UNIVERSITY OF

ILLINOIS LIBRARY

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NATURAL HIST. SURVEY

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i

Botany

Ten New Species of Erythroxylum (Erythroxylaceae)

from Bahia, Brazil

Timothy Plowman

January 30, 1987

Publication I

PUBLISHED BY FIELD MUSEUM OF NA i

Information for Contributors to Fiekliana

FIELDIANA

Botany

NEW SERIES, NO. 19

Ten New Species of Erythroxylum (Erythroxylaceae)

from Bahia, Brazil

Timothy Plowman

Associate Curator

Department of Botany

Field Museum of Natural History

Chicago, Illinois 60605-2496

Accepted for publication July 18, 1985

January 30, 1987

Publication 1373

PUBLISHED BY FIELD MUSEUM OF NATURAL HISTORY

© 1987 Field Museum of Natural History

Library of Congress Catalog Card Number: 86-82463

ISSN 00 15-0746

PRINTED IN THE UNITED STATES OF AMERICA

Table of Contents

ABSTRACT 1

INTRODUCTION 1

NEW SPECIES DESCRIPTIONS

1 . Erythroxylum andrei Plowman, sp.

nov 2

2. Erythroxylum caatingae Plowman, sp.

nov 5

3. Erythroxylum maracasense Plowman,

sp. nov 8

4. Erythroxylum mattos-silvae Plowman,

sp. nov.

12

5. Erythroxylum membranaceum Plow-

man, sp. nov 17

6. Erythroxylum petrae-caballi Plow-

man, sp. nov 20

7. Erythroxylum santosii Plowman, sp.

nov 25

8. Erythroxylum splendidum Plowman,

sp. nov 25

9. Erythroxylum stipulosum Plowman,

sp. nov 31

10. Erythroxylum tenue Plowman, sp.

nov 35

ACKNOWLEDGMENTS 40

LITERATURE CITED . . 40

List of Illustrations

1 . Erythroxylum andrei 3

2. Habitat of Erythroxylum andrei 4

3. Erythroxylum caatingae 6

4. Erythroxylum maracasense 9

5. Habit of Erythroxylum maracasense ... 10

6. Erythroxylum mattos-silvae 13

7. Habit of Erythroxylum mattos-silvae ... 14

8. Flowering twig of Erythroxylum mattos-

silvae 14

9. Erythroxylum membranaceum 18

10. Geographical distribution of Erythroxy-

lum andrei, E. caatingae, E. maracas-

ense, E. mattos-silvae, and E. membrana-

ceum 19

1 1 . Erythroxylum petrae-caballi 21

1 2. Erythroxylum santosii 24

13. Erythroxylum splendidum 27

14. Habit of Erythroxylum splendidum .... 28

1 5. Flowering and fruiting twig of Erythroxy-

lum splendidum 28

16. Erythroxylum stipulosum 30

17. Habitat of Erythroxylum stipulosum ... 31

1 8. Geographical distribution of Erythroxy-

lum petrae-caballi, E. santosii, E. splen-

didum, and E. stipulosum 32

1 9. Erythroxylum tenue 34

20. Habitat of Erythroxylum tenue 35

2 1 . Habit of Erythroxylum tenue 36

22. Flowering branch of Erythroxylum tenue 37

23. Geographic distribution of Erythroxy-

lum tenue . .38

List of Tables

Morphological comparison of Erythroxy-

lum caatingae Plowman and E. oxypetal-

um O. E. Schulz 8

Morphological comparison of Erythroxy-

lum maracasense Plowman and E. pelle-

terianum St. Hil 12

Morphological comparison of Erythroxy-

lum mattos-silvae Plowman, E. nobile O.

E. Schulz, E. compressum Peyr., E. mem-

branaceum Plowman, E. martii Peyr.,

and E. grandifolium Peyr 17

Morphological comparison of Erythroxy-

lum petrae-caballi Plowman, E. tenue

Plowman, E. distortum Mart., E. cf. mi-

kanii Peyr., and E. passerinum Mart 23

Morphological comparison of Erythroxy-

lum santosii Plowman and E. ochran-

thum Mart. . 26

iii

Ten New Species of Erythroxylum (Erythroxylaceae)

from Ba hia. Brazil

Abstract

Ten new species of Erythroxylum (Erythroxyla-

ceae) from the state of Bahia and adjacent areas

in Brazil are described and illustrated, and their

taxonomic and ecogeographic relationships are

discussed: E. andrei Plowman, E. caatingae Plow-

man, E. maracasense Plowman, E. mattos-silvae

Plowman, E. membranaceum Plowman, E. petrae-

caballi Plowman, E. santosii Plowman, E. splen-

didum Plowman, E. stipulosum Plowman, and E.

tenue Plowman.

Introduction

Since I began studies of Neotropical Erythroxy-

lum in 1974, no one region has presented such

diversity of taxa and such a challenge in their de-

limitation as the state of Bahia, Brazil. I have now

tallied 45 species from this state alone. In the Neo-

tropics, Bahia is rivaled only by Venezuela in

numbers of species of Erythroxylum in a com-

parable area (Plowman, 1982). The interesting

juxtaposition of habitats from restinga and moist

tropical forest along the coast to dry caatinga, cam-

po rupestre and cerrado in the interior has led to

a radiation of species that we are only beginning

to describe, let alone understand.

Within Bahia, the greatest concentration of

species of Erythroxylum occurs along the coast in

restinga and restinga forests on sandy soils. Other

species occur in both restinga forest and moist

tropical forest; for still others we do not yet know

the habitat within the coastal zone.

From studying recent collections from southern

Bahia and from my own fieldwork there, it has

become clear that many species of Erythroxylum

occur naturally in very small populations and are

restricted to limited areas. One example is Ery-

throxylum hamigerum O. E. Schulz collected in

1 82 1 by Riedel at "Castelnovo" (= Castelo Novo)

near Ilheus and in 1836 by Blanchet also near

Ilheus. In spite of intensive collecting around Ilh-

eus in the last 1 5 years, this species has been found

only once: in 1983 at Lagoa Encantada near Cas-

telo Novo, where Riedel first discovered it. Another

example is E. mattos-silvae Plowman, described

here, which is found locally in restinga forests south

of Ilheus but was not collected until 1969. Other

species, such as E. nitidum Sprengel, E. mem-

branaceum Plowman, or E. santosii Plowman, are

known only from single collections. The continu-

ing decimation of the forests of southern Bahia for

agriculture and housing developments has greatly

increased the probability that many of the very

rare and local erythroxylums, along with hundreds

of other species not protected in forest preserves,

will become extinct in the near future.

Our knowledge of the southern Bahian flora is

owed largely to the Centre de Pesquisas do Cacau

(CEPEC), a division of the Commissao Executiva

do Piano da Lavoura Cacaueira (CEPLAC). The

staff of this institution has been actively collecting

this area for the past 1 5 years, resulting in a sig-

nificant increase in the numbers of new species

and species records for Bahia (Mori & Mattos Sil-

va, 1979; Mattos Silva & Mori, 1981).

It is becoming increasingly clear that caatinga

and other dry habitats of northeastern Brazil are

also rich in species of Erythroxylum; but the vast,

dry areas of the interior are very poorly explored

compared with the areas of the narrow coastal belt.

PLOWMAN: NEW SPECIES OF ERYTHROXYLUM

Recent explorations of caatinga areas have been

carried out by Ray Harley and colleagues from the

Royal Botanic Gardens, Kew, by Scott Mori of the

New York Botanical Garden, and by the staff at

the Herbario Radambrasil in Salvador. Other re-

cent collectors in Bahia have also made important

contributions, especially the botanists at the Uni-

versidade Federal da Bahia in Salvador and at the

Universidade Estadual de Feira de San tana (Britto

& Noblick, 1 984). These collections suggest that,

when better collected, the dry habitats of Bahia

and other areas of the Northeast may be nearly as

rich in Erythroxylum species, including endemics,

as the moist forests.

It is the purpose of this paper to describe 10

new species of Erythroxylum, six from restinga

and restinga forest, and four from caatinga and

dry forest. This is the second contribution (see

Plowman, 1983) toward a revision of the species

of Erythroxylum in Bahia. Because of the long-

recognized difficulty in distinguishing Erythroxy-

lum species, I have provided rather long and de-

tailed descriptions, employing as many useful

characters as possible in order to avoid any future

ambiguity in interpreting these concepts.

Since my last contributions to the taxonomy of

this genus (Plowman, 1983, 1984), I have made

some modifications in terminology. Following

Schulz (1 907) and earlier workers, I formerly used

"ramenta" to designate the stem covering of con-

gested, sometimes modified cataphylls (consisting

of stipules without developed leaf blades). Since

there appear to be many intermediate conditions

between densely congested "ramenta" and scat-

tered and isolated cataphylls, I believe it is better

to abandon the rather vague term "ramenta" and

replace it with a simple description of the nature

of the cataphylls. In earlier descriptions, I did not

consider the minute, spinelike leaf rudiment usu-

ally found on cataphylls, but I now include this

character, which I have designated the "spinule."

New Species Descriptions

1. Erythroxylum andrei Plowman, sp. nov. sect.

Rhabdophyllum. Figures 1-2, 10.

Frutex ramosissimus, ramulis flexuosis. Cataphylla

disticha, stipulis foliorum similia. Stipulae foliares per-

sistentes, parvae, transverse triangulares, coriaceae, striate

nervosae, 3-setulosae. Folia breviter petiolata. persisten-

tia; laminae obovatae vel ovatae vel suborbiculares, ap-

ice obtusae vel rotundatae, retusae, basi obtusae vel ro-

tundatac vel late cuneatae, chartaceae vel subcoriaceae.

Flores e brachyblastis vel ramulis hornotinis in ramen-

torum axillis nati, brevissime pedunculati. Petal! lamina

clliptica. ungue amplo et ligula hilobata munita. Urceo-

lus stamineus calycem superans, margine integer vel mi-

nute 10-crenulatus. Styli liberi. Drupa ignota.

Shrub to 6 m tall, densely branched, with several

trunks to 3 cm diameter. Bark rather smooth, dot-

ted with small rounded or elongate lenticels or

longitudinally striate, grayish brown, ca. 2 mm

thick, the inner bark tan, the wood light tan, very

hard. Branches dense, ascending or upcurving, not

distinctly storied. Branchlets distichous, consist-

ing mainly of short shoots, flexuous, diverging 50°-

90° from axis, strongly compressed at apex, 1.5-

2 mm in diameter, dotted with rounded or elon-

gate, pale tan lenticels. Internodes 3-20 mm long

on long shoots, 0.5-1 mm long on short shoots.

Cataphylls (ramenta) distichous, produced at base

of new shoots for up to 8 mm, scarcely overlap-

ping, similar to foliar stipules, persisting, turning

black with age, the spinule (leaf rudiment) slender,

black, ca. 1 mm long. Foliar stipules persistent,

appressed to stem, diverging somewhat with age,

broadly triangular, 1.5-2 mm long, coriaceous,

striate-nerved with 5-6 parallel nerves per side,

obtuse to rounded at apex, 3-setulose, the 2 lateral

setae 0.4-0.5 mm long, the medial seta variable

in length, 0.2-0.7 mm long, the setae evanescent

with age, the keels prominent, riblike, the margin

entire. Leaves persistent, 1-2 produced at tips of

short shoots or scattered on long shoots, disti-

chous, short-petiolate, the lamina held obliquely

erect, plane, variable in shape, ovate to obovate

or suborbicular, obtuse to rounded at apex, retuse,

obtuse to rounded or broadly cuneate at base, 35-

85 mm long, 20-55 mm wide, firmly chartaceous

to subcoriaceous, adaxially medium to dark green,

drying dull, grayish green, abaxially light yellowish

green, drying yellowish or reddish brown, adaxi-

ally rather dull to shiny, abaxially shiny, elineate

with no distinct central panel, the adaxial midrib

yellowish or light green, flat or only slightly raised,

the major lateral nerves 10-14, diverging 55°-70°

from midrib, straight, more or less parallel, anas-

tomosing 2-4 mm from margin, adaxially obscure,

abaxially distinct. Petiole subterete, 2-3 mm long,

1-1.3 mm in diameter, drying very dark brown.

Flowers produced near the apex of new short shoots

or new long shoots in axils of leaves or cataphylls,

1-4 per node, whitish, subsessile or very short-

pedunculate, the peduncle flattened against stem,

0.2-1.2 mm long, diverging from axis with age.

FIELDIANA: BOTANY

1mm

FIG. 1 . Erythroxylum andrei. A, habit of flowering branch; B, habit of whole plant; C, detail of flowering twig;

D, leaf showing venation, scale as in A; E, stipule; F, short-styled flower; G, long-styled flower; H, petal. (A, C, D,

E, F, and H from Carvalho el al. 198; B from Plowman & Carvalho 12785; G from dos Santos 1287.) Drawing by

Marlene Werner.

PLOWMAN: NEW SPECIES OF ERYTHROXYLUM

FIG. 2.

12785).

Habitat of Erythroxylum andrei in shrubby restinga near Marau, Bahia, Brazil (Plowman & Carvalho

Bracteoles minute, produced at apex of peduncle,

broadly ovate, concave, membranaceous, 0.6-0.8

mm long, abruptly acuminate at apex, the seta 0.2-

0.4 mm long. Pedicel slender, 5-ribbed, 3.5-6.5

mm long, 0.5-0.7 mm in diameter, thickened at

apex into the calyx. Calyx 1.2-1.5 mm long, di-

vided '/j-'/2 its length, the lobes broadly triangular,

0.5-0.7 mm long, apically acute. Petal lamina

spreading, concave, elliptic to narrowly ovate,

rounded at apex, 2 mm long, 1-1.5 mm wide, the

claw broad and somewhat larger than the lamina,

1.5-1.6 mm long, to 1.7 mm wide, the ligule 1.4

mm long, bilobate, with a broad, inflexed, central

flap 0.7-0.8 mm long and 0.6 mm wide, the lobes

each consisting of 2 auricles, the shorter anterior

auricle 0.6 mm long, the longer posterior auricle

1-1.4 mm long. Stamina! cup 1.2-1.8 times length

of calyx, 1-1.3 mm long, entire to minutely 10-

crenulate at margin, the margin itself drying thin,

lighter brown in color, flaring slightly. Brachysty-

lous flowers: filaments 2.5-2.8 mm long, the an-

thers 0.5 mm long; styles free, 1 mm long; stigma

broadly depressed-capitate, 0.2 mm long, 0.3 mm

in diameter. Dolichostylous flowers: antisepalous

filaments 0.8 mm long, the anthers 0.5 mm long;

antipetalous filaments 1.2 mm long, the anthers

0.5 mm long; styles free, 1-1.1 mm long; stigmas

depressed-capitate, 0.2 mm long. Ovary obovoid,

rounded at apex, 1.2-1.8 times length of staminal

cup, 0.8-1.8 mm long. Mature drupe unknown.

TYPE— Brazil, Bahia, Municipo de Marau, ro-

dovia BR-030, trecho Porto de Campinhos-Mar-

au, km 1 1 , restinga, arbusto de ap. 2.0 m de altura,

folhas discolores com face superior lucida, flores

alvacentas, frutos imaturos verdes, 26 Feb. 1 980

(fl), Andre M. de Carvalho, L. A. Mattos Silva &

T. S. dos Santos 198 (holotype, CEPEC 19131; iso-

types, F 1884879, F neg. 60193, K, NY, RB).

ADDITIONAL SPECIMENS EXAMINED— BRAZIL: Ala-

goas: Municipio Marechal Deodoro: Massagueira, en-

trada para Barra de Sao Miguel, 30 Jan. 1982 (fl), M. L.

Guedes s.n. (ALCB, F). Bahia: Municipio de Marau: Es-

trada Ubaitaba-Ponta do Muta, ramal no km 71 de

Ubaitaba, entrada para o Sitio "Sao Marcos," approx.

14°04'S, 38°58'W, near sea level, 2 Feb. 1983 (fl buds),

T. Plowman & A. M. de Carvalho 12785 (ALCB, B, CEPEC,

F, G, GH, GUA, HRB, K, LE, MBM, MO, NY, P, R, RB, SP, U,

UB, UEC, us, distributed as "£". carvalhoi"), 2 Feb. 1983

(st), T. Plowman & A. M. de Carvalho 12786 (CEPEC, F,

GH, HRB, K, NY, RB, UB, us, distributed as "E. carvalhoi").

