SMITHSONIAN CONTRIBUTIONS TO BOTANY 0 NUMBER 41 

Revision of the 

Genus PsyZZocar-us (Rubiaceae) 

Joseph H. Kirkbride, Jr. 

SMITHSONIAN INSTITUTION PRESS 

City of Washington 

1979 
ABSTRACT 

Kirkbride, Joseph H., Jr. Revision of the Genus Psy llocarpus (Rubiacae). Smithsonian Contributions to Botany, number 41, 32 pages, 41 figures, 8 plates, 1979.- 

Psy Zlocarpus, a member of the Rubiaceae tribe, Spermacoceae, is separated from 

all other genera of the tribe by its capsule, which is strongly compressed parallel 

to the septum. Section Amazonica is found only in white-sands vegetation in the 

Amazon valley, and section Psyllocarpus is encountered only on savannas of the 

Planalto Central do Brasil. The two sections, of which Amaronica Kirkbride is 

described as new, are sharply delimited by their leaf structure, floral biology, number of calyx lobes, pollen morphology, and distribution. The history, morphology, 

and distribution of the genus are discussed. A key to the sections and species of 

Psy ZZocarpus, a taxonomic treatment of each taxon including nomenclature, synonymy, description, distribution and type, and an index to exsiccatae critically 

examined are given. Two species are described as new, P. cururuensis Kirkbride 

and P. goiasensis Kzrkbride, and two species P. campino?-zim (Krause) Kirkbride 

and P. psyllocarpoides (Sucre) Kirkbride, are transferred into Psyllocarpus. 

OFFICIAL PLBLICATIO\ DATE is handstamped in a limited number of initial copies and is recorded 

in the Institution's annual report, SMITHSOIIAN YEAR. SERIES COLER DESIGN: Leaf clearing from the 

katsura tree Cercidiphyllum japonzcum Siebold and Zuccarini. 

Library of Congress Cataloging in Publication Data 

Kirkbridge, Joseph H, Jr 

Revision of the genus Psyllocarpus (Rubiaceae). 

(Smithsonian contributions to botany ; v. 41) 

Bibliography: p. 

Includes indexes. 

1. Psvllocarpus. 2. Botany-Amazon \'alley. 3. Botany-Brazil. I. Title. Series: Smithsonian 

Institution. 

QKl.SZ747 TO], 41 [QK495.R85] 581'.08S [583'.52] 794631 

Smithsonian contributions to botany : v. 41 
Con tents 

Page 

Introduction ...................................................... 1 

Acknowledgments .................................... 2 

Morphology ....................................................... 2 

Habit .......................................................... 2 

............................... 2 

................................... 3 

Inflorescences ............................. ................... 4 

Flowers ......................................................... 5 

Fruits 5 

Distribution ....................................................... 5 

Psyllocarpus Martius ex Martius & Zuccarini .......................... 11 

Key to the Sections and Species of Psyllocarpus ........................ 12 

1 . Psyllocarpus campinorum (Krause), new combination ........... 13 

2 . Psyllocarpus cururuensis, new species ......................... 14 

3 . Psyllocarpus psyllocarpoides (Sucre), new combination .......... 15 

Section Psylloravpus ............................... 15 

4 . Psyllocarpus laricoides Martius ex Martius & Zuccarini .......... 16 

5 . Psyllocarpus goiasensis, new species ........................... 17 

6 . Psyllocarpus asparagoides Martius ex Martius & Zuccarini ....... 17 

7 . Psyllocarpus phyllocephalus Schumann ........................ 18 

8 . Psyllocarpus schwackei Schumann ............................ 19 

Excluded Species ................................................ 20 

Psyllocarpus thymbroides Martius ex Martius & Zuccarini ........... 20 

Literature Cited ................................................... 21 

Index to Exsiccatae ................................................ 23 

Plates ............................................................ 25 

.......................................................... 

Amazonica, new section ........................................... 13 

... 

111 

Revision of the 

Genus Psyllocarpus (Rubiaceae) 

Joseph H. Kirkbride, Jr. 

Introduction 

Psyllocarpus, a member of the Rubiaceae tribe 

Spermacoceae, is separated from all other genera 

of the tribe by its capsule, which is strongly compressed parallel to the septum. It is found on savannas in the Planalto Central do Brasil and 

white-sands vegetation in the Amazon valley. It 

was described by von Martius and Zuccarini in 

1824 from collections made by the former in 1818 

and 1819. They undoubtedly intended Psyllocarpus 

and its four new species to be published in the 

first volume of their Nova genera et species plantarum, but a few months earlier they published in 

Flora the diagnostic phrases for the taxa Psyllocarpus. Under article 11 of the International Code 

of Botanical Nomenclature (1972), their publication in Flora has clear priority over their later one 

in the Nova genera et species plantarum and must 

be cited as the place of publication. 

Subsequent authors (Sprengel, 1827; Schultes and 

Schultes, 1827; de Candolle, 1830; G. Don, 1834) 

followed von Martius and Zuccarini, and they often 

used their very phrases to describe Psyllocarpus 

and its four species. The only other revision of 

Psyllocarpus prior to this one was made by Schumann (1888) for Flora brasiliensis. He maintained 

three species in Psyllocarpus and transferred P. 

thym broides to Staelia following neotypification of 

that species. Considering the collections available 

to him at that time, he was correct in maintaining 

Joseph H. Kirkbride, Jr., Department of Botany, National 

Museum of Natural History, Smithsonian Institution, Washington, D.C. 20560, U.S.A. 

three species in Psyllocarpus, but his key to those 

species is misleading and inaccurate for some characters. 

In 1898, Schumann published two additional 

species of Psy llocarpus based upon collections sent 

to Berlin by Glaziou for identification. After 

Schumann, two more species of Psyllocarpus were 

incorrectly associated with other genera by Krause 

(1908) and Sucre (1971). Each of these species is 

found in the type of vegetation known as “campinas,” which occur on podzolized sands in Amazonian Brazil. The enormous number of new collections made in this century, as compared to those 

available to Schumann, indicated the need to reevaluate his work and to transfer into Psyllocarpus 

those species incorrectly assigned to other genera. 

Psyllocarpus has traditionally been placed in 

tribe Spermacoceae (de Candolle, 1830; G. Don, 

1834; Hooker, 1873; Baillon, 1881; Schumann, 

1888, 1891; Wernham, 1916; Hayden, 1968). Within 

the tribe, the genera are distinguished by the occurrence and the manner of fruit dehiscence. The 

opinion is again being actively presented that a 

number of genera of tribe Spermacoceae should be 

merged (Hooker, 1873; Bremekamp, 1934; Verdcourt, 1975, 1976; F. R. Fosberg, pers. comm.) because the traditional generic delimitations do not, 

in the opinions of those cited above, divide the 

species into natural groups. The traditional generic 

divisions within tribe Spermacoceae have been maintained by workers who have dealt principally with 

New World Rubiaceae (Schumann, 1888; Wernham, 1916; Standley, 1930, 1931a, 1931b, 1931c, 

1936; Smith and Downs, 1956; Anderson, 1972; 

1 
2 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

Steyermark, 1972, 1974). I believe that Psyllocarpus 

is a natural grouping of closely related species, 

and pending a worldwide revision of tribe Spermacoceae that is beyond the scope of this study, 

I am maintaining them as the genus Psyllocarpus. 

In my opinion, it is most closely related to Borreria G.F.W. Meyer (sensu stricto) because of the 

manner of dehiscence of its fruits and its habit 

and foliar development as seen in section Amazonica. 

 ACKNOWLEDGMENT.-^ am indebted to Drs. Rupert C. Barneby, Richard S. Cowan, Harold E. Robinson, and John J. Wurdack for their many helpful 

suggestions and reviews of the manuscript. 

I am also grateful to the National Science Foundation for support of my field studies of section 

Psyllocaypus through a grant for improvement of 

doctoral dissertation research, GB-31978, and to 

the Fluid Research Fund of the Smithsonian Institution for supporting my studies of section Amazonica. I also want to thank the Universidade de 

Brasilia, Brasilia, D.F., Brasil, and the Instituto 

Nacional de Pesquisas da Amazonia, Manaus, Brad, for their co-operation and support. Access to 

herbaria was permitted and loans of specimens 

were made by numerous institutions, to whose directors I express my thanks. 

Morphology 

HABrr.-The habit of Psyllocarpus ranges from 

perennial herb to shrub, but the majority of species and individuals are subshrubs. Perennial herbs 

have been found in P. laricoides and P. schwackei. 

In both species these appear to be a fecund stage 

in young individuals that will, if unaffected by 

outside forces, continue to grow and become subshrubs. The subshrubs are robust plants that, except in the case of P. schwackei, develop woody 

stems four to six times greater in diameter than 

the new herbaceous ones. Psyllocarpus schwackei 

does not generally get as tall as other species, and 

it appears to die back to ground level much more 

frequently. 

Well-developed brachyblasts are found on all 

species of Psyllocarpus except P. campinorum and 

P. schwackei, and it has not been possible to determine their presence or absence for P. psyllocarpoides since foliar structures are not known for this 

species. The brachyblasts are reduced axillary 

branchlets with the form of their structures normal 

but reduced in size with the internodes the most 

strongly reduced. Consequently, the brachyblasts 

form fascicles of reduced leaves in the axils of the 

primary leaves. Rarely short branchlets will be 

found in the leaf axils in place of the brachyblasts 

from which they differ only by the increased length 

of their internodes. 

The species of section Amazonica, P. campinorum, 

P. cururuensis, and P. psyllocarpoides, are usually 

taller than those of section Psyllocarpus. Also, SO 

far as known from a very small sample, P. campinorum and P. psyllocarpoides are less frequently 

branched that P. cururuensis and the species of 

section Psyllocarpus. The habit of the members of 

section Amazonica is probably wand-like. 

The species of section Psyllocarpus are usually of 

a lesser stature and more densely branched than 

those of section Amazonica. The branches of P. 

laricoides, P. goiasensis, and P. asparagoides are 

erect and strongly fastigiate so that the living plant 

has an obconical outline. When pressed and dried, 

it takes the form of many tightly compact, parallel 

branches. Psyllocarpus phyllocephalus has a distinct 

branching pattern: the main stem is erect and undivided until four or five cm long and then divides 

into numerous broadly spreading branches. These 

divide many times, further producing a spreading, 

flat top. When pressed and dried, specimens of P. 

phyllocephalus form a circle of tangled branches 

that seem to be divided numerous times. Psyllocarpus schwackei is also distinctive due to its short 

stature and slender stems and branches. It produces 

as many branches as other members of section 

Psyllocarpus, but they are shorter, thinner, and not 

further divided. 

