SMITIISONIAN CONTRIBUTIONS TO BOTANY 0 NUMBER 42 

A Study of the Genus Schistocarpha 

(H eli ant he a e : As t e r a c e ae) 

Harold Robinson 

SXIITHSONIAN INSTITUTION PRESS 

City of Washington 

1979 
ABSTRACT 

Robinson, Harold. A Study of the Genus Schistocarpha (Heliantheae: Asteraceae). Smithsonian Contl-ibulions to Botany, number 42, 20 pages, 25 figures, 

1979.-Schistocarpha is placed in the subtribe Galinsoginae near Oteiza and is 

not considered closely related to Nezuolaena of the subtribe Neurolaeninae. 

Sixteen species are recognized, including four newly described; S. chiapensis 

and S. matudae of Xlexico, S. pseudoseleri of Guatemala, and S. wilburii of 

Costa Rica. A progression is noted in the genus with more northern species 

having less than 20 ray flowers per head and more southern groups having more 

than 20. The widely distributed S. etipatorioides, which represents the extreme 

in the series with 40-70 rays, is regarded as South American in origin, The 

South American S. sinforosii is regarded as Central American in origin. 

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

in the Institution’s annual report, Smithsonian Year. SERIES COVER DESIGN: Leaf clearing from the 

Katsura tree Cercidiphyllum japonicum Siebold and Zuccarini. 

Library of Congress Cataloging in Publication Data 

Robinson, Harold Ernest, 1932- 

A study of the genus Schistocarpha (Heliaritheae: Asteraceae) 

(Smithsonian contributions to botany ; no. 12) 

Bibliography: p. 

1. Schistocarpha. 2. Botany-Latin America. I. Title. 11. Series: Smithsonian Institution. 

Smithsonian contributions to botany ; no. 42. 

QKl.SZi47 no. 42 [QK495.C74] 581’ .08s [583’ .55] 79-4217 
Contents 

Introduction ...................................................... Relationships ..................................................... Reliability of Characters ........................................... 

Key to the Species of Schistocarpha ................................ 

Schistocarpha eupatorioides (Fenzl) 0 . Kuntze ............... 

Schistocarpha margariiensis Cuatrecasas ...................... 

3 . Schistocarpha croatii Robinson ............................. 

Schistocarpha paniculata Klatt .............................. Schistocarpha uiilburii, new species .......................... 

Patterns in the Genus ............................................. Schistocarpha Lessing .............................................. 

The eupatorioides group ......................................... 

1 . 

2 . The paniculata group ............................................ 

4 . 

5 . The pedicellata group . . ....................................... 6 . Schistocarpha pedi lata Klatt .............................. The bicolor group .............................................. 

7 . 

8 . 

9 . Schistocarpha matudae, new species ......................... 10 . Schistocarpha seleri Rydberg ............................... The platyphylla group ........................................... 11 . Schistocarpha chiupensis, new species ........................ 

12 . Schistocarpha hondurensis Standley ,!k Williams ............... 13 . Schistocarpha liebmannii Klatt ............................. 14 . Schistocarpha platyphylla Greenman . .................... 

15 . Schistocarpha pseudoseleri, new species ...................... 16 . Schistocarpha sinforosii Cuatrecasas .......................... Excluded Species' .................................................. Literature Cited .................................................. Index .... ..................... 

Schistocarpha bicolor Lessing ............................... Sch istocarpha longiligu la Rydberg ........................... 

....... 

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111 

A Study of the Genus Schistotarpha 

(Heliantheae: Asteraceae) 

Hdrold Robinson 

Introduction 

The Neotropical genus Schistocarpha is a member of the tribe Heliantheae and is notable in the 

tribe, along with the genus Neurolaena, for having 

a pappus of many capillary bristles. The genus is 

widely distributed from Central Mexico southward through Central America into the northern 

Andes from Venezuela and Colombia to Bolivia. 

The present study was derived partly from the 

common occurrence of specimens misidentified as 

“Eupatorium,” and interest was augmented during 

efforts to purify the concept of the tribe Senecioneae 

(Robinson & Brettell, 1973). Additional isolated 

problems have been studied (Robinson 1974, 1975), 

but the lack of a roncerted effort has resulted in 

some errors. The existing partial treatment by 

Rydberg (1927) has proven less satisfactory than 

once believed, and important new information has 

accumulated since 1975. The present effort attempts 

to make necessary corrections and to survey the 

anatomy and relationships of the genus, while 

providing a preliminary treatment of the species. 