Municipio de Ilheus: estrada a Olivenca, 12 Nov. 1970

FIELDIANA: BOTANY

(fl), T. S. dos Santos 1287 (CEPEC, F); Fa/enda Barra do

Manguinho, km 12 da Rodovia (BA-001) Pontal-Oli-

ven9a, 26 Apr. 1983) (fl, fr), L. A. Mattos Silva, T. S.

dos Santos & B. Leuenberger 1823 (ALCB, CEPEC, F, MBM,

NY, RB), 27 Feb. 1985 (st), T. Plowman, L. A. Mattos

Silva & T. S. dos Santos 13971 (CEPEC, F, K, NY, RB), 27

Feb. 1985 (young fr), T. Plowman, L. A. Mattos Silva &

T. S. dos Santos 13973 (CEPEC, F, G, HRB, IPA, K, NY, RB,

UEC, us); Fazenda Guanabara, ramal com entrada no km

1 0 da rodovia Ilheus-Olivenca, lado direito 4 km a Oeste

da rodovia, 7 Mar. 1985 (fr), L. A. Mattos Silva, T. S.

dos Santos & H. S. Brito 1363 (CEPEC, F). Municipio de

Porto Seguro: estrada de Arraial d'Ajuda para Trancoso,

20 Apr. 1982 (fl, young fr), A. M. de Carvalho, S. G. da

Vinha & H. S. Brito 1249 (CEPEC, F, HRB, RB).

ETYMOLOGY— Erythroxylum andrei commem-

orates Andre M. de Carvalho, botanist and former

curator at the CEPEC herbarium. He has made ex-

cellent recent collections of the southern Bahian

flora and first brought this species to my attention.

I originally intended to name this species "£. car-

valhoi," and some of my collections were distrib-

uted under this name. I subsequently discovered

that this epithet is preoccupied by E. carvalhoi

(Engl.) Phillips, a new combination proposed in

1935 for Nectaropetalum carvalhoi Engl. from

Mozambique (Phillips, 1935).

DISTRIBUTION— Known only from the coast of

eastern Brazil from the states of Alagoas and Ba-

hia.

ECOLOGY— Erythroxylum andrei grows only in

open, low restinga formations near the sea. It may

be locally common, but is known from only four

localities. In the area south of Ilheus, Bahia, its

habitat is seriously threatened by vacation housing

developments and sand mining.

PHENOLOGY— Flowering specimens have been

collected in November, January, February, and

April. Mature fruits are unknown.

RELATIONSHIPS— Based on its having striately

nerved stipules and relatively small calyx lobes,

Erythroxylum andrei is placed in the large Neo-

tropical section Rhabdophyllum O. E. Schulz. It

superficially resembles several other species from

Bahia that also occur in restinga formations. It

differs from E. leal-costae Plowman in having more

slender twigs (1-2 mm vs. 3-4 mm), the leaves

abaxially shiny, the venation more prominent, the

flowers short-pedunculate, and the pedicels longer

(4-6.5 mm vs. 1.5-2 mm). Erythroxylum andrei

differs from E. passerinum Mart, in having a flat

adaxial midrib without a distinct, slender ridge,

well-developed short shoots, shorter (1.2-2 mm

vs. 2-3.5 mm) and broader stipules with more

pronounced nerves, thicker leaves with more

prominent and more open venation, and the sta-

minal cup exceeding, rather than shorter than, the

calyx.

2. Erythroxylum caatingae Plowman, sp. nov. sect.

Archerythroxylum. Figures 3, 10.

Arbor parva vel frutex ramosissimus, ramulis rectis,

crassis, nigrescentibus. Cataphylla pauca, stipulis foli-

aribus similia. Stipulae foliares mediocres, triangu-

lari-ovatae, estriatae, apice integrae vel brevissime

2-setulosae, fugaces. Folia petiolata, decidua; laminae

ellipticae vel oblongae, raro suborbiculares vel obovatae,

apice rotundatae, retusae, basi obtusae vel rotundatae

vel late cuneatae, firme chartaceae vel subcoriaceae. Flo-

res 1-5, e brachyblastis hornotinis in ramentorum vel

foliorum axillis nati, pedicellis brevibus, incrassatis,

5-costatis. Petali lamina ovata, ligula bilobata munita.

Urceolus stamineus calycem aequans, margine 10-cren-

atus. Styli ad basin tantum connati. Drupa endocarpio

triquetro, triloculari, duabus loculis sterilibus magnis,

loculo fertili parvo provisa.

Shrub or small tree, 1.5-4 m tall, much-branched

and with a broad crown. Branches distichous, often

at near right angles to axis, rigid, thick, becoming

smooth and black. Branchlets distichous, consist-

ing of both long and short shoots, diverging 50°-

90° from axis, compressed at apex, 1.5-2 mm in

diameter, stiff, longitudinally wrinkled, dark brown

to black, the lenticels numerous, punctate to elon-

gate, whitish, the bark becoming longitudinally

striate. Internodes 2-13 mm long on long shoots,

1-3 mm long on short shoots. Cataphylls (ramen-

ta) sparse, up to 5 scattered along base of long

shoots or alternating with leaves on short shoots,

similar in form to foliar stipules, the spinule (leaf

rudiment) 2-2.5 mm long, dark brown to black,

very early caducous. Foliar stipules distichous,

persisting for 1 or 2 seasons, then marcescent, ap-

pressed to stem, slightly diverging with age, tri-

angular-ovate, 2.5-4 mm long, firmly membra-

naceous, smooth, nonstriate, when mature drying

reddish brown, obtuse at apex, entire or briefly

2-setulose, the setae short-filamentous, 0.2-0.8 mm

long, evanescent, the keels sub-alate, the margin

entire. Leaves deciduous, scattered on long shoots

or 1-5 produced near apex of short shoots, disti-

chous, petiolate, the lamina plane or somewhat

undulate, elliptic to oblong, rarely suborbicular or

obovate, rounded or retuse at apex, obtuse to

rounded or broadly cuneate at base, 15-52 mm

long, 10-35 mm wide, firmly chartaceous to sub-

coriaceous, adaxially medium green, drying olive

green to dark brown, abaxially pale green, drying

pale olive green to pale brown, adaxially dull to

somewhat shiny, abaxially matte, elineate with no

distinct central panel, the adaxial midrib flat or

PLOWMAN: NEW SPECIES OF ERYTHROXYLUM

2 mm

1 mm

FIG. 3. Erythroxylum caatingae. A, flowering branch, 3-cm scale; B, fruiting branch, 3-cm scale; C, leaf showing

venation; D, detail of apex of flowering twig; E, stipule; F, short-styled flower; G, long-styled flower; H, petal; I,

dorsal view of endocarp; J, cross section of endocarp. (A, D, E, F, and H from Hurley et al. 16473; B and C from

Mori 13443; G from Leal Costa s.n.; I and J from Harley et al. 16928.) Drawing by Marlene Werner.

FIELDIANA: BOTANY

somewhat furrowed, the abaxial midrib drying light

straw-colored, the lateral nerves 12-14, diverging

30°-50° from midrib, sinuous, more or less par-

allel, anastomosing 0.5-2 mm from midrib, prom-

inulous on both surfaces. Petiole 3-6 mm long,

0.5-0.8 mm in diameter, subterete, narrowly can-

aliculate. Flowers in axils of leaves or cataphylls

on current season's shoots, 1-3 (5) per node, sparse

to somewhat congested, white, greenish white, or

yellowish. Bracteoles broadly ovate to orbicular,

concave, 1-1 .5 mm long, paleaceous, acute to acu-

minate at apex, the seta 0.3-0.5 mm long. Pedicel

pentagonal, 5 -ribbed, somewhat thickened, 1-1.5

mm long, 0.5-1 mm in diameter. Calyx 1.5-2.5

mm long, divided '/z-% its length, pale green, the

lobes ovate to ovate-lanceolate, 1-1.5 mm long.

Petal lamina ovate, 1.7-2.5 mm long, 1.3-1.5 mm

wide, rounded or obtuse at apex, concave, often

incurved, the claw 1.2-1.5 mm long, the ligule

bilobed, 0.7-1 mm long, each lobe consisting of a

posterior ovate auricle 0.5-0.7 mm long, some-

times with a very short, cufflike, anterior auricle

and with a broad, inflexed, medial flap between

the lobes. Stamina! cup % the length of calyx, 1.2-

1.5 mm long, the margin 10-crenate. Brachysty-

lous flowers: filaments 1 .8-2 mm long, the anthers

ovate-elliptic, 0.6 mm long; styles 1 mm long, brief-

ly connate at base; stigmas depressed-capitate, 0.3

mm long. Dolichostylous flowers: antisepalous fil-

aments 0.6-0.7 mm long, the anthers elliptic, 0.6

mm long; antipetalous filaments 1.1-1.2 mm long,

the anthers elliptic, 0.5 mm long; styles 2-2.5 mm

long, connate at base 0.5-0.8 mm; stigmas de-

pressed-capitate, 0.2-0.3 mm long. Ovary equal-

ing the staminal cup, obovoid, rounded at apex,

1 .4-1 .5 mm long. Drupe ellipsoid, rounded at apex,

8-10 mm long, 4.2-5.2 mm in diameter, reddish

orange to scarlet at maturity, the mesocarp thin,

the endocarp oblong to ellipsoid in outline, tri-

gonal in cross section with 1 wider flat side and 2

narrower, shallowly indented sides, 3-locular with

2 large, empty locules and 1 small, fertile locule;

endosperm occupying ca 1 0%-20% of fertile loc-

ule. Embryo 5.5-6.7 mm long, pale green; coty-

ledons very thin, oblong, rounded at apex, 4.2-5

mm long, 1 .5-1 .7 mm wide, 0. 1 mm thick; radicle

1.6-1.7 mm long, 0.5 mm in diameter.

TYPE— Brazil, Bahia, Raso da Catarina, vege-

tacao caatinga, arbusto de mais ou menos 2,5 m

de altura, flores pequenas, amarelo-claro, frutos

imaturos e matures (cor-de-abobora), 1 4 May 1981

(fl, fr), H. P. Bautista 445 (holotype, HRB 2356;

isotypes, CEPEC 26479, F 1912521).

ADDITIONAL SPECIMENS EXAMINED— BRAZIL: Ceara:

2-4 km south of Campos Sales, alt. 620 m, 7°14'S,

40°25'W, 15 Feb. 1985 (fl), A. Gentry. E. Zardini & A.

Fernandes 50140 (EAC, F, MO). Paraiba: Borborema,

"imbuzeiro bravo," Nov. 1936 (fl, fr), H. Zenaide 27

(SP). Pernambuco: entre Bom Nome e Sao Jose do Bel-

monte, 13 May 1971 (fr), Academia Brasileira de Cien-

cias (E. P. Heringer, D. de A. Lima, J. de P. L. Sobrinho

& A. C, Sarmento) 730 (F, IPA, RB, UB); entre Airi e Serra

Negra, 8 Feb. 1949 (st), D. de A. Lima 49-178 (IPA);

Santa Maria da Boa Vista, margem da PE-4, em direcao

a Jutai, 29 Apr. 1971 (fl), Academia Brasileira de Cien-

cias (E. P. Heringer, D. de A. Lima, J. de P. L. Sobrinho

&A. C. Sarmento) 376 (F, IPA, RB, UB); Afranio, Caboclo,

Serra do Caboclo, 21 Apr. 1971 (fl, fr), Academia Bra-

sileira de Ciencias (E. P. Heringer, D. de A. Lima, J. de

P. L. Sobrinho & A. C. Sarmento) 260 (F, IPA, RB, UB).

Bahia: Estacao Ecologico do Raso da Catarina, 26 Jun.

1982 (fl), L. Paganucci de Queiroz 380 (ALCB, F); no

entroncamento da estrada de Jorro com a de Tucano, 4

Feb. 1983 (fl), A. Leal Costa s.n. (ALCB, F). Municipio

Nossa Senhora dos Milagres: estrada para Itaberaba, km

10, estradinha para as fazendas Morros e Antonio Ro-

meu, km 2, 28 Jan. 1973 (fr), /. & G. Gottsberger 27-

28173 (F); Ia?u, Fazenda Suibra, 1 8 km a leste da cidade

sequindo a ferrovia, 12 Mar. 1985 (fr), L. Noblick &

Lemos 3581 (F, HUEFS); 14 km SW of Cansan9§o on road

to Queimadas, ca. 30°34'W, 10°47'S, alt. ca. 300 m, 22

Feb. 1974 (fl), R. M. Harley et al. 16473 (CEPEC, IPA, K,

MO, NY, p, RB, u, us); Serra do Curral Feio, 26 km NW

of Lagoinha (which is 5.5 km SW of Delfino) on side

road to Minas do Mimoso, ca. 41°23'W, 10°16'S, 7 Mar.

1974 (fr), R. M. Harley et al 16928 (CEPEC, GH, K, MO,

NY, P, RB, u, us); Serra Acurua, ca. 4 km NE from Gentio

do Ouro along road toward Central, ca. 42°30'W, 1 1°24'S,

alt. ca. 1000 m, 22 Feb. 1977 (fr), R. M. Harley et al.

18958 (CEPEC, F, K, NY, p, RB, u). Regiao da Serra do

Sincora, "Iracema" [= Iramaia], 18 Feb. 1943 (fl, young

fr), R. de L. Frdes 20183 (IAN, NY, us); km 43 da estrada

Brumado-Caetite, alt. 740 m, 14 Apr. 1983 (fl, fr), A.

M. de Carvalho, B. Leuenberger & L. A. Mattos Silva

1694 (ALCB, B, CEPEC, F, RB).

ETYMOLOGY— The specific epithet caatingae

means "of the caatinga," in reference to the ex-

clusively northeastern Brazilian habitat of this

species.

DISTRIBUTION— Erythroxylum caatingae is

known only from northeastern Brazil from the state

of Paraiba south to Bahia.

ECOLOGY— Most collections of E. caatingae

mention the habitat as "caatinga," the type of dry

vegetation characteristic of northeastern Brazil.

Only one collection (Harley et al. 18958) provides

more detailed habitat information: "caatinga on

sand and with quartzitic rocks and metamor-

phosed sandstones forming rock area with more

open vegetation including extensive areas of Vel~

lozia."

PHENOLOGY— In Pernambuco and Bahia, Ery-

throxylum caatingae appears to flower and fruit

PLOWMAN: NEW SPECIES OF ERYTHROXYLUM

TABLE 1 . Morphological comparison of Erythroxylum caatingae Plowman and E. oxypetalum O. E. Schulz.

during the rainy season, primarily from February

to April. The sole collection from Paraiba was

collected in flower and fruit in November.

COMMON NAME— "Imbuzeiro bravo" (Paraiba).

RELATIONSHIPS— Erythroxylum caatingae looks

superficially like a number of other stiff-branched

caatinga species, such as E. pungens O. E. Schulz,

E. nummularia Peyr., and E. bezerrae Plowman,

a new species from Piaui (Plowman, 1986). It is

readily distinguished from E. pungens and E. be-

zerrae in having nonstriate stipules and from E.

nummularia in having petiolate leaves.

Erythroxylum caatingae belongs to section Ar-

cherythroxylum based on its nonstriate stipules

and perfect flowers. It is most closely related to E.

oxypetalum O. E. Schulz, a rare species known

only from two collections from central (Diaman-

tina, Glaziou 12473, holotype, B, destroyed, F neg.

12634; isotypes, c, G, K, LE, p, R) and northeastern

Minas Gerais (estrada Itaobim-Joaima, Mendes

Magalhdes 17628, IAN). Morphological differ-

ences between these two species are listed in Table

1 . Although E. oxypetalum appears also to grow

in caatinga vegetation, it has a more southerly

distribution, and the two species are allopatric.

3. Erythroxylum maracasense Plowman, sp. nov.

sect. Rhabdophyllum O. E. Schulz. Figures

4-5, 10.

