, (e.g. in

Standley & Valerio 4424-0), or the whole plant may be glabrous.

The petioles are but 3^4 mm. long as against the usual length of

5-8 mm. in E. truncata. The leaves are always auriculate-cordate.

In North America the leaves in most species of Eugenia are acute

or cuneate at base, or if the margins are rounded abruptly in to the

petiole they are usually at least slightly decurrent on the petiole;

the summit of that organ may appear to be dilated or the blade may

be said to be cuneately narrowed at base. In a few species, how-

ever, the blades are sometimes or always cordate or cordate-auricu-

late, the margins seeming to merge with the ventral surface of the

petiole while curving away from its base. In several species (e.g.

E. basilaris, E. truncata) the leaves are usually truncate or sub-

cordate but may be merely rounded at base; in E. origanoides and

E. salamensis the leaves are infrequently subcordate; E. papalensis

is known only from the type, which may or may not be a represent-

ative specimen. In E. chinajensis, E. jutiapensis, E. trunciflora and

E. xilitlensis, as far as known, the leaves are always cordate-auriculate.

466 FIELDIANA: BOTANY, VOLUME 29

Flowers in elongate axillary racemes 2-5 cm. long; pubescence of appressed and

mostly dibrachiate hairs E. salamensis Donn. Sm.

Flowers glomerate or fasciculate, the racemes so much contracted that the flowers

appear to arise directly from the axils or from very short lateral spurs; pubes-

cence if present mostly of more or less erect simple hairs.

Pedicels slender, hispidulous (rarely glabrous), 1-2.5 cm. long; buds 5-7.5 mm.

long.

Hypanthium in flower glabrous or very sparingly pubescent especially at base.

Lateral veins 10-12 on each side of the midvein; foliar glands up to 0.5 mm.

in diameter, apparent on both sides of dry leaves, more conspicuous

beneath; Veracruz E. trunciflora (Cham. & Schlecht.) Berg.

Lateral veins 6-8 on each side; foliar glands mostly 0.1-0.2 mm. in diam-

eter, scarcely visible in dried leaves; Costa Rica E. truncata Berg.

Hypanthium in flower densely velutinous or hirsute; lateral veins 5-6; foliar

glands 0.1-0.2 mm. in diameter, visible from the upper surface or not

at all; El Salvador and Guatemala E.jutiapensis Standl. & Steyerm.

Pedicels very short, 3 mm. long or less, or the flowers sessile and glomerate.

Petioles 8-15 mm. long, the blades 12-20 cm. long.

Blades rounded to truncate at base; foliar glands scarcely apparent; flowers

unknown, probably large, the disk in fruit 3-3.5 mm. wide, the larger

calyx-lobes 3-4 mm. long and wide; Guatemala.

E. papalensis Standl. & Steyerm.

Blades cordate-auriculate; foliar glands prominent beneath, 0.2-0.4 mm.

in diameter; buds 10-11 mm. long; disk in flower 6-7 mm. wide, the

larger calyx-lobes 7 mm. wide, 9 mm. long; San Luis Potosl.

E. xilitlensis McVaugh.

Petioles 5 mm. long or less; blades mostly 10 cm. long or less; buds 2-4 mm.

long.

Plants uniformly and coarsely but not very densely setose, the reddish or

tawny hairs in no way obscuring the surface of the branchlets or the

lower leaf-surface; setae on the midvein near the base of the lower sur-

face of the leaf 1-1.3 mm. long; cilia of the calyx-lobes 0.5-0.7 mm.

long; eastern Guatemala E. chinajensis Standl. & Steyerm.

Plants finely and often densely soft-hirsutulous, the pale hairs often so

numerous as to cover the branchlets and principal veins; hairs of the

midvein near the base of the lower leaf-surface 0.5 (-0.8) mm. long; cilia

of the calyx-lobes mostly 0.2-0.4 mm. long.

Hypanthium narrowly urceolate; fruit 1.7-2 cm. in diameter; Costa Rica.

E. basilaris McVaugh.

Hypanthium hemispheric to subglobose; fruit 6-8 mm. in diameter;

Mexico to Colombia E. origanoides Berg.

Eugenia egensis DC. Prodr. 3: 281. 1828.

This species has not previously been reported from North Amer-

ica. It is one of the most distinctive South American species, and

I think there can hardly be any doubt of the determination of the

specimen cited below. The principal range of E. egensis is from the

McVAUGH: TROPICAL AMERICAN MYRTACEAE 467

Guianas to eastern Colombia, south chiefly in the Amazon basin to

eastern Bolivia.

Costa Rica: El General, bordes del Rio Pacuare, 600 m., 17 Sep.

1942, J. Leon 1008 (US).

Eugenia flavoviridis Lundell, Amer. Midi. Nat. 29: 479. 1943.

E. dissitiflora Lundell, Wrightia 2: 207. Sep. 1961. E. flavida Lun-

dell, Wrightia 3: 14. Dec. 1961.

This appears to be a distinct taxon of the Yucatan region, super-

ficially distinguished from other species with crowded racemes by its

long-petiolate, elliptic (rather than ovate) leaves, by the rather

sharply and narrowly impressed (rather than flat or merely de-

pressed) mid vein, by the coarse ascending pubescence of the race-

mes, and also by the very smooth upper leaf-surface in which the

veins are scarcely apparent to the eye. Material from British Hon-

duras does not seem to differ appreciably from specimens of E. dis-

sitiflora recently collected in Pete"n. Of the approximately 15 speci-

mens I have seen, all bear immature fruit. It is unfortunate that

no flowering material is available for study. Species of this affinity

are numerous in the West Indies, and it may be that the Central

American plant will eventually prove to be a synonym of one of these.

The type and only known specimen of Eugenia flavida (Guate-

mala, Pete"n, Dos Lagunas, in low secondary forest of airfield clear-

ing, E. Contreras 1693), which I have seen through the courtesy of

Dr. Lundell, is a mature leafy branch from which almost all traces

of pubescence seem to have been eroded. Vegetatively I cannot dis-

tinguish this specimen from similarly weathered leafy twigs of E.

"dissitiflora" from nearby localities in Pete"n. The fruits accompany-

ing the type specimen of E. flavida have apparently been partly

rotted while lying on the ground ; none is whole, but from the shape

of the seeds the fruit seems to have been globose or didymous, as in

E. ff dissitiflora" (E. flavoviridis). Flowering material of E. flavida is

to be sought. The type of E. flavida bears several persistent raceme

axes 2-4 mm. long, these with indurated ovate bracts. The racemes

are shorter than those of an average specimen of E. flavoviridis but

can be matched in some specimens of that species.

Eugenia hondurensis A. Molina, Ceiba 1: 261. 1951. E. crassi-

folia A. Molina I.e. 3: 169. 1953. E. nicaraguensis Amsh. Act. Bot.

Ne"erl. 5: 278. 1956.

468 FIELDIANA: BOTANY, VOLUME 29

This appears to be a distinctive species related to the series in-

cluding E. salamensis Donn. Sm., E. purpusii Stand!., E. mexiae

Standl., and E. tomentuiosa Standl., but differing from all these in

having the raceme reduced to a single (terminal) flower. There is

much variation in leaf-form in this species, even on the same plant,

the leaves on vigorous shoots often becoming large and broad.

Flowering specimens have an aspect quite unlike that of fruiting

specimens because the maturing branchlets and foliage lose most of

the reddish tomentum that is abundant and characteristic on the

young growth, and the leaves become firm and smooth.

The original specimens of E. hondurensis were from mature plants

with half -grown fruit (Molina 2559, US !) ; flowering specimens were

subsequently described under the name of E. crassifolia. Some of

the latter, including the type of crassifolia (Molina 3031, US!), bore

relatively broad obovate leaves. The Nicaraguan specimens de-

scribed under the name of E. nicaraguensis were all in flower or in

bud ; the suite included some with very large broad leaves, and some

with smaller narrow leaves. The type (Standley 94.27, F!) is a flower-

ing specimen with moderately broad leaves a little larger than those

of the type of hondurensis. There seems to be no basis for recogniz-

ing more than one species in this group, the variation being appar-

ently individual and seasonal.

Eugenia laevis Berg, Linnaea 27: 177. 1856. E. subverticillaris

Berg, Linnaea 29: 235. 1858 [type from Hispaniola, Poiteau (LE!)].

Previously unknown from the North American continent, this

rather distinctive species can now be reported from Yucatan. The

specimens cited below agree precisely with specimens from Hispani-

ola, viz. var. laevis. Especially noteworthy are the peculiarly and

minutely verruculose leaves, the pale dibrachiate hairs of the ped-

icels and the canescent hypanthium, the concentration of brownish

hairs at the tips of the rather long calyx-lobes, which are otherwise

glabrous on the inner surface.

MEXICO: Yucatan: Izamal, G. F. Gaumer in 1888 (F, ex herb.

Kew); 3 km. de El Cuyo en el camino a Colonia Yucatan, 7 Mar.

1956, 0. G. Enriquez 477 (MICH); entre Valladolid y Temozon,

7 June 1956, Enriquez 584 (MICH).

The very recently described Eugenia calciphila Lundell (Wrightia

3: 11. Dec. 1961) is known only from the type:

McVAUGH: TROPICAL AMERICAN MYRTACEAE 469

GUATEMALA: Pete*n: Dos Lagunas, in ramonal, E. Contreras

1541 (LL).

This specimen is in young bud, with a few open flowers. The

pubescence is less abundant than in the specimens from Yucatan

and Hispaniola; the pedicels are nearly glabrous and the hypanthium

merely strigose. I cannot find any other significant differences be-

tween this plant and other specimens of E. laevis, and I regard

E. calciphila as a synonym of that species.

Eugenia laevis Berg, var. gaumeri (Standl.) McVaugh, comb,

nov. E. gaumeri Standl. Field Mus. Bot. 81 28. 1930. E. lundellii

Standl. Carnegie Inst. Wash. Publ. 461: 76. 1935.

An ample series of well-prepared specimens from Pete"n, collected

by C. L. Lundell and his assistant, Elias Contreras, and made avail-

able to me through the courtesy of Dr. Lundell, makes it clear that

Eugenia gaumeri and E. lundellii are not distinguishable unless by

leaf shape, and probably not even by that feature. The original

specimens of E. gaumeri (Kancabtsonot, Yucatan, Gaumer 23843!,

the type) were described as having narrow leaves (1.3-1.8 cm. wide,

2-3 times as long as wide), the blades widest at the middle or below,

narrowed to an obtuse apex, the very tip commonly rounded. In

contrast, E. lundellii was described (Tuxpena, Campeche, Lundell

1130!, the type) as having elliptic or obovate-elliptic or rarely ob-

long-obovate leaves (1-2.5 cm. wide, 3-4 cm. long), obtuse or

rounded at the apex. I do not find these differences significant;

broadly obovate leaves may often be found on the same plant with

rather narrowly elliptic ones, and some specimens from Pete"n (e.g.

Lundell 15574, Contreras 1441) are generally narrow-leaved, the foli-

age about as in some specimens from Yucatan (e.g. Lundell 7933),

and from British Honduras (e.g. Gentle 1202, 1745).

The population named above as Eugenia laevis var. gaumeri seems

to differ from var. laevis chiefly in the shape and size of the leaves;

these in var. laevis are mostly 4.5-6.5 cm. long and rather narrowly

and prominently acuminate; in var. gaumeri they are mostly 2-5 cm.

long, at most broadly and bluntly acuminate.

Additional specimens examined:

MEXICO: Yucatan: Lundell 7416.

GUATEMALA: Pete"n: Bartlett 12801; Contreras 26, 1362, 1368,

1369, 1370; Lundell 15518, 15527, 15975, 16489, 16824.— Baja Vera-

paz: Standley 69775 (F).

470 FIELDIANA: BOTANY, VOLUME 29

Eugenia octopleura Krug & Urb. ex Urb. Bot. Jahrb. 19: 653.

1895. E. doubledayi Standl. Journ. Arnold Arb. 11: 36. 1930.

E. koepperi Standl. Field Mus. Bot. 9: 320. 1940.

The type of E. doubledayi (Honduras, Siguatepeque, Standley

56063} is a fruiting specimen collected in February. Very little

color is observable in the scanty pubescence persisting on the leaves.

Immature branchlets, however, are closely and densely coppery-

pubescent, exactly as in flowering specimens. The type of E. koep-

peri (Honduras, Mt. Cangrejal near La Ceiba, Yuncker et al. 8^00)

is a specimen with developing inflorescences; the reddish color of the

pubescence is evident on the lower leaf-surfaces. The leaves are

bluntly acuminate or even obtuse, and the petioles are 5-8 mm.

long. Superficially the specimen seems unlike specimens of E. double-

dayi in which the leaves are usually prominently acuminate and the

petioles 8-15 mm. long. Except for these features I cannot distin-

guish the two so-called species.

Certain Costa Rican specimens (Zarcero, 4500 feet, Austin Smith

A239) are seemingly identical with flowering specimens of E. double-

dayi. Other Costa Rican material obviously of the same species

(La Palma de San Ramon, 1250 m., Brenes ^625; La Palma, Prov.

S. Jos^, 1460 m., Tonduz [Herb. C. R. 12558, J. D. Smith 7382}) is

matched perfectly by two syntypes of E. octopleura: Duss 2759, 3270.

On the basis of this evidence, E. doubledayi and E. koepperi should

pass into synonymy.

3. MARLIEREA Camb.

Reference: Marlierea, Sect. Myrciopsis McVaugh, Mem. N. Y.

Bot. Gard. 10: 79. 1958.

Through an unintentional orthographic error, the name of this

well-marked section was originally published as Myrcioides (I.e. 79).

It was correctly cited as Sect. Myrciopsis on pages 84, 86 and 87 of

the article in which it was first published.

4. MYRCIA DC.

MISCELLANEOUS NOTES

Myrcia splendens (Sw.) DC. Prodr. 3: 244. 1828. Myrtus

splendens Sw. Prodr. 79. 1788. M. rufidula Schlecht. Linnaea 13:

McVAUGH: TROPICAL AMERICAN MYRTACEAE 471

416. 1839. M. costa-ricensis Berg, Linnaea 27: 104. 1855. M. dis-

color Berg, I.e. 111. M, oerstedeana Berg, I.e. 112. M. melanoclada

Berg, I.e. 113. M. plicato-costata Berg, I.e. 114. M. sartoriana Berg,

Linnaea 29: 220. 1858. M. longicaudata Lundell, Am. Midi. Nat. 29:

481. 1943. M. schippii Lundell, I.e. 482.

Additional synonymy, and some comments on the distribution

and taxonomy of this species, may be found in the Flora of Peru

(13, pt. 4: 659-661. 1958). I noted there that Myrtus splendens

Sw. perhaps was a synonym of Eugenia periplocifolia Jacq. Coll. 2:

108. 1788. I now doubt that the two are synonymous, for in the

description of E. periplocifolia the plant is said to be "tota glabra";

one of the most characteristic features of Myrcia splendens is the

abundant pubescence, readily seen with the unaided eye.

In the West Indies Myrcia splendens is represented primarily by

a form with ovate or ovate-lanceolate leaves 1.5-3 cm. wide, 3-7 cm.

long, 1.7-2.8 times as long as wide. In Trinidad, in South America

generally, and in Central America and Mexico, the leaves are pre-

vailingly elliptic-lanceolate or elliptic, 6-12 cm. long and mostly

2.5-3 (-4) times as long as wide. After the examination of a large

suite of specimens I am unable to find any consistent differences

between North American and South American material.

Through the kindness of the authorities at Copenhagen, I have

seen the types of the five Costa Rican species based by Berg upon

the collections of Oersted. With the exception of the type of M. pli-

cato-costata, none of the specimens seems in any way exceptional for

M. splendens. The Oersted specimen of M. plicato-costata (see Field

Mus. Neg. 21035) has a few veins of the longer leaves impressed on

the upper surface, a feature matched in a number of other Costa

Rican specimens. The specimens are otherwise so similar to M.

splendens that I doubt their specific distinctness. All are represented

by photographs in the Chicago Natural History Museum series ex-

cept for M. oerstedeana. This is a fruiting specimen with rather

narrow, caudate leaves, the blades about 2.5 cm. wide, 8 cm. long;

the pubescence is about that of an average specimen of M. splendens.

A related species, Myrcia mollis (HBK.) DC., known chiefly from

the eastern Cordillera of Colombia, has ovate and short-petiolate

leaves that are often rugose and have the veins impressed on the

upper surface. The pubescence is silky, more copious than usual in

M. splendens, and extending even to the inner faces of the calyx-

lobes. Similar plants occur as far north as Costa Rica; it may be

that M. plicato-costata Berg, mentioned above, is akin to M. mollis.

472 FIELDIANA: BOTANY, VOLUME 29

The whole species-complex to which M . splendens belongs [in-

cluding such species as M. fallax (Rich.) DC., M. mollis, M. acumi-

nata (HBK.) DC.] is variable with respect to the amount and qual-

ity of the pubescence. The branchlets and the inflorescence are

appressed-strigose or -silky, or in extreme forms merely sparingly

strigose, or at the other extreme almost hirsute (cf. Flora of Peru,

pt. 4: 660-661). It has been the fashion in recent years to recognize

in Mexico and Central America two species of Myrcia, one with

spreading pubescence, the other appressed; it is true that there are

extremes of pubescence, but I cannot find any correlated characters

that would seem to support a specific segregation.

In an occasional specimen from Honduras and Guatemala, about

half the known collections from British Honduras, and in most speci-

mens from Mexico, the hairs of the young branchlets and the inflores-

cence are spreading and somewhat reddish. The branchlets often

appear shaggy or velvety. Professor F. Miranda informs me that

in Chiapas the conspicuous reddish unexpanded terminal buds make

the plant easy to recognize in the field. This is Myrcia rufidula

Schlecht., which I cannot distinguish in the herbarium except in the

most subjective way from less distinctively hairy specimens. Aside

from the abundant Mexican material (from Veracruz, Tabasco, Chi-

apas) and some specimens from British Honduras, I have seen only

two specimens from Honduras, one doubtful (sterile) specimen from

Guatemala (Alta Verapaz, Coban, Standley 69529], and a recent

collection from Pete"n (Machaquila, C. L. Lundell 16^09).

The recently described Myrcia belizensis Lundell, Wrightia 2: 213.

1961, was described by the collector as a woody vine. The author

also noted the exceptionally long petioles, these 5-9 mm. long, or

about twice as long as those of M. splendens. The type, which I

have seen through the courtesy of Dr. Lundell, is Gentle 6571, from

near San Antonio, Toledo District, British Honduras. The speci-

men is not to be distinguished from other Central American specimens

of M . splendens, except that the long petioles and the relatively large

panicles (up to 10 cm. long) suggest Myrcia fallax (Rich.) DC., a

widespread and common species of tropical South America. In

M. fallax, however, the inflorescence is usually relatively stout (the

peduncle 2-2.5 mm. wide just below the first node) , whereas in the

type of M . belizensis the thickest peduncle is no more than 1.5 mm.

wide. Typical M. fallax may yet be found in British Honduras, as

have so many other Myrtaceae of the Caribbean basin, but Gentle

6571 cannot be referred to it with any assurance. On the other

McVAUGH: TROPICAL AMERICAN MYRTACEAE 473

hand there seems to be small reason to regard M. belizensis as a dis-

tinct species, at least not until after more is known about the taxon-

omy of the complex to which M . splendens and M. fallax belong.

The "vining" habit I suspect is not that of a true liana, but the

habit assumed by so many woody plants in tropical forests, viz. the

development of long trailing branches that depend for support upon

the branches of other trees and shrubs. Similar developments are

known in other species of Myrcia.

5. MYRCIANTHES Berg

SYNOPSIS OF THE GENUS (EASTERN SOUTH AMERICA EXCLUDED)

The work of Berg in the mid-nineteenth century laid the founda-

tions for all subsequent work on the American Myrtaceae, provided

a rational scheme of classification for the many taxa that had been

loosely grouped by DeCandolle in the tribe Myrteae, and established

the existence of many genera and other more inclusive taxa that had

previously been unrecognized. One such group, Berg's second major

"Subtribe," the Eugenioideae, included 15 genera having in common

a fleshy large solid embryo and a short radicle ("Embryo cotylis

carnosis, discretis v. margine v. omnino conferruminatis; radicula

abbreviata;" cf. Linnaea 27: 4. 1855).

The largest genus, Eugenia, including 473 species recognized by

Berg, was characterized by the possession of 4 calyx-lobes, and the

embryo was described as "exalbuminosus, carnosus; cotylis saepissime

margine v. omnino conferruminatis; radicula abbreviata." Within

the genus Berg recognized 8 groups (Uniflorae, Biflorae, Racemosae,

etc.) based on features of the inflorescence. The seventh such group,

Dichotomae (species numbered 355-407), was described as having

"Pedunculi bifidi v. simpliciter aut ramosissime dichotomi, 6- -mul-

tiflori, flore in dichotomiis sessili, lateralibus saepissime pedicellatis."

Most of the Dichotomae seem to have been known to Berg in flower

only, but in at least two species (E. dicrana, Linnaea 27: 259. 1856,

and E. dichotoma, I.e. 262) he described the embryo as consisting of

2 discrete unequal cotyledons.

One of Berg's new genera, Myrcianthes (Linnaea 27: 315. 1856)

was recognized by him as coordinate with Eugenia, and distinguished

from that genus chiefly by the possession of 5 (instead of 4) calyx-

lobes. The embryo of Myrcianthes was fully described: "exalbumino-

sis, cotylis 2, discretis, carnosis, piano-con vexis; radicula brevi, sub-

exserta; gemmula saepe distincta, inter cotylas latens." Myrcian-

474 FIELDIANA: BOTANY, VOLUME 29

thes was also described as having "Pedunculi axillares, 1-3-flori v.

dichotomi." The four species included by Berg in this genus were

all natives of Uruguay or southern Brazil.