The genus came to the attention of taxonomists 

late, considering that one of the species is very 

widely distributed in the Neotropical region. The 

first recognized species was described with the genus 

as Schistocarpha bicolor Lessing (183 l), based on 

Mexican material. The genus was placed with Neurolaena in the subtribe Anthemideae in the extremely broad tribal concept Senecionoideae of 

Harold Robinson, Department of Botany, National Museum 

of Natural History, Smithsonian Institution, Washington 

D. C. 20560. 

Lessing. Lessing mentioned that the pappus of 

Schistocarpha was uniseriate rather than biseriate 

as in “Neurochlaena R. BY.,” a reflection of the 

greater number of series that is found in the later 

genus. Candolle (1836) placed Schistocarpha bicolor 

in the synonymy of Peryinenium discolor, apparently without seeing material. The most widely 

distributed species of the genus did not enter the 

literature until 1850, when it was described as 

Neilreichia eupatorioides by Fenzl. 

In the Bentham and Hooker system (1873), tribes 

of the Compositae were resegregated and many 

tribes of Cassini were restored. The tribe Senecioneae, however, was allowed to retain various 

mostly epaleaceous groups having a capillary pappus, while the tribe Helenieae was establishd for 

many similar genera that lacked a capillary pappus. 

On the basis of the pappus, Schistocarpha and Neuroleana remained in the Senecioneae in the subtribe “Liabeae.” The tribal disposition was followed 

by Hoffmann (1894), where the genera were placed 

in the subtribe Senecioninae. Alost workers since 

Bentham and Hooker have accepted the placement 

of Schistocarpha in the Senecioneae. 

An early exception to the Bentham and Hooker 

placement was by Rydberg (1927), who in a highly 

individual treatment placed Neurolaena and Schist ocarpha together in a separate tribe, Neurolaeneae. 

He separated the two genera as follows: 

Heads radiate, but the ligules in some species very inconspicuous: paleae of the receptacle lacerate, often 3-cleft; 

leaves opposite .................................................. 1. Schistocarpha 

Heads discoid or rarely radiate; paleae of the receptacle 

narrow, 1-nerved; leaves alternate. ................ 2. Neurolaena 

1 
2 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

Rydberg also presented a treatment of the 10 species of Schistocarpha then recognized from Mexico 

and Central America. 

After Rydberg, nothing was done with the genera 

until Robinson and Brettell in 1973 transferred 

Schistocarpha and Neurolaena along with Alepidocline Blake from the Senecioneae to the Heliantheae on the basis of characters of the corolla hairs, 

the paleaceous receptacle, anther structure, and 

nectary shape. At that time, the genera were regarded as possibly closest to the subtribe Galinsoginae of the Heliantheae, but basic differences, 

especially in the achene striation, were noted between Schistocarpha and Neurolaena. More recently, Turner (1976) has placed the genus Alepidocline in the synonymy of Sabazia, which is placed 

in the Galinsoginae. Stuessy (1978), in contrast, has 

placed Schistocarpha and Neurolaena together in 

a separate subtribe Neurolaeninae in the Heliantheae. 

Rela tionships 

Robinson and Brettell (1973) have listed a number of differences between Schistocarpha and Neurolaena which indicate that the two are not each 

other’s closest relatives. Nevertheless, the two genera 

could have both been encompassed in the then 

prevailing broad concept of the subtribe Galinsoginae. The narrower subtribal interpretation by 

Stuessy (1973), however, places the two genera together in the segregate subtribe Neurolaeninae, a 

concept that seems to emphasize historical considerations and pappus structure. Such a use of the 

pappus seems untenable in view of the similar 

structure in Bebbia, Oteiza, Tridax, and some species of Sabazia and Selloa, which are members of 

the Galinsoginae. A more accurate placement of 

Schistocarpha and Neurolaena has required a completely new study of the problem. 

The two subtribes under consideration, Galinsoginae and Neurolaeninae, share the more obvious 

characters of usually imbricated, nonherbaceous 

involucral bracts and prismatic achenes with a 

radially symmetrical pappus. As recognized here, 

the two subtribes share the additional features of 

rays usually with three distinct apical teeth and 

the corollas containing secretory ducts, usually filled 

with yellowish resin. The secretory ducts in the 

style are almost always internal or lateral to the 

veins. The style branches each have two stigmatic 

lines. 