Frutex vel arbuscula, ramulis rectis vel subflexuosis,

lenticellis instructis. Cataphylla pauca, brevia, disticha,

stipulis foliaribus similia. Stipulae foliares persistentes,

parvae, triangulari-ovatae, striate nervosae, breviter 3-

setulosae. Folia parva, subsessilia vel brevissime pe-

tiolata. decidua; laminae late obovatae, ellipticae vel or-

biculares, apice rotundatae vel retusae, basi late cuneatae

vel obtusae, membranaceae. Flores e brachyblastis in

axillis cataphyllorum hornotinorum nati. Petali lamina

ovata vel elliptica, ungue lato et ligula bilobata munita.

Urceolus stamineus quartae vel duabus tertiis partibus

longitudinis calycis aequalis, margine 1 0-crenulatus vel

10-denticulatus. Styli liberi. Drupa endocarpio oblongo-

ovoideo, manifeste 6-sulcato provisa.

Shrub or treelet to 4 m tall. Bark rather smooth,

thin, light brown, dark brown within, the wood

very light tan. Branches more or less distichous,

ascending and spreading, straight or somewhat

flexuous, dark brown. Branchlets distichous, more

or less straight, often parallel, weakly differentiat-

ed into long and short shoots, diverging 30°-65°

from axis, somewhat compressed toward apex, 1-

1 .5 mm in diameter, dark brown to black, smooth,

dotted abundantly with minute, punctate or elon-

gate, whitish lenticels that do not break surface.

Internodes 3-25 mm long on long shoots, 0.5-2

mm long on short shoots. Cataphylls (ramenta)

distichous, produced at apex of shoots, somewhat

congested, covering stem for 3-5 mm, 1.5-3 mm

across, similar to foliar stipules, persisting, the spi-

nule (leaf rudiment) 0.6-1 mm long, broad, flat-

tened, dark brown or black. Foliar stipules per-

sistent, appressed to stem, triangular-ovate,

somewhat concave, 1.5-2.5 mm long, firmly

membranaceous, distinctly striate-nerved with 3-

5 nerves per side, pale green, turning ferruginous

with age, apically obtuse or truncate, fimbriate,

3-setulose, the setae minute, 0.2-0.5 mm long,

evanescent, the keels prominulous, sub-alate, the

margin erose to markedly fimbriate, sometimes

recurving in age. Leaves deciduous, produced along

extension shoots or 1-2 at tips of short shoots,

distichous, short-petiolate or subsessile, the lam-

ina held horizontally, plane, variable in shape,

broadly obovate, elliptic or orbicular, rounded or

retuse at apex, obtuse or broadly cuneate at base,

18-35 mm long, 12-23 mm wide, membrana-

ceous, adaxially medium green, abaxially very pale

green, dull on both surfaces, elineate with no dis-

tinct central panel, the adaxial midrib a slender,

low ridge, yellowish green, the lateral nerves 10-

12, diverging 40°-60° from midrib, straight or

somewhat sinuous, often parallel, anastomosing

1-2 mm from margin, the venation not pro-

nounced on either surface. Petiole 1-2 mm long,

0.5 mm in diameter, subterete. Flowers produced

FIELDIANA: BOTANY

II

FIG. 4. Erythroxylum maracasense. A, flowering branch, 3-cm scale; B, fruiting branch, 3-cm scale; C, stipule,

scale as in F; D, short-styled flower; E, long-styled flower; F, petal; G, endocarp; H, cross section of endocarp, scale

as in G; I, embryo, scale as in G. (A, D, and F from Gounelle s.n.; B, G, H, and I from dos Santos 3488; C from

Plowman & Carvalho 12826; E from Martins s.n.) Drawing by Marlene Werner.

PLOWMAN: NEW SPECIES OF ERYTHROXYLUM

FIG. 5. Habit of Erythroxylum maracasense growing near Maracas, Bahia (Plowman & Carvalho 12826).

in axils of cataphylls (ramenta) near tips of often

very short, axillary short shoots, 1 or 2 flowers per

node, not congested, the flower color unknown.

Bracteoles oblong-ovate, concave, 0.7-0.9 mm

long, membranaceous, the apex rounded and

markedly fimbriate, the seta obscured. Pedicel

slender, 5-ribbed, (1 .2) 4-6 mm long, 0.4-0.5 mm

in diameter, sometimes articulating with a very

short peduncle 0.5-0.7 mm long. Calyx 1 .5-2 mm

long, divided '/z-^A its length, the lobes lanceolate

to ovate-lanceolate, apically acute to acuminate,

0.7-1.8 mm long, the margins paler than the mid-

dle. Petal lamina somewhat concave, ovate to el-

liptic in outline, rounded and somewhat incurved

at apex, 1.5-2.5 mm long, 1-1.5 mm wide, the

claw 0.5-1 mm long, rather broad, the ligule 1 mm

long, bilobed, each lobe consisting of a very short

anterior auricle and a large posterior auricle, the

10

FIELDIANA: BOTANY

posterior one narrowly to broadly ovate, 0.6-0.8

mm long, with 3 broad, irregular teeth at apex.

Staminal cup '/4-2/3 times length of calyx, 0.6-1 mm

long, 10-crenulate or 10-denticulate at margin.

Brachystylous flowers: filaments 2.5-2.8 mm long,

the anthers ovate, 0.5 mm long; styles free, 1.5

mm long; stigmas depressed-captitate, 0.3 mm

long. Dolichostylous flowers: antisepalous fila-

ments 0.5-1 mm long, the anthers ovate-elliptic,

0.3-0.5 mm long; styles free, 2.5-3 mm long; stig-

ma depressed-capitate, 0.2-0.3 mm long. Ovary

ellipsoid or subglobose, rounded at apex, 1.2-2

times length of staminal cup, 1 mm long. Drupe

in dried state oblong-ovoid, 6.5-8.5 mm long, 3.0-

3.5 mm in diameter, red at maturity, the endocarp

oblong-ovoid, acute at apex, terete in outline,

markedly 6-sulcate, unilocular, the endosperm oc-

cupying ca. 50% of locule. Embryo 5.5 mm long;

cotyledons broadly ovate, rounded at apex, sub-

cordate at base, 4.5 mm long, 3 mm wide, 0.8 mm

thick; radicle 1.3 mm long.

TYPE— Brazil, Bahia, Municipio de Maracas, ro-

dovia BA-026, 1 5 km ao SW de Maracas, caatinga

arborea perturbada, folha SD-24 (14-40a), arvo-

rezinha, 4 m de altura, comum, 17 Nov. 1978 (fl),

S. A. Mori, T. S. dos Santos & C. B. Thompson

11120 (holotype, CEPEC 1 5262; isotypes, F 1 86 1 962,

F neg. 60 1 24, K NY, RB).

ADDITIONAL SPECIMENS EXAMINED— BRAZIL: Bahia:

Municipio de Cipo: entre Cipo e Ribeira do Pombal, 9

km adiante de Cipo na autoestrada, 19 Jan. 1964 (fl,

young fr), A. Leal Costa s.n. (ALCB, F). Municipio de

Maracas: "sylvis catingas ad Maracas," Oct. 1818 (fl),

C. F. P. von Martins (M, F neg. 19468, excluded syntype

of E. pungens O. E. Schulz); km 7 na estrada Maracas-

Contendas do Sincora, ca. 13°28'S, 40°29'W, alt. 850-

900 m, 9 Feb. 1 983 (st), T. Plowman &A.M.de Carvalho

12826 (CEPEC, F, o, IPA, K, NY, RB, UB, us). Municipio de

Jequie: entrada do ramal localizado ao SW do km 38 da

rodovia Jequie-Contendas do Sincora, Folha SD 24 ( 1 4-

40a), 15 Feb. 1979 (fr), T. S. dos Santos, L. A. Mattos

Silva & H. da S. Brito 3488 (CEPEC, F, NY). Goias: Near

Almas, Oct. 1839 (fl), G. Gardner 3054 (BM, K, OXF).

Minas Gerais(?): "Caraca", [ 1 885-1 890], P.-E. Gounelle

s.n. (P).

ETYMOLOGY— Erythroxylum maracasense is

named for the town of Maracas in Bahia, near

where the first and most of the subsequent collec-

tions of the species have been made.

DISTRIBUTION— This species is known primarily

from east-central Bahia. One collection is from

northern Goias, and one southerly collection by

Gounelle is from Caraca in Minas Gerais. The

latter may be erroneously labeled. Gounelle was

an engineer who made seven trips to Brazil, where

he made extensive insect collections (Millot, 1 920).

He also collected plants, mainly during 1 885-1 890,

in Minas Gerais and Bahia. That Gounelle col-

lected plants elsewhere in northeastern Brazil is

known, for example, by his collection during 1 892-

1893 of the type plant (living specimen?) ofCeph-

alocereus gounellei (Weber) Britton & Rose, in

Pernambuco (Schumann, 1903; Britton & Rose,

1937). Apparently not much care was taken in

labeling Gounelle's plant specimens, and at least

some localities (like the present case) appear to be

incorrect (A. Lourteig, in litt.). At least one other

monographer has noted discrepancies in distri-

butions of species from northeastern Brazil col-

lected by Gounelle and labeled "Cara9a" (cf. Ban-

isteriopsis calcicola Gates, cited in Gates [1982, p.

105]).

ECOLOGY— Erythroxylum maracasense is ap-

parently confined to the caatingas of eastern Brazil.

This formation is not found at Caraca in the area

of Belo Horizonte where the Gounelle collection

supposedly was made, which further supports the

possibility that this specimen is mislabeled. At

Maracas E. maracasense occurs sympatrically with

several other erythroxylums, the first three of which

are also caatinga endemics: E. betulaceum Mart.,

E. macrochaetum Miq., E. polygonoides Mart., E.

pelleterianum St. Hil., and E. subrotundum St. Hil.

PHENOLOGY— This species has been collected in

flower in October, November, and January; the

sole fruiting specimen was gathered in February.

Flowering undoubtedly follows the onset of the

seasonal rains in the caatinga.

RELATIONSHIPS— Erythroxylum maracasense

was first collected by Martius in 1818. Martius

wrote on the label of his collection at M the name

"E. asparagoides Mart.", indicating that he thought

it was a new species. Subsequently, however, he

crossed out his first annotation and identified the

plant as E. subrotundum St. Hil. Although E. sub-

rotundum belongs to a different section (sect. Ar-

chery throxylum), it is a remarkable mimic for E.

maracasense and also grows at Maracas.

Schulz (1907) included the Martius specimen as

a syntype of his E. pungens O. E. Schulz. Ery-

throxylum pungens, also from caatinga, is a very

different species characterized by thick, stiff,

pointed ("pungent") branchlets, very long pedicels

at the tips of the branchlets, and broadly ovoid,

terete endocarps. Schulz's description better ap-

plies to the second syntype, Blanchet 2771, which

collection I here designate as the lectotype of E.

pungens O. E. Schulz (lectotype, G; isolectotypes,

BM, E, G, LE, NY, w). Incidentally, Schulz also noted

PLOWMAN: NEW SPECIES OF ERYTHROXYLUM

11

TABLE 2. Morphological comparison of Erythroxylum maracasense Plowman and E. pelleterianum St. Hil.

Character

/;. maracasense

E. pelleterianum

Twig surface

Stipule shape

Stipule length (mm)

Ramenta

Dotted with lenticels

Triangular ovate

1.5-2.5

Poorly developed, to 4 mm long

Lenticels absent

or poorly developed

Lanceolate

2.5-5

Well developed,

often covering twigs

the similarity of E. pungens (apparently referring

to the leafless Martius specimen) to E. subrotun-

dum St. Hil.

Another early collection of E. maracasense,

Gardner 3054, was implicated erroneously as a

syntype of another species, E. subglaucescens Peyr.

ex O. E. Schulz. Schulz (1907) based his concept

on Peyritsch's annotations (e.g., at w) and on a

note published by Peyritsch (1878) under E. vir-

gultosum Mart, in Flora Brasiliensis. Peyritsch cit-

ed Gardner 3054 as a specimen similar to E. vir-

gultosum, but suggested it might be a distinct

species. It is now clear that Peyritsch misread the

collection number on Gardner's handwritten la-

bel; Gardner's handwritten "6's" look like "4's."

Peyritsch was actually studying Gardner 3056

which matches his diagnosis for the species. Schulz

( 1 907) published this species as E. subglaucescens

and cited two specimens as syntypes: Gardner 3054

and 3056. I here select Gardner 3056 as the lec-

totype (lectotype, w; isolectotypes, B, destroyed, F

neg. 12648, BM, CGE, F, G, K, OXF, p), since it is

this specimen with nonstriate stipules that Schulz

described and placed in section Archerythroxylum.

Erythroxylum maracasense belongs to section

Rhabdophyllum O. E. Schulz and is most closely

related to E. pelleterianum St. Hil. These two

species may be distinguished using the characters

presented in Table 2.

4. Erythroxylum mattos-silvae Plowman, sp. nov.

sect. Rhabdophyllum O. E. Schulz. Figures

6-8, 10.

Frutex vel arbuscula gracilis. Ramuli pauci, patentes,

crassi. Cataphylla sparsa, remota, stipulis foliaribus si-

milia. Stipulae foliares transverse triangulares, coriaceae,

manifeste vel obscure striate nervosae, 2- vel 3-setulosae.

Folia in ramulis dispersa, manifeste petiolata, persisten-

tia; laminae amplae, oblongo-ellipticae vel oblongo-lan-

ceolatae, plerumque ellipticae vel lanceolatae vel oblan-

ceolatae, apice abrupte acuminatae, raro acutae vel

obtusae, basi acuminatae vel cuneatae, subcoriaceae,

ubique subnitidae. Flores e ramulis hornotinis vel an-

notinis in axillis foliorum vel cataphyllorum in axibus

brevibus nati. Pedicelli valde incrassati, intus spongiosi,

pallide virides vel ochracei, cum calyce conjuncti. Petali

lamina ovata vel oblonga, cremea vel ochracea, ligula

bilobata munita; ligula inter lobos appendice obconico-

clavata, apice complanata, papillosa provisa. Urceolus

stamineus calyce longior (x 1.5-2), margine eroso-cre-

nulatus vel 5-denticulatus. In floribus brachystylis styli

liberi; in floribus dolichostylis styli usque ad medium

connati. Drupa oblongo-ellipsoidea, apice rotundata. en-

docarpio tereti, maturitate uniloculari.

Slender shrub or spindly treelet to 6 m tall,

sparsely branched. Trunk slender, to 5 cm in di-

ameter. Bark in younger plants rather smooth, light

grayish brown, rather shiny, in older trees fissured

longitudinally and transversely in concentric rings,

dark grayish brown, ca. 3 mm thick, reddish with-

in, the wood reddish tan. Branches more or less

distant, spreading, in older trees occurring only

near top and arching over, spreading, straight, red-

dish to grayish brown, more or less shiny. Branch-

lets few, thick, straight, diverging 40°-80° from

axis, compressed toward apex, soon becoming ter-

ete, 3-4 mm in diameter, longitudinally wrinkled,

shiny, reddish brown becoming light to dark gray-

ish brown, with light brown, punctate lenticels that

do not break surface. In to modes 3-25 mm long.

Cataphylls (ramenta) sparse, 3-10 scattered along

base of new shoots, similar to the stipules, the

spinule (leaf rudiment) 1.2-1.5 mm long, black,

apically curving away from axis. Foliar stipules

persistent, distichous, diverging from stem up to

45°, transversely triangular, 2.5-3.5 mm long,

prominently or obscurely striate-nerved with 5-8

nerves per side, coriaceous, pale green, turning

dark brown on drying, the apex obtuse to truncate,

12

FIELDIANA: BOTANY

I nun

2mm

FIG. 6. Erythroxylum mattos-silvae. A, flowering branch; B, stipule; C, long-styled flower on short flowering axis,

one petal removed; D, long-styled flower, one petal removed, scale as in C; E, cross section of long-styled flower

showing thickened pedicel, scale as in C; F, "short-styled" flower showing same arrangement as long-styled flower,

scale as in C; G, petal; H, drupe attached to calyx; I, cross section of drupe; J, embryo, scale as in I. (B, C, D, E, and

G from Plowman et al. 13970; A, F, H, I, and J from Plowman et al. 13965.) Drawing by Marlene Werner.