It now seems odd that Berg did not call attention to the obvious

similarities between his Dichotomae, a subgroup of Eugenia, and his

genus Myrcianthes. The inflorescence is the same in the two groups

— and quite unlike anything else in Eugenia; the cotyledons are free

and plano-convex in Myrcianthes, and also in the few species of Di-

chotomae in which Berg had seen fruit; the calyx-lobes are 5 in Myrci-

anthes (actually sometimes 4), and 4 (or sometimes 5, as pointed out

by Berg for several species) in the Dichotomae. Whatever may have

been the reason, Berg apparently did not consider the possibility

that the Dichotomae might well be removed from Eugenia into an-

other genus. Authors following Berg have at one time or another

transferred most of the species of Dichotomae to other genera, includ-

ing Anamomis Griseb., Luma A. Gray, Myrceugenella Kausel, Myrci-

anthes Berg, Pseudanamomis Kausel, and Pseudomyrcianthes Kausel.

Plants of this general affinity are frequent along an Antillean-

Andean axis, from Argentina and Bolivia to Venezuela and Colom-

bia, the West Indies, peninsular Florida, and in Central America

and Mexico north to Tamaulipas and Durango. Approximately 40

species are known from this range. There are isolated and distinct,

but clearly related groups of species in Chile, and in the South Bra-

zilian-Uruguayan-Argentinian region. Recent work indicates that

the Chilean species belong to distinct and mostly endemic genera.

I have already advocated (in the Flora of Peru, 13, pt. 4: 746. 1958)

that the concept of Myrcianthes be expanded to include numerous

Andean and West Indian species and to include Anamomis Griseb.

(1860).

There is no general agreement on the taxonomic status of the

species of eastern South America. Because of the superficial resem-

blance between the inflorescences and flowers of Myrcianthes and its

relatives, on the one hand, and those of certain genera of the sub-

tribe Pimentinae (e.g. Blepharocalyx and Pseudocaryophyllus) on

the other, it is difficult to place an unknown flowering specimen in

the proper genus. Final disposition (even to subtribe) may depend

upon the availability of mature fruit and seeds. Some species are

known from flowering material only. In recent years two Bergian

species of Eugenia have been transferred by Burret to Pseudocary-

ophyllus, and one Bergian species of Myrcianthes by the same author

to Psidium. Kausel (Ark. Bot. II. 3: 504. 1956) has proposed a new

McVAUGH: TROPICAL AMERICAN MYRTACEAE 475

genus, Pseudomyrcianthes, to include approximately 15 species origi-

nally placed in Eugenia by Cambessedes, DeCandolle, Berg and

others. On the basis of embryo characters, however, Kausel assigns

Pseudomyrcianthes to the Eugenioideae proper, proposing at the same

time a coordinate group Plinioideae1 to include Myrcianthes and its

close Andean and Antillean relatives (which he assigns to the genera

Amyrsia and Anamomis, respectively).

Legrand (Not. Syst. 15: 266-267. 1958) feels that too much sig-

nificance has been attached to the dichasium as a fundamental taxo-

nomic character. He points out that in Eugenia pyriformis (the

type species of Kausel's Pseudomyrcianthes), there is frequently a

reduction to a 1-flowered form resembling species in other subgeneric

groups of Eugenia. He feels that the occasional occurrence of a

3-flowered dichasium in a normally 1-flowered species (e.g. E. arbuti-

folia Berg, a synonym of E. obtusifolia Camb.) does not justify its

inclusion in Berg's group Dichotomae. He reminds us that Myrceu-

genia ribeireana (Berg) Legr. & Kaus. was originally placed in the

Dichotomae, in the absence of fruit, on the basis of its forked in-

florescence.

For want of adequate fruiting material of many of the species,

and in view of the lack of agreement on generic limits in the spe-

cies of eastern South America, I do not attempt to present even a

summary of them at this time. Two species of Myrcianthes, namely

M. cisplatensis (Camb.) Berg [including the genotype, M. apiculata

Berg], and M. gigantea (Legrand) Legrand, seem to be generally

accepted as members of the genus in good standing. Excluded from

Myrcianthes have been M. brunnea Berg [Psidium ovale (Spreng.)

Burret] and M. edulis Berg [considered by Legrand, An. Mus. Hist.

Nat. Montevideo II. 4, pt. 11: 59. 1936, to be related rather to the

genus Hexachlamys Berg (Eugenia subg. Myrcichlamys Legr.)].

Several species included by Berg among the Dichotomae (Linnaea

27: 246-278. 1856; Fl. Bras. 14, pt. 1: 306-312. 1857; I.e. 586-587.

1859) may be investigated as possible members of Myrcianthes; as

far as I know their generic standing has not been challenged : E. ex-

coriata Berg, E. imbricata Berg, E. ischnosceles Berg, E. itajurensis

Camb., E. subamplexicaulis DC., E. ternatifolia Camb. and E. vimi-

nalis Berg. Others (E. leandreana Berg, E. mutabilis Berg) have

been transferred to Pseudocaryophyllus by Burret, and several (E.

adamantium Camb., E. arbutifolia Berg, E. decumbens Camb., E.

1 Mentioned by name only; not properly described and not given any particu-

lar status.

476 FIELDIANA: BOTANY, VOLUME 29

kochiana DC., E. lutescens Camb., E. piresiana Camb., E. pohliana

DC., E. pruniformis Camb., E. pyriformis Camb.) have been assigned

at least provisionally to Pseudomyrcianthes by Kausel. Legrand has

pointed out (Not. Syst. 15: 269. 1958) that E. sylvatica Camb.

(Pseudomyrcianthes cambessedeana Kausel) was wrongly assigned by

Berg to the Dichotomae; the inflorescence is a loose raceme like that

of £". florida DC. and E. gardneriana Berg.

In the following pages is presented a synopsis of the species of

Myrcianthes occurring along the Andes and in northern South Amer-

ica, the West Indies and North America. The limits of the genus

as I understand them were set forth in the Flora of Peru (Field Mus.

Bot. 13, pt. 4: 746. 1958). As stated there, I believe the distinction

between Anamomis and other genera, on the basis of the presence or

absence of a plumule, to be a specious one; I have repeatedly observed

a plumule in the mature seeds of Anamomis (Myrcianthes) fragrans

and similarly in the seeds of Andean and other species of Myrcianthes.

As far as I know there is no species in eastern South America

that occurs also along the Andean-Antillean axis, except that a form

of M. cisplatensis is found in northern Argentina. The following

key summarizes the pertinent features of Myrcianthes and other

genera having the inflorescence a dichasium and the two cotyledons

distinct and plano-convex:

Radicle about as long as the cotyledons; cotyledons plano-convex, fleshy but thin;

Chile and adjacent Argentina Luma (incl. Myrceugenella).

Radicle less than half as long as the cotyledon; cotyledons plumply plano-convex.

Calyx-lobes caducous; inflorescence often irregularly dichotomous or umbelli-

form; ovules 5-6; cotyledons united about one-third their length; 1 species,

West Indies and Venezuela Pseudanamomis.

Calyx-lobes persistent; inflorescence regularly dichotomous with the terminal

flowers sessile in the forks; ovules usually more than 6; cotyledons united

at the base only.

Placenta shield-shaped with 6-12 marginal and inwardly directed ovules;

calyx-lobes 5; Chile Reichea.

Placenta wartlike or horseshoe-shaped, with usually 8-20 radially directed

ovules in a subcapitate group; calyx-lobes usually 4 Myrcianthes.

NOMENCLATURE

The name Luma A. Gray (1854) is apparently the oldest name

applied to any plant of this alliance. According to the recent studies

by Kausel on the Myrtaceae of Chile, the seven species referred by

Gray to Luma are to be distributed among three genera in two sub-

tribes (see Ark. Bot. II. 3: 503. 1956, and an earlier paper, Rev. Arg.

Agron. 9: 42. 1942). If Luma be typified by L. chequen (Mol.)

McVAUGH: TROPICAL AMERICAN MYRTACEAE 477

A. Gray, var. a, as suggested by Kausel (Lilloa 13: 127. 1948) and

accepted by McVaugh (Taxon 5: 142. 1956), then Myrceugenella

Kausel (1942) falls into the synonymy of Luma. Five of the remain-

ing six species of Luma (in the sense of Gray) are to be referred to

Myrceugenia Berg (published 1855), according to Kausel (I.e. 50.

1942). Because of this preponderance of species of "Myrceugenia"

in the original Luma, Burret in 1941 reduced Myrceugenia to the

synonymy of Luma and published a number of new combinations

and new species in the latter genus (Notizbl. Bot. Gart. Berl. 15:

522-535; Repert. Sp. Nov. 50: 50-55). Both Kausel (Lilloa 13: 125-

129. 1947) and Legrand (Darwiniana 11: 302. 1957) have argued

against the course taken by Burret.

There is no nomenclatural conflict between Luma A. Gray (1854)

and Myrcianthes Berg (1856) as long as the two genera are held to

be taxonomically distinct. Apparently a good case can be made for

their distinctness.

The species of Luma (Myrceugenella in the sense of Kausel) are

all confined to Chile, or the Chilean-Argentinian region, i.e. they are

geographically separated from all known species of Myrcianthes.

They are like many species of Myrcianthes in having the flowers

4-merous, the bracteoles caducous and the dichasium 3-flowered or

the flowers occasionally solitary. In Myrceugenella, however, the

cotyledons are described by Kausel as "piano-con vexis, carnosis sed

tenuibus," the radicle is about as long as the cotyledons (illustrated

in Rev. Arg. Agron. 9: 44, fig. 1, B. 1942), and the insertion of the

ovules is said to be basal or sub-basal (Lilloa 13: 148. 1947). On

the basis of the long radicle and the relatively thin cotyledons,

Kausel now regards Myrceugenella as a member of the Myrciinae

("Myrcioideae") (Ark. Bot. II. 3: 503. 1956). I cannot comment

upon this disposition of the genus, but in any event Myrceugenella,

or Luma, appears to be a small endemic Chilean group that for the

present may be regarded as a distinct genus.

The name Amyrsia Raf. (1838) has recently been taken up by

Kausel (Ark. Bot. II. 3: 512-514. 1956) for several Andean species

that I should refer to Myrcianthes. Rafinesque's genus Amyrsia

was a mixture (cf. Taxon 5: 137. 1956), including three species of

Myrcianthes in the present sense, and one species of Myrteola (sub-

tribe Pimentinae) . Rafinesque gave no indication of any preference

for one species or another as genotype, and my choice of lectotype

was made on the basis of the original description, which permitted

the positive identification of the Myrteola element because Rafinesque

478

FIELDIANA: BOTANY, VOLUME 29

FIG. 12. Dichasium and embryo in Myrcianthes. A, Seed of M. karsteniana,

Garcia 94 (ca. X 17); inner face of one cotyledon showing position of hypocotyl and

radicle, and small, apparently bilobed plumule at right. B, The same (ca. X 7);

outer coats removed to show plano-convex cotyledons and projecting radicle.

C, Seven-flowered dichasium in M. fragrans, Steere 2976 (X %).

compared the genus to "Pimentus," from which he separated it be-

cause of the bilocular ovary, and the many seeds "as in Myrtus."

None of the species now referred to Myrcianthes could fit this de-

scription, as ordinarily but one seed develops in each fruit. If Amyr-

sia be typified on the basis of the Myrteola element, the genus

Myrteola Berg (1856) may well be proposed for conservation, as

was done in Taxon 5: 163-164. 1956. It would seem most unfortu-

nate to preserve the poorly founded genus Amyrsia in any sense

whatever, especially since both of its constituent elements have

since become known under other names. If Kausel's proposal be

McVAUGH: TROPICAL AMERICAN MYRTACEAE 479

accepted, and Amyrsia typified on the basis of the Myrcianthes ele-

ment, then the latter name should be proposed for conservation.

Myrcianthes Berg, Linnaea 27: 315. 1856. Anamomis Griseb.

Fl. Brit. W. Ind. 240. 1860.

Small or medium-sized trees with opposite or ternate leaves; flowers in 1-7-

flowered (or sometimes 15-31-flowered) dichasia, these usually solitary and

axillary, sometimes so numerous in the upper axils as to form a compound in-

florescence, or modified into a terminal myrcioid panicle bearing the flowers

in small dichasia at the tips; central flowers of the dichasia usually sessile; calyx-

lobes and petals 4 (in some species regularly 5), or an occasional flower 5-merous;

ovary usually 2-locular, the numerous (usually 8-20) ovules radiating from a

centrally affixed placenta; seeds maturing 1 (-4); cotyledons 2, distinct, plano-

convex; radicle terete, half as long as the cotyledons or less; plumule evident

in the mature seed.

Lectotype species: M. apiculata Berg [=M. cisplatensis (Camb.)

Berg]. See Taxon 5: 143. 30 July 1956.

In the following pages are contrasted and listed all the species

known to me from continental North America, from South America

north of the Amazon River, and from Andean South America except

for the approximately 20 species treated in the Flora of Peru (Field

Mus. Bot. 13, pt. 4, no. 2: 745-775. 1958). These Peruvian species

are omitted, or in a few instances mentioned for comparison. Spe-

cies of southeastern South America are not included; West Indian

species are treated somewhat cursorily, following Urban (Bot. Jahrb.

19: 662-665. 1895).

Certain species listed below as members of the genus Eugenia

are probably to be referred eventually to Myrcianthes, but as these

are unknown in the fruiting condition any formal transfers would

seem to be premature.

Inflorescence, and often the young herbage, softly tomentose or velutinous with

rufous, brown or pale hairs; petioles stout, distinct from the blades, (4-) 10-

15 (-20) mm. long.

Calyx-lobes tomentulose on both sides.

Disk 3-4 mm. wide; buds 4-5 mm. long; flowers up to 15, the dichasium often

3 times forked; ovules 5; Bolivia M. pearcei.

Disk 5-8 mm. wide; buds 7-10 mm. long; flowers mostly 3-7; ovules 15-30.

Tomentum gray or yellowish white; disk 5 mm. wide; stamens 7-8 mm.

long; northern Peru M. lanosa.

Tomentum red; disk 6.5-8 mm. wide; stamens 10-14 mm. long.

Calyx-lobes 5, 5-9 mm. wide; Argentina M. callicoma.

Calyx-lobes 4, 4-5 mm. wide; Colombia M. borealis.

480 FIELDIANA: BOTANY, VOLUME 29

Calyx-lobes glabrous within; inflorescence irregularly 2-3 times forked; buds

5-6 mm. long; disk 4-5 mm. wide; ovules 10-11, marginal on the placenta;

Ecuador M. irregularis.

Inflorescence and herbage glabrous or often strigose or silky-strigose, the hairs often

white or very pale, most abundant and conspicuous on the outer surface of

the hypanthium; petioles often not well set off from the cuneate base of the

blade.

Leaves all or nearly all ternate; northern Andes of Venezuela and Colombia.

Coarse plants; leaves 2-3 (-5) cm. long; buds 8-9 mm. long M. crebrifolia.

Smaller-leaved plants; leaves 0.6-1.8 cm. long; buds 4-5 mm. long.

Peduncles 3-flowered; leaves spatulate M. ternifolia.

Peduncles 1-flowered; leaves obovate to suborbicular E. triquetra.

Leaves opposite [for plants of Ecuador, Peru and Bolivia, see Field Mus. Bot. 13,

pt. 4: 745-775. 1958].

Plants of the Guianas, Venezuela, Colombia, West Indies or North America.

Inflorescence decompound, many-flowered, the forking branches forming a

terminal leafless panicle.

Leaves 2-4 cm. long, rigidly coriaceous; panicles 5 cm. long or less beyond

the last leaves; southern Colombia and Ecuador E. pycnantha.

Leaves 6-12 cm. long, thinner; panicles 6-11 cm. long; Venezuela.

M. karsteniana.

Inflorescence a regular axillary dichasium, once-, twice- or thrice-forked,

the flowers thus mostly 3-7, occasionally only 1 or as many as 15.

Inflorescence stout, glabrous or essentially so, the peduncle 2-3.5 (-5)

mm. wide near the summit.

Flowers very large, the buds 1.7-2 cm. long; style 2.5 cm. long; fruit

4-5 cm. in diameter; Guianas M. prodigiosa.

Flowers smaller, the buds less than 1 cm. long; style about 1 cm. long

or less; fruit as far as known 2 cm. in diameter or usually less;

Andes from Peru northward; Panama and Costa Rica.

Leaves rounded, 3 cm. long or less, the lateral veins impressed above;

staminal disk glabrous; high mountains of ?Costa Rica and west-

ern Panama (Allen 1563) ?E. rigidissima.

Leaves elliptic to suborbicular, mostly 4-6 cm. long, the veins not

impressed; disk bristly-pubescent; style 10-11 mm. long; Andes,

Colombia to Peru M. rhopaloides.

Inflorescence more slender, variously pubescent (especially the hypan-

thium), or glabrous, the peduncle 2 mm. wide at summit or usually

less; flowers smaller, the style 8.5 mm. long or less.

Peduncles minutely bristly-hispidulous with erect hairs.

Peduncles and branchlets similarly hispidulous; leaves often elliptic,

sometimes obovate; dichasium often 7-flowered; Costa Rica and

western Panama M.fragrans var. hispidula.

Peduncles hispidulous, the branchlets glabrous; leaves mostly obo-

vate; dichasium usually 3-flowered; Venezuela. . .M. compressa.

Peduncles glabrous, appressed-pubescent or appressed-silky.

Flowers 3 in a dichasium, all sessile; hypanthium and outer surface

of the calyx-lobes whitened, silky; Santa Marta region, Colom-

bia. . . .M. sessilis.

McVAUGH: TROPICAL AMERICAN MYRTACEAE 481

Flowers 3-7 or -15, the lateral ones on definite slender branches;

calyx-lobes glabrous or essentially so without.

Hypanthium rather densely silky-strigose at least at base, often

whitened by the dense covering of hairs; inflorescence in gen-

eral somewhat strigose or hispidulous, sometimes glabrate.

Flowers usually 3; peduncle mostly 2 cm. long or less, 1.5-2.5

mm. wide at summit; style 7-8.5 mm. long; southern

Colombia and Ecuador M. hallii.

Flowers often 7, sometimes 3; peduncle 2-4 (-6) cm. long, 1.2-

1.5 mm. wide at apex; style mostly 5-6.5 mm. long; often

in lowlands, West Indies and northern Colombia and Vene-

zuela to Mexico and Florida vars. of M. fragrans.

Hypanthium glabrous, or sparingly appressed-pubescent especially

at base; peduncles glabrous; highlands, Costa Rica, Colombia

and Venezuela; Ecuador.

Peduncles 1-flowered; southern Colombia and Ecuador.

E. orthostemon.

Peduncles mostly 3-7- (-15) flowered.

Peduncles mostly 7-flowered, 3-4 cm. long; Colombia to west-

ern and northern Venezuela; see also E. rondonensis.

M. dugandii.

Peduncles mostly 3-flowered, 2.5 cm. long or less.

Costa Rica M. storkii.

Colombia (region of Bogota) M. leucoxyla.

Plants of Argentina (including southern Bolivia).

Flowers solitary; hypanthium whitened without by the dense covering of

hairs; calyx-lobes 4; style 5-8 mm. long M. mato.

Flowers 3 (occasionally 1 or 7) in a dichasium; hypanthium glabrous or

sparingly stiff-hairy without.

Calyx-lobes 5; style 3.5-5.5 mm. long M. cisplatensis.

Calyx-lobes 4; style 7-8 mm. long M. pseudo-mato.

Eugenia alaternifolia [sphalm. alternifolia] Benth. PI. Hartw.

175. 1845. Colombia. Cf. M. rhopaloides (see Flora of Peru, 13,

pt. 4: 773).

Eugenia andina Berg, Linnaea 27: 274. 1856. Peru. Unknown

(see Flora of Peru, I.e. 774).

Eugenia aromatica Berg, Linnaea 27: 263. 1856. Venezuela,

Caracas, Moritz 792 (Berlin). Unknown.

Myrcianthes borealis McVaugh, sp. nov.

Arbor rufo-tomentosa, ramulis inflorescentiisque, et foliis juvenilibus, dense

pilis contortis usque ad 0.5-1.3 mm. longis obtectis; folia elliptica vel obovata,

5-9 cm. longa, 2.5-7 cm. lata, petiolis 9-17 mm. longis, marginibus callosis,

482 FIELDIANA: BOTANY, VOLUME 29

aliquantum revolutis; flores 3 (-7), pedunculo 3-5 cm. longo, 2.5-3 mm. crasso;

alabastra ut videtur 10 mm. longa; calycis lobi 4, utraque superficie tomentulosi,

4-5 mm. lati; discus 6.5-7.5 mm. latus; stylus 7 mm. longus, pilosus; stamina

ca. 300, filamentis 10-12 mm. longis; ovula quoque loculo ca. 30.

Tree 2.5-3.5 m. high, the young leaves, branchlets and inflorescence rusty-

tomentulose, the crisped, curled hairs often more than 0.5 mm. long, some to 1.3

mm.; leaves elliptic or obovate, rounded or retuse, rounded to the base or some-

what acutely narrowed and the margins finally cuneately decurrent on the stout

channeled petiole 2-2.5 mm. thick, 9-17 mm. long; blades 2.5-7 cm. wide, 5-

9 cm. long, 1.3-2 times as long as wide, lustrous and glabrate above except the

hairy sulcate midvein, paler and somewhat pilose beneath; lateral veins 7-9

on each side, somewhat impressed above and prominent beneath, diminishing

distally to a slightly less prominent marginal vein 2-7 mm. from the margin, and

arched between the laterals; glands inconspicuous; margins callose, somewhat

revolute; flowers usually 3 (-7), the peduncle 3-5 cm. long, 2.5-3 mm. wide

near the apex, the branches 6-16 mm. long; central flower sessile, or the axis

extended 8-11 mm. above the fork; bracts narrowly triangular, acute, 1.8 mm.

wide at base, 6 mm. long; bracteoles linear, 2 mm. long, deciduous, with the

bracts, at anthesis; buds probably about 10 mm. long; hypanthium cup-shaped,

5 mm. long, 6 mm. wide at mouth; calyx-lobes 4, broadly rounded, tomentulose

on both sides, 2-3 mm. long, 4-5 mm. wide; disk 6.5-7.5 mm. wide, the staminal

ring velutinous, the glabrous center 3 mm. wide; style 7 mm. long, pilose; stamens

about 300, the filaments 10-12 mm. long; locules 2, the ovules about 30 in each

locule, radiate in a subcapitate group affixed to the central dissepiment. — Univ.

of Mich. Negs. 1331, 1347.