STIPuLES.-The stipular structure of Psy llocarpus 

is typical of tribe Spermacoceae. A membranous 

tissue or sheath connects the leaf base on each side 

of the stem forming a collar that completely surrounds the stem at each node. On the upper, free 

edge of the sheath are found three to eight setae, 

usually evenly spaced along the edge of the sheath 

beween the leaf bases. Their length varies, the 

central one being the longest, the rest gradually 

shorter outward. The apex of each seta at the apical 

nodes is glandular and can be referred to as a colleter (Lenten, 1975). At the lower nodes, the colleters are missing from the setae thus indicating 

that their secretory function is in the early develop- 
SUMBER 41 3 

ment of the apex as has been suggested for other 

members of the Rubiaceae (Krause, 1909; Uphof, 

1962; Horner and Lersten, 1968). 

LEAvEs.-The sections of Psyllocarpus are clearly 

separated by their foliar structures. The leaves of 

section Amaronica are dorsiventral (Figure 1) and 

appear grossl) to be planar in form. From herbarium specimens, their orientation appears to be horizontal. In cross-sections of the leaves of P. campinorum and P. cururuensis (Figure I), the structure 

of a typical dorsiventral leaf is seen. The epidermis 

are composed of a single layer of cells above and 

below. Stomata, as seen in cross-sections, are found 

on each face, although fewer on the adaxial one. 

The palisade parenchyma is a single layer of cells, 

elongated at right angles to the dorsal epidermis 

throughout, and more or less of uniform length. 

Immediately below it is the paraveinal mesophyll. 

This is a regular network of several-armed cells, 

lacking chlorophyll, one cell-layer thick, extending 

horizontally between the vascular bundles in the 

plane of the phloem (Fisher, 1967; Lackey, 1978). It 

has been reported from a number of unrelated 

families: Magnoliaceae (Pray, 1954), Cercidiphyllaceae (Chandrasekharem, 1972), Sapotaceae (H. E. 

Robinson, pers. comm.), Leguminosae (Debold, 

1892; Fisher, 1967; Weston and Cass, 1973; Lackey, 

1978), Sapindaceae (Radlkofer, 1890), Oxalidaceae 

(Dittmar, 1964), and Liliaceae (Pray, 1955). The 

spongy parenchyma fills the space between the 

paraveinal mesophyll and the ventral epidermis. 

Throughout both parenchyma layers there are enlarged idioblasts containing either bundles of 

raphides or druses. The raphides occur in both 

types, but the druses are only encountered in the 

palisade layer. The large main vein is clearly evident in the center of the leaf and gives rise to 

smaller pinnate secondary veins. 

Schumann (1888) described the leaves of section 

Psyllocarpus as “trigona” or “plana,” and used these 

characters to identify the species of Psyllocarpus. In 

the living state, the leaves of section Psyllocarpus 

are actually terete (Figure 2). Sometimes on the 

abaxial side of the leaf there are two or three faint 

ridges or a slight flattening. When individuals of 

this section are dried, the leaves become angled an 

irregular number of times. The orientation of the 

leaves is close to vertical, which exposes the entire 

surface of the leaf to a more uniform intensity of 

sun light. In all species of the section, the epidermis 

in cross section is composed of a single layer of cells 

with stomata distributed all around the surface. 

The palisade parenchyma is composed of elongated 

cells, of more or less equal length, placed at right 

angles to the epidermis so as to form a continuous 

layer, the individual cells being arranged radially. 

The palisade parenchyma is composed of a single 

layer of cells in all species of section PsyZZocarpus 

except P. schwackei, where it consists of two layers 

concentrically arranged. There is some spongy parenchyma, but it is much reduced in volume and 

sometimes compIeteIy absent. The leaves have one 

vein that occupies the long axis of the leaf blade. 

The bundle-sheath varies in thickness from a single 

m 

ue 

PP 

Pm 

SP 

FIGURE 1.-Cross-section of portion of leaf blade of Psvllocarpus campinorum (Bahia 32) (ue = upper epidermis: 

le = lower epidermis; pp = palisade parenchyma; pm = 

paraveinal mesophyll; sp = spongy parenchyma; m = 

midrib). 
4 S&IITHSONIAN COSTRIBUTIOSS TO BOTANY 

PP 

FIGURE 2.-Cross-section of leaf blade of Psyllocarpus Iaricoides (Anderson et al. 35761) (e = epidermis; pp = palisade parenchyma; sp = spongy parenchyma; v = vein; 

h = bundle-sheath; s = stoma). 

layer of cells to many, occupying a great deal of 

space. Enlarged idioblasts with bundles of raphides 

are found in all species, and enlarged idioblasts 

with druses were also found in the closely related 

P. asparagoides and P. phyllocepfzalus. 

INFL0RESCEsCES.-The inflorescence structures of 

Psyllocarpus are similar to those in the Labiatae, as 

described by Bentham (1835). Because of this striking similarity, Bentham’s terminology for Labiate 

has been adopted here. The most common type of 

inflorescence encountered in Psyllocarpus is the verticillastrate raceme. It is an indeterminate inflorescence with pairs or rarely, in P. Zaricoides, whorls 

of three bracts, each subtending a cymule of 1 to 

17 flowers. Each pair or whorl of bracts with their 

cymules is referred to as a verticillaster. In section 

Amazonica, the bracts are clearly differentiated from 

the leaves, and the flowers of each cymule range in 

number from 4 to 17. In the two species of section 

Psyllocarpus with verticillastrate racemes, P. 1an’- 

coides and P. goiasensis, the bracts exhibit a gradual 

change distally in form from leaf-like to bract-like, 

the lowest being very similar to true leaves. In order 

to maintain the distinction between the leaf and 

bract, Bentham’s definition (1835) has been applied 

here, “that the (bracts) are always those situated at 

the base of a verticillaster.” The cymules of the two 

species of section Psyllocarpus with verticillastrate 

racemes have one to four flowers. 

The three remaining species of Psyllocarpus exhibit two types of inflorescences: (1) capitate (P. 

asparagoides and P. phyllocephalus) and (2) solitary flowers in each leaf-axil at a few terminal 

nodes. (P. schwackei). The capitula are very tightly 

packed heads of 6 to 24 flowers. Paired bracts subtending one to three (rarely four) flowers are present within each head and are connected at base 

by a membranous tissue similar to that which connects the bases of bracts of the verticillasters and 

of the stipules. The capitula are therefore tightly 

packed concentric rings of sheathing tissues, bracts, 

and flowers. 

Psyllocai-pus schwackei has only solitary flowers, 

each subtended by a setiferous bract. The solitary 

flowers occur always in opposed pairs at a few of 

the terminal nodes. 
NUMBER 41 5 

FLOWERS.-AS is typical for family Rubiaceae, 

the flowers of Psyllocarpus have an inferior ovary, 

an actinomorphic, gamopetalous corolla, and epicorollar stamens; as is less usual a tetramerous corolla and bilocular ovary. The species of section 

Amazonica are heterostylous, and those of section 

Psyllocarpus homostylous. 

The hypanthium is turbinate, bearing a persistent calyx of either two (section Psyllocarpus) or four 

(section Amazonica) lobes. The bases of the lobes 

are either joined by a low membrane or completely 

free. Between the calyx-lobes, on the edge of the 

calyx-tube or along the distal edge of the hypanthium, there are usually one to eight setae of varying lengths, which resemble those of the stipules. 

Very rarely these setae are lacking. The hypanthium has in each of its two locules one ovule 

peltately fastened to its funiculus that arises from 

the center of the septum separating the locules. 

The corolla is small, bIue or white, and salverform or very rarely funnelform. The corolla-lobes 

are always much shorter than the tube and valvate. 

In heterostylous section Amazonica, the pin flowers have short filaments that separate from the 

corolla tube above the middle and below the apex 

so that the anthers are included, and the thrum 

flowers have long filaments that separate from the 

apex of the tube so that the anthers are exserted. 

In homostylous section Psyllocarpus, the short filaments separate from the tube wall below the middle so that the anthers are always included. 

The style varies in length, with distinct and 

much longer stigmatic lobes in section Amazonica. 

In pin flowers with exserted stigmatic lobes, the 

styles are two to three times longer than in thrum 

flowers with included stigmatic lobes. The style 

in section Psyllocarpus is always short and oneeighth to one-quarter as long as the corolla tube. 

The stigma frequently is weakly bilobate, but 

sometimes it is capitate or distinctly lobed. As a 

consequence of the relative sizes, the anthers in 

section Psyllocarpus are always positioned slightly 

above the stigma. 

The pollens of the two sections are very distinct. The species of section Arnazonica have significantly larger grains (Figures 3,5,7) and are oblatespheroidal with short spinules distributed evenly 

over the perforate sexine (Figures 4, 6, 8). In section Psyllocarpus, the grains are smaller and prolate-spheroidal (Figures 9, 11, 13, 15, 17) with short 

spinules occurring along the edges of the colpi in 

a nonrandom pattern over the smooth and nonperforate sexine (Figures 10, 12, 14, 16, 18). 

FRum.-The fruits of Psyllocarpus are capsules 

strongly compressed parallel to the septum. They 

dehisce from the apex downwards, the lines of 

dehiscence following the junction of the septum 

with the wall of the capsule on each side. Each 

unbroken valve of the capsule reflxes outwards, 

leaving the complete septum erect in the center. 

The fruits were measured at the stage just prior 

to dehiscence, and the planar shape, length, and 

width of their outline determined as well as the 

thickness of their compressed dimension. 

The seeds of Psyllocarpus are strongly compressed 

and were measured in the same manner as the 

intact fruits. They are shed from the fruit after it 

dehisces. There is a tendency towards development 

of a narrow, irregular wing by an extension of the 

testa around the edge of the seed (Figure 19, 22, 

25, 27, 31, 33, 37, 39). The abaxial surface of the 

seeds is rugulose. The surface sculpturing of the 

rugae on the seeds of section Amazonica is similar 

for all three species (Figure 20), but the rugae of 

each species of section Psyllocarpus have a distinct 

pattern of surface sculpturing (Figure 26, 29, 32, 

35, 38). The strophiole is on the adaxal side of the 

seed. Based upon data from two species, the strophiole of Psyllocarpus was described as “reduced to 

a single, round central spot” (Hayden, 1968). This 

description is very apt for all species of Psyllocarpus (Figure 21, 23, 27, 33, 39) except P. goiasensis 

(Figure 30) and P. phyllocephalus (Figure 36). In 

these two species, length and width of the strophioles are sufficiently different so that the strophioles 

become oblong to linear. 