PLOWMAN: NEW SPECIES OF ERYTHROXYLUM

13

14

FIELDIANA: BOTANY

2- or 3-setulose, the 2 lateral setae short, subulate,

0.5-0.8 mm long, the medial seta filamentous, to

1 mm long, often evanescent or entirely lacking,

the keels firm, slightly winged, the margin entire

or minutely erose. Leaves persistent, scattered along

younger branches, more or less distichous, petio-

late, the lamina held horizontally, plane or slightly

undulate, somewhat variable in shape, oblong-el-

liptic to oblong-lanceolate, sometimes elliptic,

lanceolate or oblanceolate, at apex abruptly short-

acuminate, rarely acute or obtuse, at base acu-

minate or cuneate, 90-270 mm long, 35-1 10 mm

wide, subcoriaceous or rubbery, opaque, adaxially

medium to dark green, somewhat shiny, drying

blackish green, abaxially light green, shiny, drying

dark green or ferruginous, elineate with no distinct

central panel, the adaxial midrib prominulous, flat

with a fine medial ridge or somewhat knife-edged,

pale green, the midrib on both surfaces drying fer-

ruginous, the lateral nerves 11-17 per side, di-

verging 55°-75° from midrib, straight, arcuately

anastomosing 5-10 mm from margin, adaxially

prominulous, abaxially faint to completely ob-

scure. Petiole 5-15 mm long, subterete, adaxially

canaliculate, drying dark brown or blackish. Flow-

ers fasciculate in axils of leaves or cataphylls on

current or year-old shoots; each inflorescence con-

sisting of 1-10 short, indeterminate axes to 5 mm

long and producing flowers sequentially. Brac-

teoles spiro-distichous, imbricate on flowering axes,

transversely ovate, concave, 1-1.5 mm long, red-

dish brown, paleaceous, longitudinally striate-

nerved, the apex acute to abruptly short-acumi-

nate, with a minute, apical seta 0.2-0.4 mm long.

Pedicel strongly thickened throughout its length

and fused with the calyx, 5-angled, pale green or

light yellow-ochre with pale green ribs, spongy-

parenchymatous within, the pedicel together with

calyx 3-3.5 mm long, 2.5 mm in diameter, elon-

gating somewhat in fruit. Calyx lobes coriaceous,

triangular to lanceolate, 0.5-1 mm long, the sur-

face minutely striate, the apex acute to short-acu-

minate, the tip itself thin, drying light brown. Petal

lamina spreading at anthesis, sometimes reflexed

and Ungulate, more or less plane, concave near

apex, ovate to oblong, 2-2.4 mm long, 1-1.3 mm

wide, ochreous in bud, greenish white to creamy

white or ochreous at anthesis, rounded or truncate

at apex, the claw 0.8-1 mm long, the ligule white,

1-1.2 mm long, bilobed, each lobe with a large

posterior auricle and a short anterior auricle re-

duced to a cuff at base of posterior one, the pos-

terior auricles ovate, 0.8-1 mm long, shallowly

3-lobed at apex, with a short, flaplike anterior ap-

pendage inflexed between the auricles and an erect,

fleshy appendage between and slightly behind them,

the posterior appendage obconic-clavate, flat and

papillose at apex, nearly equaling the auricles.

Staminal cup 1.5-2 times length of calyx, 1.3-2

mm long, greenish white, the margin finely erose

or with 5 short teeth alternating with antisepalous

filaments. Mesostylous(?) flowers: filaments in 2

series, the antisepalous filaments very short, su-

bulate, erect, produced from cup margin, 0.6-0.7

mm long, the anthers orbicular, 0.5-0.6 mm in

diameter; antipetalous filaments much longer,

slender, produced from inner margin of cup, 1.8-

2 mm long, inclined toward center, the anthers

orbicular, slightly smaller, 0.4-0.5 mm in diam-

eter; styles equaling or slightly longer than anti-

sepalous stamens, 1-1.5 mm long, free or connate

only at base; stigma minute, subcapitate, 0.3 mm

long. Dolichostylous flowers: antisepalous fila-

ments subulate, erect, produced from cup margin

itself, 0.5-0.7 mm long, the anthers orbicular, 0.5

mm in diameter; antipetalous filaments slender,

produced from inner margin of cup, 1.2-1.3 mm

long, inclined toward center, the anthers orbicular,

slightly smaller, 0.4 mm in diameter; styles 2.4-

2.6 mm long, connate for 0.5-1.3 mm; stigmas

minute, subcapitate, 0.2-0.3 mm long. Ovary ob-

ovoid or obconic, at apex rounded or truncate, l/2-

% times shorter than calyx, 1-1.5 mm long. Drupe

oblong-ellipsoid, rounded at apex, 11-13 mm long,

5-6 mm in diameter, crimson at maturity, the

mesocarp 0.7 mm thick, the endocarp ellipsoid in

cross section, terete, unilocular with no endo-

sperm. Embryo 10 mm long; cotyledons oblong-

ovoid, 9 mm long, 3.5 mm wide, 1 mm thick;

radicle 1.5 mm long.

TYPE— Brazil, Bahia, Municipio de Ilheus, ra-

mal novo para o povoado da Vila Brasil, com

entrada no km 28 da rodovia Ilheus-Una, inicio

do ramal (km 3), lado direito, regiao de mata li-

toranea, solo argilo-silicoso, arbusto, 3,5 m de al-

tura, flores esverdeadas, 13 Jan. 1985 (fl), L. A.

Mattos Silva, D. Daly & T. S. dos Santos 1817

(holotype, CEPEC 35860; isotypes, F 1943448, F

neg. 60194, HRB, MBM, NY, RB).

ADDITIONAL SPECIMENS EXAMINED— BRAZIL: Bahia:

Municipio de Ilheus: ramal que liga a Rodovia BR-415

(Ilheus-Itabuna) ao povoado de Japu, desvio a esquerda,

coleta a 2 km da entrada, Fazenda Sultao, 1 7 Feb. 1982

(fl), L. A. Mattos Silva, A. M. de Carvalho & T. S. dos

Santos 1570 (CEPEC, F), 26 Feb. 1985 (st), T. Plowman,

L. A. Mattos Silva & T. S. dos Santos 13958 (CEPEC, F,

NY, RB); Fazenda Barra do Manguinho, km 1 2 da rodovia

Pontal-Olivenca (BA-001), 27 Feb. 1985 (fl), T. Plow-

PLOWMAN: NEW SPECIES OF ERYTHROXYLUM

15

man, L. A. Mattos Silva & T. S. dos Santos 13970 (ALCB,

B, CEPEC, F, G, GH, HRB, IPA, K, M, MBM, MO, NY, P, RB,

SP, u, UB, UEC, us); ramal novo para o Povoado da

Vila Brasil, com entrada no km 28 da Rodovia Ilheus-

Una, lado direito, inicio do ramal (km 3), 27 Feb. 1985

(fl), T. Plowman, L. A. Mattos Silva & T. S. dos Santos

13960 (from the type tree) (CEPEC, F, K, NY, RB), inicio

do ramal (km 7), 27 Feb. 1985 (fl), T. Plowman, L. A.

Mattos Silva & T. S. dos Santos 13965 (CEPEC, F, HUEFS,

K, NY, RB, UB, us). Municipio de Una: estrada Ilheus-

Una, 19 Nov. 1983 (fl, young fr), R. Callejas, A. M. de

Carvalho & L. A. Mattos Silva 1755 (CEPEC, HBR, MBM,

NY, RB); estrada que liga a Rodovia BR 1 0 1 (Sao Jose)

com BA 215, a 17 km da entrada, 17 Jun. 1978 (fl), 5.

Mori, F. Benton & I. White 10192 (CEPEC, F, K, NY, RB);

20 km north along the road from Una to Ilheus, 39°02' W,

15°1 1'S, 23 Jan. 1977 (fl, fr), R. M. Harley et al. 18183

(AAU, CEPEC, F, K, MO, NY, P, RB, u); km 48 da estrada

Ilheus-Una, 6 Mar. 1983 (fl), A. M. de Carvalho & A.

Chautems 1646 (CEPEC, F, G, HRB, HUEFS, K, NY); Fazenda

Sao Rafael, 29 Oct. 1 969 (fl), T. S. dos Santos 450 (CEPEC,

F, NY); rodovia Una-Xapuri, Dendhevea S.A., Fazenda

Igua9ii, 29 Oct. 1971 (fl), R. S. Pinheiro 1682 (CEPEC, F).

ETYMOLOGY— It is my pleasure to dedicate this

species to Luiz Alberto Mattos Silva, curator of

the CEPEC herbarium at the Centre de Pesquisas

do Cacau, Ilheus, Bahia, Brazil. Mattos Silva has

made invaluable collections of the fast-disappear-

ing native flora of southern Bahia, including sev-

eral excellent collections of this species. He has

also provided assistance and kind hospitality for

myself and other botanists visiting southern Bahia.

DISTRIBUTION— Erythroxylum mattos-silvae is

confined to a small area along the coast of Bahia,

Brazil, from Ilheus south to Una.

ECOLOGY— This species apparently grows only

in the "restinga arborea," or restinga forest, a for-

est of low stature growing on sandy soils near the

coast. It has not been found in open restinga for-

mations. Just south of Ilheus, it grows in restinga

forest in association with the piacava palm (At-

talea funifera Mart.) in areas of disturbed or sec-

ondary vegetation. But further south along the new

road to Vila Brasil, this species occurs in a some-

what taller forest on mixed sandy/clay soil, which

possibly represents a transition from restinga to

the tropical moist forest of southern Bahia. How-

ever, E. mattos-silvae has not been found in typ-

ical "mata higrofila." The plants occur as erect,

almost columnar shrubs, or as tall, spindly trees

with thin trunks and with most of the branches

and leaves at the top. The habit may relate only

to the age of the plant or possibly to local edaphic

conditions. The plants are occasional or rare in

the few localities where they are known. Never-

theless, it is surprising that this remarkable species

was first collected only in 1969, having been over-

looked by the numerous botanists who have

worked around the city of Ilheus since the early

1800s.

PHENOLOGY— Most flowering collections of E.

mattos-silvae have been made between October

and March, but it was also found flowering in June.

Because the flowers in each cluster are produced

sequentially, it is likely that the flowering period

for a given individual or population is quite pro-

longed.

RELATIONSHIPS— In a preliminary list of Ery-

throxylum species in southern Bahian moist for-

ests (Mori et al., 1983), I identified one specimen

(Harley et al. 18183) of this species as Erythroxy-

lum aff. magnoliifolium St. Hil. of sect. Megalo-

phyllum O. E. Schulz. Closer observation of this

and subsequently collected material shows that

Erythroxylum mattos-silvae belongs to section

Rhabdophylum, based on its striately nerved stip-

ules, and is unrelated to E. magnoliifolium.

The ochre-colored pedicel, calyx, and petals of

E. mattos-silvae have been observed only in one

other species, E. tenue Plowman, which occurs

sympatrically with E. mattos-silvae and belongs

to the same section. The two species seem oth-

erwise only distantly related. The possible signif-

icance of this distinctive flower color for pollina-

tion is unknown.

With its relatively large subcoriaceous leaves

and small, subsessile flowers, E. mattos-silvae may

be confused with a number of other coastal Bahian

species of Erythroxylum. Comparisons of this

species with five others that superficially resemble

it are presented in Table 3.

In the relatively small sample of existing col-

lections of this species, two floral morphs were

found. The long-styled morph with long styles and

two shorter whorls of stamens is normal and com-

parable to long-styled flowers of most other Ery-

throxylum species. However, the second morph in

this species, observed in four collections, is in fact

a mid-styled morph with one whorl of stamens

shorter and one longer than the styles. This is very

unusual and has been reported only in one other

species, E. lucidum Moon (= E. acuminatum [Am.]

Walp.) from Sri Lanka (Burck, 1895). The mid-

styled floral morph may be only a structural anom-

aly; however, there is the possibility that a

short-styled morph also exists, in which case E.

mattos-silvae would be a tristylous species. Burck

(1895) suggested that Erythroxylum species are in

fact tristylous plants that are in evolutionary tran-

sition to distyly (Ganders, 1979). Further field-

16

F1ELDIANA: BOTANY

TABLE 3. Morphological comparison of Erythroxylum mattos-silvae Plowman, E. nobile O. E. Schulz, E. com-

pressum Peyr., E. membranaceum Plowman, E. martii Peyr., and E. grandifolium Peyr.

work is necessary to confirm or refute the existence

of a third-style morph and to determine the sig-

nificance of this deviation from the normal disty-

lous pattern of Neotropical Erythroxylum.

5. Erythroxylum membranaceum Plowman, sp.

nov. sect. Rhabdophyllum O. E. Schulz. Fig-

ures 9-10.

Frutex. Ramuli recti, lenticellis elongatis abundanter

producti. Cataphylla saepe congesta, stipulis foliaribus

similia. Stipulae foliares persistentes, parvae, manifeste

striate nervosae, 3-setulosae. Folia persistentia, petiola-

ta; laminae amplae, oblongae vel ellipticae, apice rotun-

datae vel retusae, basi late cuneatae, firme membrana-

ceae. Flores 1-3 e ramulis hornotinis in axillis

ramentorum nati, pedicellis tenuibus. Urceolus stami-

neus calycem aequans, margine 1 0-denticulatus, decem

glandulis rotundis parvis instructus. In floribus brachy-

stylis styli liberi. Drupa ovoideo-ellipsoidea, apice ob-

tusa, endocarpio oblongo-ellipsoideo, tereti, maturitate

uniloculari, endospermio nullo. Embryo magnus, coty-

ledonibus late ellipticis, apice rotundatis.

Shrub 3 m tall. Branches erect to spreading,

terete, dark brown. Branchlets erect to spreading,

straight, diverging 20°-60° from axis, compressed

at apex, 2 mm in diameter, dark reddish brown,

smooth, furnished with elongate, light tan lenti-

cels. Internodes 4-15 mm long. Cataphylls (ra-

menta) produced at base of shoots for 8-12 mm

along stem, often congested, diverging slightly from

axis, similar to foliar stipules, 1.5-2.5 mm long.

Foliar stipules persistent, appressed to stem, erect,

triangular to ovate, 2-2.5 mm long, firmly mem-

branaceous, distinctly striate-nerved with 6-8

nerves per side, at apex obtuse to acute, 3-setulose,

the medial seta 0.5 mm long, the lateral setae 0.3-

0.4 mm long, the keels prominent, entire, the mar-

gin entire. Leaves persisting 1 or 2 seasons, scat-

tered on branchlets, petiolate, the lamina plane,

oblong-elliptic, rounded at apex, sometimes brief-

ly retuse, broadly cuneate at base, 50-125 mm

long, 33-60 mm wide, firmly membranaceous,

drying grayish green adaxially, light brownish

abaxially, more or less shiny on both surfaces,

abaxially elineate with a faintly distinct central

panel demarcated by more prominent and regular

venation, the adaxial midrib furrowed, the abaxial

midrib prominent, drying dark reddish brown, the

lateral nerves 8-12 per side, diverging 50°-60° from

midrib, rather straight or arcuately curved, slightly

impressed adaxially, prominulous abaxially, the

veinlets finely reticulate, obscure adaxially, prom-

inulous abaxially. Petiole 5-6 mm long, 1-1.5 mm

in diameter, drying dark brown, subterete in cross

section, adaxially canaliculate. Flowers 1-3 in ax-

ils of last season's cataphylls, the flower color un-

known. Bracteoles persistent, broadly triangular,

concave, striate-nerved with 1 oblique keel and 2-

3 nerves per side, acute to rounded at apex, the

seta 0. 1-0.2 mm long. Pedicel slender, pentangular

and 5-ribbed, 3-7 mm long, 0.5 mm in diameter.