COLOMBIA: Norte de Santander: Eastern Cordillera, road from

Pamplona to Toledo, on the Maracaibo-Orinoco drainage, alt. 2500-

2800 m., thickets along stream, 28 Feb. 1927, Kittip & Smith 19882

(US, type; A; GH).

Myrcianthes callicoma McVaugh, sp. nov.

Arbor 6-8-metralis, rufo-tomentosa; folia elliptica vel ovata, 5-10 cm. longa,

1.5-5 cm. lata, obtusa vel obscure acuminata, glabrata, petiolis 6-10 mm. longis;

venis utroque latere 10-12, supra inconspicuis, subtus paullo elevatis, venulis

nee incrassatis; dichasio 3-floro (vel floribus solitariis), pedunculo 1-2.5 cm.

longo, 2-2.5 mm. crasso, floribus in dichotomis sessilibus, alabastris 7-9 mm.

longis; calycis lobis 5, utrinque tomentosis, 5-9 mm. latis; stylo 8 mm. longo;

disco 8 mm. lato; stamina ca. 300, filamentis 10-14 mm. longis; ovula quoque

loculo ca. 15-20.

A tree 6-8 m. high, 35 cm. in diameter, the young growth and the inflor-

escence densely tomentose with soft crisped rusty-red hairs, these very short

or on the margins of the calyx up to 1 mm. long; leaves and branchlets glabrate

or with some minute crisped hairs persisting; leaves elliptic or ovate, (1.5-)

2.5-5 cm. wide, (3-) 5-10 cm. long, 1.6-2.5 times as long as wide, bluntly pointed

or obscurely acuminate, rounded to the base, the markedly cartilaginous and

revolute margins narrowly prolonged and long-decurrent on the compressed

sulcate petiole 1.5-2 mm. wide, 6-10 mm. long; midvein impressed above, prom-

McVAUGH: TROPICAL AMERICAN MYRTACEAE 483

inent beneath; lateral veins 10-12 pairs with some additional weaker intermediate

veins, inconspicuous above, somewhat raised beneath; marginal vein weaker

than the laterals, irregularly and asymmetrically arched between them, 1-5 mm.

from the margin; blades smooth on both surfaces, concolorous, slightly more

glossy above, obscurely and finely gland-dotted; inflorescence an axillary dicha-

sium, 3-flowered (or often 1-flowered), the peduncle 1-2.5 cm. long, compressed

and enlarged distally and there 2-2.5 mm. wide; flowers sessile, or the lateral

ones in the dichasium on pedicels up to 8 mm. long; bracts and bracteoles deciduous

before anthesis; buds 7-9 mm. long, broadly obovoid or subglobose; hypanthium

broadly obconic with straight sides 4-5 mm. long; calyx lobes 5, unequal (1 or

2 smaller than the others), tomentose on both sides, broadly rounded and often

narrowed just above the base, 3-6 mm. long, 5-9 mm. wide; disk 8 mm. wide,

the staminal ring hairy, the glabrous or pubescent central portion 4 mm. wide;

style 8 mm. long; stamens about 300, 10-14 mm. long, the anthers 0.7-0.8 mm.

long; petals obovate, about 7 mm. wide, 12 mm. long, pubescent near the tips;

ovary bilocular, the ovules about 17 (15-20?) in each locule, radiate in a sub-

capitate group attached above the middle of the central partition; fruit not seen.

—Univ. of Mich. Neg. 1345.

ARGENTINA: Tucuman: Dept. Tafi, Cerro de Taficillo, 1600 m.

elevation, in quebradas, Dec. 7, 1929, S. Venturi 9859 (GH, type);

same locality, VenturilOOSS, Jan. 1930 (GH).

Myrtus coccolobaefolia HBK. Nov. Gen. & Sp. 6: 139 [folio

ed. 110]. 1823. Colombia. Referred in Flora of Peru, pt. 4: 772 to

Myrcianthes rhopaloides.

Myrcianthes compressa (HBK.) McVaugh, Field Mus. Bot.

13, pt. 4: 754. 1958. Myrtus compressa HBK. Nov. Gen. & Sp. 6: 135

(folio ed. p. 107). 1823.

Often nearly glabrous, the young shoots finely pale-strigose, the peduncles

minutely bristly pubescent (sometimes glabrate), the petals and calyx-lobes ciliate,

the hypanthium sometimes sparsely strigose; leaves obovate, up to 4 cm. wide and

6 cm. long, obtuse or rounded at the tips (on flowering branchlets often acute at

both ends and more or less elliptic), the base cuneate and the margins decurrent

on the channeled petiole 3-4 mm. long; blades lustrous above, smooth, finely

gland-dotted especially beneath; inflorescence 3-flowered (rarely 7-flowered), the

peduncle 2-3.5 cm. long, 1-1.7 mm. wide near summit, often compressed and

2-edged; style 6-7 mm. long; stamens about 175; ovules 15-20 (-25) in each locule.

The plant described above bears considerable resemblance to the

inclusive Myrcianthes fragrans, differing chiefly in the usually 3-flow-

ered dichasium, the nearly glabrous inflorescence (except that the

peduncle is usually minutely bristly-pubescent), the cuneate obovate

leaves. With some doubt I refer this to M. compressa, originally

described from near Cajamarca, Peru. Nothing like the type has

since been found in Peru, but a number of collections from the moun-

484 FIELDIANA: BOTANY, VOLUME 29

tains near Caracas, Venezuela, appear to represent the same species

and are cited below. A photograph of the type (Field Mus. Neg.

36879) shows the characteristic leaf-outline, and my notes on the

type of Myrtus compressa, made in Paris in 1954, indicated that this

specimen has the same minutely bristly peduncle, a feature unusual

in Myrcianthes. I have never seen any plants from the Ecuadorean-

Peruvian region that resembled Venezuelan plants in this respect.

It is entirely possible that the type of Myrtus compressa came from

somewhere in Venezuela and was later mistakenly supposed by

Kunth to have come from Peru.

VENEZUELA: Margen de la selva de las Flores, 1619 m., L. Wil-

liams 10095 (A, F); between Agua Negra and El Junquito, 1900 m.,

Pittier 13822 (F, US); Galipan bei Caracas, 6000 ft., 0. Kuntze 1575

(US); Caracas, H. M. Curran & M. Raman 1124 (NY, US), Karsten

s.n. (F, GH).

COUNTRY UNKNOWN: Paramo de Cisne, Andre 4307, 29 Oct. 1876

(F, GH).

Myrcianthes crebrifolia (Steyerm.) McVaugh, comb. nov. Eu-

genia crebrifolia Steyerm. Fieldiana, Bot. 28: 1011. 1957.

As suggested by Steyermark, and as indicated in the key above,

this species is to be compared with Eugenia ternifolia Berg. It is

also strikingly similar in most respects to M. rhopaloides, from which

it seems to differ chiefly in the ternate arrangement of the leaves

and in the very short fleshy petioles 2-4 mm. long and about as thick.

In addition to the two collections cited by Steyermark, I should

refer to this species the following:

COLOMBIA: Santander: Vicinity of La Baja, 3200 m., Killip &

Smith 18154 (US). — Norte de Santander: Between Mutiscua and

Pamplona, 3100 m., Killip & Smith 19743 (GH).

Myrcianthes dugandii (Standl.) McVaugh, comb. nov. Eu-

genia dugandii Standl. Trop. Woods 52: 28. 1937.

Nearly glabrous, the young herbage somewhat appressed-pubescent, the

hypanthium glabrous or minutely appressed-pubescent, the calyx-lobes densely

appressed-silky within, the disk hairy; leaves rigidly coriaceous, smooth and

glossy above, 2-4.5 cm. wide, 3.5-7.5 cm. long, 1.3-2 times as long as wide, obo-

vate or elliptic, rarely ovate, rounded or retuse at tip or obtusely pointed, the

base abruptly or gradually rounded, the margins finally often cuneately decurrent

on the stout petiole 1-2 mm. thick, 3-8 mm. long; inflorescence mostly 7-flowered,

varying to 3-flowered or 15-flowered on the same plant; peduncles (1.5-) 3-4 (-5)

McVAUGH: TROPICAL AMERICAN MYRTACEAE 485

cm. long, 1-1.7 (-2) mm. wide near summit, the lateral branches up to 1 cm. long

(or nearly 2 cm. long in 15-flowered dichasia); calyx-lobes 4; disk about 3 mm.

wide; style 6.5 mm. long; locules 2; ovules (12-) 15-20 (-25); fruit orange; cotyle-

dons 2, plano-convex; plumule present. — Univ. of Mich. Neg. 1334.

Few specimens that can definitely be referred to Myrcianthes fra-

grans have been collected in South America. Many specimens from

the Andes of Colombia and Venezuela, however, are similar to M.

fragrans except that they are glabrous or essentially so throughout,

and the dichasia tend to be 7- or 15-flowered, rather than 3-7-flow-

ered as in most populations of M. fragrans. This glabrous and rather

abundantly flowered plant is Eugenia dugandii Standl. It is known

mostly at middle elevations, from 1500 to 2200 meters.

COLOMBIA: Antioquia: Medellin, A. Dugand G. 904 (F, type);

Bello, W. A. Archer 233 (US); Copacabana, Bro. Daniel 269 (US).—

Tolima: Rio Paez, Lehmann 5818 (GH, US).— Huila: Cordillera

Oriental, 25 km. southeast of La Bodega, E. L. Little 9056 (MICH).

VENEZUELA: Me"rida: Aristeguieta 3332 (MICH), Bernardi 283

(NY), John 708 (US), 1049 (GH, US), Steyermark 55955 (F), Ta-

mayo 2431 (US). — Sucre: Headwaters of Rio de Amana, Steyermark

62734 (F).— Monagas: Cerro Guacharo, Steyermark 62338 (F). In

the specimens from Sucre and Monagas, the leaves are thinner and

more ovate than usual, in these features suggesting West Indian

specimens of M. fragrans.

Myrcianthes fragrans (Sw.) McVaugh, comb, nov., var. fra-

grans. Myrtus fragrans Sw. Prodr. 79. 1788. Eugenia punctata Vahl,

Symb. 3: 65. 1794 (type from Santa Cruz, von Rohr, West). E. fra-

grans (Sw.) Willd. Sp. PI. 2: 964. 1800. Anamomis fragrans (Sw.)

Griseb. Fl. Brit. W. Ind. 240. 1860. E. triflora Sesse* & Moc. Nat-

uraleza II. 1: app. 83. 1888 (type from Cuernavaca, Morelos). M.

biflora, sensu Sess<§ & Moc. Fl. Mex. ed. 2. 125. 1894 (from Cordoba,

Veracruz). Myrcia seleriana Donn.-Sm. Bot. Gaz. 27: 332. 1899

(type from Chacula, Guatemala, Seler 3169). E. steyermarkii Standl.

Field Mus. Bot. 22: 360. 1940 (type from Guatemala near Volcan

de Tacana, Steyermark 36210). E. lopeziana A. Molina, Ceiba 3:

170. 1953 (type from Honduras, Molina 3007).

Leaves nearly glabrous, elliptic to ovate or obovate, 2.5-6.5 cm. long, usually

about twice as long as wide, short-petioled; inflorescence more or less appressed-

pubescent, the hypanthium often densely white-hairy and contrasting with the

green glabrous calyx-lobes; dichasia 3- or often 7-flowered, the peduncles 2-4 (-6)

486 FIELDIANA: BOTANY, VOLUME 29

cm. long, (0.7-) 1.2-1.5 mm. wide at apex; flowers small (buds 3-4.5 mm. long,

styles 6.5 mm. long or less, disk 3-3.5 mm. wide, calyx-lobes 2.5-3.5 mm. wide

at base).

What appears to comprise a single species, Myrcianthes fragrans,

is centered geographically in the Caribbean region. The nomen-

clatural type was from Jamaica. Members of the complex occur

naturally on all the Greater Antilles, and to a lesser extent on other

islands farther east; in the Bahamas and Florida; rather abundantly

in eastern Mexico from Tamaulipas to Veracruz and in the Yucatan

peninsula, in Central America and in northern Venezuela. A few

collections from western and southwestern Mexico (state of Mexico

to Durango) apparently belong to the same species-complex but are

hard to classify because of the small number of specimens available.

Additional small populations in eastern Mexico (San Luis Potosi

and Tamaulipas), in Florida (Eugenia dicrana Berg, Anamomis

simpsonii Small), in the Bahamas (Anamomis lucayana Britton),

and larger populations in the Greater Antilles, are evidently inter-

related in a way not yet understood. It would seem premature to

assign varietal or subspecific names to most of these populations

until their taxonomy can be more adequately worked out. Urban

said of this species many years ago, "Typus polymorphus in futuro

in varietates dividendus nee vix in species distinctas disjungendus."

In accordance with the letter of the nomenclatural Code, a pedant

may argue that Myrtus fragrans Sw. must be rejected because it was

superfluous when published and therefore illegitimate. It is true

that in the Prodromus Swartz cited in the synonymy of M. fragrans,

without any question or qualification, "Eugenia montana Aubl. guian.

495. t. 195." In a later publication, however (Fl. Ind. Occ. 2: 914.

1800), Swartz included Eugenia montana Aubl. in the synonymy of

M. fragrans with a query, and with the statement on p. 916: "Eu-

genia montana Aubl. (I.e.) meae [i.e. to Myrtus fragrans] simillima

est; quam vero color fructus hujus mini sit ignotus, nihil pro certo

affirmare possum." I think it may be reasonably argued that Swartz

never was really sure of the identity of Eugenia montana Aubl., and

that even in the Prodromus his intention was merely to compare

with his own new species an Aublet species that he thought might

be the same. Limitations of space in the Prodromus presumably

precluded the full explanation given later in the Flora. I do not see

any indication that Swartz intended to replace Aublet's name by a

new one; his early judgment, and also his more matured opinion,

was that the Jamaican Myrtus fragrans was an independent species.

I do not think that a well-established name should be upset on a

McVAUGH: TROPICAL AMERICAN MYRTACEAE 487

legalistic interpretation of a technicality, perhaps merely because

Swartz, in ignorance of the International Code of Nomenclature,

neglected in 1788 to insert the question-mark that he included in 1800.

Among the insular populations in the West Indies, it seems there

are two principal tendencies: One toward a 3-flowered dichasium

and smooth, elliptic, broad and obtusely pointed leaves with short

petioles (e.g. in Jamaica) , and another toward a 7-flowered dichasium

and glandular-verrucose, oblong-obovate, obtuse or rounded or re-

tuse leaves with longer petioles (e.g. Eugenia granulata Berg, in

Cuba and Hispaniola). Many exceptions to these generalizations

may be observed, and it is evident that a revision of West Indian

Myrcianthes is still to be desired. Most specimens of the fragrans

group from Central America or Mexico, however, can be referred

to one or the other of the above types.

A majority of the specimens from the continent of North America,

and a few from northern South America, bear a strong resemblance

to the plant of Jamaica (i.e., M. fragrans var. fragrans) except the

petioles are longer (often 4-6 mm. as against 2-4 mm. in the Jamaican

plant) and the dichasia are often 7-flowered (usually 3-flowered in

the Jamaican plant); the leaf -blades in Central American plants

may be narrower than those of Jamaica, where the blades are often

1.3-2 times as long as wide.

The following specimens are of the general type of M. fragrans

var. fragrans:

MEXICO: Veracruz: Rancho La Palmilla, Purpus 16350 (A, F);

Zacuapan, Purpus 7518 (A); Mirador, Liebmann 3971 (US). — Chi-

apas: Mt. Pasitar [Paxtal], Matuda 1*68 (MICH, US); Pinabeto,

Motozintla, Matuda 154-81 (F, an exceptionally lanose form).

BRITISH HONDURAS: Guatemala boundary, Camp 33, Schipp 1240

(A, F, GH, MICH).

GUATEMALA: Huehuetenango: Maxbal, Steyermark 48904. (A, F);

Chacula, Seler 3169 (a narrow-leaved form, type of Myrcia seleriana;

US). — San Marcos: Volcan Tacana, Steyermark 36210 (F, type of

E. steyermarkii) .

HONDURAS: Comayagua, Edwards P592 (A, F, US). — Morazan:

Rio Guarabuqui, Molina 3007 (F; US, type of E. lopeziana; Univ.

of Mich. Negs. 1326, 1327).

NICARAGUA: Near Granada, summit Mt. Mombacho, Grant 835

(A, F).

488 FIELDIANA: BOTANY, VOLUME 29

COSTA RICA: Alajuela: Zarcero, Austin Smith 608, 2687, 4,173,

4185 (all F); Zapote, Smith NY 57 9 (A, F); Llano Bonito, Smith

P2391 (UC, A). Copey, Tonduz 11741 (GH). Piedra Blanca,

Iscasu, F. Solis R. 480 (F).

COLOMBIA: Sta. Marta, near Escalera de los Indies, H. H. Smith

434. (GH); near Masinga, Smith 402 (A, F, GH, MICH, PH, US).—

Norte de Santander: Pamplona-Toledo, on divide, Killip & Smith

19917 (A, GH, US).

VENEZUELA: Paraguana Peninsula, Cerro Sta. Ana, Curran &

Haman 703 (GH, NY, US).

Plants resembling Eugenia granulata Berg (i.e. like many speci-

mens from Cuba and Hispaniola) are not commonly found in con-

tinental North America. A few collections, however, are note-

worthy, for their resemblance to the many-flowered West Indian

populations with oblong-obovate glandular-verrucose leaves:

MEXICO: Quintana Roo: Cozumel Island, Steere 2976 (MICH).—

Tres Marias: Maria Madre, Nelson 4306 (F, US), Maltbyl47 (F, US).

In Costa Rica and Panama there has been some confusion be-

tween what I take to be M. fragrans in the broad sense, and an

endemic species, Myrcianthes (Eugenia) storkii. After the publica-

tion in 1930 of Eugenia storkii, Standley routinely referred to that

species all specimens of "cymosely flowered" eugenias from Costa

Rica and Panama. Miss Amshoff, in her treatment of the Myrta-

ceae of Panama (Ann. Missouri Bot. Gard. 45: 182. 1958), referred

all the Panamanian material to E. fragrans, and relegated E. storkii

to synonymy without comment. Some comment seems necessary.

Most specimens from both Costa Rica and Panama, at least from

elevations up to 1650-1700 meters, I should refer to Myrcianthes

fragrans in the broad sense. Two principal races, however, are dis-

tinguishable. One of these is like typical fragrans of Jamaica, and

specimens of it are cited above under M. fragrans var. fragrans;

the pubescence in this race is usually appressed, and the hypan-

thium rather strongly and noticeably white-hairy. In the second

race the pubescence on the branchlets and inflorescence is bristly

rather than appressed, but the hypanthium is only sparingly stri-

gose. This is var. hispidula, described below. At higher elevations

(1900-2850 meters) in Costa Rica, Myrcianthes fragrans is appar-

ently replaced by another species, with somewhat larger flowers,

stouter glabrous inflorescence, and broader and more obtuse leaves;

this is Eugenia storkii Standl., q.v. below.

McVAUGH: TROPICAL AMERICAN MYRTACEAE 489

Myrcianthes fragrans, var. hispidula McVaugh, var. nov.

Ramuli inflorescentiaeque hispiduli; hypanthium basi sparsim vel vix sericeo-

strigosum; folia elliptica vel obovata, 2.5-6 cm. longa, (1.6-) 2-3-plo longiora

quam latiora, apice obtusa vel rotundata, basi acuta vel cuneata; pedunculus

2-3 (-4) cm. longus, ad apicem 1.2-1.5 (-1.7) mm. crassus, flores 3-7 gerens;

alabastra 3-4.5 mm. longa; discus diametro 2.5-3 mm., hirsutus; ovula ovarii

quoque locula 16-20. — Univ. of Mich. Neg. 1351.

Costa Rica and Panama, in woods, savannahs and pastures, at

middle elevations (1000-1700 meters), flowering March-June, fruit-

ing July-August.

COSTA RICA: San Ramon, vicinity of La Palma, A. M. Brenes

5497 (F), 5503 (F), 5659 (F), 15142 (A; F, type), 16194 (F); Cerro

Jucosal, H. E. Stork 1228 (F, paratype of Eugenia storkii); Las Con-

cavas, C. H. Lankester K99 (F, paratype of E. storkii); Navarrito-

Cartago, R. Torres R., 18 Oct. 1926 (US); Prov. Cartago, Dulce

Nombre, Standley 35929 (US).

PANAMA: Chiriqui: Between El Boquete and Caldera, H. Pittier

3326 (US) ; Boquete, M. E. Davidson 460 (A, F, US), 774 (A, F, US),

821 (F), 1062 (F); valley of Rio Chiriqui Viejo, G. White 108 (F,

GH); valley of Rio Chiriqui Viejo near El Volcan, P. White 211

(F, GH), 217 (F, GH).

Myrcianthes irregularis McVaugh, sp. nov.

Arbor velutina vel sparse hirsuta, pilis fulvo-flavidis usque ad 0.8 mm. longis

instructa; folia rigide coriacea, suborbicularia, ovalia vel obovata, 2-5.5 (-11) cm.

longa, 1.5-5 (-9) cm. lata, apice rotundata retusave, basi rotundata truncata vel

subcordata, petiolis hirsutis 4-8 mm. longis, venis lateralibus utroque latere 5-6,

subtus prominiusculis, inter se arcuatis sed venam marginalem vix formantibus;

inflorescentia irregulariter 2-3-dichotoma, pedunculo 2.5-3.5 cm. longo, 2-2.5 mm.

lato; discus 4-5 mm. latus, annulo staminali molliter pubescente; stamina ca. 150;

ovarium biloculare, loculis intus hirsutis, ovulis quoque loculo 10-11, e marginibus

placentae erectae hippocrepiformis radiantibus.