Distribution 

The genus Psyllocarpus is found only within the 

confines of modern-day Brazil. Its distribution 

parallels its subgeneric classification. Section Amazonica is now known from the extreme eastern 

end of Amazonas, the western end of Rondonia 

and the southwestern corner of Para in the Amazon 

valley (Figure 40). The herbarium label data indicate that the three species of section Amazonica 

grow only in “campina” vegetation. This is a form 

of low, woody vegetation found on podzolized 

white-sand soils. Campinas have a low species di- 
6 SUITHSONIAN CONTRIBUTIONS TO BOTANY 

FIGURES 3-18.-Psyllocarpus pollen, SEM. Psyllocarpus campinorum: 3, polar view, X 1350; 

4, surface, X 3750 (3, 4, Silva 4346). Psyllocarpus cururuensis: 5, Polar \iew, x 1250; 6, 

surface, X 3750 (5, 6, Anderson et al. 10779). Psyllocarpus psyllocarpoides: 7, equatorial view, 

X 1200; 8, surface, X 3750 (7, 8, Froes 25262). Psyllocarpus laricoides: 9, equatorial view, x 2500; 

10, surface, X 5000 (9, 10, Anderson et al. 35750). Psyllocarpus goiasensis: 11, equatorial view, 

X 2500; 12, surface, X 5000 (11, 12, Anderson et al. 7161). Psyllocaipus asparagoides: 13, equatorial \iew, x 2550; 14, surface, x 7500 (13, 14, Irwin et al. 22247). Psyllocarpus phyllocephalus: 

15, equatorial view, X 2500; 16, surface, x 5000 (15, 16, Imin et al. 34434). Psyllocarpus 

schwackei: 17, equatorial view; x 2000; 18, surface, x 5150 (17, 18, Duarte 11381) 
NUhlBER 41 7 

FIGURES 19-24.-Section Amazonica seeds, SEM. Psyllocarpus campinorum: 19, abaxial view, 

X 20; 20, abaxial surface, x 270; 21, adaxial view, x 22 (19, 20, 21, Bahia 32). Psyllocarplc 

cururuensis: 22, abaxial view, x 20; 23, adaxial view, x 19 (22, 23, Anderson et al. 10779). 

Ps)llocarpus pspllocarpoides: 24, abaxial view, x 22 (24, Froes 25262). 

versity, and their flora contains a number of endemic 

species (Lisboa, 1975). Judging from the number of 

herbarium collections, the species of section Amazonica are not common. 

Section Psyllocarpus is known from the Distrito 

Federal, Goias, Bahia, and Minas Gerais in southeastern Brazil (Figure 41). The distribution of the 

species falls into two groups: two species, P. goiasensis and P. phyllocephalus, which are not closely 

related, are found on the system of chapadas that 

extends from Rios Tocantins and Parana to the 

north and to the Rios Corumba and Sao Marcos to 

the south, approximately centered on the Distrito 

Federal. Psyllocarpus goiasensis is found on its 

higher northern end, where occur a larger concentration of sand-based soils that are probably 

lithosols. Psyllocarpus phyllocephalus is found 

mainly on the lower center and southern end of 
8 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

F~CURES 25-33. Section Psyllocarpus seeds, SEM. Psyllocarpus laricoides: 25, abaxial view, 

X 25; 26, abaxial surface, x 340; 27, adaxial view, x 25 (25, 26, 27, Mexia 5755). Psyllocarpus 

goicrrensis: 28, abaxial view, x 14; 29, abaxial surface, x 340; 30, adaxial view, x 14 (28, 

29, 30, hin et al. 15531). Psyllocarpus asparagoides: 31, abaxial view, x 23; 32, abaxial surface, X 480; 33, adaxial view, x 23 (31, 32, 33, Irwin et al. 21998). 
NUMBER 41 9 

FIGURES 34-39.-Section Psyllocarpus seeds, SEM. Psyllocarpus phyllocephalus: 34, abaxial 

Tiew, x 34; 35, abaxial surface, x 360; 36, adaxial view, x 35 (34, 35, 36, Pires et al. 9143). 

Psyllocarpus schwackei: 37 abaxial view, x 22; 38, abaxial surface, x 360; 39, adaxial Liew, 

x 30 (37, 38, 39, Macedo 2960). 

the system. From my observations and the herbarium data, it seems to grow on soils that are regosols 

and latosols. 

Three species of section Psyllocarpus, P. laricoides, 

P. asparagoides, and P. schwackei, are found on the 

series of highlands that run up through Minas 

Gerais and Bahia. This series of mountains and 

chapadas forms the eastern watershed of the Rio 

Sao Francisco and separates the bulk of the savanna 

vegetation referred to as “cerrado,” sensu lato, from 

the coastal forests. Psyllocarpus laricoides and P. 

asparagoides are generally distributed throughout 

this region and frequently encountered. From my 

experience in the southern portions of this area 

and the herbarium label data, these two species are 

found on the thin quartzite lithosols that in this 
10 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

Psyllocarpus 

section Amazonica 

0 campinorum 

0 cururuensis 

t psyllocarpoides 

FIGURE 40.-Known distribution of section Amazonica in Amazonas, Para and Rondonia, Brazil, 

region typically support ‘campos rupestres’ (Eiten, 

1972, 1978), and P. laricoides is also encountered 

on regosols and latosols at the eastern side of the 

valley of the Rio Sao Francisco. Psyllocarpus 

schwackei is a narrow endemic that was described 

from Serra do Cipo and was known only from there 

for many years. I have examined it in the type 

locality, where it is also found in ‘campos rupestres’ 

over quartzite lithosols. Mello Barreto made two 

collections (nos. 1935 and 1939) of this species and 

labelled them as coming from Santa Luzia, which 

is near Belo Horizonte, in a much lower area with 

regosols and latosols typical of “cerrado” vegetation, sensu lato. It must be assumed that there is 

some edaphically suitable habitat near Santa Luzia 

in which P. schutackei can flourish. 

It has been independently suggested by Kirkbride 

based upon Declieuxia (1976) and by Harley based 

upon Eriope (1 976) that Pleistocene climatic changes 

have been an important factor in speciation in 

southeastern Brazil. These climatic changes have 

caused cyclical range expansions and contractions 

of the species in this area. In section Psyllocarpus, 

there are two vicariant pairs of species, P. laricoidesgoiasensis and P. asparagoides-phyllocephalus, a5 

indicated by their inflorescence forms and the presence or absence of druses. One member of each 

vicariant pair of species is found on either side of 

the drainage of the Rio Sao Francisco at higher 

elevation. The most parsimonious explanation 

would be that in a period of broader ranges, the 

ancestral populations of each species pair were in 

contact or continuous across the basin of the Rio 

Sao Francisco and that in a period of smaller ranges 

the populations were separated, giving impetus to 

their speciation through natural selection and 

genetic drift. It is also possible that the populations 

were established on each side of the basin of the 

Rio Sao Francisco by long range dispersal, but this 

seems less likely because the distribution of the seeds 

of Psyllocarpus is probably by gravity with only 

minor assistance by the wind. 
NUMBER 41 11 

FIGURE 41.-Known distribution of section PsIlZocarpus in the Distrito Federal, Goias, Bahia, 

and Ifinas Gerais, Brazil. 

Psyllocarpus Martius ex Martius & Zuccarini 

Psyllocarpus Martius ex Martius & Zuccarini, Flora 7(1), 

suppl. (4):130, 25 Apr - 7 Jul 1824; Nov. Gen. Sp. P1. 1:44, 

Oct 1824. [Not Psyllocarpus Pohl ex de Candolle, Prodr. 

4:479, 1830 (published in synonymy)]. 

Diodois Pohl ex Weldon, Flora 8(12):183, 1825 [invalid puhlication]. 

DEsCRIPTIo~.-ShrubS, subshrubs, or rarely perennial herbs, erect with a single, erect main stem, the 

secondary and tertiary stems, when present, either 

erect and sometimes fastigiate giving rise to an 

obconical habit, or spreading (P. phyllocephalus) 

and giving rise to a flat-topped habit, the stems 

quadrate to terete with 4 wings or ridges, often very 

narrow (section Psyllocarpus), the bark glabrous 

to puberulent, splitting and peeling from the larger, 

older stems, many tissues with raphides. 

Stipular sheaths uniting the leaf bases to form a 

collar about the stem, the edge of the sheaths each 

bearing 3-8 fimbriate appendages, the appendages 

narrowly triangular to subulate-linear, at apex 

narrowly acute and glandular, the central appendage longest, the lateral ones decreasing gradually 

towards each leaf base. 

Leaves opposite or sometimes ternate, sessile, the 

blade either terete with the apex narrowly conical 

or planar and narrowly to very narrowly elliptic 

with the apex narrowly acute, sometimes subtending 
12 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

axillary brachyblasts or short branchlets with 1-10 

pairs of reduced, fasciculate leaves. 

Inflorescences verticillastrate racemes or indeterminate capitula terminal and pedunculate or reduced to solitary flowers in a few distal leaf axils. 

Flowers heterostylous or homostylous, pedunculate or sessile, subtended by few or numerous setae 

or by a setiferous bracteole; hypanthium turbinate, 

glabrous or puberulous above the middle or with 

puberulous stripes subtending the calyx lobes, 

biloculate with a discoid placenta peltately attached 

to the center of the septum and bearing one peltately attached ovule; calyx 4- or 2-lobate, when 

2-lobate 1 lobe above each locule, the calyx-lobes 

triangular to narrowly triangular, narrowly acute, 

glabrous to glabrate and sometimes sparsely ciliate, 

sometimes joined at base into a tube, each charged 

with (0-)I-8 setae evenly spaced on the rim of 

the calyx-tube or of the hypanthium between the 

calyx lobes, the setae narrowly triangular, at apex 

narrowly acute and glandular, all equal or slightly 

unequal in length or rarely half as long as others; 

corolla white or blue, salverform to very rarely 

funnelform, the tube externally glabrous or variously pubescent, and internally on the upper 

y2-?,$ sericeous to sparsely sericeous and glabrate 

below, the lobes ovate or elliptic, narrowly to 

broadly acute, externally glabrous or variously 

pubescent, internally glabrous or papillate and 

sometimes apiculate at the apex; exserted stamens 

(in heterostylous species) separating from the tube 

at the apex, included stamens (in homostylous 

species) separating below the middle or (in heterostylous species) at various heights, the filaments 

glabrous, the anthers with the filament attachment 

below the middle, dorsifixed or rarely versatile, 

oblong to narrowly oblong, glabrous, the pollen 

grains oblate- or prolate-spheroidal, 5- to 8-colporate, 

in polar diameter 15-42 pm and equatorial diameter 

15-45 pm, the sexine perforate and finely and 

evenly spinulose or nonperforate with spinules 

along each side of the colpi; style terete, glabrous, 

the stigma deeply to weakly bilobate to rarely 

capitate, the lobes minutely papillate or scabrous 

or glabrous. 

Fruit a pedicellate capsule crowned by persistent 

calyx-lobes, elliptic to circular or obovate to broadly 

obovate in outline, glabrous or sometimes puberulous above the middle to varying heights, strongly 

compressed parallel to the septum, dehiscent from 

the apex downwards along the edge of the septum, 

the valves both free and spreading, the septum 

entire and erect; seeds shed from the capsule after 

dehiscence, elliptic to rarely broadly elliptic in outline, strongly compressed, weakly and irregularly 

winged, colliculate to verruculose, adaxially concave, abaxially convex, glabrous, the striophiole 

narrowly to broadly elliptic or linear to narrowly 

oblong and the ends obtuse to truncate or (P. asparagoides) the base sagittate and the lobes spreading, broadly acute. 