Calyx 1.5 mm long, divided '/2-% its length, the

lobes triangular-ovate, 0.9-1 mm long, acute to

acuminate at apex. Petals not seen. Staminal cup

equaling the calyx, 1 mm long, bearing 10 small,

round, surface glands alternating with the fila-

ments, the margin 1 0-denticulate. Brachystylous

flowers: filaments 2-2.3 mm long, the anthers not

seen; styles free, 1 mm long; stigma minute, de-

PLOWMAN: NEW SPECIES OF ERYTHROXYLUM

17

5mm

4mm

FIG. 9. Erythroxylum membranaceum. A, fruiting branch; B, stipule; C, detail of fruiting twig; D, endocarp; E,

cross section of endocarp, scale as in D; F, embryo, scale as in D. (All from the type, Belem & Magalhaes 932.)

Drawing by Pollyanna Quasthoff.

18

FIELDIANA: BOTANY

FIG. 10. Known geographical distribution of new Erythroxylum species in eastern Brazil. Solid stars = E. andrei;

solid circles = E. caatingae; open stars = E. maracasense; small, 8-pointed stars = E. mattos-silvae; solid square =

E. membranaceum.

PLOWMAN: NEW SPECIES OF ERYTHROXYLUM

19

pressed-capitate, 0.2 mm long. Dolichostylous

flowers: not seen. Ovary: not seen. Drupe ovoid-

ellipsoid, obtuse at apex, 9-10 mm long, 5-6 mm

in diameter, the mesocarp ca. 0.2 mm thick, the

endocarp ovoid, terete, obtuse or broadly acute at

apex, unilocular, with little or no endosperm. Em-

bryo 7.3 mm long; cotyledons broadly elliptic,

rounded at apex, subcordate at base, 6.7 mm long,

4.2 mm wide, 1.3 mm thick; radicle 1-1.5 mm

long.

TYPE— Brazil, Bahia, Municipio Aurelino Leal,

rodovia Ubaitaba-Lages [= Lage do Banco], 8 km

de Ubaitaba, mata virgem e perigosa [sic], planta

de 3 m de altura, frutos verdes, 24 Apr. 1965 (fr),

R. P. Belem & M. Magalhaes 932 (holotype, CEPEC

1057, F neg. 60130; isotypes, F 1622274, F neg.

60188, IAN 1 19424, NY, UB).

ETYMOLOGY— From Latin membranaceus

meaning "membranaceous," in reference to the

thin leaves of this species.

DISTRIBUTION— Southern Bahia, known only

from the type collection.

ECOLOGY— This species is apparently a local en-

demic in the moist forests of southern Bahia, where

at least 10 other endemic species ofErythroxylum

are known to occur (Mori et al., 1983).

PHENOLOGY— The sole collection was gathered

with fruit in late April.

RELATIONSHIPS— Erythroxylum membrana-

ceum belongs to section Rhabdophyllum O. E.

Schulz. It appears to be related to a number of

species of eastern Brazil in which the striately

nerved stipules are relatively small and persistent.

It differs from E. distortum Mart, and E. tenue

Plowman, both of which are endemic in the same

general area as E. membranaceum, in the much

larger leaves and stouter branchlets. Erythroxy-

lum compressum Peyr., known from moist forests

south of Ilheus, differs in the strongly compressed

branchlets, subsessile, oblong-lanceolate, apically

acuminate leaves, and a staminal tube much short-

er than the calyx. Erythroxylum affine St. Hil. non

A. Rich. (syn. E. blanchetii O. E. Schulz) is known

from Bahia (without locality) south to Espirito

Santo and Cabo Frio in the state of Rio de Janeiro.

It differs from E. membranaceum in having thick-

er, shorter (40-78 mm long vs. 50-125 mm long),

apically acuminate leaves, in which the adaxial

midrib is prominent and markedly acute in cross

section. In E. membranaceum the adaxial midrib

is a furrow sunken in the leaf surface. Erythroxy-

lum membranaceum is compared with other large-

leaved species from southern Bahian moist forests

in Table 3.

6. Erythroxylum petrae-caballi Plowman, sp. nov.

sect. Rhabdophyllum O. E. Schulz. Figures

11, 18.

Frutex vel arbor parva, ramulis rectis vel flexuosis,

tegumento cereo obtectis et lenticellis instructis. Cata-

phylla pauca, disticha, stipulis foliaribus similia. Sti-

pulae foliares persistentes, parvae, triangulari-ovatae,

striate nervosae, breviter 3-setulosae. Folia parva, bre-

viter petiolata, decidua; laminae ellipticac vel oblongae,

raro lanceolatae, apice obtusae vel rotundatae, mucro-

nulatae, basi acutae vel obtusae, chartaceae, supra niti-

dae, subtus impolitae. Flores e ramulis annotinis vel

hornotinis in axillis foliorum vel ramentorum nati. Petali

lamina oblongo-ovata, ligula bilobata munita. Urceolus

stamineus quam calyx dimidio brevior, margine integer.

Styli liberi. Drupa endocarpio ampullaceo, tereti, uni-

loculari, endospermio nullo, embryone magno et crasso

pro visa.

Shrub 1.5-3 m tall or tree to 9 m tall, densely

or openly branched. Branchlets more or less dis-

tichous, undifferentiated into short shoots, di-

verging 40°-70° from axis, more or less flexuous,

compressed at apex when young, 1-1.5 mm in

diameter, smooth, reddish brown, soon becoming

covered with a thin, whitish wax coating which

sloughs off with age, the lenticels punctate or elon-

gate, whitish, the bark becoming light brown, lon-

gitudinally striate with age. In tern odes 0.5-15 mm

long. Cataphylls (ramenta) distichous, produced

at base of new twigs, few, scattered, in form similar

to foliar stipules, rarely more developed, overlap-

ping and covering stem for 3-10 mm, 2.5 mm

wide, the spinule (leaf rudiment) 0.8-2 mm long,

flattened, black, sharp-attenuate at apex. Foliar

stipules persistent, appressed to stem, diverging

only slightly with age, triangular-ovate, 1.7-2.5

mm long, subcoriaceous, striate-nerved with 4-6

nerves per side, light green, turning ferruginous in

age, obtuse or acute at apex, 3-setulose, the setae

thin-filamentous, turning black and spreading, 0.2-

0.7 mm long, the medial seta a little longer or

shorter than 2 equal lateral ones, the setae eva-

nescent, the keels sub-alate, the margin erose.

Leaves deciduous or briefly persistent, scattered

along branchlets, distichous, short-petiolate, the

lamina plane, elliptic to oblong, rarely lanceolate,

obtuse to rounded at apex, mucronulate, acute to

obtuse at base, 20-65 mm long, 10-30 mm wide,

chartaceous, adaxially drying gray-green, shiny,

abaxially drying very pale green to ochreous, matte,

elineate with no distinct central panel, the adaxial

midrib shallowly furrowed, rarely flat, the abaxial

midrib drying straw-colored or ochreous, the lat-

eral nerves 9-12, diverging 45°-80° from midrib,

20

FIELDIANA: BOTANY

I cm

1 mm

B

FIG. 1 1 . Erythroxylum petrae-caballi. A, flowering branch; B, leaf showing venation; C, stipule, scale as in E; D,

short-styled flower; E, petal; F, petal, scale as in E; G, drupe, scale as in H; H, cross section of endocarp; I, embryo,

scale as in H. (A, B, D, and E from Grupo Pedra do Cavalo 967; C and F from Grupo Pedra do Cavalo 987; G, H,

and I from Noblick & Hahn 3361.).

PLOWMAN: NEW SPECIES OF ERYTHROXYLUM

21

rather straight or crooked, anastomosing 1-4 mm

from margin, scarcely evident on either surface.

Petiole 2-4 mm long, 0.5-0.7 mm in diameter,

suberect, broadly canaliculate. Flowers 1-3, pro-

duced sequentially in axils of cataphylls or leaves

on current or last season's twigs, rather sparse,

"yellow." Bracteoles transversely ovate or trans-

versely triangular, concave, 0.8-1 mm long, ob-

tuse or acute at apex, the seta 0.1-0.3 mm long.

Pedicel slender, 2-1 1 mm long, 0.5 mm in di-

ameter, subterete or pentagonal, 5-ribbed, in fruit

sometimes articulating with a short, persistent pe-

duncle 0.5-1.5 mm long. Calyx 1.3-1.8 mm long,

divided '/z-3/! its length, the lobes ovate to trian-

gular, 0.7-1 . 1 mm long, acute to attenuate at apex,

the tip itself drying dark brown. Petal lamina ob-

long-ovate, concave and rounded at apex, 1.2-1.5

mm long, 0.8-1 mm wide, the claw 0.5-1 mm

long, the ligule with 2 simple, ovate auricles, 0.5-

0.6 mm long, with a short, sometimes inflated

posterior-medial lobe and a broad, truncate, an-

terior flap inflexed between the auricles. Staminal

cup l/2 the length of calyx, 0.6-0.8 mm long, the

margin entire. Brachystylous flowers: all filaments

subequal, ca. 1.5 mm long; or antisepalous fila-

ments shorter, 1-1.2 mm long, the antipetalous

filaments 1.1-1.7 mm long; anthers orbicular, 0.4

mm long; styles free, 0.5-0.7 mm long; stigma

depressed-capitate, 0.2 mm long. Dolichostylous

flowers: not seen. Ovary ovoid or ellipsoid, round-

ed at apex, 1.5-1.8 times length of staminal cup,

0.8-1.2 mm long. Drupe flask-shaped, 9 mm long,

5.5 mm in diameter, red, the mesocarp ca. 0.3 mm

thick, the endocarp ovoid, flask-shaped, acute at

apex, terete, smooth, unilocular, lacking endo-

sperm. Embryo 6.5-8 mm long; cotyledons ellip-

tic, rounded at apex, cordate at base, 5.5-7 mm

long, 4.5-5 mm wide, 1.5 mm thick; radicle 1.4

mm long.

TYPE— Brazil, Bahia, Cachoeira, Vale dos Rios

Paragua9u e Jacuipe, ca. 39°05'W, 1 2°32'S, alt. 40-

120 m, Morro Belo, arvore con cerca de 9 m de

altura; folhas eliticas e com frutos, Dec. 1980 (fl,

imm fr), Grupo Pedra do Cavalo (Scardino, Nob-

lick, Paranhos, Guedes, Queiroz, Paganucci, Nas-

cimento, Suely, Quaglia) 967 (holotype, CEPEC

26980; isotypes, ALCB 8103, BAH 2984, BOTU, F,

HRB 7253, HUEFS 1026, MBM, RB, SP).

ADDITIONAL SPECIMENS EXAMINED— BRAZIL: Bahia:

Municipio de Cachoeira: Cachoeira, Vale dos Rios Par-

aguacu e Jacuipe, ca. 39°05'W, 12°32'S, alt. 40-120 m,

Esta<;ao da Mata, Jun. 1980 (imm fr), Grupo Pedra do

Cavalo 293 (ALCB), same locality, Jun. 1980 (fl bud),

Grupo Pedra do Cavalo 811 (ALCB, BAH, BOTU, CEPEC, F,

HRB, HUEFS, RB, SP); Morro Belo, Aug. 1980 (fl), Grupo

Pedra do Cavalo 504 (ALCB, BAH, BOTU, CEPEC, F, HRB,

MBM, RB), same locality, Dec. 1980 (fl bud), Grupo Pedra

do Cavalo 987 (ALCB, BAH, CEPEC, HRB, HUEFS). Muni-

cipio de Feira de Santana: Serra de Sao Jose, Fazenda

Boa Vista, 12°15'S, 38°58'W, 24 May 1984 (fr), L. R.

Noblick 3271 (CEPEC, F, HUEFS), same locality, 9 Jun.

1984 (fr), L. R. Noblick & W. J. Hahn 3361, (CEPEC, F,

HUEFS, K, MO).

ETYMOLOGY— The specific epithet "petrae-ca-

balli" is a Latin rendering (genitive form) of the

Portuguese Pedra do Cavalo, "horse's rock." This

is the name taken by the group of botanists who

collected most of the material of this species in

the area of the new hydroelectric dam on the Rio

Paragua9u known as Pedra do Cavalo. The name

Pedra do Cavalo originally referred to a rock out-

crop, now destroyed, that extended into the Rio

Paragua9u, where cowboys used to take their horses

to drink and to be washed (Geraldo Pinto and

Larry Noblick, in litt.). The Pedra do Cavalo col-

lectors, based at the Universidade Federal da Ba-

hia in Salvador and headed by Lecticia Scardino

Scott Faria, have conducted a floristic-ecological

study of the area to be inundated by the Pedra do

Cavalo Dam.

DISTRIBUTION— Erythroxylum petrae-caballi is

known only from the municipalities of Feira de

Santana and Cachoeira in eastern Bahia.

ECOLOGY— This species grows in seasonally dry,

deciduous forest, especially in forest clearings and

along small seasonally dry creeks in the forest.

Larry R. Noblick of the Universidade Estadual de

Feira de Santana supplied the following descrip-

tions of the habitats of this species in two areas

where it has been collected.

1) Serra de Sao Joao, ca. 22 km north northwest

of Feira de Santana: "This is a seasonal, deciduous

forest that usually undergoes a definite dry season,

especially from the end of September through Jan-

uary. Within this forest, E. petrae-caballi grows

preferentially in the valleys of temporary streams.

Annual rainfall is around 800 mm. The main for-

est is on a north-facing slope. The forest canopy

is about 20-25 m tall. Some dominant tree species

are Cavanillesia arborea, Cnidoscolus marcgravii,

Goniorrhachis marginata, Cedrela odorata, Gal-

lesia gorazema, and Chorisia sp. The shrubby

understory includes, in addition to Erythroxylum

petrae-caballi, Urera baccifera, Cordia superba,

and species of Phyllanthus, Psychotria, and Clavi-

ja. The herbaceous layer includes the grass Raddia

portoi and species of Tripogandra, Tinantia, and

Dichorisandra of the Commelinaceae. A few cac-

tus species are scattered here and there; scattered

22

FIELDIANA: BOTANY

TABLE 4. Morphological comparison of Erythroxylum petrae-caballi Plowman, E. tenue Plowman, E. distortum

Mart., E. cf. mikanii Peyr., and E. passerinum Mart.

ferns are found on the moister upper slopes. The

soil is shallow and extremely rocky; the soil itself

is sandy mixed with a significant amount of hu-

mus."

2) Pedra do Cavalo area: "The forests of the

Pedra do Cavalo area are essentially the same.

Morro Belo contains a seasonal, deciduous forest

with Cavanillesia arborea as one of the most con-

spicuous tree species. The 'Estasao de Mata' lo-

cality is perhaps a bit more humid since it is closer

to the coast and with soil containing a bit more

clay. Nevertheless, Cavanillesia arborea was pres-

ent here, as were Cnidoscolus marcgravii and

Erythrina velutina, etc.

"All of these areas have the same type forest:

seasonally deciduous to semi-deciduous with a

mixture of tree species from the Atlantic coastal

forest and the caatinga" (L. Noblick, in litt.).

PHENOLOGY— Flowering specimens have been

collected in June, August, October, and Decem-

ber; fruiting specimens, in May and June.

RELATIONSHIPS— Erythroxylum petrae-caballi

belongs to sect. Rhabdophyllum. It resembles sev-

eral other species in this large and diverse section;

these species form a poorly defined, complex group

whose center of diversity is the state of Bahia. They

all have small, relatively thin leaves, small, striate-

ly nerved stipules, and small flowers produced on

the new twigs. The species differ mainly in the size

and form of stipules and number of setae, form of

cataphylls, leaf persistence, shape and form of the

adaxial midrib, relative lengths of calyx and sta-

minal tube, form of the petal ligule, connation of

the styles, and form of the endocarp. In spite of

striking similarities in one or more of these char-

acters, the species of this group usually are eco-

logically specialized in different vegetation types,

soils, and microclimates.

Three species that appear most closely related

to E. petrae-caballi are E. distortum Mart., E. te-

nue Plowman (described below), and E. mikanii

Peyr. The last species is positively known only

from the sterile unicate type (preserved at w) col-

lected by Mikan in the state of Rio de Janeiro at

"Tocaja" (cf. Mikan, 1837). "Tocaja" is an old

spelling corresponding to Rio Itocaia, Pedra de

Itaocaia, or Itaocaia, all of which are located near

the present-day town of Itaipu-Acu. This is an area

of restinga and restinga forest, now largely de-

stroyed for beachfront housing, on the coast south-

east of the city of Niteroi.