A tree, densely velutinous or sparsely hirsute with yellowish-brown bristly hairs

up to 0.8 mm. long; leaves rigidly coriaceous, glabrate above (at first bristly along

the broadly sulcate midvein), suborbicular to broadly oval or obovate, 1.5-5 (-9)

cm. wide, 2-5.5 (-11) cm. long, rounded or retuse, rounded truncate or subcordate

at base, the margins abruptly narrowed to stout short hirsute petioles 1-2.5 mm.

thick, 4-8 mm. long; lateral veins on each side 5-6 in addition to some intermediate

ones, inconspicuous above in mature leaves, prominent and somewhat elevated

beneath, diminishing distally, not forming a strong marginal vein but each over-

arching 2-6 mm. from the margin and joining the adjacent lateral; glandular dots

inconspicuous; immature leaves conspicuously reticulate-veined in drying; inflores-

cence irregularly forked 2 or 3 times; peduncle 2.5-3.5 cm. long, 2-2.5 mm. wide

near summit, compressed, the flowers usually 3 in small dichasia at the tips of

490 FIELDIANA: BOTANY, VOLUME 29

the 2-3 branches; central flowers sessile; hypanthium subglobose, thinly hirsute,

2.5 mm. wide, somewhat contracted to a neck; calyx-lobes 4, nearly equal, fleshy,

concave, rounded or obscurely triangular, 1.5-1.9 mm. long, 3-3.5 mm. wide,

glabrous or essentially so within; disk quadrate, 4-5 mm. wide, the staminal ring

soft-hairy; style soft-hairy at base (not seen complete); stamens about 150; petals

broad-clawed, finely pubescent near base, about 4 mm. broad and long; ovary

bilocular, the locules hirsute on the inner surfaces, the ovules 10-11 in each locule,

radiating in one plane from an axile horseshoe-shaped placenta; fruit "round,

yellow," according to Acosta-Solfs, not seen. "Mate-mate."

ECUADOR: Loja: Bosque de "La Mira," Hda. La Hamaca, Cata-

cocha, alt. 2400-2600 m., 17 Apr. 1944, Acosta-Solis 7923 (F, type).

The generic position of this species may be questioned by some.

The disposition of the ovules and the irregularly forked flowering

branches are somewhat suggestive of Psidium, but against this are

the bilocular ovary and the 4-parted calyx. If the fruit is actually

yellow, as stated by the collector, this may indicate a relationship

to Psidium. Fruiting specimens should be sought for final disposi-

tion of this species.

Myrcianthes karsteniana (Berg) McVaugh, comb. nov. Eu-

genia karsteniana Berg, Linnaea 27: 277. 1856. Myrcia karsteniana

(Berg) Steyermark, Fieldiana, Bot. 28: 1017. 1957.

A recent collection, Garcia 94, is the first-known specimen in the

fruiting condition. In the slightly immature fruit the beanlike em-

bryo separates easily into 2 plano-convex plump cotyledons; the

radicle is short and erect, the plumule immature but perceptible.

On the basis of this embryo-structure, the species is to be referred

to Myrcianthes as I understand it. In two flowering specimens,

Fendler 2321 and Allart 413, the ovules are 15-20 in each of two

locules, radiate in a subcapitate group affixed to the central dissepi-

ment; this seems to rule out any possibility that the species belongs

with Myrcia. I have not seen the syntypes (Karsten, Moritz in herb.

Berlin), but duplicates photographed in Vienna (Field Mus. Negs.

31578, 31579) certainly are the same species as the Fendler, Allart

and Garcia specimens.

The inflorescence in this species is not exactly like that of other

species of Myrcianthes; it is in fact a myrcioid panicle (see Fieldiana,

Bot. 29: 158-160. 1956) 6-11 cm. long, each main branch with 5-6

decussate nodes from which arise shorter branches, the strongly

compressed internodes 2.5-5 mm. wide or the distal ones narrower,

the flowers numerous, mostly in 3- or 7-flowered dichasia at the tips

of the branches.

McVAUGH: TROPICAL AMERICAN MYRTACEAE 491

In Eugenia pycnantha Benth., a species of western Ecuador and

southern Colombia (also erroneously referred to Myrcia by Steyer-

mark, I.e. 1018), the inflorescences are corymbosely aggregated

toward the ends of the branches, the individual dichasia are 3 or 4

times dichotomously forked with flat joint-like more or less appressed

internodes 1 cm. long or less. This is probably also best referred to

Myrcianthes, but certain disposition of it must await the discovery

of fruiting material. The ovules are 10-15 in each of two locules,

definitely excluding this species from the Myrciinae.

Myrcianthes leucoxyla (Ort.) McVaugh, comb. nov. Myrtus

leucoxyla Ort. Dec. 129. t. 17, fig. 2. 1800. Myrtus foliosa HBK.

Nov. Gen. & Sp. 6: 134 (folio ed. p. 106). 1823. Myrcianthes foliosa

(HBK.) McVaugh, Field Mus. Bot. 13, pt. 4: 758. 1958.

The name Myrtus leucoxyla Ort. seems to have been overlooked

both by DeCandolle and by Berg in their general revisions of the

American Myrtaceae. The type-locality according to Ortega was

"in novo Regno Granatensi cum Hopea terniflora" [i.e. "in Sanctae-

fidensi Australis Americes Regno, ad urbem principem, aliisque

passim locis," or in other words near Bogota, Columbia, where the

local name was said to be palo bianco]. Ortega's plate is a very

good representation indeed of the plant long known as Myrtus foli-

osa or as Eugenia foliosa (HBK.) DC., a species common in the moun-

tains about Bogota. The description calls for a glabrous plant with

short-petiolate, ovate, subcordate leaves with short-acuminate tip;

3-flowered axillary peduncles and short-pedicellate flowers; four-

parted calyx with subrotund concave ciliate lobes reflexed in flower,

and subglobose fruit. There can be little doubt that Myrtus foliosa

HBK. and M. leucoxyla Ort. are conspecific.

Eugenia mariquitensis Berg, Linnaea 27: 267. 1856.

This was based on a specimen from Mariquita, Colombia, Lin-

den 1102. I have seen isotypes in the herbaria of Harvard University

and the Naturhistorisches Museum in Vienna, and I should refer

this species to the synonymy of Myrcianthes rhopaloides (HBK.)

McVaugh.

Myrcianthes mato (Griseb.) McVaugh, comb. nov. Eugenia

mato Griseb. Abh. Kgl. Ges. Wiss. Goett. 24: 125. 1879.

I have not seen authentic material of this species, but Legrand

studied a "cotype" (Lorentz & Hieronymus 880), and on the basis

492 FIELDIANA: BOTANY, VOLUME 29

of this and other material published a detailed description and an

illustration contrasting this species with another previously unde-

scribed, E. pseudo-mato (Legrand, D., Contribution al conocimiento

de tres arboles argentinos de la familia de las mirtaceas. Lilloa 10:

471-482, with plate. 1944). The embryo of E. mato is described by

Legrand as follows (I.e., p. 476) : ". . . cotiledones grandes, algo con-

vexos, separados, con radicula diminutas exserta."

This plant is very like Eugenia [Myrcianthes] pseudo-mato and

other undoubted species of Myrcianthes, except that the peduncles

are one-flowered. This is apparently not a particularly fundamental

character, as one-flowered peduncles may be found occasionally in

various species of Myrcianthes that are normally 3-flowered (see the

key in Flora of Peru, 13, pt. 4: 747-749).

In addition to the specimens cited by Legrand, the following may

be mentioned:

ARGENTINA: Tucuman: Venturi 1991 (A), 10035 (A, GH).

Myrcianthes minimifolia (McVaugh) McVaugh, comb. nov.

Eugenia minimifolia McVaugh, Fieldiana, Bot. 29: 213. 1956.

This was treated in the Flora of Peru, 13, pt. 4: 764-765, as a

species of Myrcianthes, but the formal nomenclatural transfer was

not made because this species has one-flowered peduncles and at that

time I had seen none except flowering specimens. In 1959 Dr. Ra-

mon Ferreyra sent me fruiting specimens; in the somewhat imma-

ture embryo there are two large fleshy separate cotyledons and a

short radicle. This would appear to justify the transfer made above.

The following additional specimens have been seen:

PERU: Arequipa: Prov. Caraveli, Lomas de Pongo cerca de Acari,

700-750 m., Ferreyra 13423 (MICH), 13*81 (MICH).

Eugenia orthostemon Berg, Linnaea 27: 179. 1856.

This was based on Linden 807, from near Fusagasuga, Colombia.

Because of the one-flowered peduncles it was placed by Berg in his

large group Biflorae, not with the Dichotomae. On the basis of most

of its morphological characteristics, I believe it belongs with the other

Andean species of Myrcianthes, but fruit is needed to confirm this.

It may be related to such species as Eugenia punicifolia (HBK.) DC.

Myrcianthes prodigiosa McVaugh, sp. nov.

Arbor glabra 5-15-metralis; folia rigide coriacea, elliptica vel elliptico-ovata

vel -obovata, 10-23 cm. longa, 5-10 cm. lata, brevi-acuminata, basi rotundata

McVAUGH: TROPICAL AMERICAN MYRTACEAE 493

usque ad acuta, petiolis 1.5-2.5 cm. longis, nervo medio supra impresso, venis

lateralibus numerosis inconspicuis; inflorescentia crassa, plerumque triflora, pe-

dunculo compresso 3-8 cm. longo, versus apicem 4-5 mm. lato; alabastra turbinata

1.7-2 cm. longa; calycis lobi 4 (-5), suborbiculares, 5-10 mm. longi latique; discus

12-15 mm. latus; stylus 2.5 cm. longus; stamina ca. 250; ovarium triloculare, loculis

duobus minoribus, ovulis quoque loculo 10-20; fructus globosus oblatusve, dia-

metro 4-5 cm.; semina 2 (-1), cotyledonibus 2, plano-convexis inaequalibus.

A tree 5-15 meters high, 20-30 cm. in diameter, glabrous except for stiff sordid

hairs on the vegetative buds, and for the bristly staminal ring; leaves rigidly cori-

aceous, elliptic or elliptic-ovate or -obovate, 5-10 cm. wide, 10-23 cm. long, 2-2.5

times as long as wide, abruptly short-acuminate at tip, the base unequal-sided and

rounded to acute, the revolute margins decurrent on the narrow inner angles of

the stout petioles 2-3 mm. thick, 1.5-2.5 cm. long; midvein narrowly impressed

above, elevated most of its diameter beneath; lateral veins numerous, 12-20 pairs

including some intermediate ones, inconspicuous on both sides or somewhat ele-

vated above when dry; marginal vein like the lateral ones and somewhat arched

between them, 2-5 mm. from the margin; blades lustrous above and finely resinous-

dotted (sometimes impressed-punctate), dull below, opaque and nearly featureless;

inflorescences in leaf-axils, or a pair opposite at the lowermost (bracteate) nodes

of a short terminal branch; peduncles stout, 3-flowered or by reduction 2-flowered,

3-8 cm. long, compressed toward the summit and there 4-5 mm. wide; terminal

flower sessile or its pedicel 3-5 mm. long, the lateral pedicels 5-15 mm. long;

bracteoles falling before the buds mature; buds turbinate, 1.7-2 cm. long; calyx

4- to 5-merous, the lobes 4 or 5 and subequal, or 4 with one additional smaller

lobe; lobes suborbicular, fleshy with thin hyaline short-ciliate margins, 5-10 mm.

long and wide, soon reflexed at anthesis; hypanthium 1 cm. long, attenuate to the

base; disk 12-15 mm. wide (10-12 mm. wide in fruit), the glabrous center 6-8 mm.

wide; style 2.5 cm. long; stamens about 250, about as long as the style; petals

white; ovary trilocular (one locule sometimes about as large as the other two

combined), the ovules 10-20 in each locule, in compact somewhat elongate groups

radiating from the central axis; fruit green, globose or oblate, 4-5 cm. in diameter;

seeds 1 (-2), up to 4 cm. long, with leathery testa wrinkled in drying, and 2 unequal

plano-convex cotyledons.

BRITISH GUIANA: Kaieteur Plateau, savanna and savanna forest,

Kaieteur Falls to Mure-Mure Savanna, elev. 1400 ft., 12 Mar. 1962

(fr.), R. S. Cowan & T. R. Soderstrom 2147 (MICH).

SURINAM: Tafelberg, near summit, margin of North Ridge Creek,

mixed high forest, 8 Sep. 1944 (bud & fl.), B. Maguire 24657 (MICH,

type); Tafelberg, 5 km. northeast of Savanna No. 2, mixed high

forest, 1850 ft., 13 Sep. 1944 (old fl.), Maguire 24711 (MICH).

This extraordinary species can hardly be referred to any genus

except Myrcianthes, but it is unique in that genus by virtue of its

very large flowers and fruits and its trilocular ovary.

Myrcianthes pseudo-mato (Legrand) McVaugh, comb. nov.

Eugenia pseudo-mato Legrand, Lilloa 10: 477. 1944. Pseudomyrci-

anthes pseudo-mato (Legrand) Kausel, Ark. Bot. II. 3: 505. 1956.

494 FIELDIANA: BOTANY, VOLUME 29

I have not seen the type, but from the photograph of it published

by Legrand, and from his excellent description and illustration, there

can be no doubt of its identity. It is clearly a species of Myrcianthes;

actually some Argentinian and Bolivian material can scarcely be dis-

tinguished from M. discolor of the Peruvian Andes. In West's

no. 8312, cited below, the cotyledons are distinct, plano-convex,

the radicle half as long, the plumule evident, silky, 1.5 mm. long.

The following specimens have been seen:

BOLIVIA: Tarija: Rincon de la Victoria, 15 km. southwest of

Tarija, J. West 8312 (GH).

ARGENTINA: Tucuman: Dept. Chicligasta, Estancia Las Pavas,

P. Jorgensen 13 (GH; this number also cited by Legrand). Dept.

Monteros, Quebrada de Caspiuchango, Schreiter 5765 (Herb. Lillo

6672) (F, GH), Venturi 9595 (A, GH, US). Dept. Tafi, Cerro de

Taficillo, Venturi 9723 (A, GH), 9998 (US).

The genus Pseudomyrcianthes was distinguished by Kausel be-

cause it was said to have the placenta arising near the apex of the

locule, the embryo sub-homogeneous ("Embryo homogeneus later-

aliter paullo retusus, fissuram interembryonariam longitudinalem

praebens") and splitting only partially. My observations on speci-

mens of what I take to be Myrcianthes pseudo-mato do not confirm

those of Kausel on the structure of the embryo as described for

Pseudomyrcianthes. I cannot attest to the importance of apical

versus central placentation, but the distinction between these would

seem to be somewhat subjective.

Eugenia pycnantha Benth. PI. Hartw. 174. 1845. Colombia

(the type from the province of Popayan, Hartweg 976) and Ecuador.

This is probably a species of Myrcianthes, in general appearance

not unlike M . bifurcata, and M. osteomeloides (see Flora of Peru, 13,

pt. 4: 750, 752, 768, 773), and somewhat suggesting M. karsteniana

(q.v. above). Fruiting material is needed for proper generic dis-

position.

Eugenia rigidissima Cufodontis, Archivio Bot. 9: 198. 1933.

Costa Rica, the type from Volcan de Turrialba at 2500 m., Porsch

758.

Standley (Field Mus. Bot. 18: 774. 1937) referred this species to

the synonymy of E. storkii Standl. I have not seen the type of

E. rigidissima, but it may be a distinct species. Amshoff (Ann.

McVAUGH: TROPICAL AMERICAN MYRTACEAE 495

Missouri Bot. Card. 45: 184. 1958), in writing of P. H. Allen 1563,

from Cerro Punta at 2000 m., Chiriqui, Panama, refers this speci-

men to E. fragrans but says: "... a rather distinct looking specimen

with suborbiculate, very rigid leaves with impressed nerves agreeing

with Eugenia rigidissima Cuf . ..."

Allen's specimen (F, US) may be described as follows:

Tree 30 m.; young growth with silky-matted hairs, the year-old branchlets

strigose; leaves very broadly rounded, 1.5-2.8 cm. wide, 1.8-3 cm. long, rounded

or obtuse at both ends, very coriaceous, 3-5 pairs of the lateral veins impressed

above, stronger than any intermediate veins and prominent beneath; buds gla-

brous, 5-6 mm. long; inflorescence glabrous, the peduncles 3-flowered, 1-2 cm. long,

2-2.5 mm. wide at apex; disk glabrous, 3.5-4.5 mm. wide; stamens 175-200;

ovules about 30 in each locule.

On the basis of characteristics used elsewhere in separating spe-

cies of Myrcianthes, this would deserve recognition as a distinct

species.

Eugenia rondonensis Steyerm. Fieldiana, Bot. 28: 1013. 1957.

Venezuela, the type from Mt. Roraima, Steyermark 58956 (F).

This is a nearly glabrous ample-leaved plant looking superficially

like some form of M. fragrans or M. dugandii. The type is a speci-

men in very immature bud (Univ. of Mich. Neg. 1337). The pedun-

cles are relatively long and very slender (5-6 cm. long), all 3-flowered,

the hypanthium glabrous like the rest of the inflorescence, the ovules

about 12 in a subcapitate group in each of two locules. In the key

above, this specimen would be identified as M. dugandii, except that

the peduncles are 3-flowered and unusually long and slender. It

does not appear to be a very distinctive species, but the available

material is too immature to permit a judgment.

Myrcianthes sessilis McVaugh, sp. nov.

Arbor 12-metralis, innovationibus gemmisque, et petiolis, pilis appressis seri-

ceis, pallidis, usque ad 0.5 mm. longis, rare praeditis; hypanthio et loborum calycis

superficie exteriori similiter sed densiuscule sericeis; foliis 4-8 cm. longis ellipticis

obtusis vel obtuse acuminatis, venis inconspicuis; dichasio 3-floro, pedunculo 2-

3 cm. longo, apicem versus 2-2.5 mm. lato, floribus sessilibus; stylo 7-8 mm. longo.

A tree to 40 feet high (Smith), the youngest branchlets and petioles, and the

leaf-buds thinly appressed-silky with white hairs up to 0.5 mm. long; hypanthium

and the outer surface of the calyx-lobes similarly but conspicuously and densely

pubescent; leaves coriaceous, lustrous above, elliptic, 2-3.5 cm. wide, 4-8 cm. long,

1.8-2.2 times as long as wide, the tip obtuse to acute or bluntly acuminate, the

base acute, the margins decurrent on the sulcate petiole 1-1.5 mm. thick, 8-10 mm.

long; midvein impressed above, prominent beneath; lateral veins 10-12 pairs, in-

496 FIELDIANA: BOTANY, VOLUME 29

definite in number with some intermediate veins, in drying raised a little on both

surfaces; marginal vein about equaling the laterals and arched between them,

1-2 mm. from the margin; inflorescence an axillary dichasium, 3-flowered, the

flowers all sessile, the peduncle 2-3 cm. long, compressed and enlarged distally,

where 2-2.5 mm. wide; bracts and bracteoles deciduous before anthesis; buds

5-6 mm. long, obovoid; hypanthium 3 mm. long, funnelform or somewhat rounded

at the base; calyx-lobes in 2 nearly equal pairs, rounded, glabrous or silky on the

inner surface, 3-5 mm. wide, 2.5-3.5 mm. long; disk 4-5 mm. wide, the broad

staminal ring hairy, the style-base conic, 1 mm. wide; style 7-8 mm. long; stamens

150-175, to 8-9 mm. long, the anthers 0.7-0.8 mm. long; petals oblong-obovate,

ciliate-f ringed and silky on outer surface near base, 6 mm. wide and 10 mm. long;

ovary bilocular, the ovules 8 in each locule, somewhat biseriate on a centrally

affixed placenta; fruit not seen.

COLOMBIA: Masinga, [Sierra de] Santa Marta, rare below 1500

feet, generally in wooded valleys, Dec. 22 [1898-1901], Herbert H.

Smith 776 (A; F; GH, type; MICH; PH; US); Santa Marta, Smith

1735 (F; PH; US).

Apparently a most distinctive species, easily recognized from the

limited material now available by the abundant silky pubescence,

the sessile flowers and the long-petiolate, lustrous, and coriaceous

leaves.

Myrcianthes storkii (Standl.) McVaugh, comb. nov. Eugenia

storkii Standl. Field Mus. Bot. 8: 143. 1930.

Young growth thinly silky-strigose, the 1-year-old branchlets thinly strigose

or glabrate; leaves obovate or rounded to broadly elliptic, 1.5-3.5 cm. wide, 2-5

cm. long, 1-1.7 (-2.5) times as long as wide [on shoots up to 2.5-4.5 cm. wide,

4.5-7 cm. long, 1.5-1.75 times as long as wide]; blades coriaceous, rounded or

bluntly pointed at apex, cuneate to obtuse (or rounded) at base, the veins not

impressed above but 3-7 pairs of the laterals much stronger than the intermediate

ones and prominent beneath; buds 4-6 mm. long, glabrous; inflorescence glabrous,

the peduncle 3-flowered, 2-2.5 cm. long, 1.5-1.7 mm. wide at the apex; disk hir-

sute, 3.5-4 mm. wide; stamens ca. 150; ovules ca. 20 in each of 2 locules.

Costa Rica, in potreros and about edges of woodlands, 1900-

2850 m., flowering in April and May.

COSTA RICA: Potrero below Poas, H. E. Stork 2501 (F, type);

east of Irazu, Stork 2074 (F, paratype; MICH); Alajuela, south

slope of Volcan Poas, Austin Smith P2401 (A, UC) ; Alajuela, Pal-

mira, Smith NY 676 (A), H676 (F), P2627 (F); El Tablazo, Echever-

ria 377 (F, UC); Prov. Heredia, Cerro de las Caricias, Standley &

Valeria 52193 (US).

Of the four paratype collections of E. storkii, only one (Stork

2074) can be confidently referred to this species. Two (Stork 1228

McVAUGH: TROPICAL AMERICAN MYRTACEAE 497

and Lankester K99) are cited above under M. fragrans var. hispidula.

The fourth paratype (Standley & Torres 47426, from Fraijanes, Ala-

juela, at 1500-1700 m.) is sterile (F, US) and indeterminable. Two

other collections from the same general locality (Standley and Torres

47619, 47725, both US) are similarly sterile and are likewise inde-

terminable.

For discussion of the taxonomic position of this species, see above

under M. fragrans var. hispidula.

Myrcianthes ternifolia (Berg) McVaugh, comb. nov. Eugenia

ternifolia Berg, Linnaea 27: 246. 1856. Venezuela, the type "in

montibus Sierra Nevada," Moritz 1183.