Key to the Sections and Species of Psyllocarpus 

1. Leaves strongly planar, the blade 1.5-10 mm wide (leaves of P. psyllocarpoides unknown); 

flowers heterostylous, the calyx 4-lobate, the stamens separating from the corolla tube wall 

above the middle, the pollen grains oblate-spheroidal, in polar diam 33-42 pm and in 

equatorial diam 33-45 Gm, the sexine perforate and finely and evenly spinulose, the stigma 

deeply bilobate; Amazonas, Para and Rondonia ... .......... Section Amazonica 

2. Bracts of inflorescence ovate to narrowly ova calyx lobes 0.5-1.4 mm 

long; fruit 3.848 x 3.2-3.8 mm. 

3. Leaf blade 6-13 mm wide; without brachyblasts; bracts 3.7-10 mm wide, subtending 

13,-17 flowers; drainages of the Rios Marmelos and Aripuana, Amazonas, and Rio 

Cururu, Para .......................... 1. P. campinorum 

3. Leaf blade 1.5-2.8 mm wide; brachyblasts with 1-3 pairs of small leaves; bracts 

0.6-3 mm wide, subtending 4-7 flowers; drainage of the Rio Cururu, Para, and near 

Porto Velho, Rondonia .............................................. 2. Bracts of inflorescence linear, 35-47 mm long; calyx lobes 1.W mm long; fruit ca. 

Leaves terete or, vary rarely (in P. Zuricoides), slightly flattened, the blade 0.1-0.5(-1) mm in 

diam; flowers homostylous, the calyx 2-lobate, the stamens separating from the corolla tube 

wall well below its middle, the pollen grains prolate-spheroidal, polar diam 15-24 pm and 

in equatorial diam 15-21 Fm, the sexine nonperforate and spinulose along each side of the 

3.2 X 1.9 mm; drainage of the Rio Urubd, Amazonas ............................ 3. P. psyllocarpoides 

1. 
NUMBER 41 13 

colpi, the stigma weakly bilobate to rarely (in P. schwackei) capitate; Minas Gerais, Bahia, and 

4. Leaf axils with brachyblasts or short branchlets; inflorescences multi-flowered verticillastrate racemes or capitula of 6-77 flowers. 

5. Inflorescences verticillastrate racemes. 

Goias ........................................................................... ................. Section Psyllocarpus 

6. Hypanthium glabrous; calyx tube 0.1-0.5 mm long; fruit glabrous: strophiole 

elliptic to broadly elliptic; Serra do Espinhaco, hlinas Gerais, and Serra do Sincora, 

Bahia ...................................................................................................... 4. P laricoih 

Hypanthium with the upper ‘/2 puberulous and lower ‘/2 glabrous; calyx tube 0; 

fruit with the upper sparsely puberulous and lower 2/9 glabrous; strophiole 

narrowly oblong; eastern half of the central region of Goih ............ 5. P. goiastnsis 

7. Habit erect and slightly spreading, the branches appearing fastigiate when 

pressed; calyx lobes 1.2-2 mm long; corolla tuh 1.1-1.5 mm long; strophiole elliptic to narrowly elliptic, at base sagittate with the lobes spreading, broadly acute; 

Serra do Espinhaco, Minas Gerais, and Chapada Diamantina, Bahia 

Habit spreading and flat topped, branches radially spreading and tangled 

when pressed; calyx lobes 2.2-3.6 mm long; corolla tube 2.2-2.6 mm long; strophiole linear to narrowly oblong, at base obtuse to truncate; Distrito Federal and 

Most leaf axils without brachyblasts or pairs of smaller leaves; flowers solitary in a few 

distal leaf axils; Serra do Cipo and Santa Luzia near Belo Horizonte, Minas Gerais ........ 

6. 

5. Inflorescence capitate. 

........................................................... ......................................... 7. 

eastern half of the central region of Goias ................................ 7. P. phyllocephalus 

...................................................................................................... 8. P. schwackei 

4. 

Ammonica, new section 

DIAGNOSIS.-A sect, Psy llocarpo foliis valde compressis lamina 1.5-10 mm lata (Psyllocarpi psyllocarpoidis foliis ignotis), floribui heterostylis calyce 

quadrilobo, staminibus e corollini tubi parieti 

supra medium secedentibus, pollinis granis oblatis 

sphaeroideis diametro polari 33-42 pm, diametro 

equatoriali 33-45 pm, sexinio perforato subtiliter 

aequaliter spinuloso, stigmate profunde bilobato 

differt. Sect. Amazonicae plantae in Brasiliae Amazonarum Rondoniae Paraeque civitatibus tantum 

crescun t. 

DESCRIPTIoN.-Shrub or subshrub, the stems quadrangular or older, larger ones terete, the wings welldeveloped. 

Leaves (unknown for P. psyllocarpoides) opposite, 

the blade planar, narrowly to very narrowly elliptic, 

narrowly acute at apex, acute at base, with or 

without brachyblasts or short branchlets in the 

axils. 

Inflorescences verticillastrate racemes, terminal, 

pedunculate, with opposite decussate bracts, each 

pair joined at base by a sheath. 

Flowers heterostylous, pedicellate, the terete pedicels subtended by a few setae; calyx lobes 4, joined 

at base to form a tube, the rim of the tube bearing 

between the lobes 1-3 evenly spaced setae; corolla 

white, the tube externally glabrous; exserted stamens separating from corolla tube wall at the 

mouth, the included ones above the middle, the 

anthers versatile or dorsifixed, the pollen grains 

oblate-spheroidal, 6- to 8-colporate, in polar diameter 33-42 Prn and in equatorial diameter 33-15 

pm, the sexine perforate and finely and evenly 

spinulose; stigma deeply bilobate, the lobes glabrous, 

scabrous or papillate; strophiole elliptic to narrowly so with the ends obtuse to subtruncate. 

TYPE.-Psyllocarpus cururzrensis, new species. 

D1sTRIBuTIoN.-Known only from “campina” 

vegetation presumably in extreme eastern Amazonas 

and the southeastern corner of Para (Figure 40). 

DrscussIoN.-Section Amazonica is well separated 

from section Psyllocarpus by its laminar leaf blades, 

heterostylous flowers with quadrilobate calyx, and 

larger, more highly sculptured pollen grains. It is 

probably the less specialized of the two sections since 

its vegetative structures more closely resemble those 

of closely related Borreria G.F.W. Meyer. 

1. Psyllocarpus campinorum (Krause), 

new combination 

PLATE I 

Borreria campinorum Krause, Verh. Bot. Vereins Prov. 

Brandenburg, 50(2):118, 1908. [Type: Uk 6102 (lectotype 

HBG; isotype K), Mar 1902, “Campina an den Cachoeiras 

des Marmelos,” Amazonas, Brazil.] 
14 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

DESCRIPTIoh..-~hrub 50-1 00 cm tall, the stems 

and branches quadrangular with wings 0.2-0.4 mm 

wide, glabrous to sparsely puberulous between the 

wings, the internodes 14-60 mm long, 1-3 mm in 

diameter. 

Stipular sheath 1.5-2.8 mm long, 2.4-4.2 mm wide 

at top, glabrous, bearing 3-7 setae, narrowly triangular, narrowly acute, 0.4-4.4 x 0.2-0.8 mm with 

the central seta longer than the lateral ones, 

glabrous. 

Leaves including blade 24-68 X 6-13 mm, glabrous, with stomata on the abaxial surface, without 

axillary brachyblasts or short branchlets. 

Inflorescences 20-40 cm long, 1-4.5 cm in diameter, 150-600-flowered, with 6-22 pairs of bracts, 

each bract subtending 13-17 flowers with a few setae 

at the base of each flower, the bracts ovate to narrowly ovate, narrowly long-acuminate or narrowly 

acute, 6.5-26 X 3.7-10 mm, glabrous, the peduncle 

quadrangular with wings 0.2-0.4 mm wide, glabrous 

to sparsely puberulous. 

Pedicel 0.1-0.3 mm long; hypanthium 0.8-1 mm 

tall, 0.6-0.8 mm in diameter, glabrous or glabrate 

to sparsely puberulous on upper v2; calyx tube 0.2- 

0.4 mm high, the rim of the tube bearing 1-3 evenly 

spaced setae between the lobes, the setae 0.1- 

0.3 X ca. 0.05 mm, glabrous or glabrate, the calyx 

lobes 0.6-1.3 X 0.2-0.6 mm; corrolla-tube 2-2.7 mm 

long, ca. 0.6 mm in diameter at base, its orifice 

0.8-1.0 mm in diameter, the lobes ovate, 1.5- 

1.9 X 0.8-1.3 mm, externally glabrous; filaments 

of exserted stamens ca. 1 mm long, of included ones 

ca. 0.2 mm long, the anthers dorsifixed, 1.1-1.8 mm 

long; style 1.2-1.4 or ca. 3.8 mm long, ca. 0.04 mm 

in diameter, the stigma 0.3-0.6 mm long, the lobes 

glabrous. 

Fruit in outline broadly elliptic to circular or 

broadly obovate, 3.8-4.4 x 3.3-3.8 mm, 1.5-1.8 mm 

thick, glabrate to sparsely puberulous above the 

middel and glabrous below, the seed 2.7-3.2 X 2.2- 

2.4 mm, ca. 0.4 mm thick, the strophiole 1-1.4 X 

0.6-0.8 mm. 

DISTRIBUT0N.-Known with certainty from the 

campinas along the Rio Marmelos and along the 

Rio Aripuana near the Transamazonian Highway, 

Amazonas, and from sandy areas along the Rio 

Cururu, Para (Figure 40). 

DrscussIoN.-This species differs from other members of section Amazonica by its wider leaves and 

inflorescence bracts, lack of brachyblasts, and large 

number of flowers. 

2. Psyllocarpus cururuensis, new species 

PLATE 2 

DIAGNOSIS.-A P. campinorum foliorum laminis 

1.5-2.8 mm latis, brachyblastis e foliorum minorum 

paribus 1-3 compositis; inflorescentiarum bracteis 

0.6-1.6 mm latis 4-7 flores subtendentibus et a P. 

psyllocarpoide inff orescen tiarum bracteis anguste 

ovatis 1.8-7.7 mm longis, fructibus 4-4.8 X 3.2-3.6 

mm differt. 

DEScRIPTIoh’.-Shrub or subshrub to 50 cm tall, 

the stems and branches quadrangular becoming 

terete with age, with wings 0.1-0.2 mm wide, puberulous to sparsely puberulose or glabrous between 

the wings, and the internodes 13-28 mm long, 1-4.2 

mm in diameter. 

Stipular sheath 0.4-0.5(-2) mm long, 1.2-2.4 mm 

wide at the top, puberulous, bearing 5 setae evenly 

spaced, the setae narrowly triangular to subulatelinear, 0.6-5 X 0.2-0.4 mm, narrowly acute and 

glandular, glabrous, the longest in the center and 

gradually decreasing in length on each side. 

Leaves including blade 19.7-38 X 1.5-2.8 mm, 

glabrous or muricate above, glabrous beneath, with 

stomata on both surfaces, subtending axillary brachyblasts or short branchlets with 1-3 pairs of smaller 

leaves. 