There is a species locally common in coastal

Bahian restinga forests (cf. Araujo 22, Plowman &

Carvalho 12817, Plowman et al. 13959, 13969)

that closely resembles E. mikanii, but its identity

has not been established with certainty. This plant

also may prove to be a distinct undescribed species.

Erythroxylum petrae-caballi is compared to these

species (including the Bahian plants of E. cf. mi-

kanii) in Table 4.

PLOWMAN: NEW SPECIES OF ERYTHROXYLUM

23

1 mm

1 mm

D

FIG. 12. Erythroxylum santosii. A, flowering branch, 3-cm scale; B, detail of flowering axillary short-shoot; C,

leaf showing venation, 3-cm scale; D, short-styled flower; E, petal; F, stipule. (All from the type, dos Santos 1618.)

Drawing by Marlene Werner.

24

FIELDIANA: BOTANY

7. Erythroxylum santosii Plowman, sp. nov. sect.

Archerythroxylum O. E. Schulz. Figures 1 2,

18.

Frutex. Cataphylla pauca, stipulis foliaribus similia.

Stipulae foliares persistentes, late ovatae, subcoriaceae,

estriatae, brunnescentes, 2(-3)-setulosae. Folia breviter

petiolata; laminae oblongae vel lanceolatae vel oblan-

ceolatae, apice acutae vel breviter acuminatae, ipso apice

obtusae, basi cuneatae vel obtusae, firme chartaceae,

opacae. Flores e ramulis hornotinis in axillis cataphyl-

lorum nati, pedicellis brevibus, recurvatis. Petali lamina

oblonga. Urceolus stamineus tribus quintis partibus lon-

gitudinis calycis aequalis, margine irregulariter 10-den-

ticulatus. Styli liberi. Drupa ignota.

Shrub 2 m tall. Branchlets weakly differentiated

into long and short shoots, diverging 40°-80° from

axis, somewhat compressed at apex, 2 mm in di-

ameter, light brown, smooth, becoming lightly

suberose-verruculose but without lenticels break-

ing surface. Ink-modes on long shoots 5-25 mm

long, on short shoots 1-1.5 mm long. Cataphylls

(ramenta) 3-4, scattered along base of long shoots

or congested at tips of short shoots, similar to foliar

stipules, the spinule (leaf rudiment) 1.5-2 mm long,

black. Foliar stipules persistent, appressed to stem,

diverging slightly in age, broadly ovate, 3-4 mm

long, subcoriaceous, smooth, nonstriate, drying

dark brown or black, rounded at apex, 2(-3)-set-

ulose, the 2 lateral setae 0.5-1 mm long, some-

times with a short, medial seta to 0.5 mm long,

evanescent, the keels sub-alate, the margin entire.

Leaves persistent, scattered along long shoots or

1 (2) produced at tip of short shoots, distichous,

short-petiolate, the lamina plane, oblong, lanceo-

late or oblanceolate, acute or short-acuminate at

apex, the apex itself blunt, cuneate to obtuse at

base, 40-90 mm long, 20-35 mm wide, firmly

chartaceous, opaque, drying adaxially chestnut-

brown, abaxially ochreous to ferruginous, dull on

both surfaces, elineate with no distinct central

panel, the adaxial midrib yellowish brown with a

slender, slightly raised, knife-edged medial ridge,

the lateral nerves 9-11, diverging 50°-80° from

midrib, curved and arching upward, anastomosing

3-4 mm from margin, the veinlets obscure. Petiole

2-3 mm long, 1 mm in diameter, amply canalic-

ulate. Flowers in axils of cataphylls on mature ex-

tension shoots or short shoots, the petals yellow.

Bracteoles lanceolate in outline, cymbiform, 1.5

mm long, acute at apex, the seta 0.5-0.7 mm long.

Pedicel recurved, 5-angled, 1.5-2 mm long, 0.8

mm in diameter, slightly thickened at apex. Calyx

2.5 mm long, divided nearly to base, drying black,

the lobes lanceolate, 2 mm long, narrowly acute

at apex. Petal lamina somewhat concave, oblong,

rounded at apex, 2.5 mm long, 1.5 mm wide, the

claw 1.5 mm long, the ligule 1 mm long, bilobed

with a pair of suborbicular auricles 0.8 mm long,

with an erect, medial lobe or flap, nearly equaling

the auricles between and immediately posterior to

the auricles. Stamina! cup % as long as calyx, 1.5

mm long, drying dark brown with 1 0 light-colored,

longitudinal markings, the margin minutely and

irregularly 1 0-denticulate. Brachystylous flowers:

filaments white, 2.5 mm long, the anthers subor-

bicular, 0.5-0.6 mm long; ovary equaling staminal

cup, ovoid, 1.2 mm long, at apex rounded-trun-

cate; styles free, 1.7 mm long; stigmas depressed-

capitate, 0.4 mm long. Dolichostylous flowers: un-

known. Drupe: unknown.

TYPE— Brazil, Bahia, Vale do Rio Alcobaca

[= Rio Itanhem], Teixeira de Freitas, arbusto de

2 m de alt, flor amarelada, estames brancos, mata,

12 May 1971 (fl), T. S. dos Santos 7<57S(holotype,

CEPEC 6910; isotype, F 1839581, F neg. 60125).

ETYMOLOGY— Erythroxylum santosii is named

for Talmon S. dos Santos, who collected the type

and only known specimen of the species. Dos San-

tos, who works at the CEPEC herbarium, is an ex-

cellent collector who has been instrumental in doc-

umenting the flora of southern Bahia. His keen eye

and his intimate knowledge of Bahia forests have

resulted in discovery of numerous previously un-

known species. A number of these, like Erythroxy-

lum santosii, are apparently quite rare and have

not been found by other collectors. He has also

been an able assistant and amiable field compan-

ion to many botanists visiting southern Bahia.

DISTRIBUTION— Known only from the type col-

lection.

ECOLOGY— This species grows in the understory

of the tropical moist forest in the Itanhem (for-

merly Alcobaca) River valley in southern Bahia.

PHENOLOGY— The only known collection was

flowering in May.

RELATIONSHIPS— Erythroxylum santosii be-

longs to section Archerythroxylum and is clearly

related to E. ochranthum Mart., a rarely collected

species known only from coastal Bahia. Ery-

throxylum santosii can be separated from E.

ochranthum using characters listed in Table 5.

8. Erythroxylum splendidum Plowman, sp. nov.

sect. Megalophyllum O. E. Schulz. Figures

13-15, 18.

Frutex vel arbuscula gracilis, virgata. Ramuli recti,

adscendentes. Cataphylla sparsa, remota, stipulis foli-

PLOWMAN: NEW SPECIES OF ERYTHROXYLUM

25

TABLE 5. Morphological comparison of Erythroxylum santosii Plowman and E. ochranthum Mart.

Character

/ . santosii

E. ochranthum

Leaf shape

Leaf surface

Pedicel shape

Pedicel length (mm)

Calyx lobe shape

Ovary vs. staminal cup

Oblong, lanceolate, or oblanceolate

Dull, opaque

Recurved

1.5-2

Lanceolate

Equal

Elliptic to oblong

Somewhat shiny, translucent

Straight

2.5-8

Ovate

Twice as long

aribus similia. Stipulae foliares persistentes, triangulari-

ovatae, estriatae, 2-setulosae. Folia in ramulis dispersa,

disticha, petiolata, persistentia; laminae ovatae vel ob-

longae vel lanceolato-ovatae vel ellipticae, apice obtusae

vel rotundatae, basi breviter acuminatae, coriaceae,

ubique nitidae. Flores e ramulis hornotinis vel annotinis

in axillis foliorum vel cataphyllorum nati. Petali lamina

elliptica vel late ovata, ligula bilobata munita. Urceolus

stamineus tribus quartis partibus longitudinis calycis ae-

qualis vel calycem aequans, margine integer. In floribus

brachystylis styli liberi. Drupa ellipsoidea, parum fal-

cata, apice rotundata, endocarpio paulo compresso,

manifeste curvato-falcato, triquetro, triloculari, loculis

vacuis duobus magnis, loculo fertili parvo, endospermio

abundante.

Slender, virgate shrub or treelet, 1-5 m tall.

Trunk solitary, to 2 cm in diameter. Bark smooth,

longitudinally finely striate, light brown, ca. 1 mm

thick; wood light tan. Branches all near top on

older individuals, ascending, straight, brown,

"knobby" at the nodes. Branchlets distichous,

straight, often parallel, diverging 50°-70° from axis,

compressed when very young, soon becoming ter-

ete, 1.5-2 mm in diameter, dark reddish brown,

furnished abundantly with pale tan, punctate or

elongate lenticels, the branchlets becoming "stub-

by" at apex through abscission of terminal bud.

Internodes 5-20 mm long. Cataphylls (ramenta)

3-6, sparse, distichous, scattered along base of new

shoots, similar to foliar stipules, the spinule (leaf

rudiment) 0.8 mm long, erect, straight, black. Fo-

liar stipules persistent, erect, straight, triangular-

ovate, 1.5-2.3 mm long, subcoriaceous, smooth,

nonstriate, drying dark brown or black, obtuse at

apex, 2-setulose, the setae filamentous, 0.3-0.6 mm

long, evanescent, the keels prominulous, sub-alate,

the margin straight, entire. Leaves persistent, scat-

tered along younger branches, distichous, petio-

late, the lamina plane or somewhat undulate, ovate

to oblong, lance-ovate or elliptic, obtuse to round-

ed at apex, rarely retuse, short-acuminate at base,

40-105 mm long, 20-45 mm wide, subcoriaceous

to coriaceous, adaxially shiny, dark green, abaxi-

ally less shiny, lighter green, drying ferruginous on

both surfaces, elineate with no distinct central

panel, the adaxial midrib flat, rarely with a fine

medial ridge, the lateral nerves 8-12, diverging

50°-70° from midrib, straight, eventually arching

and anastomosing 2-3 mm from margin, promi-

nulous on both surfaces, the veinlets obscure. Pet-

iole 5-15 mm long, 1-1.2 mm in diameter, sub-

terete. Flowers fasciculate in axils of leaves or

cataphylls on new and seasoned shoots, 1-6 flow-

ers produced per node, 1-3 flowers at anthesis at

each flowering node per day. Bracteoles persisting,

giving nodes a "knobby" appearance, orbicular,

concave, 1.2-1.8 mm long, paleaceous, rounded

at apex, the seta 0.2-0.5 mm long. Pedicel short,

thick, 5-angled or 5-ribbed, 1-2 mm long, 1 mm

in diameter. Calyx 1.5-2.5 mm long, divided to

near base, pale green, the lobes strongly incurved,

narrowly ovate to lanceolate, 1.5-2.2 mm long,

narrowly acuminate at apex. Petal lamina suberect

or spreading at anthesis, creamy white or greenish

white, concave, elliptic to broadly ovate, rounded

at apex, the apex incurved, 1.5-3 mm long, 1.2-

1.5 mm long, the claw 0.8-1.2 mm long, the ligule

bilobed, 0.7-1.5 mm long, the posterior auricle

well developed, facing sideways, ovate, 0.5-1.2

mm long, entire, the anterior auricle greatly re-

duced or absent. Staminal cup 3/4 as long as or

equaling the calyx, 1-1.5 mm long, entire at mar-

gin. Brachystylous flowers: filaments 2-3 mm long,

the anthers suborbicular, 0.4—0.5 mm in diameter;

styles 1-1 .5 mm long, connate 0.3-0.5 mm; stigma

depressed-capitate, 0.2-0.3 mm long. Dolicho-

stylous flowers: antisepalous filaments 0.8-1 mm

long, the anthers orbicular, 0.45-0.50 mm in di-

ameter; antipetalous filaments 1.5-2 mm long, the

anthers orbicular, 0.40-0.45 mm long; styles 2.2-

2.3 mm long, connate at base 0.6-1 mm; stigma

subglobose, 0.2-0.4 mm long. Ovary ellipsoid,

shallowly 3-sulcate, rounded at apex, a little short-

er to a little longer than staminal cup, 1.2-1.8 mm

long. Drupe ellipsoid, somewhat falcate, rounded

at apex, 10-13 mm long, 6 mm in diameter, shiny,

26

FIELDIANA: BOTANY

FIG. 13. Erythroxylum splendidum. A, flowering branch; B, detail of apex of flowering twig; C, detail of fruits on

twig; D, stipule, scale as in G; E, short-styled flower; F, long-styled flower, scale as in E; G, petal; H, cross section

of drupe, scale as in B; I, embryo. (A, B, D, E, and G from Belem & Pinheiro 2466; C, H, and I from Mattos Silva

et al. 1262; F from Belem & Pinheiro 3170.) Drawing by Pollyanna Quasthoff.

PLOWMAN: NEW SPECIES OF ERYTHROXYLUM

27

28

FIELDIANA: BOTANY

reddish orange at maturity, the mesocarp 0.5-0.7

mm thick, the endocarp 9-12 mm long, com-

pressed, strongly curved-falcate, unequally trigo-

nal with 2 shorter obtuse lobes and 1 longer, nar-

rowly acute lobe, acute at apex, 3-locular with 2

larger, empty locules and 1 small, fertile locule;

endosperm occupying ca. 80% of fertile locule.

Embryo 7.2-9.5 mm long; cotyledons linear-lan-

ceolate, somewhat falcate, 5.5-7 mm long, 1 mm

wide, 0.1-0.2 mm thick; radicle 2-2.5 mm long,

0.5 mm in diameter.

TYPE— Brazil, Bahia, Municipio de Valenca, km

9 na estrada Valenfa-Guaibim, approx. 13°18'S,

39°00'W, arboreal restinga on white sand near sea

level; slender treelet 5 m tall growing in disturbed

restinga, flowers creamy white, 6 Feb. 1983 (fl, fr),

T. Plowman & A. M. de Carvalho 12818 (holo-

type, CEPEC 32439; isotypes, F, F neg. 60186, G,

HRB, K, NY, RB, SP, UB, US).

ADDITIONAL SPECIMENS EXAMINED— BRAZIL: Bahia:

Municipio de Valenca: ramal a esquerda da rodovia que

liga Valenca ao Guaibim (literal), com entrada no km

9, 1 1 Dec. 1980 (fr), L. A. Mattos Silva, A. M. de Car-

valho & J. L. Hage 1262 (CEPEC, F, HRB, HUEFS, MBM,

RB), 6 Feb. 1983 (st), T. Plowman & A. M. de Carvalho

1281 9 (CEPEC, F, K, RB), same locality, km 8, 27 Jul. 1981

(fl, fr), A. M. de Carvalho & J. Gatti 815 (CEPEC, F, HRB,

K, NY, RB), same locality, km 10, 8 Jan. 1982 (fl, fr), A.

M. de Carvalho & G. P. Lewis 1 125 (ALCB, CEPEC, F, SPF),

same locality, km 15, 25 Feb. 1986 (fr), J. L. Hage, L.

Andersson & M. Hagberg 1953 (CEPEC, F), 7955 (CEPEC,

F). Municipio de Cairu: rodovia Nilo Pecanha-Cairu,

km 14-18, 29 Apr. 1980 (fr), T. S. dos Santos, L. A.

Mattos Silva & E. B. dos Santos 3594 (CEPEC, F). Mun-

icipio de Marau: Marau, 1 8 Jan. 1 967 (fl, fr), R. P. Belem

& R. S. Pinheiro 3170 (CEPEC, F, NY, UB). Municipio de

Itacare: 65 km NE of Itabuna at mouth of the Rio de

Contas on the N bank opposite Itacare, 39°00'W, 14°16'S,

sea level, 30 Jan. 1977 (fl), R. M. Harley et al. 18417

(CEPEC, F, K..). Municipio de Belmonte: Belmonte, 30 Jun.

1966 (fl), R. P. Belem & R. S. Pinheiro 2466 (CEPEC, F,

IAN, NY, UB); ca. 26 km SW of Belmonte along road to

Itapebi and 4 km along side road toward the sea, ca.

39°02'W, 16°03'S, alt. sea level, 25 Mar. 1974 (fr), R.