A little-known species of the mountains of northeastern Colom-

bia and adjacent Venezuela. The leaves are sometimes partly oppo-

site, usually ternate; the flowers in all the specimens I have seen are

aggregated near the tips of the branches in abundantly produced

but 3-flowered stout axillary dichasia; the total effect is that of a

series of small corymbose panicles suggesting the inflorescences of

Eugenia pycnantha, q.v.

Berg distinguished this species from Eugenia triquetra by the

spatulate leaves and the "pedunculis saepissime 3-floris." A photo-

graph of a presumed isotype, however, Moritz 1183 (G-DEL), shows

the peduncles chiefly 1-flowered (Field Mus. Neg. 23594). Possibly

Moritz 1183 was a mixture. Berg described the ovary as bilocular

and many-ovulate.

Eugenia triquetra Berg, Linnaea 27: 141. 1856. Colombia and

Venezuela, the syntypes from "inter Pamplona et Tapagua," Linden

726 and "in montibus Sierra Nevada, in Venezuela," Moritz 1184-

A poorly understood species, treated in the Flora of Peru, 13, pt. 4:

774 with Myrcianthes. It seems to be still unknown in the fruiting

condition.

Eugenia turumiquirensis Steyerm. Fieldiana, Bot. 28: 1015.

1957. Venezuela, the type from Sucre, Cerro Turumiquire, Steyer-

mark 62548.

The type is in young bud. It suggests a moderately or perhaps

more than ordinarily pubescent form of Myrcianthes fragrans. If it

had come from Jamaica or Central America I should have referred

it without question to M. fragrans. — Univ. of Mich. Neg. 1338.

498 FIELDIANA: BOTANY, VOLUME 29

6. CERTAIN EUGENIOID GENERA

There is need for some clarification of generic lines among cer-

tain taxa that seem to represent specialized lines of evolution among

the Eugeniinae. In these the flowers are morphologically racemose

but the axis of the raceme is usually so much reduced in length that

the flowers appear to be in small axillary fascicles or glomerules; the

number of ovules in each locule of the ovary is reduced at least to 4

and usually to 2; the hypanthium is prolonged into a tube or cylin-

der extending well above the level of the summit of the ovary; and

the combined structures of the calyx and hypanthium are otherwise

specialized: The calyx is closed or nearly closed in the bud, and

usually splits irregularly down toward the summit of the ovary at

flowering time, and in many species the tubular part of the hypan-

thium is circumscissile and cleanly dehiscent from the summit of

the ovary.

In recent years explorations in northern South America have

brought to light a number of species, mostly as yet unstudied and

undescribed, that according to current interpretations might be as-

signed (and sometimes have been assigned) to Marlierea, Myrciaria,

Plinia or Siphoneugena. A considerable number of flowering speci-

mens is available, but fruiting specimens are poorly represented in

herbaria, and it has not often been possible to associate flowering

and fruiting material of the same species. Characters of the embryo

are especially important in the delimitation of the genus Plinia as

understood by Amshoff (Fl. Suriname 3: 97-99. 1951) and McVaugh

(Flora of Peru, 13, pt. 4: 775-780. 1958). As a result of the incom-

pleteness of material now available, it is sometimes impossible to

assign flowering specimens to genus. As I understand the genera

listed above, they may be distinguished as follows (Marlierea not

included, as its relationships are with the Myrciinae; it resembles

the other genera only in having 2 ovules. The key that follows is

particularly applicable to plants of northern South America; the

numerous West Indian species of Plinia described by Urban have

not been considered, as they are mostly vegetatively very different

and few of them are known except from sterile specimens) :

Hypanthium circumscissile at base, cleanly deciduous (with the perianth and

androecium) at about the time of anthesis, leaving a circular scar on the

ovary and fruit; calyx shortly four-lobed, the lobes ciliate, probably always

imbricate but often obscurely so.

Ovary below the deciduous part of the hypanthium prolonged, fusiform or cam-

panulate, usually contracted into a neck in bud and in flower; bracteoles

scarcely connate, much shorter than the ovary; embryo as far as known

McVAUGH: TROPICAL AMERICAN MYRTACEAE 499

with 2 distinct plano-convex cotyledons; ovules usually 4 in each locule;

flowers mostly pedicellate; wet mountain forests, known chiefly from the

Guayana region of Venezuela; Guadeloupe Siphoneugena.

Ovary broadened from the base upward, not contracted below the line of de-

hiscence of the hypanthium; after dehiscence broad and flat, scarcely longer

than the involucre formed by the broad connate bracteoles; pedicels almost

none, or very stout and scarcely longer than the bracteoles; embryo as far

as known undivided ("eugenioid"); ovules usually 2 in each locule; wide-

spread Myrciaria.

Hypanthium not circumscissile; calyx closed in bud or sometimes lobed, in an-

thesis splitting by irregular longitudinal fractures down to the summit of the

ovary, some remnants usually persisting in fruit; embryo, as far as known,

with 2 free plano-convex cotyledons, one above and one below; ovules 2 in

each locule.. . .Plinia.

REVISION OF MYRCIARIA BERG, IN NORTH AMERICA AND

NORTHERN SOUTH AMERICA

The genus typified by Myrciaria tenella (DC.) Berg (cf. Taxon

5: 143. 1956) is a very natural and closely related group, more easily

recognized as a distinct genus than any other American group of the

Eugeniinae. Berg, unfortunately, included in the genus as he first

characterized it a number of species that obviously were unrelated

to the rest — in fact, mostly species of Myrcia — and this error on the

part of Berg may have led Bentham and Hooker to disregard the

very distinct and homogeneous generic group that remains after

these anomalous species have been removed. At any rate these

authors, in the Genera Plantarum, relegated Myrciaria to the synon-

ymy of Eugenia. Niedenzu, in the Naturlichen Pflanzenfamilien,

accepted Myrciaria as a distinct genus but apparently did not un-

derstand it well, delimiting it more or less in the original sense of

Berg to include several species of Myrcia. Beginning with the work

of Chodat and Hassler on the Myrtaceae of Paraguay (Bull. Herb.

Boiss. II. 7: 807-808. 1907), in which the authors recognized 11 spe-

cies of Myrciaria, the genus has been rather generally accepted by

students of the American flora.

The following key is intended to be applicable to all known spe-

cies of Myrciaria in South America north of the Amazon River, in

Central America and Mexico, and in the West Indies.

Plants glabrous except the ciliate-margined bracts and bracteoles, and the inner

faces of the perianth-parts; bracts membranous, ovate, sometimes nearly

concealing the buds; bracteoles membranous, distinct, orbicular or sometimes

pointed, 1.5-2.5 mm. wide and about as long; leaves elliptic-oblong or -ovate,

500 FIELDIANA: BOTANY, VOLUME 29

2-4.5 cm. wide, 5.5-13.5 cm. long, acuminate and acutely attenuately pointed,

broadly rounded at base, on stout terete petioles 4.5-8 mm. long; glands in-

conspicuous or not seen in mature leaves; veins apparent but inconspicuous

above, the lower surface opaque, the veins obscure; northern Venezuela;

Panama; Costa Rica M. vexator.

Plants sometimes glabrous, usually at least the petioles (ventrally) and the young-

est branchlets pubescent; bracts inconspicuous, much shorter than the brac-

teoles; bracteoles somewhat fleshy or coriaceous, usually wrinkled in drying

like the hypanthium, connate by the basal margins and forming an involucre-

like cupule beneath the flower; leaves various.

Leaves cordate or subcordate at base, the sinus between the basal auricles defi-

nite but sometimes shallow; blades ovate to oblong, 4-8 cm. long.

Leaves rounded or obtusely pointed at tips, prominently cordate, subsessile,

the very stout petioles 2-2.5 mm. thick, 2-4 mm. long from the basal

sinus of the blade to the base of the petiole; midvein about equally con-

vex on both surfaces; plants as far as known quite glabrous; style more

than 7 mm. long; Mt. Roraima M. cordata.

Leaves gradually acuminate, the tip attenuate and acutely pointed; blades

broadly subcordate; petioles pubescent, 1 mm. thick, 2-3 mm. long;

midvein flat or somewhat convex above, convex beneath; styles probably

6-7 mm. long (ex Berg); British Guiana; Suriname M. vismeifolia.

Leaves acute or cuneate or sometimes slightly rounded to base, sometimes with

minute auricle-like folds at the summit of the petiole, never cordate; blades

various, often elliptic to lanceolate.

Leaves at apex acute (usually not prominently acuminate), mostly 6-10 cm.

long; blades when dry often discolorous, the lower surface smooth and

opaque, the upper dotted with large convex dark or amber-colored resi-

nous glands; lateral veins near the base of the blade leaving the midvein

at an angle less than 45°; small anastomosing veinlets almost none, the

small veins all nearly parallel and ascending; margins at the base of the

blade abruptly incurved into the petiole, often forming a pair of slightly

raised auricle-like folds; style (8-) 10-11 mm. long; plants glabrous or

the petioles (ventrally) and the youngest branchlets pubescent; widely

distributed in the Amazon basin and the upper Orinoco drainage, from

eastern Peru to Para (Santarem), north to southern Venezuela . .M. dubia.

Leaves at apex mostly prominently and often caudately acuminate, incon-

spicuously if at all glandular above, concolorous or essentially so; mar-

gins at the base of the blade cuneately decurrent or at least narrowly

acute (except in M. ibarrae with style 5-6 mm. long); style in all species

(as far as known) 7 mm. long or less.

Leaves (1.5-) 2.5-5 cm. wide, (4.5-) 7-14 cm. long, gradually long-acumi-

nate; plants glabrous or the petioles ventrally pubescent; glands in-

conspicuous or scarcely apparent on both surfaces of the leaves; lateral

veins near the base of the blade leaving the midvein at an angle of 45°

or more; small irregular anastomosing veinlets frequent; open flowers

unknown; Para, Brazil, near the mouth of the Amazon. .M. amazonica.

Leaves 0.7-2 (-3) cm. wide, 2-6 (-8) cm. long; branchlets and petioles

usually abundantly and finely pubescent; glands inconspicuous, or

small convex glands rather prominent beneath; lateral veins near the

base of the blade varying in direction, the small veinlets few or very

inconspicuous; style 4.5-6 mm. long.

McVAUGH: TROPICAL AMERICAN MYRTACEAE 501

Leaves of an ovate type, mostly 2-2.5 times as long as wide, sharply

acuminate and often cuspidate, broadly rounded at base, the mar-

gins little if at all decurrent on the petiole; branchlets and petioles

coarsely and rather sparsely hispidulous; flowers sessile; bracteoles

very broadly rounded and imbricate, connate at very base only,

1.5-2 mm. long and wide, much exceeding the ovary; Guatemala

(Pete"n) M. ibarrae.

Leaves of a lanceolate type, mostly 2.5-3 times as long as wide, variously

acuminate but the very tip usually blunt or merely acute, the base

of the blade somewhat rounded below the middle, finally usually

acute or cuneate with the margins cuneately decurrent on the splayed

summit of the petiole; branchlets and petioles finely pubescent or

rarely glabrous; pedicels usually 0.5-1 mm. long; bracteoles not im-

bricate (or very slightly so in the bud stage), united by their proxi-

mal margins into an oval bilobed or finally explanate involucre 2 mm.

long; West Indies; eastern Mexico and Central America; Colombia;

lowlands of the Guianas; Orinoco and Amazon lowlands from south-

ern Venezuela to Peru and eastern Brazil (Maranhao).

M . floribunda.

Myrciaria amazonica Berg, in Mart. Fl. Bras. 14, pt. 1: 374.

1857.

An imperfectly known species, known to me from two nearly

sterile specimens only, both of these from Brazilian Amazonia:

Pard: Belem, B. E. Dahlgren & E. Setta 398 (F); Insula Colares,

Poeppig s.n. (W, type). The leaves seem to be distinctive as indi-

cated in the key.

Myrciaria cordata Berg, Linnaea 27: 337. 1856.

Apparently known only from the original collections by Schom-

burgk. The type was seen by Berg at Berlin, and was cited by him

as from "Guiana Anglica (Rich. Schomburgk, coll. no. 957)"; another

collection, not cited by Berg, is from Mt. Roraima, Schomburgk 608

(F, W).

Myrciaria dubia (HBK.) McVaugh, comb. nov. Psidium du-

bium HBK. Nov. Gen. & Sp. 6: 152 [folio ed. p. 121]. 1823. M. para-

ensis Berg, in Mart. Fl. Bras. 14, pt. 1: 364. 1857. M. caurensis

Steyerm. Fieldiana, Bot. 28: 1020. 1957.

The type of Psidium dubium HBK., from near Atures on the

Orinoco, is surely a Myrciaria. This is indicated by the description

("bracteis . . . ovarium subaequantibus eique adpressis. . . . Calyx

superus, hemisphaericus, glaber, glanduloso-punctatus, margine quad-

rilobus. . . . Stamina . . . calyci medium versus inserta . . . ovarium

502 FIELDIANA: BOTANY, VOLUME 29

. . . biloculare; . . . ovula duo in quolibet loculo, apposita . . . fructus

. . . edulis"), and is apparent from a photograph (Field Mus. Neg.

36872). The species is not identifiable from the plate (t. 547bis)

accompanying the original description of P. dubium.

The type of P. dubium appears to represent a relatively narrow-

leaved race, frequent in the Orinoco drainage, of a species found also

in the Amazon drainage. The type of M. paraensis Berg has the

somewhat broader leaves characteristic of most specimens from

the Amazon basin, but I believe M. dubia and M. paraensis are

conspecific. The following may be cited as representing the species

in the Orinoco drainage or at its upper limits.

VENEZUELA [possibly COLOMBIA]: Prope Atures (Missiones del

Orinoco), Bonpland (P, not seen). — Bolivar: La Prision, Medio

Caura, L. Williams 11691 (F, type of M. caurensis); Cerro Guaiqui-

nima, Rio Paragua, Maguire 33135, 33139 (both MICH).— Ama-

zonas: Rio Sanariapo, H. M. Curran 1833 (NY); Capibara, Canal

del Casiquiare, Holt & Gehriger 289 (NY).

Myrciaria floribunda (Willd.) Berg, Linnaea 27: 330. 1856.

This, the most widely distributed species of the genus, has a

considerable synonymy. The principal synonyms are listed by Ur-

ban (Bot. Jahrb. 19: 657-658. 1895) and by Amshoff (Fl. Suriname

3: 108-109. 1951, and Ann. Missouri Bot. Gard. 45: 177. 1958).

An additional synonym is apparently M. maragnanensis Berg, in

Mart. Fl. Bras. 14, pt. 1 : 372. 1857, of which I have seen an isotype

(Gardner 6023, W).

Two species described by Berg, Myrciaria verticillata Berg and

M. divaricata (Benth.) Berg, were based entirely or in part on

Schomburgk 958. This collection seems actually to have been a mix-

ture. A sheet of no. 958 (MICH) has the leaves apparently of

M. floribunda and is presumably to be referred to that species, al-

though the styles are about 7 mm. long. This is presumably the

plant described by Berg as M . verticillata ("foliis . . . utrinque angus-

tatis, apice longissime lineari-acuminatis, basi elongato-acutis. . . .")•

Another sheet of no. 958 (F) has the leaves of M. dubia (HBK.)

McVaugh, but the flowers are smaller than usual in that species;

I have not seen the type of Eugenia divaricata Benth., but the leaves

are described as being rounded at base, i.e., therefore more like dubia

than floribunda. Apparently Berg had the same plant, for he de-

McVAUGH: TROPICAL AMERICAN MYRTACEAE 503

scribed M. divaricata as with "foliis . . . ovatis v. ovato-oblongis, . . .

acute acuminatis, basi rotundatis v. breviter acutis. ..."

The plant from Jamaica described as Eugenia polyneura Urb.

Symb. Antill. 5: 446. 1908 (E. pycnoneura Urb. Symb. Antill. 6: 25.

1909, not E. polyneura Koord. & Val. 1855) is apparently not spe-

cifically different from M. floribunda. I have seen an isotype (F)

or perhaps the actual type (Jamaica below Vinegar Hill, W. Harris

7448, ex herb. Krug et Urb.). The specimen is nearly glabrous,

with but traces of pubescence on the petioles; the petioles them-

selves are 7-8 mm. long on the mature leaves; the blades are 1.5-

2.5 cm. wide, elliptic-ovate and mostly acutely long-acuminate.

The combination of nearly complete lack of pubescence, broad sharp-

tipped leaves and long petioles is most unusual in M. floribunda,

but the individual features are matched in many other South Amer-

ican and West Indian specimens.

I have not seen specimens of Eugenia asa-grayi Krug & Urb.

[Plinia asa-grayi (Krug & Urb.) Urb.], nor of E. I'heritieriana DC.,

both of which species were associated by Urban (1895) with his

Eugenia floribunda. The former was from eastern Cuba (Wright

1610) and the latter supposedly from the island of Tobago.

Myrciaria ibarrae Lundell, Wrightia 2: 213. 1961.

This recently described species is known to me only through the

original specimens cited by Lundell. It seems to be abundantly

distinct from M. floribunda, which is known from the same gen-

eral area.

Myrciaria phillyraeoides Berg, Linnaea 27: 326. 1856.

This plant, based on a collection by Humboldt in "America me-

ridionali," and seen by Berg in the Willdenow herbarium (No. 9574),

I do not recognize from the description. It may not be a Myrciaria.

If it belongs to that genus, I suspect it is M. dubia.

Myrciaria vexator McVaugh, sp. nov. M. piltieri Burret ex

Badillo in Pittier et al. Cat. Fl. Venez. 2: 198, nomen. 1947. Eugenia

palmarum Standl. & L. O. Williams ex Allen, Rain For. Golfo Dulce

201, nomen. 1956.

Arbor 10 m. alta, bracteis bracteolisque, et perianthii segmentis, ciliatis, peri-

anthii segmentis intus tomentulosis; folia elliptico-oblonga vel ovata, 2-4.5 cm.

lata, 5.5-13.5 cm. longa, acuminata, apicibus attenuatis acutis; lamina basi ro-

tundata, marginibus plusminusve decurrentibus, petiolis teretibus crassis anguste

504 FIELDIANA: BOTANY, VOLUME 29

profundeque sulcatis 4.5-8 mm. longis; glandulae vix prominulae; bracteae ovatae,

membranaceae, usque ad 3 mm. latae, usque ad 4-4.5 mm. longae, saepe alabastra

fere occultantes; bracteolae rotundae, distinctae, membranaceae, 1.5-2.5 mm.

longae lataeque; hypanthium supra germen 2 mm. productum; discus 1.3-1.5 mm.

latus, glaber; stylus 8 mm. longus; calycis lobi 2 mm. lati, 1.5 mm. longi; petala

ovalia 2 mm. longa lataque; stamina ca. 50, antheris 0.6-0.8 mm. longis; fructus

globosus, atropurpureus, carnosus, diametro 2.5 cm.

Attributed to Burret by Badillo, as a new species, but quite

without description; described in English by Allen, under the name

of Eugenia palmarum.

This species is known mostly from cultivation, but has appar-

ently been collected in the wild both in Costa Rica and in northern

Venezuela. Specimens examined are:

COSTA RICA: Puntarenas, forested hills above Palmar Norte, alt.

450 m., P. H. Allen 6331 (F).

PANAMA: Summit Gardens, C. Z., J. E. Higgins 287 (type, US);

Summit, J. Zetek 3520 (F; MO, sterile, probably this species; see

below) .

VENEZUELA: Lara: Bosques de Chirgua, entre Nonavana y Bar-

quisimeto, Jose Saer 515 (F). Carabobo: Valencia, in gardens,

Pittier 8154 (US); Valencia, often cultivated, Pittier 8899 (US).

Myrciaria vismeifolia (Benth.) Berg, Linnaea 27: 336. 1856.

The type was from British Guiana; Amshoff cites a number of

collections from Suriname. In the Flora of Panama (Ann. Missouri

Bot. Gard. 45: 179. 1958) Miss Amshoff refers to this species a single

specimen from the Canal Zone, Zetek 3520, remarking at the same

time that this differs from the typical plant of the Guianas by its

"firmer leaves which are never (?) cordate at the base, its longer

petioles and its slightly larger flowers." The Zetek specimen, which

I have seen (MO), is sterile, but can hardly be referred to M. vismei-

folia. In that species the petioles are 3-4 mm. long, ventrally some-

what pubescent and flat or shallowly sulcate; the blades are defi-

nitely cordate-auriculate, the auricles extending mostly 1-1.5 mm.

beyond the sinus at the insertion of the petiole. In the Zetek speci-

men the petioles are 5-8 mm. long, quite glabrous (like the rest of

the specimen), nearly terete but with a narrow sharply indented

ventral furrow; the blades are rounded at base, but not auriculate.

The leaves are in fact strikingly similar to those of Myrciaria vexator,

and the specimen probably belongs to that species. In any event

M. vismeifolia should be excluded from the known flora of Panama.

McVAUGH: TROPICAL AMERICAN MYRTACEAE

A NEW SPECIES OF PLINIA L., FROM COSTA RICA

505

There seem to be no reports of any species of this genus from

continental North America. A few species are known in South

America, and a number have been reported from the West Indies.

FIG. 13. Plinia salticola, from the type. A, Open flower; B, Bud (X 2.5);

C, Enlarged tip of bud, seen from above ( X 5). The open flower is seen from the

adaxial side; the observer is looking down at the tip of the raceme-axis. Opposite

the open flower may be seen the bract and bracteoles of the other flower of the

proximal pair; at right angles to this may be seen the bracts and bracteoles of the

second (distal) pair of flowers. The small bud is seen from the abaxial side, with

bract at base and the bracteoles at the sides. In the enlarged terminal view of

the bud the outer calyx-lobes are shown above and below the central opening; the

inner lobes overlap and are concealed; compression of the tissues in the young bud

has given rise to curved ridges at right and left of the outer lobes.

The following species resembles in many respects such South Amer-

ican species as P. duplipilosa McVaugh, Fieldiana, Bot. 29: 224.

1956, but differs in detail from that and all other species known

to me.

Plinia salticola McVaugh, sp. nov.