Inflorescences 2.5-20 cm long, 1-2.5 cm in diameter, 80-340-flowered, with 6-28 pairs of bracts, each 

bract subtending 4-7 flowers and some setae surrounding each flower, the bracts narrowly ovate, at 

apex mucronate, 1.8-22 X 0.6-3 mm, decreasing 

gradually in size from the lower primary to the 

uppermost ones, glabrous or weakly scabrous above, 

glabrous below, the sheath ca. 0.2 mm long, glabrous, the peduncle terete, 8-18 mm long, sparsely 

puberulous or glabrous. 

Pedicel ca. 0.2 mm Ion?; hypanthium 0.4-0.8 

mm tall, 0.5-0.9 mm in diameter, glabrous; calyx 

tube 0.2-0.4 mm high, its rim bearing 0, 1, or 2 

evenIy spaced setae between the lobes the setae 0.05- 

0.2 X ca. 0.05 mm, glabrous, the lobes 0.5-1.4 X 

0.2-0.5 mm; corolla-tube 1.9-2.4 mm long, 0.6-1 

mm in diameter at base, its orifice 1.2-1.8 mm in 

diameter, the lobes ovate, 1.6-2.8 X 1-1.2 mm, externally glabrous; filaments of exserted stamens ca. 
SUhIBER 41 15 

1.3 mm long, of included ones ca. 0.4 mm long, the 

anthers 0.8-1 .I mm long, the exserted ones versatile, 

the included ones dorsifixed: style ca. 1.5 or 3.2-4.2 

mm long, 0.040.08 mm in diameter, the stigma 

0.5-1 mm long, included stigma with scabrous lobes 

and exserted stigma with papillate lobes. 

Fruit elliptic in outline, 44.8 X 3.2-3.6 mm, ca. 

1.2 mm thick, glabrous, the pedicel 0.2-0.4 mm long, 

glabrous, the seed 3.2-3.4 X ca. 2.4 mm, ca. 0.1 mm 

thick, its strophiole ca. 1.6 X 0.8 mm. 

TYPE.-Anderson, da Fonseca, Reis dos Santos 

and Soztza 10779 (holotype IAK; isotypes NY, US), 

11 February 1974, region of village of Pratati, ca. 

8" S, 57"5' W. 6-8 km S of village, Para Brazil. 

DIsTRIBuTIoN.-Known from sandy habitats, possibly with campina vegetation, along the Rio Cururu, Para, and near Porto Velho, Rondonia (Figure 

40). 

DIscussIoN.-Psyllocarpus cururuensis is separated 

from P. campinorum by narrower leaves and inflorescence bracts, presence of brachyblasts, and 

fewer flowers. It is distinguished from P. psyllocarpoides by its shorter, ovate inflorescence bracts, 

shorter calyx lobes, and larger fruits. 

3. Psyllocarpus psyllocarpoides (Sucre), 

new combination 

PLATE 3 

Staelia psyllocarpoides Sucre, Rodriguesia 26(38):255, 1971. 

[Type: Froes 25262 (holotype IAK), 17 Sep 1949, Rio 

Urubu, Amazonas, Brazil.] 

DESCRIPTIoN.-$ubShrub 30-35 cm tall, the stem 

leafless at and following anthesis or indistinguishable from axis of the inflorescence. 

Inflorescences 15-18 cm long, 5-9 cm in diameter, 

90-300 flowered, with 11-22 pairs of bracts, each 

bract subtending 4-7 flowers, the bracts linear, with 

the apex narrowly acute, 35-47 X 3-4 mm, glabrous, 

the sheath 2-3 mm long, puberulous to sparsely 

puberulous. 

Pedicels 0.1-0.2 mm long; hypanthium ca. 1 mm 

tall, ca. 0.6 mm in diameter, glabrous: calyx tube 

ca. 0.2 mm high, its rim bearing 1 or 2 evenly 

spaced setae between the lobes, the setae ca. 0.15 X 

0.04 mm, glabrous and sometimes sparsely ciliate, 

the lobes 1.8-2 X 0.4-0.6; corolla-tube (fide Sucre, 

1971) 1.5 mm long, the lobes narrowly triangular: 

anthers (fide Sucre, 1971) 0.3 mm long; stigmatic 

lobes (fide Sucre, 1971) papillate. 

Fruit elliptic in outline, ca. 3.2 mm X 1.9 mm, 

ca. 1.9 mm thick, glabrous, the pedicel ca. 0.2 mm 

long, glabrous, the seed 2.4-2.9 X 1.4-2 mm, ca. 

0.4 mm thick, the strophiole 1-1.6 X 0.5-0.7 mm. 

DISTRIBUTION.-Known only from the type locality, campinarana vegetation near the Rio Urubu 

Amazonas (Figure 41). 

DIscussIoN.-This is easily separated from the 

other members of section Amazonica by its longer, 

linear inflorescence bracts, longer calyx lobes and 

smaller fruit. 

Section Psyllocarpus 

DEscRIPTIoN.-shrub, subshrub, or perennial herb, 

the stems terete or quadrangular to weakly quadrangular sometimes with ridges or small wings. 

Leaves opposite or sometimes ternate, the blade 

terete with apex conical to very rarely (in P. Zaricoides) planar and linear with apex narrowly acute, 

subtending axillary brachyblasts or short branchlets 

with 2-10 pairs of smaller leaves or very rarely (in 

P. schwackei) with 1 or 2 pairs of smaller axillary 

leaves. 

Inflorescences terminal verticillastrate racemes or 

capitulate or (in P. schwackei) solitary flowers in a 

few distal leaf axils. 

Flowers homostylous, sessile or pedicellate, subtended by setae or 1 or 2 setiferous bracteoles, the 

pedicel terete and glabrous when present; calyx 

lobes 2, free or joined at base to form a tube, the 

rim of the calyx tube or of the hypanthium bearing 

0-8 evenly spaced setae between the lobes; corolla 

blue or white, the tube externally glabrous or variously pubescent; stamens included, the filaments 

separating from corolla tube wall well below the 

middle, the anthers dorsifixed, the pollen grains 

prolate-spheroidal, 5- to 7-colporate, in polar diameter 15-24 pm and in equatorial diameter 15-21 pm, 

the sexine nonperforate with spinules along each 

side of the colpi; stigma weakly bilobate to rarely 

(in P. schzvackei) capitate, minutely papillate; 

strophiole narrowly to broadly elliptic or linear to 

narrowly oblong with the ends obtuse to truncate, 

or (P. asparagoides) sagittate at base with the lobes 

spreading, broadly acute. 

TYPE.-Psyllocarpus laricoides Martius ex Martius & Zuccarini (Lectotype). 
16 SMITHSONIAN CONTRIBUTIOSS TO BOTANY 

DISTRIBUTION.-Known from the chapadao and 

its extensions west of the basin of the Rio Sao Francisco in Distrito Federal and Goias, and from the 

upper and western slopes of the Serra do Espinhaco, 

Minas Gerais, and of Chapada Diamantina, Bahia, 

east of the basin of the Rio Sao Francisco (Figure 

41). 

DIscussIoN.-section Psy llocarpus is readily distinguished from section Amaronica by its ericoid 

foliage or needle-like, terete leaves, homostylous 

flowers with bilobate calyx, and smaller, less sculptured pollen grains. 

4. Psyllocarpus laricoides Martius ex 

Martius & Zuccarini 

PLATE 4 

Psyllocarpus laricoides Martius ex Martius & Zuccarini, 

Flora 7(1), suppl. (4):131, 25 Apr - 7 Ju1 1824; Nov. Gen. 

Sp. PI. 1:45, pl. 28, Oct 1824. [Type: Martius s.n. (lectotype M, plant farthest left on the herbarium sheet; presumed isotype M), 1818, “habitat sumo Brasiliae monte 

ItamM,” Minas Gerais, Brazil.] 

Psyllocarpus ericoides Martius ex Martius & Zuccarini, Flora 

7(1) suppl. (4):131, 25 Apr - 7 JuI 1824; Sov. Gen. Sp. 

P1. 1:45, pl. 28, Oct 1824. [Type: Martius 1295 (holotype 

M; presumed isotype M), 1818, “habitat in campis ad 

Bandeirinha, propr Tijuco,” hlinas Gerais, Brazil.] 

Psyllocarpus laricoides var. p densifolius Martius & Zuccarini, 

Nov. Gen. Sp. P1. 1:46, Oct 1824. [Type: Martius s.n. 

(holotype hl, plant farthest right on the herbarium sheet), 

1818, “habitat sumo Brasiliae monte Itambe,” Minas 

Gerais, Brazil.] 

Psyllocarpus Iaricinus Sprengel, Caroli Linnaei, Syst. veg., 

16th edition, 4(2):39, 1827 [superfluous name]. 

Psyllocarpus laricoides var. y longicornu Schumann, Mart. 

F1. bras. 6(6):33, 1888. [Type: Pohl 1317(852) (lectotype 

W; isotypes K, W), 1820, Ouro fino, Minas Gerais, Brazil.] 

DEsCRIPTIoN.-shrub, subshrub, or perennial herb 

22-120 cm tall, obconical in outline, the stems and 

branches erect and fastigiate to slightly spreading, 

moderately to sparsely branched, quadrangular to 

weakly quadrangular and terete with age, with narrow wings 0.05-1 mm wide, puberulous to sparsely 

puberulous or glabrous, the bark thin, splitting and 

peeling from the older stems and branches, the 

internodes 3-28 mm long, 0.4-5.4 mm in diameter. 

Stipular sheath 0.6-1.4 mm long, (0.9-)1.4-2 mm 

wide at the top, puberulous to glabrate or glabrous, 

bearing 4-6 setae evenlv spaced, these narrowly triangular to linear-subulate, narrowly acute, (0.2-) 

0.3-1.4 X 0.04-0.28(-0.4) mm, all equal in length or 

with some half as long as other, scabrous or glabrous 

and ciliate to minutely ciliate. 

Leaves opposite or ternate, the leaf blades (4-) 

5-16(-21) mm long, 0.2-1 mm in diameter or wide, 

scabrous to glabrate or glabrous, subtending axillary 

brachyblasts with 2-7 pairs of leaves forming dense 

fascicles. 

Inflorescences verticillastrate racemes, terminal, 

pedunculate, 2.1-8.5 cm long, 0.8-1.2 cm in diameter, 8-48-flowered, with opposite and decussate or 

rarely ternate bracts in 4-13 pairs or whorls, with a 

sheath joining the bases or each pair or trio of 

bracts, each bract subtending 1 or sometimes 2 

flowers, the bracts decreasing upward gradually, narrowly oblong to linear, narrowly acute, 1.2-7.3 X 

0.3-0.8 mm, scabrous to glabrate or glabrous, the 

sheath gradually decreasing in length from the 

lowermost to the uppermost, 0.61 mm long, puberulous to glabrate or glabrous, the peduncle terete, 

1.4-19 mm long, puberulous to sparsely puberulous 

or glabrous. 