M. Harley 17402 (CEPEC, K, NY, RB). Without exact lo-

cality: ca. 1832, J. S. Blanchet (BM, G, F neg. 26398,

photos F, GH, NY, distributed as "£. grandifolium").

ETYMOLOGY— From Latin splendidus, meaning

"shining" or "splendid," in reference to the hand-

some, glossy leaves of this species.

DISTRIBUTION— Erythroxylum splendidum oc-

curs only along the coast of Bahia, Brazil, from

Valenca south to Belmonte, covering a distance of

only 1 50 km.

ECOLOGY— This species grows only in arboreal

or shrubby restinga vegetation ntar the sea on sandy

soil. It appears to be nowhere very abundant, but

some large areas of the restingas of Bahia remain

to be explored botanically.

PHENOLOGY— Erythroxylum splendidum has

been collected in flower or fruit every month from

December to July, but no collections have been

made between August and November. This may

reflect a true flowering period or merely a collect-

ing accident.

RELATIONSHIPS— The first collection of E. splen-

didum was made by Blanchet in Bahia about 1 832;

the plant was not collected again until 1 966. Schulz

(1907) identified the Blanchet specimen, which has

abnormally pointed leaves, as E. grandifolium

Peyr., and this is indeed the closest relative of E.

splendidum. Erythroxylum splendidum differs from

E. grandifolium primarily in having dark reddish

brown (vs. light tan or whitish) bark on the twigs,

much smaller (40- 105 mm vs. 1 10-250 mm long),

ovate to oblong (vs. elliptic) leaves with obtuse or

rounded (vs. acuminate) apices.

Schulz ( 1 907) placed E. grandifolium in section

Megalophyllum O. E. Schulz. This group includes

four species from southeastern Brazil with large

leaves, nonstriate stipules, and numerous, glom-

erate, sessile flowers with more or less connate

styles. Three of the species are endemics from Rio

de Janeiro. Schulz thought that E. grandifolium

grew in Rio and Bahia, and noted that the Bahian

specimen of Blanchet had much smaller leaves.

The type of E. grandifolium is Riedel 774 collected

at "Esperanca," a locality reported to be in the

state of Rio de Janeiro (Peyritsch, 1878). How-

ever, this species has never been recollected in the

state of Rio, but is known from several collections

from southern Bahia, where most probably it is

endemic in the coastal forests. "Esperanca" is most

likely an old name for a fazenda or farm in the

area around Ilheus where Riedel spent consider-

able time.

One of the most distinctive features ofE. gran-

difolium and E. splendidum is the endocarp which

has one small, fertile locule and two larger, empty

locules. This character is found in another species

with nonstriate stipules from Bahia, E. caatingae

(and probably in the related E. oxypetalum O. E.

Schulz) of section Archerythroxylum. However,

trilocular endocarps are found also in some species

with striate-nerved stipules of section Rhabdo-

phyllum (cf. E. laetevirens O. E. Schulz and E.

steyermarkii Plowman) as well as in some Old

World species (Schulz, 1907).

PLOWMAN: NEW SPECIES OF ERYTHROXYLUM

29

1)

FIG. 16. Erythroxylum stipulosum. A, leafless flowering branch, 3-cm scale; B, sterile branch with mature leaves,

3-cm scale; C, detail of flowering twig; D, stipule; E, long-styled flower; F, short-styled flower, scale as in E. (A, C,

and D from Luetzelberg 26459; B from Black 55-17977; E from Luetzelberg 26187; F from Luetzelberg 26464.)

Drawing by Pollyanna Quasthoff.

30

FIELDIANA: BOTANY

FIG. 17. Habitat of Erythroxylum stipulosum at Chapada de Araripe, Ceara (Plowman & Copula 12716). Ery-

throxylum stipulosum is marked with arrow. Erythroxylum vacciniifolium is tall, leafy shrub forming background

(Plowman & Cagula 12717). The man is Jose "Arnaldo" Tavares Ca9ula, resident guard of the Parque Nacional de

Araripe.

9. Erythroxylum stipulosum Plowman, sp. nov.

sect. Rhabdophyllum O. E. Schulz. Figures

16-18.

Frutex dense ramosus, ramulis nigrescentibus, longi-

tudinaliter striate fissuratis. Cataphylla conspicua, con-

gesta, disticha, paleacea, stipulis foliaribus similia. Stip-

ulae foliares oblongo-lanceolatae, striate nervosae, longe

3-setulosae, margine manifeste villoso-fimbriatae, mar-

cescentia. Folia brevissime petiolata, decidua; laminae

ellipticae, apice et basi rotundatae, firme chartaceae,

opacae. Flores solitarii e ramulis annotinis in axillis ca-

taphyllorum nati. Urceolus stamineus calyce dimidio

brevier, margine integer. Styli liberi. Drupa ignota.

Much-branched, erect shrub to 3 m tall with

single trunk to 2 cm diameter. Bark smooth, dark

brown, longitudinally striate, ca. 1 mm thick, red-

dish brown within, the wood light tan. Branches

short, ascending and spreading, light brown.

Branchlets dense, rather straight, diverging 55°-

90° from axis, compressed at apex when new, more

or less longitudinally wrinkled, becoming terete,

1-1.2 mm in diameter, black or dark brown,

smooth, sometimes with a few, elongate, light tan

lenticels, becoming longitudinally striately fis-

sured. Internodes 3-20 mm long. Cataphylls (ra-

menta) large, conspicuous, congested, at apex of

twigs forming a terminal bud, covering stem

through 5-15 mm, 4-6 mm wide, diverging from

stem up to 30°, distichous, chaffy, light brown,

persisting but marcescent, similar in form to foliar

stipules, the spinule (leaf rudiment) appressed to

cataphyll, flattened, 1 mm long, acute at apex,

black. Foliar stipules appressed to stem, oblong-

lanceolate, 3.5-6 mm long, chaffy, straw-colored,

markedly striate-nerved with 5-8 closely parallel

nerves per side, acute to attenuate at apex, 3-set-

ulose, the 2 lateral setae 0.7-1.5 mm long, the

medial seta 0.3-1 mm long, the apex of the stipule

becoming ragged with age, the keels prominent

with a slender medial ridge, fimbriate, the margin

densely villose-fimbriate when new. Leaves 1-3

scattered on new twigs, deciduous, distichous,

short-petiolate, the lamina plane, broadly to nar-

rowly elliptic, rounded and mucronulate at apex,

PLOWMAN: NEW SPECIES OF ERYTHROXYLUM

31

FIG. 18. Known geographical distribution of new Erythroxylum species in eastern Brazil. Solid stars = E. petrae-

caballi; open star = E. santosii; solid circles = E. splendidum; solid squares = E. stipulosum.

32

FIELDIANA: BOTANY

rounded at base, 38-55 mm long, 22-26 mm wide,

firmly chartaceous, opaque, drying adaxially dark

brown, abaxially pale brown, somewhat shiny

adaxially, dull abaxially, elineate with the central

panel barely distinct, the adaxial midrib a low,

slender, medial ridge, the lateral nerves 10-12,

diverging 50°-70° from midrib, straight, then arch-

ing irregularly, anastomosing 2-3 mm from mar-

gin, prominulous on both surfaces, obscured

somewhat by prominent, reticulate veinlets. Pet-

iole 2 mm long, 1 mm in diameter, subterete.

Flowers produced singly in axils of last season's

ramenta, sparse to somewhat congested. Brac-

teoles lanceolate in outline, cymbiform, 2.5-3 mm

long, chaffy, acute to attenuate at apex, the seta

0.3-1 mm long. Pedicel pentangular, 5-ribbed, 1.5-

3.5 mm long, 0.5 mm in diameter. Calyx 1.5-2

mm long, divided nearly to base, the lobes lan-

ceolate, acute to acuminate at apex, incurved, 0.8-

1.8 mm long. Petal: specimen lost. Stamina I cup

'/2 length of calyx, 0.6-0.8 mm long, entire at mar-

gin. Brachystylous flowers: filaments 2.2-2.5 mm

long, the anthers oblong-ovate, 0.5 mm long; styles

free, 1.1 mm long; stigma depressed-capitate, 0.3

mm long. Dolichostylous flowers: antisepalous fil-

aments 1.3-2 mm long, the anthers oblong-ovate,

0.4-0.5 mm long; antipetalous filaments 2-2.5 mm

long, the anthers oblong-ovate, 0.4-0.5 mm long;

styles free, 2.8 mm long; stigmas depressed-capi-

tate, 0.3-0.5 mm long. Ovary ovoid, becoming

trigonal in young fruit, rounded at apex, 1.2 times

length of staminal cup, 1 mm long. Drupe not seen.

TYPE— Brazil, Ceara, Serra do Araripe, Bom

Fim, carasco [sic], 17 Jan. 1935 (fl), P. von Luetz-

elberg 26187 (holotype, M, F neg. 60126).

ADDITIONAL SPECIMENS EXAMINED— BRAZIL: Ceara:

Serra do Araripe: Araras, 1 1 Feb. 1935 (fl), P. von Luetz-

elberg 26464 (NY, F NEG. 16464); Baixa Rasa, 1 1 Feb.

1935 (fl), P. von Luetzelberg 26459 (M, F neg. 60127).

Municipio de Crato: Parque Nacional do Araripe, access

road E of federal road BR-122, ca. 20 km SW of Crato,

ca. 7°20'S, 39°35'W, alt. 860 m, 18 Jan. 1983 (st), T.

Plowman & J. Tavares Copula 12716 (ALCB, CEPEC, EAC,

F, O, HRB, IPA, K, M, MBM, MO, NY, P, R, RB, SP, U, UB, US).

Bahia: regiao de Barreiras, Chapadao da Panair, Serra

do Mimo, Grameal da Onca, 1 Jan. 1 955 (st), G. A. Black

55-17977 (IAN, F neg. 60129).

ETYMOLOGY— The specific epithet is derived

from Latin stipulosus, "having very large stip-

ules," in reference to the large, conspicuous stip-

ules.

DISTRIBUTION— Erythroxylum stipulosum is

known only from the Serra do Araripe on the bor-

der of Ceara and Pernambuco states, and from

one collection near Barreiras in western Bahia.

ECOLOGY— This species inhabits an open vege-

tation type best described as "closed shrubby caa-

tinga" (Eiten, 1983). There is little agreement on

the classification of dry vegetation types in north-

eastern Brazil, and different authors vary widely

in their opinions. In the Serra do Araripe, E. sti-

pulosum was said to be growing in carrasco, a term

which locally is used for closed caatinga scrub

vegetation on sandy soil. It is noteworthy that the

species was not found growing in adjacent habitats

of cerradao or "cerrado woodland."

Three other species of Erythroxylum were found

growing alongside E. stipulosum in the carrasco

vegetation on the Chapada do Araripe: E. vacci-

niifolium Mart., E. rosuliferum O. E. Schulz (if

distinct from E. betulaceum Mart.), and E. loef-

grenii Diogo. Of these only E. vacciniifolium is

widespread, occurring from Ceara south to Santa

Catarina in a diversity of habitats. The other two

species are restricted to dry formations in north-

eastern Brazil. None of these four species occurred

in adjacent cerrado woodland where E. barbatum

O. E. Schulz and E. suberosum were found. Nei-

ther of the latter was found in carrasco. Clearly

the distribution of species in these related yet ap-

parently different habitats needs to be studied more

closely.

PHENOLOGY— Erythroxylum stipulosum flowers

apparently from mid-January to mid-February

with the advent of the winter rains. Since rains are

notoriously unpredictable in northeastern Brazil,

the flowering time may vary appreciably from year

to year. Fruits are unknown in this species.

RELATIONSHIPS— Erythroxylum stipulosum is a

very distinct species, easily distinguished by its

long, fimbriate stipules. It belongs to section Rhab-

dophyllum, but within that large group has no ap-

parent close relatives. It may be related to E. pel-

leterianum St. Hil. and E. subracemosum Turcz.,

both of which have somewhat fimbriate but much

shorter stipules. Erythroxylum stipulosum might

also be confused with the poorly known E. stro-

bilaceum Peyr. (syn. E. warmingii Peyr.) from Mi-

nas Gerais, but in that species the stipules are en-

tire (vs. fimbriate), the enlarged cataphylls of the

ramenta are caducous (vs. persistent and marces-

cent), and the pedicel is much longer (9-1 1 mm

vs. 1.5-3.5 mm).

Additional collections of E. stipulosum are nec-

essary to furnish better material of mature leaves

(especially from Chapada do Araripe), flowers, and

PLOWMAN: NEW SPECIES OF ERYTHROXYLUM

33

FIG. 19. Erythroxylum tenue. A, flowering branch; B, habit of entire plant; C, detail of twig apex; D, stipule, scale

as in F; E, leaf showing venation; F, short-styled flower; G, long-styled flower, scale as in F; H, petal, scale as in F;

I, immature fruit. (A, B, C, D, E, F, and H from Plowman & Carvalho 12812; G from Pinheiro 2046; I from dos

Santos 1712.) Drawing by Marlene Werner.

34

FIELDIANA: BOTANY

FIG. 20. Restinga forest habitat of Erythroxylum tenue at type locality near Itubera, Bahia (Plowman & Carvalho

12812). The palm at left is pia9aba (Attalea funifera), which is characteristic of this habitat in southern Bahia.

Erythroxylum mucronatum Benth., E. martii Peyr., E. nobile O. E. Schulz, and E. \ cuspidifolium Mart, were also

found growing with E. tenue in this same restinga forest.

most importantly fruits, in order to determine its

relationships with other species.

10. Erythroxylum tenue Plowman, sp. nov. sect.

Rhabdophyllum O. E. Schulz. Figures 19-23.

Frutex gracillimus. Rami et ramuli recti, tenues. Cata-

phylla pauca, remota, stipulis foliaribus similia. Stipulae

foliares persistentes, manifeste striate nervosae, 2-setu-

losae. Folia in ramulis dispersa, breviter petiolata, per-

sistentia; laminae anguste ovatae vel lanceolatae, apice

late vel anguste acutae, basi obtusae vel acutae, mem-

branaceae, supra aliquantum nitidae, subtus impolitae.

Flores e ramulis hornotinis vel annotinis in axibus brev-

issimis in axillis foliorum vel ramentorum nati. Petali

lamina lingulata. armeniaca vel ochracea, ovata vel ob-

longa, ligula bilobata munita, utroque lobo usque ad

medium valde incrassato et auricula brevi proviso. Ur-

ceolus siam i nous calyce longior (x 1.2-1.5), margine

eroso-crenulatus, quinque dentibus antisepalis promi-

nulis munitus. Styli usque ad medium connati. Drupa

ellipsoidea, apice rotunda ta et subexcentrica, endocarpio

obovoideo, apice subexcentrico, tereti, longitudinaliter

nervoso, maturitate uniloculari.