Arbor vel frutex puberula, foliorum nervo medio subtus ad basin copiose pilis

pallidis appressis 1-2 mm. longis obsito; folia elliptica, 3.5-5 cm. lata, 11-17 cm.

506 FIELDIANA: BOTANY, VOLUME 29

longa, acute acuminata, basi subrotundata, petiolis 8-10 mm. longis, nervo medio

supra convexo, nervis lateralibus utrinque 7-12; inflorescentia abbreviata, glom-

erulis 4-floris, sessilibus, bracteatis; bracteis 2-seriatis, bracteolis membranaceis

obtusis ciliatis 3 mm. longis; alabastra pyriformia 9-10 mm. longa, basi sericeo-

strigosa, apice breviter 4-lobata, hypanthio supra germen 5-7 mm. producto,

stylo 16 mm. longo, staminibus ca. 150; ovarium biloculare, ovulis quoque lo-

culo duobus.

A tree or shrub, the branchlets and petioles thickly puberulent with short pale

ascending hairs 0.3-0.5 mm. long; hairs at the base of the midvein on the lower

leaf-surface appressed and longitudinally oriented, up to 2 mm. long; leaves ellip-

tic, 3.5-5 cm. wide, 11-17 cm. long, 2.5-3 times as long as wide, about equally

narrowed and rounded to a slenderly and sharply acuminate tip 8-10 mm. long

and to the obtusely rounded base, the curved margins meeting at about right angles

and passing abruptly to the nearly terete petiole 8-10 mm. long, 1-1.3 mm. thick;

midvein broadly convex above, persistently puberulent; lateral veins 7-12 on each

side including some intermediate ones, inconspicuous and sometimes inscribed

above, more prominent beneath, the proximal ones arcuate-ascending, those

toward the middle of the blade straight, connected by asymmetrical arches 5-

8 mm. from the margin, a distinct but weaker submarginal vein 1-3 mm. from

the margin; blades dark green and somewhat lustrous above, paler and finely

dark-dotted beneath; inflorescence at leafless axils on old wood, the flowers in

4's, in sessile bracteate decussate pairs; bracts membranous, scarious, ciliate,

ovate, blunt, 1.5-2 mm. wide, 2.5-3 mm. long; bracteoles like the bracts, ellip-

soid-oblong, blunt, 1 mm. wide, 3 mm. long, probably deciduous after anthesis;

buds 9-10 mm. long, pyriform, silky-strigose at base, rounded and obscurely

4-lobed at apex, the fusiform-ellipsoid base 2 mm. thick, 3 mm. long, the distal

part turbinate, 7 mm. long and about 5 mm. in diameter, formed by the much-

prolonged tube of the hypanthium; hypanthium at anthesis splitting irregularly

between the calyx-lobes, and the divisions recurving and bearing out the petals

and portions of the staminal ring; calyx-lobes in the bud short and broadly

rounded, about 1 mm. long, appressed, the inner ones relatively shorter, 2 mm.

wide, the outer ones bluntly ovate, 1.5 mm. wide at base; style 16 mm. long,

much exserted; stamens about 150, borne in a zone about 1.5 mm. wide at the sum-

mit of the hypanthium, strongly incurved in the bud, the anthers 1-1.5 mm. long;

petals 4, broad-based, subrotund. about 4 mm. long and wide; ovary bilocular,

the ovules collateral, 2 in each locule.

COSTA RICA: Cataratas de San Ramon, 21 Feb. 1931, A. M.

Brenes 1SW (F; NY, type).

At the present time I should refer to Plinia any eugenioid spe-

cies having 2 or 4 ovules in each locule of the ovary, the hypanthium

somewhat prolonged beyond the ovary but not circumscissile, and

the additional features outlined in the key above. The inflorescence

is usually, but not invariably, glomerate with relatively large bracts

and bracteoles simulating an involucre (cf. Amshoff in Rec. Trav.

Bot. Ne"erl. 42: 11. 1950). More complete data on fruit-types, em-

bryo-structure, and number of ovules per locule are needed before

the limits of the genus can be firmly established. The species now

McVAUGH: TROPICAL AMERICAN MYRTACEAE 507

assigned to Plinia on the basis of flower-characters seem to represent

a somewhat heterogeneous assemblage, as I have pointed out (Field

Mus. Bot. 13, pt. 4: 775. 1958), and it may be that some reconsidera-

tion of generic lines will be indicated when new data become available.

THE GENUS SIPHONEUGENA BERG

The type species of Siphoneugena is S. densiflora Berg, described

from somewhere in southeastern Brazil (cf. Taxon 5: 146. 1956; see

also Field Mus. Neg. 31474). According to Berg the genus was

distinguished from Myrciaria by having the locules of the ovary

multiovulate, and from other multiovulate genera with a prolonged

hypanthium by the circumscissile nature of that structure. The

three original species were all Brazilian in origin, one from Minas

Gerais and the others from unspecified localities. All three species

have at one time or another been transferred to other genera; the

genus Siphoneugena, as a matter of fact, hardly became known to

botanists and was given short shrift. By Bentham and Hooker it

was united with Eugenia; these authors regarded it as a connecting

link between Eugenia and Myrciaria. By Niedenzu it was merged

with Calycorectes and given the rank of a subgenus. No new taxa

in the genus were described for a little more than a century follow-

ing the work of Berg, until in 1957 Steyermark reported the discovery

of what he took to be a new variety of Siphoneugena densiflora from

the mountains of Venezuela. It now transpires that members of the

same taxon occur on a number of the high mountains (tepuis) of

Venezuela; in similar ecological situations in northern Venezuela;

and in the Lesser Antilles. All the specimens from northern South

America and the West Indies, specimens of the newly described

S. reitzii Legrand (a native of southern Brazil; cf. Sellowia 8: 78.

1958), and the specimens shown in the type-photograph of S. densi-

flora Berg, are so similar in general appearance and in structural

details that they could all belong to the same species. Surely they

are congeneric. The affinities of the taxon appear to be with Myrci-

aria rather than with Plinia or with Eugenia as a whole. In view of

the distinctive morphological features of Siphoneugena, and in view

of its distinctive geographical distribution as now demonstrable, I

suggest that it is best regarded as an independent genus, represent-

ing an evolutionary offshoot from eugenioid stock, parallel to Myr-

ciaria but in some respects less specialized. It is interesting phyto-

geographically that Myrciaria is primarily a lowland genus, wide-

508 FIELDIANA: BOTANY, VOLUME 29

spread both north and south of the Amazon Basin, whereas Siphon-

eugena is exclusively a montane genus, occupying a disjunct range

that spans the Amazon lowland.

Although Siphoneugena densiflora was reported by Berg as multi-

ovulate, this is open to question. Modern specimens of the genus

usually have 3 or 4 ovules in each of the two locules of the ovary.

The locules vary from 1 to 3 or possibly 4; the ovules may be as few

as 2, or occasionally as many as five. It is suggested that this vari-

ation in the number of ovules reflects a less specialized condition

than that in Myrciaria, in which the number of ovules is consist-

ently two.

There would appear to be little real affinity between this very

homogeneous and relatively invariable taxon, and the more hetero-

geneous assemblage that is called Plinia. The two have little in

common except that in each the number of ovules is reduced. The

South American species of Plinia, at least, are superficially dissim-

ilar to those of Siphoneugena, and perhaps are more closely related

to Calycorectes and to some of the larger-flowered species of Eugenia,

differing from the former in the reduced number of ovules, and from

the latter in the closed or nearly closed calyx. This feature of the

closed or nearly closed calyx has led some workers to confuse the buds

of Plinia with those of Siphoneugena. In the latter genus the dis-

tinctive feature of the flower is the circumscissile hypanthium, which

in most species terminates in four small but definite calyx-lobes.

These lobes are in unequal pairs, closely incurved in bud so that to

the casual observer the bud may appear closed. The lobes in the

opening flower are usually not perceptibly imbricate, but the differ-

ence in the form and position of the inner and outer pairs makes it

clear that they are not valvate. The inner pair are somewhat

broadly rounded, with membranous, long-ciliate margins that are

appressed to the opening petals; the outer pair are smaller, deltoid-

ovate, with thicker, often nearly erect and poorly fringed apex.

These differences are subtle, in lobes that may be no more than half

a millimeter long, but to one familiar with the imbricate calyx of

Myrciaria it is apparent that in this feature there are strong simi-

larities between the two genera.

In 1895 Krug and Urban described from the mountains of Guade-

loupe a species they called Marlierea dussii. Urban later transferred

this to Plinia. An isotype (Duss 2750, K) is in my judgment con-

specific with the Venezuelan Plinia fruticosa Steyermark, described

from the States of Sucre (Steyermark 62611, F, type) and Anzoategui

McVAUGH: TROPICAL AMERICAN MYRTACEAE 509

(Steyermark 61658 & 61620, both F). Neither species has anything

to do with the other species of Plinia, and both are typical members

of Siphoneugena.

The question of specific limits in Siphoneugena is a difficult one,

complicated by the peculiar isolation of individual small populations

on the tepuis of Venezuela, on other similarly isolated mountain

ranges in northern South America, and in the West Indies. Leaves

vary from plant to plant: from small to large, from ovate to elliptic

or obovate, from rounded to acute at apex, from rounded to acute

to cuneate at base. A few specimens, like those of Marlierea dussii

and Plinia fruticosa, have sparingly pubescent branchlets and /or

young foliage. Pedicels and buds vary somewhat in length from

plant to plant. These features make the assemblage of specimens

superficially diverse, but, as indicated in the description below, the

specimens have so many — and such specialized — features in common

that their relationship is apparent. I believe these isolated but not

excessively varied populations comprise a single species as indicated

below:

Siphoneugena densiflora Berg, in Mart. Fl. Bras. 14, pt. 1:

379. 1857. Calycorectes densiflorus (Berg) Ndzu. in Nat. Pflanzen-

fam. 3, Abt. 7: 82. 1893. Marlierea dussii Krug & Urb. ex Urb. Bot.

Jahrb. 19: 590. 1895. Plinia dussii (Krug & Urb.) Urb. Repert.

Sp. Nov. 15: 413. 1919. Paramitranthes densiflora (Berg) Burret,

Notizbl. Bot. Gart. Berlin 15: 541. 1941. Plinia fruticosa Steyer-

mark, Fieldiana, Bot. 28: 1023. 1957. Siphoneugena densiflora var.

tepuiensis Steyermark, I.e. 1024.

A shrub up to 3 m. high (rarely a tree up to 12-13 m. high), nearly glabrous,

the branchlets and petioles (and seldom the pedicels) sometimes strigose or pubes-

cent; inner faces of the petals and calyx-lobes silky tomentose with crisped matted

nearly white hairs; bracts and bracteoles ciliate, the inflorescence otherwise gla-

brous; leaves coriaceous, rigid when dry, lustrous and dark green above, dull and

paler beneath; blades mostly elliptic-ovate, short-petiolate, opaque and nearly

featureless except that the midvein is prominently convex above and somewhat

less so beneath; upper surface sometimes obscurely impressed-punctate, the lower

often with small convex translucent glands; lateral veins close and parallel, the

number indefinite, mostly 15-20, visible as fine convex lines on the lower surface

of dry leaves; blades up to 10 cm. long, mostly 3-5 cm., mostly about twice as

long as wide but sometimes as wide as long, or up to 2.7 times as long as wide;

petioles fleshy, dorsiventrally compressed, (1.5-) 3-4 (-6) mm. long, ventrally

concave, 1.5-2.5 mm. wide; inflorescence a much abbreviated raceme, the axis

up to about 7 mm. long, usually much shorter, bearing up to 5 approximate de-

cussate pairs of flowers; bracts broadly ovate-rounded, persistent, scarious; pedi-

cels up to 5 mm. long, sometimes much shorter, strongly dorsiventrally compressed

510 FIELDIANA: BOTANY, VOLUME 29

or even 2-edged, up to 1 mm. wide; bracteoles persistent, scarious, distinct, ovate,

blunt or acute, convex on the back, 0.7 mm. long, much shorter than the fusiform,

urceolate or campanulate ovary; hypanthium prolonged 1.5-2.5 mm. above the

ovary, usually notably flaring above the summit of the ovary and then contracted

to the mouth; bud as a whole 3.5-4.5 mm. long; calyx-lobes in unequal pairs,

scarcely imbricate, the outer pair deltoid-ovate, scarcely fringed, more erect, the

inner lying flatter, broadly rounded, copiously fringed, the free parts 1 mm. wide

or a little more, 0.5-0.7 mm. long; hypanthium funnelform at anthesis, splitting

two-thirds of its length on one side, or into 3-4 irregular lobes, its margins finally

rolled back (including most of the staminal ring) so the calyx and petals are re-

flexed, the whole ultimately circumscissile at base; style 4-6 mm. long; stamens

75-100, borne on the margin of the hypanthial tube; petals white, obovate, small;

locules of the ovary 2 (rarely 1 or 3), the ovules usually 4 (sometimes 3 or 5) in

each locule, ascending from near the base of the locule; fruit globose or nearly so,

purplish-black, soft and edible, probably 1 cm. in diameter or less; seed reniform,

the 2 cotyledons distinct or fused along the margins only, the radicle very small.

The type of Siphoneugena densiflora (as shown in Field Mus.

Neg. 31474) is a flowering specimen with leaves 2-3.2 cm. wide,

5-7 cm. long; the pedicels are up to about 5 mm. long, and the

styles 7 mm. long. The species was described by Berg as glabrous.

In most of its parts this plant is larger than the majority of plants

from north of the Amazon. The following, however, may be com-

pared with it except that the pedicels are shorter:

VENEZUELA: Amazonas: Serrania Yutaje, Maguire 35397, 35477

(both MICH); Cerro Sipapo (Paraque), Maguire & Politi 28228

(MICH, fr. only). — Dist. Federal: Cordillera del Avila, Steyermark

55604 (F; leaves as above, flowers smaller).

Specimens in the groups listed below conform to the general de-

scription given above for this species, but are divided into groups

characterized primarily by features of the leaves.

The following are glabrous; the leaves are elliptic or obovate,

mostly obtuse, occasionally obscurely acuminate, 2.5-5 cm. long:

VENEZUELA: Bolivar: Chimanta massif, Steyermark & Wurdack

729, 857, 1061 (all MICH); Ptari-tepui, Steyermark 59711 (F, type

of var. tepuiensis) ; Sororopan-tepui, Steyermark 60157 (F).

The following are somewhat pubescent; the leaves resemble those

of the preceding group but include some specimens with more ovate

leaves and some in which acuminate leaves are not rare; the flowers

tend to be nearly sessile, often with a few on each plant pedicellate:

GUADELOUPE: Duss 2750 (K, isotype of Marlierea dussii; most

leaves acuminate).

McVAUGH: TROPICAL AMERICAN MYRTACEAE 511

VENEZUELA: Anzoategui: Cerro Peonia above Santa Cruz, Steyer-

mark 61658 (F; paratype of Plinia fruticosa; leaves mostly acumi-

nate), Steyermark 61620 (F; paratype of P. fruticosa; leaves mostly

obtuse). — Sucre: Cerro Turumiquire, Steyermark 62611 (F; type of

P. fruticosa; leaves broader, ovate, obtuse to acuminate) .

The above data support the hypothesis that there are regional

populations, and perhaps local populations within these, of a wide-

ranging species.

7. PIMENTA Lindl.

The allspice of commerce, Pimenta dioica (L.) Merrill, is supposed

by some authors to be confined to the West Indies. There is some

evidence that the West Indian plants have smaller flowers and more

slender panicles, and smaller fruits with a greater proportion of vola-

tile oil, than do those of Central America. Because of these differ-

ences, Standley proposed to recognize the Central American plant

as a variety [P. dioica var. tabasco (Schlecht. & Cham.) Standl.

Ceiba 3: 172. 1953], and Lundell recognized it as a species distinct

from P. dioica [P. tabasco (Schlecht. & Cham.) Lundell, Wrightia 2:

58. I960]. Both of these names are based nomenclaturally on Myrtus

tabasco Schlecht. & Cham. Linnaea 5: 559. 1830, presumably with

the assumption that the name M. tabasco was based also on a Cen-

tral American plant. Examination of the protologue of M. tabasco,

however, shows that this was not the case.

In reporting on the Mexican plants collected by Schiede and

Deppe, Schlechtendal and Chamisso described a number of new spe-

cies; most of these naturally enough were Mexican in origin. The

protologue of Myrtus tabasco, however, begins as follows:

"542 [the serial number under which Schiede's collections were

reported]. Myrtus Tabasco H[erb.] W[illd.] n[o.] 9580. (Specimen

Humb. 'Thermidor an 7. Cumana. n. 113. Myrtus Pimenta. Ta-

basco').— Myrtus Pimenta, Pimenta de Tabasco Hispano-Mexicano-

rum Schiede in schedula."

It seems clear that Schlechtendal was merely taking up a name

by Willdenow based on a Humboldt specimen from Venezuela, and

identifying with this the new specimens from Mexico. Schiede's

collection cannot be regarded as the type of Myrtus tabasco, and

until and unless the identity of the Humboldt specimen can be de-

termined, it should not be assumed that Central American popula-

tions of allspice are members of the taxon to which the specimen

belongs. Berg, who seems to have seen both the Humboldt speci-

512 FIELDIANA: BOTANY, VOLUME 29

men and those of Schiede and Deppe, regarded them as varietally

different (cf. Linnaea 27: 424-425. 1856). His Pimento, officinalis /3

cumanensis, which he saw in Willdenow's herbarium, was presum-

ably based on the very Humboldt specimen that formed the basis

for Myrtus tabasco Schlecht. & Cham., and P. officinalis e tabasco

(Schlecht. & Cham.) Berg was based nomenclaturally, but not know-

ingly, on the same specimen. Berg described the Venezuelan plant

as having petioles 3-4 lines long; the Central American 9-12 lines;

his diagnoses are not otherwise very comparable. The leaves of

cumanensis are described as "approximato-tenuissime venosis," a

characterization that seems unlikely for the Central American pop-

ulation.

8. PSIDIUM L.

DELIMITATION OF THE GENUS

As discussed in an earlier article (Fieldiana, Bot. 29: 172-173.

1956), the two large pimentoid genera Psidium and Myrtus have

been separated traditionally upon the basis of calyx morphology.

The two have in common a C-shaped or uncinate embryo and hard,

bony seeds. The delimitation of Myrtus is particularly difficult, as

pointed out by Bentham in an important paper many years ago

(Jour. Linn. Soc. Bot. 10: 101-166. 1868), because the concepts of

earlier authors included numerous species-groups now known to

belong to distinct genera (even members of other sub-tribes); be-

cause fruits and seeds are unknown in many supposed species of

Myrtus; and because there is no sharp distinction between the num-

ber of locules in the ovary of Myrtus (2-3) and in that of the closely

related Psidium (3-5 or sometimes only 2). Since the time of

Bentham, taxonomic opinion has favored the continuing dismem-

berment of Myrtus, which at the time of the publication of the

Genera Plantarum of Bentham and Hooker (1865) included more

than 100 species in South America, tropical North America, Austra-

lia, New Zealand, and one (the type species) in the Mediterranean

region.

As defined by Berg in his classic papers on the American Myrta-

ceae (Linnaea 27: 397. 1856; Mart. Fl. Bras. 14, pt. 1: 413. 1857),

the genus Myrtus was impossibly heterogeneous by modern stand-

ards. The Chilean species, approximately one-third of the total,

have all been transferred to other genera. About half the remainder

were South American species that have recently been transferred

by Burret to Psidium (Notizbl. Bot. Gart. Berlin 15: 483-485. 1941).

McVAUGH: TROPICAL AMERICAN MYRTACEAE

513

FIG. 14. Floral morphology in Psidium. A, Bud of P. mouririoides, Gentle

type (X 5). B, Axillary flower of P. salutare, showing open lobed calyx,

Haught 2602 (X 2.5). C, Diagrammatic section of seed of P. guineense, showing

curved embryo unshaded in center (X 10). D, Bud and old flower of P. anglo-

hondurense, Bartlett 13060 (X 5). E, Lateral and terminal views of bud and old

flower of P. biloculare, Gentle 3382 ( X 5).

According to Burret, Myrtus is represented in the American flora

by about 14 species only, these in Florida and the Greater Antilles.

The type species of Myrtus, M. communis of Europe, has a 3- or

sometimes 2-locular ovary, and most of the known West Indian spe-

cies seem to have the ovary 2- or sometimes 3-locular. The generic

limits of Myrtus are by no means agreed upon even yet, but the

number of locules in the ovary, in the genus approximately as under-

stood by Burret, appears usually to be 2 or 3, as against 3 or more

in Psidium. Species having bi- or trilocular ovary, therefore, cannot

514 FIELDIANA: BOTANY, VOLUME 29

be assigned forthrightly to one genus or the other on this character

alone. Bentham reached essentially this conclusion nearly a cen-

tury ago (Jour. Linn. Soc. Bot. 10: 152. 1868).

The generic limits of Myrtus being somewhat uncertain, at least

in the American flora, it seems more profitable to concentrate upon

Psidium, to determine what limits, if any, can be assigned to it,

and to set forth criteria by which it may be distinguished from

Myrtus or Myrtus-\ike plants. There are apparently several diverse

evolutionary lines represented in the large American genus Psidium,

and morphological variation among the species is extreme.

Many species of Psidium known to the earlier authors had con-

spicuously large flowers, the calyx closed or nearly closed in bud,

and the ovary 3- to 5-locular. The best example of this is the com-

mon guava, P. guajava, widespread in tropical regions through cul-

tivation from the earliest times following the discovery of America

by Europeans. The earliest concepts of the genus were based upon

this species and a few others like it. Unfortunately none of the

above characteristics is consistent in the genus, and none is distinc-

tive. The range in size of flowers in Psidium is about the same as it

is in other large genera (e.g., Eugenia, Syzygium, Calyptranthes) ; the

calyx is perhaps more often open than closed; and a bilocular ovary

occurs in a considerable number of species.

Too much taxonomic importance has apparently been attributed

to the most "unusual" feature of Psidium flowers, the completely

closed calyx which opens at anthesis by irregular longitudinal splits.