Pedicels 0.1-0.4 mm long, subtended by 1 or 2 

setiferous bracts 0.2-0.4 mm long or by numerous 

setae; hypanthium 0.8-1.4 mm tall, 0.8-1.4 mm in 

diameter, glabrous; calyx lobes 1-2.8 X 0.2-0.8 

mm, joined at base to form a tube 0.1-0.5 mm high, 

glabrous, its rim bearing 3-7 evenly spaced setae 

between the lobes, the setae 0.1-0.8 X 0.01-0.2 mm, 

glabrous and minutely ciliate: corolla blue, the tube 

(2.4-)2.9-6.7 mm long, 0.7-1.2 mm in diameter at 

base, the orifice 1.2-2 mm in diameter, externally 

scabrous or glabrous, the lobes ovate to broadly SO 

or elliptic to very broadly elliptic, 1.2-3.2 X 0.8-2.2 

mm, externally glabrous or scabrous, internally 

papillate to minutely papillate; filaments 0.2-0.4 

mm long, the anthers 0.9-1.3 mm long; style 0.3-0.7 

mm long, 0.1-0.3 mm in diameter, the stigma 

weakly bilobate, 0.2-0.3 mm tall, 0.2-0.4 mm in 

diameter. 

Fruit elliptic to broadly elliptic in outline, 2.8- 

4.4 X 1.8-3.2 mm, 1-1.6 mm thick, glabrous, the 

pedicels 0.2-0.6 mm long, glabrous, the seed 2.6-3.6 

X 1.7-2.6 mm, ca. 0.1 mm thick, its strophiole 

elliptic to broadly elliptic, obtuse at both ends, 

0.9-2 X 0.4-1.1 mm. 

DIsTRIBuTIoN.-~nown from the higher regions 

of Serra do Espinhaco, Minas Gerais, and of Serra 

do Sincora, Bahia. 

DIscussIoN.-Psy~~ocarpus laricoides is most closely 

related to P. goiasensis, from which it is set apart 
NUMBER 41 17 

by its glabrous hypanthium and fruit and by the 

presence of a calyx tube. These are vicarious species 

whose ranges are separated by the lower portions 

of the basin of the Rio Sao Francisco. From my own 

field experiences with P. laricoides and the data in 

the herbaria, this species appears to be found in 

the vegetation referred to by Eiten (1972) as 

“campos rupestres” that are characteristically formed 

on thin lithosols of quartz sand. 

5. Psyllocarpus goiasensis, new species 

PLATE 5 

DIACNOSIS.-~~ P. laricoide hypanthiis fructibusque supra medium puberulis infra medium glabris, calycisque tub0 nullo differt. 

DESCRIPTIoN.-Subshrub 3 1-63 cm tall, stem and 

branches erect and slightly spreading, moderately 

branched, terete to subquadrangular with ridges 

or small wings 0.05-1 mm wide, puberulous, the 

bark thin, splitting, and peeling from the older 

stems and branches, the internodes 421 (-24) mm 

long, 0.5-3.7 mm in diameter. 

Stipular sheath (0.6-)0.8-1 mm long, 1.2-1.6 mm 

wide at the top, puberulous, bearing 5-8 setae 

evenly spaced, these narrowly triangular, narrowly 

acute, 0.6-2 X 0.1-0.3 mm, all equal in length or 

with some half as long as others, scabrous. 

Leaves opposite, the leaf blade (5.5-)7.3-12.5 mm 

long, 0.2-0.3 mm in diameter, scabrous, subtending 

axillary brachyblasts with 4-9 pairs of smaller leaves 

forming dense fascicles. 

Inflorescences verticillastrate racemes, terminal, 

pedunculate, 1.5-6.5 cm long, 0.8-1.7 cm in diameter, 16-77-flowered, with opposite and decussate 

bracts in 5-11 pairs and a sheath joining the bases 

of each pair of bracts, each bract subtending 2-4 

flowers surrounded by setae, the bracts decreasing 

upward gradually in size, linear, narrowly acute, 

2.8-5.2 X 0.3-0.5 mm, scabrous, the sheath upward 

gradually decreasing in length 0.4-1.4 mm long, 

puberulous to sparsely puberulous, the peduncle 

terete, (2-)5-17 mm long, puberulous. 

Flowers sessile, the hypanthium 1.6-1.7 mm tall, 

1-1.2 mm in diameter, with the upper y2 puberulous 

and the lower glabrous or glabrate; calyx lobes 

2.4-3 X 0.7-2 mm; 4-8 setae evenly spaced along 

the apical rim of the hypanthium between them, 

the setae 0.2-1.4 X 0.04-0.24 mm, glabrous and 

minutely ciliate; corolla blue or white, the tube 

3.6-4.6 mm long, externally glabrate to sparsely 

puberulous, ca. 0.8 mm in diameter at base, the 

orifice 1-1.2 mm in diameter, the lobes ovate to 

broadly ovate, 1.8-2 X 1.2-1.3 mm, externally glabrous, internally minutely papillate, tipped with an 

apicule ca. 0.1 mm long; filaments 0.2-0.3 mm long, 

the anthers 0.8-1 mm long; style 0.4-0.6 mm long, 

ca. 0.1 mm in diameter, the stigma bilobate, 0.1-0.2 

mm tall, ca. 0.2 mm in diameter. 

Fruit elliptic in outline, 3.4-4(-4.8) X 2.3-2.4 

(-3) mm, 1.4-1.7 mm thick, the upper ‘/9 sparsely 

puberulous and the lower ?,$ glabrous, the seed 2.2- 

4.3 x 0.8-2.7 mm, 0.04-0.28 mm thick, the strophiole narrowly oblong 1.6-2.4 X 0.3-0.5 mm, its ends 

truncate. 

TYPE.-Zrwin, Harley, and Smith 33082 (holotype 

UB; isotypes NY, US), 24 Rlarch 1971 ca 10 km N 

of Alto Paraiso, Gioas, Brazil. 

DIsTRIBuTI0N.-Known from the Chapada dos 

Veadeiros and Serra Geral do Parana as far south 

as Formosa, east-central Goias (Figure 41). 

DIscussIoN.-This species is most closely related 

to P. laricoides, from which it is separated by the 

hypanthium puberulous above the middle, the absence of a calyx tube, and the fruit with the upper 

third sparsely puberulous. As was pointed out earlier, this species forms with P. laricoides a vicarious 

pair whose ranges are separated by the lower basin of 

the Rio Sao Francisco. The Chapada dos Veadeiros 

has many areas in its higher reaches that have soils 

composed of quartz sands that are probably lithosols. The scanty herbarium data would indicate that 

P. goiasensis is found on this type of soil as is P. laricoides in the Serra do Espinhaqo. 

6. Psyllocarpus asparagoides Martiue ex 

Martiue & Zuccarini 

PLATE 6 

Psyllocarpus asplragoides Martius ex Martius & Zuccarini, 

Flora 7(1), suppl. (4):131, 25 Apr - 7 Jul 1824; Nov. Gen. 

Sp. P1. 1:46, Oct 1824. [Type: Martius s.n. (holotype M; 

presumed isotype M), 1818, “habitat in campis altis termini Minarum Novarum prope Piedade,” Minas Gerais, 

Brazil.] 

DEScRIPTIoN.-.%bshrub 15-65 cm tall, the stems 

and branches erect and slightly spreading, terete, 

sometimes with ridge ca. 0.05 mm high, puberulous 
18 SMITHSONIAN CONTRIBUTIOSS TO BOTANY 

to sparsely puberulous on older stems, the bark 

thin, reddish, splitting, decidous from older stems, 

the internodes 3-35(-65) mm long, 0.3-3 mm in 

diameter. 

Stipular sheath 0.5-0.8 mm long, 0.6-2.4 mm wide 

at top, glabrous to puberulous, bearing 3-8 setae 

evenly spaced, these narrowly triangular to subulatelinear, narrowly acute, 0.1-0.8 X 0.03-0.15 mm, all 

equal in length or sometimes with shorter ones 

x-fi as long as the others, glabrous to minutely 

sparsely scabrous. 

Leaves opposite, the blade 8-19 mm long, 0.15- 

0.3 mm in diameter, glabrous, with 3 faint parallel 

ridges running from base to apex on the abaxial 

side, subtending axillary brachyblasts with (3-)4-10 

pairs of shorter leaves forming dense fascicles. 

Inflorescences capitulate, terminal, pedunculate, 

5-15 mm in diameter, I6-flowered, with 4 decussate 

pairs of bracts, each pair joined by sheaths to form 

concentric rings within the inflorescence, glabrous 

to sparsely puberulous, each bract subtending 2 

(very rarely 3 or 4) flowers surrounded by numerous 

setae, the primary bracts linear to narrowly oblong, 

narrowly acute, 3-6.7 X 0.3-0.5 mm, with sheath 

0.8-2.2 mm long, the secondary ones narrowly oblong, narrowly acute, 2-4.4 X 0.4-0.5 mm, with 

sheath 0.8-1.2 mm long, the tertiary ones narrowly 

oblong, narrowly acute, 1.2-2.3 X 0.2-0.4 mm, with 

sheath 0.4-0.8 mm long, the quaternary ones narrowly triangular, narrowly acute, 0.4-1.3 X 0.1-0.2 

mm with sheath 0.2-0.4 mm long, the peduncle 

terete, (0.3-)1.84.8 cm long, glabrous to sparsely 

puberulous. 

Pedicels 0.1-0.2 mm long: hypanthium 1-1.2 mm 

tall, 0.7-0.9 mm in diamcter, glabrous; calyx lobes 

1.2-2 X 0.3-0.9 mm, with 0-5 setae evenly spaced 

along the rim of the hypanthium between the lobes, 

the setae 0.1-0.2 X 0.04-0.08 mm, glabrous; corolla 

white, the tube 1.1-1.5 mm long, externally granulate, the orifice 0.9-1.2 mm in diameter, 0.5-0.9 

mm in diameter at base, the lobes broadly to very 

broadly ovate, 0.7-1.5 X 0.6-1.2 mm, externally 

granulate, internally papillate; filaments 0.1-0.2 

mm long, the anthers 0.4-0.6 mm long; style 0.2-0.4 

mm long, ca. 0.1 mm in diameter, the stigma weakly 

bilobate, 0.1-0.2 mm tall, ca. 0.2 mm in diameter. 

Fruit obovate to broadly obovate in outline, 

3-3.7 X 2.2-3 mm, 0.8-1.3 mm thick, glabrous, the 

pedicel 0.3-0.5 mm long, glabrous, the seed 2.7- 

3.4 X 1.8-2.8 mm, ca. 0.2 mm thick, its strophiole 

elliptic to narrowly elliptic, the apex obtuse, the 

base sagittate with spreading, broadly acute lobes 

0.6-0.9 X 0.3-0.4 mm. 

DISTRIBUTION.-Known from higher elevations of 

the Serra do Espinhaco, Minas Gerais, and of the 

Chapada Diamantina, Bahia (Figure 41). 