Shrub to 3 m tall with 1 slender trunk ca. 2 cm

in diameter. Bark thin, smooth, more or less shiny,

dark brown with scattered brown lenticels, ca. 1.5

mm thick; wood light tan. Branches plagiotropic,

horizontal, somewhat storied, straight, without

short shoots. Branchlets distichous, slender,

straight, diverging 30°-90° from stem, a little com-

pressed at apex, soon becoming terete, 1 mm in

diameter, smooth, dark brown, somewhat shiny,

without lenticels. Inter nodes 1-15 mm long. Cata-

phylls (ramenta) few, scattered along base of new

shoots, the spinule (rudimentary leaf) straight,

brown, 0.3-0.7 mm long, caducous. Foliar stipules

persistent, appressed to stem, diverging somewhat

with age, broadly ovate to broadly triangular, 1-

2 mm long, markedly striate-nerved with 4-6

nerves per side, pale green, becoming chaffy with

age, obtuse to rounded at apex, 2-setulose, the

setae minute, filamentous, 0.2-0.3 mm long, eva-

nescent, the keels prominent, slightly alate, the

margin entire. Leaves persistent, scattered along

branchlets or sometimes clustered toward apex,

distichous, petiolate, the lamina plane, narrowly

ovate to lanceolate, broadly to narrowly acute at

apex, rarely obtuse or acuminate, obtuse to acute

at base, 24-67 mm long, 9-28 mm wide, mem-

branaceous, adaxially medium to dark green, shiny,

PLOWMAN: NEW SPECIES OF ERYTHROXYLUM

35

FIG. 21. Slender habit of the type plant of Erythroxylum tenue (Plowman & Carvalho 12812).

drying dark, grayish green, abaxially dull, very pale

green, drying ochreous or ferruginous, elineate and

with no distinct central panel, the adaxial midrib

flat with a very slender, low, medial ridge, the

lateral nerves 9-1 2 per side, diverging 50°-70° from

midrib, sinuous, obscure on both surfaces when

fresh, appearing slender and faint when dried, the

veinlets fine, open-reticulate. Petiole 2.5-4 mm

long, subterete, broadly canaliculate. Inflores-

cences produced in the axils of leaves or cataphylls

on current or last season's shoots on a short axis

to 2.5 mm long, the axis itself very short-pedun-

culate, the peduncle flattened, appressed to stem,

0.5-0.7 mm long. Flowers ca. 3 per node, ap-

36

FIELDIANA: BOTANY

FIG. 22. Flowering branch of the type plant of Erythroxylum tenue (Plowman & Carvalho 12812).

pearing sequentially, one at a time. Bracteoles per-

sistent, minute, spirally imbricate on short axis,

broadly ovate, concave, 0.6-1 mm long, mem-

branaceous, sparingly nerved, obtuse or rounded

at apex, the seta 0.1-0.4 mm long. Pedicel terete,

5-ribbed, 1.2-4 mm long. 0.5-0.7 mm in diame-

ter, slightly thickened toward apex. Calyx 1-1.5

mm long, divided '/j-'/z its length, pale green, be-

coming ochreous, the lobes triangular to ovate,

acute to obtuse, 0.5-0.7 mm long. Petal lamina

Ungulate, rather thick, convex with slightly de-

pressed midrib, recurving at anthesis, pale orange

to ochreous, ovate to oblong, rounded or truncate

at apex, 1.2-2 mm long, 0.8-1.2 mm wide, the

claw 0.7-1 .5 mm long with a pair of gibbous bulges

at base, the ligule bilobate, thickened, 0.6-0.7 mm

long, creamy white, each lobe with a single, re-

duced auricle 0.5-0.6 mm long, separated by a

swollen, Ungulate, indexed flap. Stamina! cup 1.2-

1.5 times length of calyx, 1-1.3 mm long, erose-

crenulate at margin with 5 short, protruding, an-

tisepalous "teeth." Brachystylous flowers: fila-

ments somewhat variable in length, 1-1.7 mm long;

anthers 0.4-0.5 mm long; styles 1-1.3 mm long,

connate for half their length; stigmas depressed-

capitate, 0.2 mm long. Dolichostylous flowers: an-

tisepalous filaments 0.8 mm long, the anthers 0.5

mm long; antipetalous filaments 1.2 mm long, the

anthers 0.4 mm long; styles 3 mm long, connate

for 1 mm; stigma depressed-capitate, 0.3 mm long.

Ovary obovoid-obconic, broadly rounded at apex,

Vz-'/j the length of staminal cup, 0.6-0.9 mm long.

Drupe ellipsoid, slightly excentric at apex, round-

ed, 10 mm long, 6 mm in diameter, red at ma-

turity, the mesocarp 0.7 mm thick, the endocarp

narrowly obovoid, slightly excentric at apex, nar-

rowed at both ends, terete, with ca. 1 2 longitudinal

nerves, unilocular with 2 collapsed rudimentary

locules at base; endosperm occupying ca. 50% of

locule. Embryo 7 mm long; cotyledons oblong, the

apex rounded, slightly excentric, basally subcor-

date, 6.5 mm long, 3 mm wide, 0.3 mm thick;

radicle 1.3 mm long.

TYPE— Brazil, Bahia, Municipio de Itubera, km

1 1 na estrada Itubera- Valen9a, 1-2 km no ramal

de acesso a Estacao da Telebahia, ca. 13°40'S,

39°07'W, near sea level, slender, erect single-

stemmed shrub 3 m tall, calyx yellowish brown,

petals dirty yellowish, ligule white, 4 Feb. 1983

PLOWMAN: NEW SPECIES OF ERYTHROXYLUM

37

FIG. 23. Known geographic distribution of Erythroxylum tenue in Bahia in eastern Brazil.

38

FIELDIANA: BOTANY

(fl), T. Plowman &A.M.de Carvalho 12812 (ho-

lotype, CEPEC 32462; isotypes, ALCB, F 1916664, F

neg. 60187, G, GH, HRB, IPA, K, LE, MBM, MO, NY,

p, R, RB, SP, u, UB, us).

ADDITIONAL SPECIMENS EXAMINED— BRAZIL: Bahia:

Municipio de Camamu: Acarai, estrada ao lado sul, 1

Jul. 1971 (fl, imm fr), T. S. dos Santos 1712 (CEPEC, F).

Municipio de Marau: rodovia BR 030, trecho Ubaitaba-

Marau, 45-50 km a leste de Ubaitaba, ca. 50 m alt.,

14°H'S, 30°01'W, 12-13 Jun. 1979 (fr), S. Mori el al.

1 1 965 (CEPEC, F, K, NY, RB). Municipio de Itacare: Itacare,

6 Jan. 1967 (fl, imm fr), R. P. Belem & R. S. Pinheiro

3009 (CEPEC, F, K, NY, RB). Municipio de Itabuna: Fa-

zenda Pirataquice, 30 Nov. 1970 (fl), L. Emygdio de

Mello Filho 3022 & M. Emmerich 3560 (CEPEC). Mu-

nicipio de Ilheus: prope Ilheus, Feb. 1 822 (fl), L. Riedel

612 (F, fragment, K, LE, F neg. 599 1 7, p, excluded syntype

of E. bongardianum C. A. Meyer ex Peyr.); km 35 na

estrada Ilheus-Serra Grande, 22 Oct. 1983 (fl), A. M. de

Carvalho el al. 2007 (CEPEC, F); Fazenda Barra do Man-

guinho, ramal com entrada no km 10 da rodovia Pontal-

Olivenca, lado direito, 3 km a oeste da rodovia, 5 Feb.

1982 (fl), L. A. Mattos Silva, T. S. dos Santos & B. Boom

1443 (CEPEC, F); entre km 5-8 da estrada Olivenca-Ma-

ruim, alt. 30-50 m, 14 Oct. 1983 (fl), M. P. M. de Lima

et al. 22 (CEPEC, RB), same locality, km 7, 16 Apr. 1986

(fl), L. A. Mattos Silva, T. S. dos Santos & J. L. Hage

2043 (CEPEC, F); estrada que liga Olivenca a Vila Brasil,

km 7, 16 Feb. 1982 (fr), L. A. Mattos Silva, A. M. de

Carvalho & T. S. dos Santos 1528 (ALCB, CEPEC, F, HBR,

RB); estrada que liga a Estacao Hidromineral de Olivenca

ao Povoado de Vila Brasil, 5 km ao sudoeste de Olivenca,

8 Feb. 1982 (fl, imm fr), L. A. Mattos Silva, T. S. dos

Santos & G. P. Lewis 1487 (CEPEC, F, HRB), 26 Apr. 1983

(fl), L. A. Mattos Silva, T. S. dos Santos & B. E. Leuen-

berger 1688 (B, CEPEC, F, HUEFS, RB); ramal novo para o

povoado da Vila Brasil, com entrada no km 28 da ro-

dovia Ilheus-Una, lado direito, inicio do ramal km 3,

27 Feb. 1985 (fl), T. Plowman, L. A. Mattos Silva & T.

S. dos Santos 13963 (CEPEC, F, K, MO, RB), same locality,

km 7, 27 Feb. 1985 (fl, imm fr), T. Plowman, L. A.

Mattos Silva & T. S. dos Santos 13964 (CEPEC, F, GH,

NY, RB, SP). Municipio de Una: estrada Ilheus-Una, 1 9

Nov. 1983 (fl), R. Callejas, A. M. de Carvalho & L. A.

Mattos Silva 1757 (CEPEC, F, MBM, NY). Municipio de

Itapebi: Rodovia Itapebi a Camaca, 17 Oct. 1969 (fl), J.

A. de Jesus 465 A (RB). Municipio de Porto Seguro: km

7 da estrada Porto Seguro a Santa Cruz Cabralia, Tapera-

pua, 20 Apr. 1982 (fl, fr), A. M. de Carvalho, S. G. da

Vinha & E. S. Brito 1228 (CEPEC, F, RB, SPF); estrada

Porto Seguro-Eunapolis, km 13, 18 Oct. 1969 (fl), /. A.

de Jesus 465B (CEPEC, RB); Parque Nacional do Monte

Pascoal, 18 Jan. 1974 (fl, imm fr), T. S. dos Santos 2725

(CEPEC, F, K); rodovia para povoado de Trancoso, km 5

depois de Arraial d'Ajuda, 5 Nov. 1983 (fl), R. Callejas,

A. M. de Carvalho & L. A. Mattos Silva 1688 (CEPEC, F,

MBM, NY). Municipio de Guaratinga: km 10, Guaratinga

a Sao Paulinho, 29 Mar. 1973 (fl), R. S. Pinheiro 2046

(CEPEC, F). Municipio de Ibicoara: regiao da Serra do

Sincora, 24 Feb. 1943 (fl), R. L. Froes 20092 (IAN, NY).

ETYMOLOGY— The specific epithet is derived

from Latin tenuis, "thin" or "fine," referring to

the slender trunk and branches of this species.

DISTRIBUTION— Erythroxylum tenue grows only

in moist coastal forests in southern Bahia, Brazil,

from Valenca south to Parque Nacional do Monte

Pascoal. The Froes collection from Serra do Sin-

cora appears to be somewhat out of range. How-

ever, forest habitats do occur in this area (R. Har-

ley, in litt.), and other species from the Atlantic

forest, such as Ruellia affinis (Mart.) Lindau and

Brunfelsia clandestine! Plowman (Plowman, 1981),

are known to occur here.

ECOLOGY— This species grows primarily in the

understory of restinga forest on sandy soils near

the coast, but has also been collected in areas of

mata higrofila or tropical moist forest further in-

land, presumably on clay soils. It may be occa-

sional to relatively abundant at any one locality.

PHENOLOGY— Erythroxylum tenue flowers and

fruits throughout the year. The flowers are pro-

duced sequentially such that only a few flowers on

an individual are open at a given time, but the

flowers appear over a relatively long period.

RELATIONSHIPS— Erythroxylum tenue was first

collected by Riedel at Ilheus in 1822. Riedel's col-

lection (no. 612) was cited as one of two syntypes

of E. bongardianum C. A. Meyer ex Peyr. The

second element (Langsdorff s.n.), from Espirito

Santo, was the basis for the manuscript name "£".

bongardianum C. A. Meyer" appearing on one

sheet at LE. C. A. Meyer had worked with F. E. L.

Fischer at LE on a general monograph of Ery-

throxylum, continuing earlier studies of the Bra-

zilian species by A. G. H. Bongard. Although nei-

ther of these studies was ever published, O. E.

Schulz, in preparing his worldwide monograph for

Das Pflanzenreich, obtained the handwritten

manuscripts and some of the lithographed plates

(Schulz, 1907). Schulz cited in synonymy many

nomina nuda from Bongard's and Fischer and

Meyer's manuscripts. The original manuscripts

have not been located at LE, and possibly were

retained by Schulz at B and subsequently destroyed

during World War II.

I here designate one duplicate of the Langsdorff

collection at LE as the lectotype of E. bongardian-

um C. A. Meyer ex Peyr., for which I give the

following reasons. The description published by

Peyritsch applies principally to this collection.

Peyritsch annotated many Erythroxylum speci-

mens at Leningrad and may have had access to

Meyer's notes. Meyer annotated only the Langs-

dorff collection as "E. bongardianum," a duplicate

of which Bongard had previously annotated as "E.

ovatum"; Meyer annotated the Riedel collection

at LE as "E. minutiflorum." Last, Peyritsch himself

PLOWMAN: NEW SPECIES OF ERYTHROXYLUM

39

stated in a note appended to his description that

the Riedel collection strongly suggested a separate

species. From this analysis, it is clear that the

Langsdorff collection must be chosen as the lec-

totype of E. bongardianum. The Langsdorff col-

lection, however, in fact belongs to E. passerinum

Mart., so that E. bongardianum must be placed

in synonymy under that species. In spite of this

confusion, the new name E. tenue is correctly ap-

plied to Riedel 612.

Erythroxylum tenue belongs to section Rhabdo-

phyllum. Within this large and diverse complex,

E. tenue belongs to the species group discussed

above under E. petrae-caballi. Of the several sim-

ilar species in this group that occur in Bahia, Ery-

throxylum tenue is the most distinct. The com-

bination of a slender habit, the lack of short shoots,

the small, narrowly ovate to lanceolate leaves, the

brief elongation of the flowering axis, the thick-

ened, reduced petal ligule, and the connate styles

all serve to distinguish E. tenue from any other

species. Table 4 gives the characters by which E.

tenue differs from its closest congeners in Bahia.

Acknowledgments

This paper is dedicated to Jack Staehle who has

supported my fieldwork in Brazil. I deeply appre-

ciate his generosity and continuing interest in Field

Museum's programs of botanical exploration in

the Neotropics.

I would like to thank the curators of the follow-

ing herbaria who kindly lent or allowed me to

study specimens cited in this study: AAU, ALCB,

B, BAH, BM, BOTU, C, CEPEC, CGE, E, EAC,

F, G, GH, GUA, HRB, HUEFS, IAN, IPA, K,

LE, M, MBM, MO, NY, OXF, P, R, RB, SP, SPF,

U, UB, UEC, US, and W. The following institu-

tions were especially generous in providing me

with valuable duplicates of their Brazilian collec-

tions: ALCB, CEPEC, EAC, GUA, HRB, HUEFS,

IPA, K, MBM, NY, RB, SP, and UB.

Fieldwork in Brazil leading to this paper was

greatly facilitated by a number of institutions and

individuals. Primary among these is CEPEC, Ita-

buna, which I have visited on three occasions and

where I received generous hospitality and collab-

oration. Paulo Alvim, Roberto J. Pereira, and most

recently, Walny Sou/a da Silva made the excellent

botanical facilities of CEPEC available to me and

arranged for visits to several key localities. In field

and herbarium work at CEPEC, I was ably assisted

by Luiz A. Mattos Silva, Talmon S. dos Santos,

Sergio da Vinha, and Andre M. de Carvalho; both

Carvalho and Robert Voelks graciously provided

hospitality in Ilheus. In Salvador, I received kind

assistance from Hortensia P. Bautista, Geraldo

Pinto, and Jorge C. A. Lima at Herbario Radam-

brasil; from Ivomar C. Brito and Lecticia Scott

Scardino Faria of the Universidade Federal da Ba-

hia; and from Pedrito Silva of CEPLAC. Larry

Noblick of the Universidade Estadual de Feira de

Santana supplied valuable data on localities and

habitats, especially of Erythroxylum petrae-ca-

balli. Field studies at the Chapada do Araripe in

Ceara were facilitated by Luiz Bezerra de Oliveira

of the Institute Brasileira de Desenvolvimento

Florestal. Afranio Fernandes and the late Prisco

Bezerra of the Universidad Federal de Ceara (EAC)

provided hospitality in Fortaleza and helped ex-

pedite fieldwork at Serra do Araripe. I am also

indebted to Jose "Arnaldo" Tavares Ca9ula, res-

ident guard at the Parque Nacional de Araripe, for

transportation and field assistance.

Rupert Barneby, Scott Mori, Sandra Knapp, and

Michael Huft offered helpful scientific and tech-

nical comments for improving the manuscript. Al-

icia Lourteig and Bronwen Gates provided infor-

mation on Gounelle's collections, and Dorothy

Araujo was instrumental in locating Mikan's lo-

cality "Tocaja." Rolf Singer patiently corrected

my Latin diagnoses. Christine Niezgoda provided

technical assistance with the figures. Finally, I am

grateful to Marlene Werner and Pollyanna Quast-

hoff for preparing the excellent line drawings.

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