This prejudice seems to stem in part from the original concepts of

Linnaeus and his contemporaries, but chiefly from the work of Berg,

who (perhaps inadvertently) emphasized this one morphological fea-

ture (Linnaea 27: 347 et seq. 1856; Mart. Fl. Bras. 14, pt. 1: 382 et

seq. 1857). Although more than two-thirds of the species of Psidium

known to Berg (58 out of 84 in his treatment of 1856) were assigned

by him to infrageneric groups having the calyx open and normally

toothed or lobed in bud, he nevertheless in his keys to genera omitted

all mention of toothed or lobed calyces, saying without any qualifi-

cation: "Calyx ante anthesin omnino clausus v. apice pertusus."

[italics mine]. By thus highlighting unduly the characteristic of the

closed calyx, Berg seems to have biased the interpretations of later

authors. Bentham, for example, 20 years after the work of Berg

(Gen. PI. 1: 694, 716, 720. 1865), groups Psidium in his key with

those genera having the "calycis limbus in alabastro clausus v. sub-

imbricato-lobatus" ; after this concession to the species with open

McVAUGH: TROPICAL AMERICAN MYRTACEAE 515

calyx, however, he describes Calycorectes and Marlierea as having

"Calyx Psidii." His idea of the "calyx of a Psidium" may be ascer-

tained from the descriptions of the individual genera: Of Calycorectes

he says later "Calycis limbus in alabastro clausus"; of Marlierea he

says "Calycis tubus ... in alabastro clausus v. apice brevissime

apertus." It may be inferred that Bentham thought of Psidium as

a genus having the calyx typically closed in bud, even though he

included in the genus some species with lobed or toothed calyx.

A similar bias seems apparent in the work of Niedenzu (Natiirl.

Pflanzenfam. 3, pt. 7: 64, 67. 1893), who describes the calyx of

Psidium as gamosepalous ["verwachsenblattrig"], and who separates

Psidium from Myrtus in the key because the calyx-lobes of the former

are united ["verwachsen"] in the bud, as against "already free in the

bud." Like Berg, however, Niedenzu stressed the feature of the

closed calyx, and then assigned a large group of species to an intra-

generic group (in this instance a new section, Apertiflora), charac-

terized by having an open calyx. It seems clear that this one feature

of the calyx is not to be thought of as having diagnostic significance

in the determination of generic limits.

A second characteristic of the calyx (or hypanthium), however,

provides the single most dependable feature by means of which spe-

cies of Psidium can be recognized. This feature was known to Berg,

and mentioned by Bentham and by Niedenzu, but has perhaps been

insufficiently emphasized. Berg's descriptions, not his keys, provide

the following distinctions between Psidium and Myrtus:

Hypanthium (calycis tubus auct.) supra germen productum . . ., in calycem

abiens. . . . Calyx ante anthesin indivisus claususque v. apice pertusus v.

4-5-dentatus, demum longitudinaliter usque ad verticem germinis 2-5-par-

titus, ex margine hypanthii libero oriundus Psidium.

Hypanthium (calycis tubus auct.) supra germen . . . inferum vix productum;

sepala 4-5, ex margine angusto, libero hypanthii oriunda, in alabastro globo

petalorum incumbentia, . . . sinu acuto separata, serius non profundius

partita Myrtus.

The important point here is not the difference between a closed

calyx and a calyx with 4-5 sepals, as has often been assumed, but

(in Psidium) the initial prolongation of the hypanthium or calyx-

tube beyond the ovary, and the subsequent irregular longitudinal

splitting of these tissues down to or beyond the staminal ring and

often to the very summit of the ovary. In Myrtus, on the other

hand, the calyx-lobes are separate from the beginning, develop at

about the level of the summit of the ovary, and fail to separate

farther at anthesis, remaining contiguous so that the entire calyx

516 FIELDIANA: BOTANY, VOLUME 29

may persist as a unit on the fruit. Berg's words as quoted above

may now be paraphrased:

Hypanthium produced beyond the summit of the ovary, the stamens borne on the

inner wall of the tube; calyx (whether closed or open at tip in bud) appearing

tubular at base, in anthesis splitting longitudinally to the summit of the ovary

into 2-5 somewhat irregular divisions, the individual lobes usually rather

widely separated and often deciduous after anthesis Psidium.

Hypanthium hardly if at all produced beyond the summit of the ovary, the sta-

mens borne on a perigynous disk; calyx-lobes 4-5, distinct, fully formed in

the bud, separated by acute sinuses, but not separating farther at anthesis

or in fruit, the calyx not splitting between the lobes and often persisting as

a unit at the summit of the fruit Myrtus.

The validity of the above distinction seems to have been ac-

cepted by Bentham, who in the Genera Plantarum used it in com-

bination with other calyx-characters to separate the principal groups

of genera in the Myrteae. The same distinction was apparently

familiar to Urban, who, after Berg, had perhaps the best knowledge

of the American Myrtaceae as a whole. In describing Psidium

leiophloeum, for example, Urban wrote (Symb. Antill. 9: 460. 1928):

"Ad seriem earum specierum pertinens, quae calyce ab initio aperto

et lobis liberis sub anthesi dirumpendo paullo profundius separatis

gaudent ideoque a Myrto vix v. ne vix quidem discernendae sunt."

In another place (Symb. Antill. 9: 78. 1923) Urban stressed the

importance of the same feature: "Myrtus et Psidium . . . inter sese

arete affinia nee nisi aut calycis lobis ab initio liberis aut sub anthesi

(supra basin saltern) diruptis in speciebus nonnullis antillanis dis-

cernenda sunt."

As Urban points out in the same place, species of Eugenia may

be confused with Myrtus or with Psidium in the absence of seeds.

The last two genera are quite different from Eugenia in the structure

of the testa and that of the embryo, but in vegetative characters

and those of the flowers, including those of the placentation, there

may be no discernible differences. This brings us to the matter

of the importance, if any, of the number of locules in the ovary of

Psidium and related genera.

The three principal groups of American Myrtaceae differ among

themselves in the constancy of the number of locules. In two of

the subtribes, the Myrciinae and the Eugeniinae, the number of

locules is almost always 2, occasionally 3 or 4. In the Pimentinae,

to which belong Psidium and Myrtus, the number is far more variable

and often more than 2. Some genera, according to Berg, are in-

deed always bilocular (e.g., Blepharocalyx) . In others (e.g., Caly-

McVAUGH: TROPICAL AMERICAN MYRTACEAE 517

colpus, Abbevillea, Campomanesia, Acrandra, Britoa and various small

genera), the number of locules is said to be always 4 or more, some-

times as many as 11. In the two large genera having bony seeds

and an uncinate-curved embryo, namely Psidium and Myrtus, the

number of locules is said to vary from 2 to 7, and 2 to 4, respectively.

In most of the species of Psidium known to Berg the ovary was

said to be 3- or 4-locular (about 50 species); about twelve species

were described as sometimes or always with more than 4 locules;

and eight species as having but 2, 2-3, or 2-4 locules. Berg appar-

ently knew but four species of Psidium having consistently a bi-

locular ovary (cf. Linnaea 27: 359, 376. 1856; Mart. Fl. Bras. 14,

pt. 1 : 604, 605. 1857) . He seems not to have regarded these species

as in any way atypical. Among the species of Psidium described

more recently by Urban are several with bilocular ovary. The

number of locules in Psidium may thus be stated as usually 3 or

4, occasionally as few as 2 or as many as 7.

Perhaps correlated with the number of locules in the ovary

(i.e., with respect to 2- and 3-locular ovaries) is the type of placenta-

tion. Altogether too little is known of the details of placentation

in the American Myrtaceae, but in analyzing nearly 600 Australian

species Bentham found that the "best generic characters" were

afforded by the modifications of the ovules and their placentation,

"provided too much weight be not attached to the precise form of

the placenta." (Jour. Linn. Soc. Bot. 10: 119. 1868.)

Berg seems to have attached little importance to the details

of placentation in the subtribes Myrciinae and Eugeniinae, in which

the ovary is generally bilocular. Usually he did not mention plac-

entation in his descriptions of genera. Of Eugenia he said merely:

"ovula sporophoris centralibus aflixa." He was impressed, however,

by what he considered to be the parietal origin of the ovule-bearing

structures ["sporophora"] among the Pimentinae. In Psidium he

described these as "parietal, 2-7, reaching the axis, there connate

or discrete and bilamellate, the lamellae reflexed, more or less pro-

duced [i.e., radially into the locule, away from the axis], ovule-

bearing on the inner surface ["intus"], connate in pairs, simulating

central bilamellate sporophores."

These structures were illustrated by Berg, for species having

respectively 3-, 4- and 5-locular ovary (Mart. Fl. Bras. 14, pt. 1: pis.

41, 4.1 a, 4-2, 1857). Similar structures, apparently derived in the way

described by Berg, can be seen in various species with multilocular

518 FIELDIANA: BOTANY, VOLUME 29

ovary. Even in some species with bilocular ovary (e.g., Psidium

pedicellatum; see Flora of Peru, pt. 4: 799) the ovules occur in a double

incurved row about the margins of an apparently peltate placenta.

In other genera of the Pimentinae (e.g., Blepharocalyx) having

bilocular ovary, Berg made no attempt to demonstrate a parietal

origin for the sporophores. In the description of Blepharocalyx in

the Flora Brasiliensis he said merely "ovula sporophoro centrali

affixa" (p. 420). Berg's concept of the placentation in Psidium

(and in the Pimentinae as a whole) seems to have been based pri-

marily upon study of species with 3-locular or multilocular ovary.

As pointed out above, he seems to have known few species of Psidium

with bilocular ovary, and these from few specimens. In his descrip-

tions of P. moritzianum and P. cordatum (both described as bilocu-

lar), he described the sporophores as "bilamellate," but did not

elaborate upon this.

The ovules of Myrtus were described by Berg as "saepissime

plurima, sporophoris bilamellatis, spurie centralibus affixa" [italics

mine}. This seems to me to point up the question of the difference

between the type of placentation in Psidium and in Myrtus, and

between 2- and 3-locular ovaries. I suppose Berg meant that in

Myrtus (especially in the 3-locular types) the sporophores were actu-

ally parietal in origin, as in Psidium, but seemed to arise directly

from the central axis. Nowhere did he specifically say, however,

that the centrally attached placenta in the bilocular ovary was

parietal in origin. We still do not have the information that will

permit us to speculate upon the derivation of the placentae when

these structures, even if bilamellate, seem to arise directly from

the central dissepiment of a bilocular ovary. Comparative develop-

mental studies are much needed. In the absence of such studies,

variation in placental structure must be interpreted with caution.

No correlation has been demonstrated between characters of the

placenta and other characters in the Psidium-Myrtus complex, ex-

cept that in species having a bilocular ovary the ovules are often

attached in a single close sessile group near the middle of the central

dissepiment, exactly as in many species of Eugenia.

In summary, the genus Psidium may be recognized by its pi-

mentoid C -shaped or uncinate embryo; hard or bony seeds; calyx

(whether originally open or closed) splitting between the lobes at

anthesis; ovary 3-4- (rarely 2-7-) locular; placentae usually bila-

mellate but in bilocular ovaries the ovules often in a single cluster.

McVAUGH: TROPICAL AMERICAN MYRTACEAE 519

ADDITIONS TO THE GENUS PSIDIUM

The above review of the generic character of Psidium was occa-

sioned by the discovery in Central America, in the region bordering

the Gulf of Honduras, of a group of five species that are surely

members of the Pimentinae, although all were originally described

as species of Eugenia. These five species are vegetatively similar,

with moderately large, coriaceous, glabrous or glabrate subsessile

or short-petiolate leaves; the flowers are sessile or short-pedicellate,

borne in small axillary clusters at leafless (or sometimes leafy) nodes

on old wood; the calyx varies from completely closed in the bud

to somewhat prominently lobed, but the lobes in all species sep-

arating at anthesis, the base of the calyx splitting down to the

ovary, which is bilocular (or in one species trilocular and in another

unilocular); the ovules are attached in a subcapitate group to the

central dissepiment (or a bilamellate placenta in the trilocular ovary) ;

the seeds (known in three species only) are reniform, compressed,

osseous, the embryo curved into a nearly complete ring.

These plants, evidently very closely interrelated, are quite un-

like any other species known from Central America, superficially

suggesting in appearance some species of Psidium from the West

Indies. They combine in a unique way several features that per-

haps suggest evolutionary tendencies in the Pimentinae. The pro-

duction of flowers at leafless nodes may be regarded as a character

more specialized than the production of leafy or bracteate inflor-

escences on young shoots. In these five species, otherwise so similar,

may be seen an almost completely closed calyx (in anglohondurense) ;

the calyx-lobes united their whole length but not closing the bud

(mouririoides) ; or the calyx-lobes about half as long as the tube

(biloculare) . The ovary is trilocular (in anglohondurense), or bilocular

(in the other species), or sometimes unilocular (in mouririoides).

It may be supposed that the bilocular condition is more specialized

than the trilocular; the unilocular condition, if normally developed

in this species, is presumably even more specialized. The unilocular

condition is well known in Old World Myrtaceae, perhaps having

resulted from a reduction series (cf. Bentham, Jour. Linn. Soc.

Bot. 10: 119. 1868). It is of interest to note that the placentae

are bilamellate in anglohondurense, associated with a trilocular ovary;

in the other species the closely grouped ovules are associated with

a bilocular ovary. Any generic distinction based on these char-

acters, among species that are otherwise so strikingly alike, would

seem to be specious.

520 FIELDIANA: BOTANY, VOLUME 29

The above five Central American species, which are apparently

properly transferred to Psidium, may be separated as follows:

Branchlets acutely 4-angled, the angles becoming winglike in the distal parts of

the internodes and terminating in the stipular positions at the sides of the

petioles; flowers sessile or nearly so, the pedicels 1 mm. long or less; ovary

bilocular.

Plants, including the inflorescence, glabrous or essentially so.

Leaves subsessile, rounded to acute at base, 3-7 (-12) cm. long, mostly 1.6-

2.3 times as long as wide; style 6-8 mm. long; stamens 60-75.

P. biloculare.

Leaves on petioles 3-5 mm. long, rounded or subauriculate at base, 12-20 cm.

long, 3-3.5 times as long as wide; style 9-10 mm. long; stamens about 200.

P. musarum.

Young leaves and branchlets, and the inflorescence, coarsely silky- or velutinous-

tomentose; leaves subsessile, abruptly rounded or subcordate at base, 6-

10 cm. long, 2-3 times as long as wide; style 9-10 mm. long; stamens about

125 P. apodanthum.

Branchlets terete or compressed, the nodes enlarged; plants glabrous; flowers

pedicellate; calyx closed in the bud or with a small apical opening; style 5.5-

7 mm. long.

Petioles 3-6 mm. long; pedicels 10-18 mm. long; buds 5-6 mm. long, apiculate,

closed except a minute subapical pore; locules of the ovary 3 (-2), the

ovules about 20 (-45) in each locule P. anglohondurense.

Petioles 1-3 mm. long; pedicels 3-4 mm. long; buds 3-4.5 mm. long, subglobose

or pyriform, the calyx-lobes united their whole length but leaving an apical

opening 1.5-2 mm. across; locules 2 (-1), the ovules about 50 in each locule.

P. mouririoides.

Psidium apodanthum (Standl.) McVaugh, comb. nov. Eu-

genia apodantha Standl. Field Mus. Bot. 17: 380. 1938.

HONDURAS: Comayagua: Known only from the type collection,

near Siguatepeque, Yuncker et al. 5766 (F, type; MICH).

Psidium biloculare McVaugh, nom. nov. Eugenia gentlei Lun-

dell, Carneg. Inst. Wash. Publ. 478: 216. 1937. Not Psidium gentlei

Lundell, 1943.

GUATELAMA: Izabal: Rio Chacon, Harry Johnson 1191 (F, US).

BRITISH HONDURAS: Belize Dist., Gracie Rock, Sibun River,

P. H. Gentle 1684 (MICH, type). Known also from several other

localities: Stann Creek Dist., W. A. Schipp S-141 (F), Gentle 1887,

3382, 8552; Belize Dist., Gentle 9479, O'Neill 8761; Toledo Dist.,

Gentle 4062B, 4157, 4200, 4719. A specimen from Toledo Dist., near

Condemned Branch Pine Ridge, Gentle 5358 (LL), is apparently

this species but the calycine lobes are pubescent on the inner sur-

McVAUGH: TROPICAL AMERICAN MYRTACEAE 521

face and the styles are 8-9 mm. long, almost as in P. apodanthum.

An imperfect specimen, Montana del Mico, Dept Izabal, Guatemala,

Steyermark 38544, is similar (F).

Psidium anglohondurense (Lundell) McVaugh, nom. nov.

Eugenia schippii Standl. Field Mus. Bot. 11 : 137. 1932. Not Psidium

schippii Standl., 1931. Eugenia anglohondurensis [sphalm. anglohon-

duransis] Lundell, Wrightia 2: 123. 1961.

BRITISH HONDURAS: Stann Creek Dist.: Seine Bight, W. A.

Schipp 669 (F, type of E. schippii; MICH); Commerce Bight,

P. H. Gentle 8354 (LL, isotype of E. anglohondurensis). — El Cayo

Dist.: Little Mountain Pine Ridge, H. H. Bartlett 13060 (F, MICH).

—Toledo Dist.: Monkey River, Jenkins Creek, Gentle 4063 (F,

LL, MICH).

The type of Eugenia schippii was a specimen with half-grown

fruit; Gentle 4063 and 8354 include mature fruit, and the latter

has flowers as well; Bartlett's collection has flowers only. It is

noteworthy that Gentle gives the common name "Wild Guava"

for both his collections.

Psidium mouririoides (Lundell) McVaugh, comb. nov. Eu-

genia mouririoides Lundell, Am. Midi. Nat. 29: 480. 1943.

BRITISH HONDURAS: Toledo Dist.: Monkey River, Jenkins Creek,

Gentle 4142 (MICH, type). The one known specimen of this species

is in some ways intermediate between P. biloculare and P. anglo-

hondurense; e.g., the pedicels are of intermediate length and the

calyx is in some ways intermediate (fig. 14). In view of the fact

that all three species were collected by Gentle at the same local-

ity within a period of about a month, they presumably grow to-

gether or in associated habitats, and it may be that P. mouririoides

is a hybrid.

Psidium musarum (Standl. & Steyerm.) McVaugh, comb. nov.

Eugenia musarum Standl. & Steyerm. Field Mus. Bot. 22: 358. 1940.

GUATEMALA: Izabal: Rio Juyama, about 15 miles southwest of

Bananera, Steyermark 39165 (F, type). Known also from Alta

Verapaz.

The only known specimens of P. musarum are sterile, or bear

flowers just past anthesis. Confirmation of the generic status of

522 FIELDIANA: BOTANY, VOLUME 29

this species must await the discovery of fruiting specimens, but I

believe a transfer to Psidium is justified on the basis of general

similarity to the three species in which fruits and seeds are known.

SPECIES DESCRIBED BY STANDLEY FROM BRITISH HONDURAS

Four species of Psidium described by Standley in 1931, from a

single locality near the coast of British Honduras, were all based

on fruiting specimens. Their author, in differentiating them, empha-

sized their differences in leaf-form, but did not compare them with

P. guineense, to which they all bear more than a casual resemblance.

Perhaps the most distinctive is the following, which seems to be

known only from the type collection, Schipp 596 (F!).

Psidium chrysobalanoides Standl. Field Mus. Bot. 8: 319.

1931.

Plant nearly glabrous, the leaves and the strongly 4-angled branchlets merely

sparingly pale-strigose; blades mostly obovate, abruptly deltoid-acuminate with

acute tip, mostly cuneate at base, 2.5-5 cm. wide, 4-7.5 cm. long, 1.25-2 times as

long as wide, on petioles 2 mm. long; midvein convex above; lateral veins 4-6 on

each side, strongly ascending, forming with the small veinlets a prominent ele-

vated reticulum on both surfaces; peduncles 1- or 3-flowered, 1.5-2 cm. long;

immature fruit 1 cm. long, probably pyriform.

In an area that has been rather well collected, it seems odd

that this plant has not been found again. It is conceivable that

it represents some complex hybrid involving principally P. guineense

and perhaps also P. guajava and even P. salutare. In the absence

of better material, any disposition of it can be little better than

a guess.

Psidium schippii Standl. Field Mus. Bot. 8: 319. 1931.

This species, also apparently known from the type collection

only, seems to have been based on a vigorous shoot of P. guineense

Sw. The leaves in the type (Schipp 595, F!) are obovate and

somewhat elongated toward the tip of the shoot. A 3-flowered

and apparently normal inflorescence near the base of the specimen

seems typical of P. guineense.

Psidium rotundifolium Standl. Field Mus. Bot. 8: 318. 1931.

Standley noted in the protologue of P. rotundifolium that the

distinctive leaves were "usually large, rounded and nearly as broad

as long." In his own later travels in Central America from 1940

McVAUGH: TROPICAL AMERICAN MYRTACEAE 523

to 1951, Standley collected (or annotated for others) approximately

30 specimens of what he took to be P. rotundifolium. Most of these

were from Honduras; others came from Guatemala, Nicaragua, or

Costa Rica. All tended to have large stiff leaves (Standley noted

in one instance that the leaves were used as spoons). The blades

are very broadly elliptic or obovate, up to 10-12 cm. wide and 12-16

cm. long, resembling superficially those of P. guineense but the

red pubescence not obscuring the lower surface and the small veins

forming an elaborate raised reticulum. Specimens with the smallest

leaves look much like guineense, and the young growing tips when

present are indistinguishable from those of guineense. Most sig-

nificantly, to my mind, all the specimens that Standley called ro-

tundifolium appear to be vigorous shoots, i.e., shoots bearing large

fast-growing leaves; of the 35 sheets so named by Standley, 24

are sterile terminal shoots; 3 of the others seem to be ordinary

flowering specimens of P. guineense; the other 8, including the type

of P. rotundifolium (Schipp S85, F!), bear nothing that may be

considered a normal inflorescence, but only scattered axillary pe-

duncles. In the absence of distinctive flowering and fruiting speci-

mens P. rotundifolium can hardly be recognized as a valid species;

the evidence suggests that it is a vegetative form of P. guineense.