DIsCussIoN.-Ps~~locarpus asparagoides is most 

closely related to P. phyllocephalus, from which it 

can be distinguished by its erect, fastigiate habit 

and shorter calyx lobes and corolla tube. The degree of similarity and therefore probable kinship 

between P. asparagoides and P. phyllocephalus is 

not nearly as close as that between P. Zaricoides and 

P. goiasensis, but they are also a pair of vicarious 

species with similar distributions again separated by 

the lower drainage of the Rio Sao Francisco. 

7. Psyllocarpus phyllocephalus Schumann 

PLATE 7 

Psyllocarpus phyllocephalus Schumann, Bot. Jahrb. Syst. 

25(3), suppl. (60):17, 19 Jul 1898. [Type: Glazious 21504 

(lectotype P; isotypes BR, C, G, K, LE, S), “entre Paranaua et le Rio Torte,” Distrito Federal, Brazil.! 

DEscRrPTIoN.-Subshrub 12-50 cm tall, flattopped, the stem erect, terete with ridges ca. 0.05 mm 

high, puberulous to sparsely puberulous, diffusely 

and extensively branched at the apex, the branches 

terete, puberulous, the bark thin, splitting, decidous 

from the older portions of the stem and branches, 

the internodes 4.2-8.3 mm long, 0.3-2.7 mm in 

diameter. 

Stipular sheath 0.8-1.2 mm long, 0.8-1.6 mm wide 

at top, puberulous to sparsely puberulous, bearing 

3-6 setae evenly spaced, these narrowly triangular 

to sublinate-linear, narrowly acute and glandular at 

apex, 0.3-1.2 X 0.08-0.1 mm, all equal in length or 

occasionally some ca. fi as long as the rest, glabrous 

to sparsely puberulous. 

Leaves opposite or ternate, the left blade 4.5-10 

mm long, ca. 0.2 mm in diameter, weakly scabrous, 

with a narrow, flat, abaxial face running from the 

base to the apex, subtending axillary brachyblasts 

with (3-)4-8 pairs of smaller leaves forming dense 

fascicles. 

Inflorescences capitulate, terminal, sessile or 

pedunulate, 5-1 7 mm in diameter, 6-24-flowered, 

with 3-7 sets of opposite and decussate or rarely 

ternate bracts, each pair or trio joined by sheaths to 
NUMBER 41 19 

form concentric rings within the inflorescences, each 

bract subtending 1-3 flowers surrounded by numerous setae, the bracts decreasing upward gradually 

in size, linear, narrowly acute, 3.3-8.3 X 0.2-0.4 

mm, glabrous or scabrous, the sheath gradually decreasing upward in length, 0.6-1.5 mm long, glabrous to sparsely puberulous, the peduncle if pressent terete, 1.5-9 mm long, puberulous to sparsely 

Flowers sessile, the hypanthium 0.8-1.2 mm tall, 

0.6-1.1 mm in diameter, glabrous or with a puberulous stripe subtending each calyx lobe or with the 

upper 14 glabrate; calyx lobes 2.2-6 X 0.4-0.6 mm, 

with 3-5 setae evenly spaced along the rim of the 

hypanthium between them, the setae 0.3-0.8 X 

0.04-0.16 mm, sparsely puberulous; corolla blue or 

white, the tube externally puberulous or scabrous, 

2.2-2.6 mm long, 0.4-0.7 mm in diameter at base, 

the orifice 0.5-1 mm in diameter, the lobes ovate 

to narrowly ovate, 0.8-1.2 X 0.4-0.7 mm, externally 

puberulous to scarbrous, internally papillate, apiculate at apex; filaments 0.1-0.3 mm long, the anthers 

0.4-0.5 mm long; style 0.2-0.5 mm long, ca. 0.1 mm 

in diameter, the stigma weakly bilobate, 0.2-0.3 

mm tall, 0.1-0.3 mm in diameter, 

Fruit obovate in outline, 2.2-2.9 X 1.2-2 mm, 

1.1-1.5 mm thick, glabrous or upper ‘/s sparsely 

puberulous, the seed 1.7-2.2 X 0.9-1.6 mm, 0.2-0.3 

mm thick, the strophiole linear to narrowly oblong, 

0.7-1.4 X 0.1-0.3 mm, its ends obtuse to subtruncate. 

DIsTRIBUTION.-Frequent in the Distrito Federal 

and known from the Serra dos Cristais, the vicinity 

of the Serra dos Pirineus and the southernmost 

flanks of the Chapada dos Veadeiros in east-central 

and southeastern Goias (Figure 41). 

DIscUssIoN.-This species is related to P. asparagoides, from which it is separated by its spreading, 

frequently branched, flat-topped habit that appears 

as a circular, tangled mass when dried for the 

herbarium and by the longer calyx lobes and corolla 

tube (see page 17). 

so. 

8. Psyllocarpus schwackei Schumann 

PLATE 8 

Psyllocarpus schwackei Schumann, Bot. Jahrb. Syst. 29(3). 

suppl. (60):18, 19 Ju1 1898. [Type: Anderson, Stieber, and 

Kirkbride 36254 (neotype US; isotypes NY, UB), 18 Feb 

1972, Serra do Cipo, elev. ca. 1125 m, Minas Gerais, 

Brazil.] 

DESCRIPTIoN.rubshrubs or perennial herbs 15- 

60 cm tall, the stems and branches erect and slightly 

spreading, terete, rarely with faint ridges ca. 0.02 

mm high, glabrous, the internodes (4-)7-25(-27) mm 

long, 0.1-3.3 mm in diameter. 

Stipular sheathes 0.6-1 mm long, 0.5-1.5 mm wide 

at top, glabrous, bearing 3-5 setae evenly spaced, 

these narrowly triangular, narrowly acute, 0.2-1 X 

0.04-0.2 mm, all equal in length or sometimes the 

central one or ones up to twice as long as the lateral 

ones, glabrous or sometimes sparsely and minutely 

ciliate, 

Leaves opposite, sometimes subtending 1 or rarely 

2 pairs of smaller axillary leaves, the leaf blade 8-24 

mm long, 0.2-0.5 mm in diameter, glabrous. 

Inflorescences of solitary flowers in a few distal 

leaf axils. 

Pedicels 0.2-0.5 mm long, subtended by a setiferous bracteole 0.7-1 mm long, the setae Y3-G as 

long as the bracteole; hypanthium 0.8-1.3 mm tall, 

0.8-1.3 mm in diameter, glabrous; calyx lobes 1-1.8 

x 0.3-0.6 mm, joined at base to form a tube 0.15- 

0.25 mm high, glabrous, the rim of the tube bearing 

3-6 setae between the lobes, the setae 0.2-0.6 X 

0.1-0.2 mm, sparsely and minutely ciliate but 

facially glabrous; corolla blue, externally minutely 

and densely papillate, the tube 3.8-6.1 mm long, 

0.7-1.1 mm in diameter at base, the orifice 1.2-1.8 

mm in diameter, the lobes elliptic to broadly elliptic, 1.6-3.7 x 1-2.2 mm, externally minutely and 

densely papillate, internally glabrous, tipped with 

a setae standing at right angles to its surface; filaments 0.2-0.5 mm long, the anthers 0.9-1.3 mm 

long; style 0.1-0.6 mm long, 0.08-0.25 mm in diameter, the stigma capitate to weakly bilobate, ca. 0.2 

mm tall, 0.1-0.3 mm in diameter. 

Fruit elliptic in outline, 2.7-4.4 X 1.8-2.7 mm, 

0.7-1.1 mm thick, glabrous, the pedicel 0.44.8 mm 

long, glabrous, the seed 1.5-2.5 X 1-1.6 mm, ca. 

0.1 mm thick, the strophiole elliptic, ca. 0.7 X 

0.4-0.5 mm, its end obtuse. 

DIs’mIsumoN.-Known from Serra do Cipo in the 

Serra do Espinhaco and from Santa Luzia near Belo 

Horizonte on the wester slope of the Serra do 

Espinhaco, Minas Gerais (Figure 41). 

DIsCUSSIoN.-In his publication of P. schwackei 

(1 898), Schumann cited the following collection, 

“in Brasiliae civitate Minas Geraes, locis arenosis in 

Serra do Cipo: Schwacke n, 8089, floret Aprili.” The 
20 SXIITHSONIAN COSTRIBUTIOh’S TO BOTANY 

specimen studied by Schumann at Berlin was destroyed in the Second World War, and I have been 

unable to locate any duplicates of this collection. In 

the Field Museum of Natural History type photograph series, there is a photograph of the specimen 

that existed in Berlin prior to the war, negative 

number 896. Based upon the description, the photograph of the Berlin specimen, and my experience at 

the type locality, Serra do Cipo, I have selected my 

topotypical collection, Anderson, Stieber, and Kirkbride 36254, as neotype. 

Psyllocarpus schwackei is easily separated from 

the other species of section Psyllocarpus by its lack 

of brachyblasts and only rare occurrence of 1 or infrequently two pairs of smaller leaves in leaf axils 

and an inflorescence structure of only a few solitary 

flowers in some distal leaf axils. The relationships 

of P. schwackei are very difficult to assess because of 

its reduction in these characters. In view of its lack 

of capitular inflorescence development, it would 

seem to be related to the species pair P. laricoidesgoiasensis, but not closely. 

Excluded Species 

Psyllocarpus thymbroides Martius ex 

Martius & Zuccarini 

Psyllocarpus thyrnboides Martius ex Martius & Zuccarini, 

Flora 7(1), suppl. (4):131, 23 Apr - 7 Ju1 1824; Nov. Cen. 

Sp. PI. 1:46, Oct 1824. [= Staelia thymbroides (Martius 

ex Martius & Zuccarini) Schumann, Mart. F1. bras. 6(6):77, 

1888.1 

DIscussIoN.-Martius and Zuccarini described 

Psyllocarpus thym broides from a collection made by 

Martius “in campis montosis ad montium tractum 

Serra do Gran Mogol adamantibus divitem, Provinciae Mina Geraes” in July of 1818. They gave a 

diagnostic phrase of one sentence, one line of general description, the collection locality and time of 

flowering, but omitted those characteristics of the 

fruit that are essential for determining the generic 

affinity. I have not been able to locate any specimens 

of the collection upon which they based their species. 

Schumann in the Flora brasiliensis (1888) transferred the epithet to Staelia, citing, however, in 

place of the type collection, Martius s.n. (in alpinis 

ad Rio das Contas, Bahia, Brasil) and Blanchet 

3605 (in Serra da Jacobina, Bahia, Brasil). Apparently, he too was unable to locate the type collection. Schumann’s extensive description matches the 

scanty protologue in all particulars except that it 

fails to mention fasciculate leaves, but the collections that he cited do exhibit this character. In my 

opinion, Schumann neotypified Psyllocarpus thymbroides by the citation of the above neosyntypes and 

transferred it to Staelia based upon this typification. 