A search for differences has emphasized Neurolaena and Galinsoga as representatives of their 

respective subtribes. Three significant characters 

have been found that correlate closely through the 

whole series of related genera. 

ACHENE TVALL.-The lack of clear striations in 

the carbonized achene wall of Neurolaena was first 

noted by Robinson and Brettell (1973). A similar 

lack has been seen now in Brasilia Barroso, Calea 

L., Staurochlamys Baker, and Tyleropappus 

Greenm. In contrast, Galinsoga Ruiz & Pavon has 

striations in the walls between the veins, a character shared by Alloispernium Willd., Aphanactis 

Wedd., Bebbia Greene, Cuchumatanea Seidenschnur & Beaman, Cymopliora B. L. Robins., 

Jaegeria H.B.K., Oteiza La Llave, Sabazia, Cass., 

Selloa H.B.K., Tetragonotheca L., and Tridax L. 

CARPOPODIUM.-h Neurolaenn and Galinsoga 

and all associated genera, the base of the prismatic 

achene is asymmetrical, apparently reflecting the 

basically conical receptacle to which they are attached. The outer side, consisting of distinctly sclerified cells, extends downward and curves inward. 

The inner side consists of radially arranged cells, 

usually forming a cushion. In Neurolaena and associated genera, the cells of the inner cushion are 

firm in texture. In Galinsoga and the related series, 

the inner cushion consists of thin-walled cells that 

collapse to leave the outer side of the carpopodium 

as an isolated, sclerified plate. 

SECRETORY DucTs.-Ducts can be seen in Neurolaena most clearly in N. intermedia and N. macrophy lla Greenm., both of which have yellowish 

resin present. In these species the ducts are broad 

and single along the inner surface of the veins in 

the throat of the disk corolla. At the sinus, the 

ducts split and diverge with the veins along the 

margins of the corolla lobes. The ducts have the 

same positions in Calea, Staurochlamys, and related genera. In the Galinsoga series, the ducts of 

the disk corolla throat are narrower and usually 

paired, with one along each side of the vein. The 

ducts pass into the lobes usually on the inside of 

the veins and they are often widely separated from 

the margin. In some genera with reduced amounts 

of resin, such as Aphanactis and Cuchumatanea, 

only a single duct may occur per vein, but the duct 
NUMBER 42 3 

is narrow and is lateral to the vein, and ducts from 

adjacent lobes do not normally join at the sinus. 

These three characters are found in a cohsistent 

pattern that can be used as a basis for subtribal 

distinction. Additional correlated characters are 

the nonpapillose inner surfaces of the corolla lobes 

and the trend for elongate lobes in the Neurolaena 

series, versus the slightly to strongly papillose inner 

surfaces and short lobes in the Galinsoga series. 

The Neurolaena series is here recognized as the 

subtribe Neurolaeninae, while the Galinsoga series 

is recognized as the Galinsoginae. 

When compared with the revised subtribal concepts, the genus Schistocarpha shows all the characters of the Galinsoginae and obviously is not 

closely related to Neurolaena. The closest relative 

of Schistocarplza seems to be the genus Oteiza La 

Llave (1832). One of the species of the latter, 0. 

raucophila (J. D. Smith) Fay, recently was unnecessarily redescribed as S. steyermarkii H. Robins. because it had a habit and involucral bract form 

identical to that in Schistocarpha. Oteiza shares 

even the distinctive four secretory ducts of the 

style seen in Schistocarpha. Oteiza has been placed 

in the synonymy of Perymenium by most treatments, but recently it has been reestablished for 

two species by Fay (1975, 1976). The strong resemblance of Oteiza raucophila to Schistocarpha 

might suggest a broadened concept for the latter 

genus, which has priority by one year, but the 

type-species of Oteiza, 0. acuminata La Llave, 

does not share the aspect and would seem an unwise addition. Also, both species of Oteiza do differ 

from Schistocarphu by having nonlacerate paleae, 

having glabrous lobes of the disk corollas, having 

extremely deciduous pappus setae more fragile than 

any in Schistocarpha, and by lacking resin in the 

lower parts of the secretory ducts in the disk corollas and styles. Thus, the species of Oteiza seem 

clearly outside of Schistocarpha and more closely 

related to each other, but differences between the 

two species are greater than any seen within the 

larger genus Schistocarpha. 

Reliability of Characters 

A number of characters appear in species of 

Schistocarpha that vary in reliability. A few are 

worth special note. 