Psidium Xhypoglaucum Standl. Field Mus. Bot. 8: 320, pro.

spec. 1931. P. guineense Sw. X P. guajava L. (?).

Most individuals of this so-called species bear a strong super-

ficial resemblance to P. guineense, seeming to differ from it chiefly

in the gray pubescence of the lower leaf-surface, and in the thinly

pubescent, angled branchlets. Each of these latter features can

be matched precisely in some individual specimens of P. guajava.

The usual habitat of P. Xhypoglaucum is in disturbed places

where other species of Psidium are growing. At one locality in

Honduras, for example (Las Mesas, Dept. Morazan, at an elevation

of about 900 meters), P. Xhypoglaucum was collected with P.

guineense (Williams & Molina 1007 '4 includes both) at a spot where

P. guajava also grew, and where the collectors obtained the so-

called P. rotundifolium, P. salutare, and a pubescent form of P.

salutare that bears some resemblance to P. guineense.

The hybrid nature of P. Xhypoglaucum is suggested by its

usual habitat and by its morphological intermediacy between two

species that often grow together.

524 FIELDIANA: BOTANY, VOLUME 29

Specimens examined:

BRITISH HONDURAS: All Pines, W. A. Schipp S99 (F, type).

GUATEMALA: Jutiapa: Cerro Colorado just west of Jutiapa,

Standley 76173, 76190 (both F).

HONDURAS: Morazan: Standley 846, 12148, 22187, 22198, 22251,

J. Valeria R. 520 (all F).

MISCELLANEOUS NOTES ON PSIDIUM

Psidiumxoerstedeanum Berg, Linnaea 27: 360, pro. spec.

1856. P. chiapasense Lundell, Wrightia 2: 204. 1961.

The type locality of P. oerstedeanum was given as "In provincia

Guanacarte [sic], floret Martio (Dr. Oersted), nee non in monte

Rincon in Guatemala (Friedrichsthal, coll. no. 1226)." The Fried-

richsthal specimen, according to Standley (MSS.), probably came

from Volcan Rincon de la Vieja, Guanacaste, Costa Rica, not from

Guatemala.

Central American plants referred to P. oerstedeanum by Standley

and others seem to differ from Psidium salutare (HBK.) Berg chiefly

in having slightly broader and sometimes obovate or elliptic leaves,

strongly pubescent stems and somewhat pubescent leaves, canescent

buds and strongly pubescent peduncles, and somewhat more prom-

inent veins in the mature leaves. Presumably the material upon

which Berg based P. oerstedeanum was all of this white-pubescent

plant. The Oersted specimen cited by Berg ("in hb. Oersted, no.

16"), which I have seen through the kindness of the authorities

at Copenhagen, is of this description and so apparently is the speci-

men of Friedrichsthal 1226 at Vienna (Field Mus. Neg. 31430).

I find no significant differences between the type of P. chiapasense

Lundell (Matuda 5769, an isotype, kindly lent by Dr. Lundell)

and the syntype specimens of P. oerstedeanum.

Very few specimens in herbaria resemble the original ones of

Psidium oerstedeanum; the ones that do have the open calyx of

oerstedeanum and salutare, and the canescent herbage and buds of

oerstedeanum, are otherwise so diverse they can hardly represent

an independent species. Two Mexican specimens from near Comitan,

Chiapas (Matuda 15769, Carlson 1892) suggest by their appearance

small-leaved gray-pubescent forms of P. guineense, a supposition

strengthened by the reddish color of the pubescence of immature

shoots. Lundell noted that the peduncles of his P. chiapasense

McVAUGH: TROPICAL AMERICAN MYRTACEAE 525

were sometimes 2-flowered, a feature that also suggests some con-

nection with P. guineense.

A clue to the origin of the so-called Psidium oerstedeanum seems

to lie in its frequent close association with P. salutare, a species

inhabiting lowland savannahs from Mexico to the Guianas. Ordi-

narily P. salutare is glabrous, with mostly lanceolate or lance-ovate

narrowly pointed leaves. In various localities in Central America

and southern Mexico, however, the plants are somewhat pubescent,

but otherwise seemingly unchanged. At some localities, moreover,

the narrow-leaved glabrous plants and narrow-leaved pubescent

plants grow intimately associated with canescent plants that have

even broader leaves (i.e. the "oerstedeanum" type). The type of

Psidium gentlei Lundell, for example (British Honduras, Monkey

River, Jenkins Creek, Gentle 4062, MICH), is a narrow-leaved plant

with glabrous leaves and with but scanty pubescence on the branches;

I should refer it without hesitation to P. salutare. Two other collec-

tions from the same locality and taken apparently at the same

time, Gentle 4062C and 4062D, have quite a different aspect, viz.

that of small-leaved specimens of P. guineense; they are pubescent

and broad-leaved and would be referred to P. oerstedeanum if that

species were to be recognized.

Two collections from Honduras (Standley 21236 and Williams

& Molina 10484) are similarly mixed; each collection includes one

branchlet with narrow glabrous leaves and glabrous twigs, and one

or more with broader, pubescent leaves and pubescent inflorescence.

These are from a disturbed pastured area to which allusion has

already been made above (under P. Xhypoglaucum) , where natural

hybridization is suspected.

It seems likely that the broad-leaved, somewhat variable but

always densely pubescent or canescent plants called Psidium oer-

stedeanum have resulted from hybridization between P. salutare and

some other species, perhaps P. guineense, which is common and

weedy and variable throughout Central America.

Psidium sartorianum (Berg) Ndzu. in Engl. & Prantl, Natiirl.

Pflanzenfam. 3, Abt. 7: 69. 1893. Mitranthes sartoriana Berg, Lin-

naea 29: 248. 1858.

At the time of the publication of the genus Mitranthes it was

assigned by Berg to the subtribe Eugenioideae. One wonders why

such an astute student of the Myrtaceae as Berg did not suspect

some relationship between Mitranthes and Psidium, in view of his

526 FIELDIANA: BOTANY, VOLUME 29

descriptions of the several species of Mitranthes as having a closed,

circumscissile and operculate calyx, a multiovulate 3-locular ovary,

and the "ovulis sporophoris bilamellatis afRxis." Apparently, how-

ever, he never saw fruits or seeds of any of the original eight species

of Mitranthes, and thus could never have been entirely sure of the

systematic position of the genus (see Linnaea 29: 248-249. 1858,

for the complete list of these species, including M. sartoriana).

Most recent authors have agreed that Mitranthes belongs taxo-

nomically with Psidium rather than with Eugenia, but there has

been no general agreement as to the exact generic status of the

group. Niedenzu (in the Natiirlichen Pflanzenfamilien) proposed

a revised classification of Psidium and related genera, erecting a

new infra-generic group, Psidium sect. Calyptropsidium (Berg)

Ndzu., to include the one species referred by Berg to the genus

Calyptropsidium, and five species, including P. sartorianum, that

Berg had referred to Mitranthes.

Urban (Bot. Jahrb. 19: 571. 1895) re-established Calyptropsidium

as a genus, and transferred to it Mitranthes sartoriana. Standley,

in the Trees and Shrubs of Mexico (Contr. U. S. Nat. Herb. 23:

1035. 1924), followed Niedenzu in treating this species as a Psidium,

and in this he has been followed by most recent American workers.

Burret, however, in 1941 (Notizbl. Bot. Gart. Berlin 15: 486),

created a new genus, Mitropsidium, based primarily on Psidium

sect. Calyptropsidium (Berg) Ndzu. but including three additional

species with operculate calyx, and excluding the type species of

Calyptropsidium Berg, C. friedrichsthalianum, on the grounds that

in that species the calyx opens irregularly and not by a true calyptra.

Burret argued that Mitropsidium constituted a taxon closely re-

lated to Psidium, the two genera perhaps not entirely monophyletic,

and separated somewhat artificially by the one character of the

calyx; he used for support of his argument the universally recog-

nized genera Calyptranthes and Marlierea, separated artificially by

the same character.

The most recent student of this particular group, Miss G. J. H.

Amshoff, pointed out that the character of the circumscissile calyx

"has to be used with circumspection as e.g. in Psidium (Calyptro-

psidium) Friedrichsthalianum (Berg) Ndz. the calyx is sometimes

calyptrate and sometimes vertically dehiscent" (Act. Bot. Ne"erl. 5:

277. 1956). In the Flora of Panama (Ann. Missouri Bot. Gard. 45:

198. 1958) Miss Amshoff treated Mitranthes-Calyptropsidium-Mitro-

psidium sartorianum as a species of Psidium, with no comment as

McVAUGH: TROPICAL AMERICAN MYRTACEAE 527

to its generic position but noting in her description that the calyx

usually splits irregularly, "with 1 large more or less lid-like deciduous

segment and 2-3 other minute persistent segments," and stating

specifically that a neatly circumscissile calyx is exceptional.

It seems probable that too much emphasis has been placed upon

this one feature of the calyx in determining generic lines. It is

at best a variable feature, even in individual species; furthermore

it has not been shown that the species having in common the cir-

cumscissile calyx have also other shared attributes to set them

apart from Psidium. There seem to be no sharp distinctions in

Psidium between groups of species with closed buds and those

with open buds. Two Amazonian species, for example, P. guianensis

Pers. and P. densicomum DC., are essentially indistinguishable ex-

cept that the former has a closed bud and the latter an open, shal-

lowly lobed calyx.

The specific limits of Psidium sartorianum are somewhat in

doubt also. Several so-called species have been distinguished from

it; most of these have been based partly upon features of the calyx,

but in part also on leaf-characteristics. The entire species-complex

should be reviewed. The leaves are usually elliptic and short- or

obtusely pointed, but in some parts of the range, particularly in

Mexico from Jalisco to Guerrero, in the Yucatan Peninsula, and

in Honduras, many or most individuals have long-acuminate or at

least long-acute leaves, often of a lanceolate type. This local popu-

lation may be appropriately designated as follows:

Psidium sartorianum (Berg) Ndzu., var. yucatanense

McVaugh, var. nov.

A var. sartoriano foliis lanceolatis vel ovato-lanceolatis longiuscule acutis acu-

minatisve differt.

This variety is typified by a specimen from British Honduras,

P. H. Gentle 9 (MICH), which is also the type of Psidium yucatanense

Lundell (Contr. Univ. Mich. Herb. 7: 35. 1942). I cannot dis-

tinguish specimens of P. yucatanense from Cuban specimens de-

scribed as Calycorectes protractus Griseb., the type a specimen col-

lected by Charles Wright (isotype, US!) (Griseb. Cat. PI. Cub. 284.

1866). If these Cuban and Central American plants together con-

stitute a distinct species, it will be necessary to provide a new specific

epithet in the genus Psidium, the name Calycorectes protractus having

76 years' priority over P. yucatanense. If, as in the present paper,

this taxon is regarded as a variety of P. sartorianum, the name var.

528 FIELDIANA: BOTANY, VOLUME 29

yucatanense may be used. It should be noted that in accordance

with the International Code of Botanical Nomenclature (Art. 60, 72)

this is to be treated as a new varietal name, not a new combination

based on P. yucatanense Lundell, which is regarded as a taxonomic

(not nomenclatural) synonym of Calycorectes protractus and therefore

undesirable as a basis for a name in a new status.

Dr. Lundell contrasted his new species with P. sartorianum, em-

phasizing especially the point that in P. yucatanense the basal part

of the calyx "above the ovary splits into five irregular lobes at

anthesis." I do not find this feature of diagnostic value, for in

most specimens of undoubted P. sartorianum the calyx splits in

a similar fashion at or after anthesis. There is no doubt, however,

that all specimens from the Yucatan Peninsula have a distinctive

appearance because of the slender-pointed lanceolate or lance-ovate

leaves. The following belong here:

CUBA: Santa Clara: Dist. Cienfuegos, Cieneguita, Combs 201 (F).

YUCATAN: Chiche"n Itza, Steere 1293; Tizimin, Swollen 2502;

Yakdzonoot, Lundell 7494 (all MICH).

BRITISH HONDURAS: Belize Dist., Belize-Sibun Road, P. H.

Gentle 9 (F; MICH, type, and type of P. yucatanense Lundell).

GUATEMALA: Pete"n: Tikal National Park, E. Contreras 1340

(LL, MICH).

This local population of the Yucatan Peninsula hardly seems

specifically different from P. sartorianum of the Mexican mainland.

A paratype of P. yucatanense, for example (Steere 1293}, is a good

match for a Karwinsky specimen identified by Berg as Mitranthes

sartoriana (Field Mus. Neg. 19745). In western Mexico, moreover,

where P. sartorianum is abundant, individual plants of the "yuca-

tanense" type, i.e., with acute and more or less lanceolate leaves,

are frequent but do not seem to differ otherwise from the plants

with more obtuse elliptic leaves. Some specimens are cited below:

MEXICO: Jalisco: Near Bahia de Tenacatita, McVaugh 20965

(MICH) ; 4 miles north of Bahia Navidad, McVaugh 20832 (MICH).

—Guerrero: Acapulco, Palmer 357 (F), MacDaniels 149 (F).

Another plant that seems to have close affinities with P. sar-

torianum is P. molinae Amsh. (Act. Bot. Ne"erl. 5: 277. 1956) (Hon-

duras, the type from Dept. Morazan, Standley 21255, F!). This

was contrasted by its author with P. sartorianum, with the state-

ment that the latter species is at once distinguishable "from our

new one by its calyptrate calyx and shortly acuminate leaves"

McVAUGH: TROPICAL AMERICAN MYRTACEAE 529

(P. molinae having "alabastra . . . apice minute aperta, brevissime

4-loba. Calyx ... in flore adulto irregulariter 4-fissus," and the

leaves obtuse). The type of P. molinae bears a few buds that open

in the way described by Amshoff, i.e., the calyx is 4-lobed, open at

the tip, with thick incurved-deltoid lobes 1 mm. long. One other

specimen, Williams & Molina 14143 (F), also in bud, is similar

to the type. The other specimens cited by Amshoff are sterile

or in the fruiting condition, and apparently were selected on the

basis of leaf-shape only. The leaves are indeed somewhat obtuse,

but not more so than in about half the available specimens through-

out the range of P. sartorianum. Miss Amshoff points out that

a specimen from Oaxaca (Makrinius 609} has "the flowers of P.

molinae and the leaves of P. sartorianum."

A plant similar to P. molinae is P. microphyllum Britton (Sci.

Surv. P. R. & Virgin Is. 6: 555. 1930), based on a cultivated plant

from the Agricultural Experiment Station at Mayaguez, Puerto

Rico (J. B. McClelland, July 1930, NY, type!). The leaves are

relatively obtuse, and the buds apparently are never fully closed;

there is an apical pore about 0.3 mm. in diameter, and minute lobe-

like hairy appendages surround the pore; the calyx in an thesis splits

irregularly, one lobe often much larger than the others but not

forming a calyptra.

I do not know how much importance is to be attached to the

single feature of the calyptrate vs. non-calyptrate calyx. Super-

ficially the two seem very different, but, as pointed out above, the

calyx in a normally calyptrate species may sometimes open by

irregular longitudinal splits. Superficial examination of a number

of specimens of P. sartorianum shows a great deal of individual

variation in the "neatness" with which the calyptra is cut off;

sometimes it is very irregular, resembling a large calyx-lobe opposing

the smaller irregular lobes around the circumference of the hypan-

thium. I think it probable that this calyx-character is not a very

stable one, genetically speaking, and that a form with open calyx

might well appear in a large population like P. sartorianum. Addi-

tional series of specimens of plants in bud and in flower are needed

to establish the taxonomic position of P. molinae and P. micro-

phyllum.

INDEX

New names in bold -face type

Anamomis fragrans, 485

Calycorectes densiflorus, 509

Calyptranth.es apoda, 412

aromatica, 411

costa-ricensis, 408

chytraculia, 403

var. americana, 404

var. chytraculia, 403

5 pauciflora, 407

euryphylla, 405

hernandezii, 404

johnstonii, 405

mexicana, 409

millspaughii, 405

pallens, 406

var. mexicana, 409

var. pallens, 407

var. williamsii, 408

paxillata, 410

schiedeana, 411

schlechtendaliana, 411

tenuipes, 411

tovarensis, 412

vexata, 412

williamsii, 408

zuzygium, 412

Eugenia acapulcensis, 421

alaternifolia, 481

alnifolia, 425

andina, 481

anglohondurensis, 521

antiquae, 421

apodantha, 520

argyrea, 426

aromatica, 481

asa-grayi, 503

avicenniae, 427

axillaris, 428

bartlettiana, 421

baruensis, 421

basilaris, 464

biflora, 428

bonplandiana, 421

bracteolosa, 421

buxifolia, 441

calciphila, 468

campechiana, 421

capuli, 432

carthagenensis, 421

/3 baruensis, 421

casearioides, 433

chepensis, 434

chinajensis, 449

choapamensis, 435

cocquericotensis, 446

colipensis, 436

comitanensis, 421

contrerasii, 432

conzattii, 446

crassifolia, 467

crebrifolia, 434

crenularis, 436

culminicola, 437

deltoidea, 421

dissitiflora, 467

divaricata, 502

doubledayi, 470

dugandii, 484

egensis, 466

escuintlensis, 421

farameoides, 439

fieldingii, 428

fiscalensis, 454

flavida, 467

flavifolia, 439

flavoviridis, 467

florida, 446

foetida, 453

18 maleolens, 454

7 rhombea, 453

fragrans, 485

gaumeri, 469

gentlei, 520

granulata, 487

guadalupensis, 421

guanacastensis, 465

guatemalensis, 439

hintonii, 436

hiraeifolia, 455, 457

hondurensis, 467

hypargyrea, 439

inconspicua, 440

itzana, 425

jutiapensis, 465

karsteniana, 490

karwinskyana, 440

koepperi, 449

laevis, 468

var. gaumeri, 469

var. laevis, 468

530

INDEX

531

ledophylla, 430, 440

leptppa, 453

1'heritieriana, 503

liebmannii, 441

lindeniana, 442

lopeziana, 485

lundellii, 469

macrocarpa, 442

mariquitensis, 491

maritima, 421

mato, 491

mexiae, 455

mexicana, 442

michoacanensis, 443

micrantha, 433

minimifolia, 492

mirandae, 444

monticola, 432

var. latifolia, 432

mosquitensis, 421

mouririodes, 521

musarum, 521

nicaraguensis, 467

oaxacana Berg, 445, 455, 456, 458

oaxacana Standl., 455, 456, 458

octopleura, 470

oerstedeana, 446

oreinoma, 465

origanoides, 448

orthostemon, 492

ovatifolia, 421

pallens, 406

palmarum, 503

paplensis, 465

periplocifolia, 471

petenensis, 446

pittieri, 433

pleurocarpa, 449

polyneura, 503

praeterita, 450

principium, 451

pseudo-mato, 493

pueblana, 452

punctata, 485

purpusii, 455, 456

pycnantha, 494

pycnoneura, 503

rekoi, 453

rhombea, 453

rigidissima, 494

roepperi, 470

rondonensis, 495

rubella, 449

salamensis, 455

var. hiraeifolia, 457

var. rensoniana, 456

var. salamensis, 455

schiedeana, 432

schippii, 521

sibunensis, 421

sinaloae, 457

standleyi, 458

steyermarkii, 485

storkii, 496

subverticillaris, 468

symphoricarpos, 458

tabascensis, 429

teapensis, 459

tenuissima, 434

ternifolia, 497

tikalana, 435

tomentulosa, 455

triflora, 485

triquetra, 497

truncata, 465

trunciflora, 460

turneri, 460

turumiquirensis, 497

vincentina, 446

xalapensis, 461

xilitlensis, 462

yautepecana, 463

yucatanensis, 429

Marlierea dussii, 509

Mitranthes sartoriana, 525

Myrcia arpmatica, 411

belizensis, 472

costa-ricensis, 471

discolor, 471

fallax, 472

karsteniana, 490

longicaudata, 471

melanoclada, 471

mollis, 471

oerstedeana, 471

plicato-costata, 471

rufidula, 470

sartoriana, 471

saxicola, 431

/3 grandifolia, 431

schippii, 471

seleriana, 485

splendens, 470

tovarensis, 412

Myrcianthes biflora, 485

borealis, 481

callicoma, 482

compressa, 483

crebrifolia, 484

dugandii, 484

foliosa, 491

fragrans, 485

var. fragrans, 485

var. hispidula, 489

irregularis, 489

karsteniana, 490

leucpxyla, 491

mariquitensis, 491

mato, 491

minimifolia, 492

prodigiosa, 492

pseudo-mato, 493

rhopaloides, 481, 483

sessilis, 495

storkii, 496

532 FIELDIANA: BOTANY, VOLUME 29

ternifolia, 497 xalapensis, 461

Myrciaria amazonica, 501 zuzygium, 412

caurensis, 501

cordata, 501 Paramitranthes densiflora, 509

divaricata, 502 Pimenta dioica, 511

dubia, 501 tabasco, 511

floribunda, 502 Plinia asa-grayi, 503

ibarrae, 503 dussii, 509

maragnanensis, 502 fruticosa, 509

paraensis, 501 salticola, 505

phillyraeoides, 503 Pseudomyrcianthes pseudo-mato, 493

pittieri, 503 Psidium apodanthum, 520

verticillata, 502 anglohondurense, 521

vexator, 503 biloculare, 520

vismeifolia, 504 chiapasense, 524

Myrteola acerosa, 431 chrysobalanoides, 522

microphylla, 432 dubium, 501

Myrtus acerosa, 430 gentlei, 520

axillaris, 428 guajava, 523

biflora, 428 guineense, 523

casearoides, 433 hyppglaucum, 523

capuli, 432 molinae, 528

chytraculia, 403 mouririodes, 521

coccolobaefolia, 483 musarum, 521

compressa, 483 oerstedeanum, 524

ehrenbergii, 442 rensonianum, 455, 456, 457

fragrans, 485 rotundifolium, 522

foliosa, 491 sartorianum, 525

ledophylla, 430 var. yucatanense, 527

leucpxyla, 491 schippii, 522

maritima, 421 yucatanense, 527

micrantha, 433

oaxacana, 458 Siphoneugena densiflora, 509

splendens, 470 var. tepuiensis, 509

trtmciflora, 460 reitzii, 507

Publication 968

UNIVERSITY OF ILLINOIS-URBANA