I have examined Schumann’s neosyntypes, and 

they are both members of the genus Staelia. They 

show what may be significant differences from each 

other that may necessitate their specific or subspecific segregation. I now have underway a revision 

of the genus Staelia that will resolve this question. 
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Tribu XVIII. Spermacoceae. In T. Lasser, Flo~a de 

Venezuela, 9(3): 1834-1978. 

1974. 

Sucre, D. 

1971. Estudo das Rubiaceae Brasileiras, 111: Cinco novas 

especies da Tribo Spermacoceae. Rodriguesia, 26(38): 

253-260. 

Uphoff, J.C.T. 

1962. Plant Hairs. In M’. Zimmermann and P. G. Ozenda, 

editors, Encyclopedia of Plant Anatomy, IV(5): 

292 pages. Berlin: Gebruder Borntraeger. 

Verdcourt, B. 

1975. Studies in the Rubiaceae, 1: Rubioideae ior the 

“Flora of Tropical East Africa.” Keul Bulletin, 

30(2):247-326. 

1976. Rubiaceae, 1. In R. M. Polhill, Flora of Tropical 

East Africa. 414 pages. London: the IYhitefriars 

Press Ltd. 

FVelden, F. L., yon 

It‘ernham, H. F. 

1825. 11, Correspondenz. Flora, 8(12):182-190. 

1916. Tropical American Rubiaceae, VII. Journal of 

Botany, British and Foreign, 54:322-334. 

Weston, G. D., and D. D. Cass 

1973. Observations on the Development of the Paraveinal Mesophyll of Soybean Leaves. Botanical 

Gazette, 134232-235. 
Index to Exsiccatae 

(Numbers in parentheses refer to species treated in this paper) 

Anderson, IV. R., M. Arroyo, S. R. Hill, R. Reis dos Santos, 

and R. Souza 6437 (5); 7161 (5); 7860 (7) 

Anderson, W. R., S. G. da Fonseca, R. Reis dos Santos, and 

R. Souza 10779 (2); 10918 (1) 

Anderson, W. R., P. A. Fryxell, S. R. Hill, R. Reis (10s 

Santos, and R. Souza 8075 (7); 8374 (4); 8637 (6) 

.Inderson, W. R., M. Stieber, and J. H. Kirkbride, Jr. 35075 

(7); 35107 (4); 35230 (4); 35382 (4); 35595 (4); 35750 (4); 

35761 (4); 36216 (4); 36254 (8) 

Badini, J. 3285 (4) 

Bahia, T. R. 32 (1) 

B,irl)oza, H. s.ti. (4) 

Baring, C. s.71. (4) 

Barreto, hf. 1935 (8); 1939 (8); 3454 (4); 4038 (4); 4039 (4); 

Barieto, &I. 1128, and A. C. Brade 14736 (8) 

Barreto, Xl. 1130, and A. C. Brade 14737 (4) 

Barreto, M., and A. P. Viegas 6349 (4) 

Blanchet, J S. 2534 (6); 3332 (6); s.n. (6) 

Brade, .4. C. 13448 (4); 13838 (6); 13839 (4); 13847 (4); I3988 

Bunbniy, C J.F. s.n. (4) 

8460 (4); 9071 (4); 9968 (4); 9995 (4); 10726 (8) 

(4) 

Campos, R. F. s.n. (4) 

Castellanos, A. 25970 (6) 

Casaretto, G. 2713 (4) 

Claussen, P. 5 (4); 64 (4); 162 (4); I82 (4); I83 (pro parte) 

(4); 264 (4); 336 (4); 603 (4); s.n. (4) 

Daniazio, L. 234 (4); 1647 (4); s.n. (4) 

Dawson, E. 9. 14169 (5) 

Duarte, A. P. 2113 (4); 2595 (4); 2655 (8); 2730 (4); 8528 

Duarte, .i. P., and G. Barroso 7931 (4) 

Duarte, .I. P. 9199, and E. Pereira 10109 (6) 

(4); 9697 (8); 11381 (8); 11580 (4) 

Egler, W. A. I238 (2) 

Eiten, G., and L. T. Eiten 6707 (4); 6799 (8) 

Froes, R. L. 25262 (3) 

Gardner, G. 4735 (4); 4736 (4); 4737 (4) 

Glaziou, A.F.M. 8470 (4); If930 (6); 14931 (4); 14932 (4); 

17641 (4); I9428 (7); 19429 (7); 19430 (6); 19431 (4); 

19432 (4); 19432~ (4); 19540 (8); 20350 (4); 21504 (7) 

Gounelle 41 (4) 

Harley, R. M., S. A. Renvoize, C. M. Erskine, C. A. Brighton, 

Hatschbach, G., W. R. i\nderson, R. Barneby, and B. Gates 

Hatschhach, G., and P. Pelanda 27900 (6); 28001 (4) 

Hatschbach, G., L. B. Smith, and E. Avensu 29059 (4) 

Heringer, E. P. 10683 (7); I1451 (5); 13078 (7); s.n. (7) 

Heringer, E. P., and A. Castellanos 22080 (8); 22204 (4) 

Hoehne, F. C. 4914 (4); 4937 (4) 

and R. Pinheiro 15055 (6); 15355 (6) 

36224 (7); 36340 (5); 36410 (5); 36475 (6) 

Irwin, H. S., 7.V. R. Anderson, M. Stieber, and E. Y.-T. Lee 

Irwin, H. S., S. F. da Fonseca, R. Souza, R. Reis dos Santos, 

and J. Ramos 26662 (7); 27090 (4); 27490 (4); 27643 (6): 

27724 (4); 28097 (4); 28313 (4); 28316 (6); 28493 (6) 

Irwin, H. S., J. W. Grear, Jr., R. Souza, and R. Reis dos 

Santos 12221 (7); 12870 (5); 13445 (7); 13656 (7) 

Irwin, H. S., R. M. Harley, and E. Onishi 28875 (4); 28967 

(4); 29135 (4); 29254 (4); 29634 (4); 30430 (4) 

Irwin, H. S., R. XI. Harley, and G. L. Smith 30890 (6); 32482 

(6); 33082 (5) 

Irwin, H. S., H. Maxwell, and D. C. TVasshausen 18633 (7); 

19449 (7); 19888 (4); 19976 (8); 20642 (4); 20659 (4) 

Irwin, H. S., R. Reis dos Santos, R. Souza, and S. F. da 

Fonseca 21850 (4); 21998 (6); 22085 (4); 22247 (6); 22352 

(4); 22359 (4); 22597 (4); 22720 (4); 23118 (6); 23466 (4); 

23572 (4); 24745 (5) 

34434 (7) 

Irwin, H. S., and T. R. Soderstrom 5143 (7); 5655 (7) 

Irwin, H. S., R. Souza, J. W. Grear. Jr., and R. Reis dos 

Irwin, H. S., R. Souza, and R. Reis dos Santos 8224 (7); 

Santos 15531 (5) 

8993 (7); 10678 (7) 

Kithlmann, J. G. s.n. (4) 

Langsdorff, G. H. s.n. (4) 

Lanstyak, L. s.n. (4) 

Lima, A. 58-2957 (7) 

Lund, P. W. s.n. (4) 

Macedo, A. 2067 (4); 2827 (4); 2960 (8) 

Magalhaes, hl. 1281 (4); 1785 (4); 2364 (4); 2586 (4); 16909 

hfarkgraf, F. 3406, M. Barreto 12011 and A. C. Brade(4) 

Martius, K.F.P. von 998 (4); I295 (4); s.n. (4); s.n. (6) 

Mexia, Y. 5755 (4) 

Mota, C. D., and L. Coelho 209 (2) 

(4); 17981 (4) 

23 
24 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

Oliveira, J. E. de s.n. (8) 

Palacios, Balegno, and Cuezzo 3976 (4) 

Pereira, E. 1566 (6); 7385 (7); 8892 (8); 9070 (7) 

Pereira, E. 2526, and G. Pabst 3362 (4) 

Pereira, E. 2834, and G. Pabst 3670 (4) 

Philcox, D., and E. Onishi 4815 (7) 

Pires, J. M., X. T. Silra, and R. Souza 9143 (7) 

Pohl, J. E. 1317(851) (4); 3445 (4); 3477 (4); s.n. (4) 

Prance, G. T., and N. T. Silva 58197 (5) 

Ramariz, D. 0309 (4) 

Reinhardt, J. T. s.n. (4) 

Riedel, L. 39 (4); 376 (4); 1238 (6); 1335 (4); 1385 (4); 1884 

(4): 2884 (4); SA. (6) 

Saint-Hilaire, A. de Bl 835 (4); Bl 989 (4); B1 1157 (4); 

s.n. (4) 

Silva, N. T. 4346 (1) 

Sampiao, A. J. de 6828 (4) 

Santos, G. s.n. (4) 

Schwacke, C.A.\t7. s.n. (4) 

Segadas-Vianna, F. 6044 (4) 

Segadas-Vianna, F., and J. Loredo 1081 (4) 

Sellow, F. 1167 (4); s.n. (4) 

Smith, L. B. A-20 (7); 15040 (7) 

Sucre, D. 279 (7) 

Ule, E. 2576 (4); 6102 (1) 

Vauthier 230 (4); 231 (4); 233 (8); 570 (4); s.n. (4) 

l'idal, J. I1 5981(953) (4); II 6038(953) (4); IV 1172 (4); s.n. 

(4) 

Warming, E. s.n. (4) 

Weddell, H. A. 1738 (4); s.n. (4) 

Williams, L. O., and V. Assis 6977 (4): 7918 (4); 8150 (4) 
NUMBER 41 

PLATE I.-Lectotype of Psvllocarpus campinorum with an insert showing a verticillaster (C’le 

6101. HBG). 

25 
26 SMITHSONIAN COSTRIBUTIOXS TO BOTANY 

PLATE 2.-Holotype of Psyllocarpus cuiuruensis with an insert showing the distal portion of 

an inflorescence (Anderson et al. 10779, IAN). 
NUMBER 41 

PL~TE J.-Holotype of Ps>llocarpzrs ps~lloca~poides with an insert showing a Lerticillaster with 

a dehisced fruit (Frdes 25262, IAN). 

27 
28 ShlITHSONIAS COSTRIBUTIONS TO BOTANY 

PLATE 4.-Lectotype of Psyilocarpus Zaricoides, farthest left, and holotype of P. laricoides var. 

densifolius, farthest right, with an insert showing the distal portion of an infructescence of 

P. [aricoides with late flowers (Martius s.n., M). 
NUMBER 41 

PLATE 5.--Isotype of Psyllocarpus goiasensis with an insert showing an infructescence (Irwin 

et al. 33082, US). 

29 
30 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

PLATE 6.-Lectotype of Psyllocar$~us asparagoides with an insert showing an infructescence 

(,\fortius s.n., M). 
PIATE 7.-Lectot)pe of Ps?l[ocarpus phvllocephalus with an insert showing an infructescence 

(Glaziou 21504, P). 

31 
32 SVITHSOSIAN CONTRIBUTIONS TO BOTANY 

PLATE 8.-h'eotype of PSI llocarpus schwackei with an insert showing two flowers (Anderson 

et al. 36294, US).