Density of pubescence on leaves and stems has 

been used to distinguish some species but has been 

regarded in this study as too unreliable to be used 

without other supporting evidence. Two species 

previously distinguished only by pubescence are 

reduced to synonymy: S. velutina under S. bicolor 

and S. kellermanii under S. platyphylla. 

Distinctive broadly winged petioles are highly 

developed in S. bicolor but are not present in all 

specimens. The narrow-winged forms scarcely can 

be distinguished from the slightly winged condition present in all petiolate members of the Asteraceae. 

Presence of stipitate glands on the pedicels and 

involucre was used by Rydberg (1927) to distinguish S. paniculata and S. pedicellata from other 

prominently rayed species. Only in S. pedicellata 

do glands appear to be a consistent feature. In 

S. paniculata, the glands are consistent in any one 

plant but plants from the same area that are distinguishable by no other feature lack glands. A 

smaller form of gland that is easily overlooked has 

been seen on some South American specimens of 

S. eupatorioides and on one specimen of S. sinforosii. 

Completely elimbate rays occur in some specimens of S. eupatorioides and S. platyphylla, but 

small limbs are common in both species. The two 

species represent two basically different groups in 

the genus, indicating a parallel development. All 

of the S. platyphylla group except S. sinforosii are 

characterized by limbs less than 6 mm long. Reduced limbs also are characteristic of S. margaritensis, S. croalii, and S. ntatudue, which shows that a 

tendency toward limb reduction exists throughout 

the genus. 

The pappus of all species of Schistocarpha is deciduous, with a slight variation in degree. It is 

most persistent in S. eupatorioides, where it remains on most mature achenes. Some species of 

the S. platyphylla group have a more fragile pappus, with mature achenes often retaining no setae. 

No species of Schistocarpha has a pappus as persistent as that of Neurolaena nor as fragile as that 

of Oteiza. 

The paleae of the receptacle are characteristically lacerate in the genus. In the S. eupatorioides 

and S. paniculata groups and in S. bicolor, a generally trilobed condition is obvious, with a long, 

narrow central lobe often projecting above the 

flowers in the immature heads. Members of the 
4 SiMITHSONIAN CONTRIRUTIONS TO BOTANY 

S. platyphylla-S. liebmannii group and other members of the S. bicolor group, such as S. seleri and 

S. longiligula, have a characteristically unequal 

laceration that usually lacks the well-defined narrow central lobe. 

The disk corollas of Schistocarpha often have a 

secretory duct running up the center of each lobe, 

in addition to those along the veins. The expression of the character is variable, but even when 

a duct is not evident there is sometimes a partial 

row of hairs in the area. The only species that 

seem to characteristically lack the duct are S. eupatorioides and the closely related S. margaritensis. 

Other species in which the duct is weak or not 

observed, such as S. longiligula, S. pedicellata, and 

S. pseudoseleri, are represented by too few specimens for the character to be known reliably. 

The form of the hairs on the corolla proves to 

be one of the two most valuable characters in the 

genus. Slender hairs with elongate cells (Figure 1) 

are characteristic on the disk corolla lobes of S. 

eupatorioides and S. margaritensis. Thicker hairs 

with mostly 3-6 cells (Figures 2, 3) are characteristic of S. croatii, S. paniculata, S. wilburii, and 

S. longiligula. Shorter hairs with mostly 2-3 cells 

(Figures 5, 6) are found in most species of the S. 

bicolor and S. platyphylla groups. The evenly distributed hairs on the throat and lobes of the disk 

corolla are a distinctive feature of S. hondurensis. 

On the ray corollas, the lax pilosity on the tubes 

in S. eupatorioides is distinctive of the species and 

markedly different from the densely hispidulous 

condition in S. croatii, S. paniculata, and S. wilburii. There is considerable variation in pubescence 

density on the ray tubes of some other species, such 

as S. liebmannii and S. seleri. 

The most useful character in Schistocarpha is 

the number of rays in the head. The S. eupatorioides and S. paniculata groups each have over 20 

rays per head, while the remaining species all have 

fewer. Of the remaining species, only S. pedicellata 

comes close to the higher number, with some heads 

having about 18 rays. The basic subdivision is accentuated by the evident multiseriate condition 

associated with the greater number of rays. A further subdivision involves the S. platyphylla group, 

with 8-10 or rarely 12 rays, and the S. pedicellata 

and S. bicolor groups, with 12-18 rays in the head. 

This correlates with the number of involucral 

bracts, the former group having 16-20 and the 

latter groups 20-30. The seeming overlap of number is no obstacle to the delimitation of the groups, 

since numerous heads are available on specimens 

and estimates based on counts of exposed halves 

are adequate. The primary need for caution derives from the deciduous nature of the rays, and 

for this reason younger heads serve best. 

Patterns in the Genus 

On even superficial examination, the basic uniformity of Schistocarpha as defined in this paper 

is most obvious. Previous taxonomic treatments of 

the genus have been unsatisfactory because they 

utilized certain obvious but often unreliable characters. All species have been subject to misidentification. Geographic patterns have not been evident 

previously for the foregoing reasons, but in the 

present study distribution has proven a valuable 

guide to the understanding of species and groups 

of species. This has been aided by the fact that 

most closely related species occur in adjacent areas, 

but they rarely overlap in distribution. The combination of distribution, number of ray and disk 

flowers in the head, number of involucral bracts, 

and form of hairs on the corollas has led to the 

recognition of the following five groups. 

1. The eupatorioides group, including S. eupatorioides and S. margaritensis. The group is Andean, with one species having extended its distribution throughout the entire range of the genus. 

The heads contain 25-30 involucral bracts and 

22-70 rays in 2-3 series. Tubes of the rays are 

laxly pilosulous, and the lobes of the disk corollas 

have slender hairs. 

2. The parziculata group, including S. croatii, 

S. paniculata, and S. wilburii. The group is restricted to Costa Rica and western Panama. The 

heads contain 20-40 involucral bracts and 20-60 

rays in 2-3 series. Tubes of the rays are densely 

hispidulous and the lobes of the disk corolla have 

stout, tapering 3-6-celled hairs. 

3. The pedicellata group, including S. pedicellata. The group is restricted to the eastern escarpment in central Mexico. The plants consistently 

bear numerous glandular hairs. The heads have 

30-40 externally reddish involucral bracts and 

15-18 rays in 1 series. Tubes of the rays are minutely hispidulous, and the lobes of the disk corollas have mostly 1- or 2-celled hairs. 
NUMBER 42 5 

4. The bicolor group, including S. bicolor, S. 

longiligula, S. rnatudae, and S. seleri. The group 

is restricted to the eastern or northern ranges or 

escarpments in hlexico and Guatemala, with the 

exception of S. matudae on hlt. Ovando in Chiapas. 

The heads contain 20-25 involucral bracts and 

12-15 rays in 1 series. Tubes of the rays are sparsely 

to densely hispidulous, and the lobes of the disk 

corollas have short 1-3-celled hairs, except for longer 

3-5-celled hairs in S. longiligula. 

5. The platyphylla group, including S. chiapensis, S. hondurensis, S. liebmannii, S. platyphylla, 

1 

S. pseudoseleri, and S. sinforosii. The group is concentrated in southern Guatemala and adjacent El 

Salvador and Honduras, with S. liebmannii on the 

eastern escarpment in central hlexico and S. sinforosii in the northern Andes in South America. 

The heads contain 16-20 involucral bracts and 8-12 

rays in 1 series. Tubes of the ra)s are glabrous to 

hirtellous or hispidulous and the disk corollas have 

short, 1-3-celled hairs on the lobes. All species except S. sinforosii have limbs of the rays less than 

6 mm long. 

An overall pattern can be seen in distribution. 

5 

B 

d 

0 dbrn Lbd P n 

8 

8 

3 

6 

FICL~RE~ 1-6.-Hairs of disk corolla lobes: 1, Schistocarpha eupatorioides; 2, S. pani~ulata; 3, 

S. longiligula; 4. S. sinforosii; 5, S. pedicellata; 6, S. seleri. 
6 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

FIGURES 7-14.-Heads of Schistocarpha (scale in mm): 7, S. eupatorioides, Asplund 8859; 8, S. 

margaritensis, Pennell 8694; 9, S. croatii, Davidson 337; 10, S. paniculata, Wilbur 14341; 11, 

S. wilburii, Wilbur Q Almeda 16097; 12, S. pedicellata, Pringle 6134; 13, S. bicolor, King 4144; 

14, S. longiligula, Lundell Q Contreras 21164. 
NUMBER 42 7 

FIGURES 15-22.-Heads of Schistocarpha (scale in mm): 15, S. matudae, Matuda 0709; 16, S. 

seleri, Breedlove 9313; 17, S. chiapensis, Matuda 2018; 18, S. hondurensis, Molina 7729; 19, 

S. liebmannii, Purpus 7763; 20, S. platyphylla, Skutch 189; 21, S. pseudoseleri, Williams 26256; 

22, S. sinforosii, Archer 1513. 
8 SMITHSONI.4N CONTRInUTIOSS TO BOTANY 

The pattern initially is obscured by the ranges of 

S. eupatorioides and S. sinforosii, which represent 

opposite extremes in the genus. The first is distributed throughout the range of the genus and 

the latter is found in Colombia and Peru. Both 

distributions are interpreted here as secondary extensions from the evident places of origin of the 

groups to which the species are related. When 

places of origin are considered, the following simple trend can be recognized in the genus. The more 

northern groups of species have a smaller number 

of rays (8-18), and S. sinforosii is seen as a recent 

introduction of these groups into South America. 

Farther south in Costa Rica, the S. paniculata 

group has a greater number of rays (20-60). Finally, 

S. eupatorioides, the most distinctive member of 

the genus with up to 70 rays, would seem to be 

at the extreme of the progression, with origins in 

South America, where it has most of its variation 

and where its only close relative, S. margaritensis, 

occurs in Colombia. The wide occurrence of S. 

ezipatorioides is regarded as a range expansion 

from a South American point of origin. 

Schistocarpha Lessing 

Schistocarpha Less., Linnaea, 13409, 1831. [Type: S. bicolor 

Less.] 

Neilreichia Fenzl, Denkschrift. der Kaiserlich. Akad. der 

Ll’issenschaft., 13258, 1850. [Type: N. eupatorioides Fenzl.] 

Z!co?ln 0. Kuntzc, Rev. Gen. Pl., 1:373, 1891. [Type: 2. oppositifolin 0. Kuntze.] 

Large herbs or subshrubs to 4 m tall, sparsely 

branching. Stems subterete, striated, pith solid; 

internodes 7-20 cm long. Leaves opposite, sometimes alternate above, long-petiolate with wing on 

at least distal half; lamina chartaceous, ovate to 

broadly subdeltoid, base abruptly acuminate and 

decurrent on petiole, trinervate from near base, 

apex short-acuminate, surfaces usually scabrid to 

velutinous. Inflorescence corymbose-paniculate or 

with corymbose branches, pedicels mostly 2-20 mm 

long. Heads broadly campanulate, ca. 6-10 mm 

high; involucre subimbricate, 16-40 unequal bracts 

in 24 series, mostly chartaceous to coriaceous, with 

6-8 striations, mostly greenish to yeilowish brown; 

receptacle short-conical; paleae scarious, rather deciduous, trifid to irregularly laciniate, enclosing 

outer half of achene. Rays 8-70 in 1-3 series, pistillate, usually white, with tube about as long as 

pappus, limb sometimes lacking, when fully developed being papillose above and with 3 small 

distinct teeth at tip; disk flowers 5-75, hermaphroditic, yellow; basal tube slender, glabrous to densely 

pilosulous or hispidulous; throat cylindrical; lobes 

5, triangular, slightly longer than wide, with 

slightly papillose inner surface, outer surface with 

numerous stiff hairs; secretory ducts with yellowish 

resin, paired along veins of tube and throat, extending to base of corolla; anther collar cylindrical, 

lacking more crowded cells in lower part; thecae 

acute at base, usually not reaching level of base 

of collar, exothecium pale to greenish, endothecial 

cells shortly oblo,ig with 1 or 2 nodular thickenings on transverse walls; appendage ovate, concave 

abaxially, with radially oriented marginal cells, 

without glands; style partially immersed in nectary, 

with distinct basal node sclerified in lower half at 

maturity, shaft with 2 veins and with 4 collateral 

secretory ducts, ducts of branches inside of veins, 

resin prominent; branches with 2 stigmatic lines; 

appendage small, short-acute; achenes of ray and 

disk flowers alike, prismatic to terete, shortly obovate, 1-2 mm long, with carbonized, minutely 

punctate walls interrupted by numerous broad 

striations; cells of seed coat irregular with undulating to angular walls, with transverse linear thickenings across inner surface; carpopodium bilateral, 

outer side a sclerified plate, inner side a cushion 

of clustered thin-walled cells; pappus born on distinct callus, radially symmetrical, with 25-35 slender capillary setae 2.5-5.5 mm long, mostly in 1 

series, rather deciduous, tips not or slightl) broadened, apical cells sharply acute. Pollen grains ca. 

27 Fm in diameter, sharply spinose. Chromosome 

number n=8. 

The most useful distinguishing characters of the 

genus are the trifid or laciniate paleae of the receptacle and the pappus of 25-35 slender capillary 

setae. 

In the following treatment of the species, the 

most common species are provided with only general distribution data and citations of only particularly significant collections. 

Key to the Species of Schistocarpha 

1. Heads with more than 20 ray flowers in 2 or 3 series. 

2. Tubes of ray flowers densely hispidulous with short hairs; 1-ays rvhite; lobes of disk 

yellow: lobes of disk corollas with slender, cylindrical hairs 10-13 pm wide. 
NUMBER 42 9 

3. Heads with 25-30 rays, with limbs mostly near 3 mm long: disk flowers 2540; 

pedicels mostly 10-20 mm long (Colombia) ............ S. margaritemis 

3. Heads with 40-70 rays, with limbs usually less than 1 mm long and often totally 

lacking; disk flowers 5-15; pedicels mostly 2-10 mm long (Mexico, Central America, 

2. Tubes of ray flowers densely hispidulous with short hairs: rays white; lobes of disk corollas 

with stout, tapering hairs usually 30-40 pm wide at base. 

4. 

4. Heads with 20-30 rays, with limbs 5-10 mm long. 

5. Involucral bracts not strongly convex at maturity; central awn of paleae slender 

and chartaceous (central Costa Rica) S. paniculata 

5. Involucral bracts strongly convex and coriaceous at maturity, with prominent 

n of paleae broad and indurated (southern Costa Rica) .... .......................................................................................... S. wilburii 

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

South America) .................................................................................................... S. eupatorioides 

Heads with 40-60 rays, with limbs ca. 2 mm long (Panama) ............................ S. croatii 

1. Heads with fe rs in 1 series. 

6. Heads with 12-18 rays, with 20-40 involucral bracts. 

7. Stems, pedicels, and involucral bracts with gland-tipped hairs; heads 7-9 mm high 

with reddish or reddish-tinged outer surfaces of involucral bracts; limb of rays 8-12 

7. Plants without gland-tipped hairs: heads 6-7 mm high; outer surfaces of involucral 

bracts greenish to brownish; limb of rays 1.5-7.0 mm long. 

8. Rays with limb 1.5-2.0 mm long (southern Chiapas) . S. matudae 

8. Rays with limb 4-7 mm long. 

9. Petioles winged in at least upper half, with wing often broad to the base and 

continuous across the node; paleae with slender central awn; outer surface 

of involucral bracts pilose with few to many hairs (Hidalgo, Puebla, San Luis 

9. Petioles winged only in distal third; paleae usually without slender central awn; 

involucral bracts glabrous on outer surface or sparsely puberulous near tips. 

10. Stems hirsute; lobes of disk corollas with mostly 3- to 5-celled hairs (cen10. Stems glabrous to irtellous: lobes of disk corollas with mostly 

long (Veracruz) ............................................................................................................. S pedicellata 

Potosi, Veracruz) ..................................................................................... S. bicolor 

tral Guatemala) .... ............................................................. S. longiligula 

1- or 2-celled hairs (northern Chiapas) 

6. Heads with mostly 8-10 rays, rarely 12, with 16-20 involucral bracts. 

11. Rays with limbs mostly 6-11 mm long (Colombia, Peru) 

11. Rays with limbs less than 6 mm long (Mexico, Central America). 

12. Rays with limbs narrowly vestigial or lacking, less than 2 mm long (southern 

12. Rays with limbs 2-5 mm long, broad in short 

13. Stems sparsely to densely hirsute. 

14. Heads with 8-10 disk flowers; disk corollas with hairs denser on lobes 

than on throat (western Guatemala) 

14. Heads with ca. 20 disk Rowers: disk c 

throat and lobes (Honduras, h'icaragu 

13. Stems glabrous to slightly puberulous. 

15. Heads with ca. 12 disk flowers: leaf blades glabrous above, distinctly 

15. Heads with ca. 25 disk flowers; es pilose above with stiff hairs, 

not distinctly paler below (southern Chiapas) .S. chiapensis 

Guatemala, El Salvador, Chiapas, Oaxaca) ...................... S. platyphylla 

paler below (Veracruz) ......... ........................... S. liebmannii 

The eupatorioides group Akad. der Wissenschaft., 1:258, 1850. [Type: 

Peru. Poepoig 74, holotye IV?, not seen.] 

Neurolaena ' iSchistocarpha) lindeni Sch.-Bip. 

Schistocarpha lindeni J. Donn. Smith, Enum. P1. Guat., 1:24, 

1. Schistocmpha eupatorioides (Fenzl) Kuntze 

FIGURES 1, 7 

proc. Amer. Acad., 5:185, 1861. [Nomen nudum.] 

1881. [Nomen nudum.] 

near Cuchero, 

ex A. Gray, 

Schistocarpha eupatorioides (Fenzl) 0. Kuntze, Rev. Gen. Pl., 

Neilreichia eupatorioides Fenzl, Denkschrift. der Kaiserlich. 

Zycona oppositifolia 0. -Kuntze, Rev. Gen. Pl., 1:373, 1891. 

S (2): 170, 1898. [Type: La Guayra, Venezuela. 0. Kuntze s.n., holotype 

NY.] 
10 ShIITHSONIAN CONTRIBUTIONS TO BOTANY 

Schistocarpha hoflmanii 0. Kuntze, Rev. Gen. PI., 3 (2):170, 

1898. [Type: Rio Juntas, Bolivia. 2600 m. 0. Kuntze s.n., 

holotype and isotype NY.] 

Yehistocarpha oppositzfolza (0. Kuntze) Rydb., North American Flora, 34 (4):306, 1927. 

Stems usually brownish yellow to castaneous, 

densely puberulous to pilose, somewhat glabrescent. 

Petioles 1-5 cm long, winged only in distal half; 

lamina ovate to broadly ovate, mostly 10-20 cm 

long and 6-17 cm wide, margin closely to remotely 

serrate. Inflorescence thrysoid to pyramidally panicdate with densely corymbose branches; pedicels 

mostly 2-10 mm long, puberulous to pilosulous, 

sometimes with intermixed minute stipitate glandular hairs. Heads (Figure 7) ca. 8 mm high; involucral bracts 25-30 in 3-4 series oblong to linearlanceolate, 1.5-6.0 mm long, 1.0-1.5 mm wide, 

apices mostly rounded with minute fringe of hairs, 

outer surface glabrous; paleae trifid with filiform 

central awn, 5-6 mm long, Rays 40-70 in 3-4 

series, yellow to greenish yellow; tube 4-5 mm long, 

usually sparsely pilosulous; limbs 1 mm or less 

long, sometimes lacking. Disk flowers 5-1 1, rarely 

to 18; corollas 5.0-6.0 mm long; tube 2.5-3.0 mm 

long, glabrous or with a few hairs at apex; lobes 

with slender cJlindrica1 hairs on outer surface 

(Figure 1); resin ducts not present in middle of 

lobes; anther thecae 1.3-1.5 mm long. Achenes 

1.3-1.5 mm long; pappus setae ca. 30, 4-5 mm 

long, moderately deciduous, not broadened above. 

The species is common througout the range of 

the genus, extending farther north in Mexico and 

farther south in South America than any other 

species of the genus. Numerous specimens have 

been seen from Mexico, from every country of Central America except El Salvador, and from Colombia, Venezuela, Ecuador, Peru, and Bolivia in 

South America. 

Fenzl originally cited an additional specimen 

from Rio de Janeiro, but the species does not seem 

to occur naturally in Brazil. There is an amazing 

uniformity of structure throughout the range of 

the species. Two slight exceptions involve the 

glabrous tubes of the disk corollas, which seem to 

be most common in Central American specimens, 

and minute glandular hairs, which Qccur in some 

specimens from the Andes. Fenzl reported 11-17 

disk flowers in the head. A brief survey has shown 

usually 5-10, with 11 seen twice and 18 one time. 

I find no basis for higher numbers often cited. 

2. Schistocarpha margaritensis Cuatrecasas 

FIGURE 8 

Schistocarpha margaritensis Cuatr., Rev Acad. Colomb, 

Cienc., 9:244, 1954. [Type: Pichinde, El Valle, Colombia. 

Cuntrecasas 18544, holatype F, not seen, isotype CS.] 

Stems dark greenish brown, stiffly short-hirsute. 

Petioles mostly 3-8 cm long, winged in distal half; 

lamina ovate, 6-18 cm long and 4-11 cm wide, 

margin closely serrulate. Inflorescence corymbosepaniculate; pedicels mostly 10-20 mm long, pilosulous with nonglandular hairs. Heads (Figure 8) 

ca. 8 mm high; involucral bracts ca. 25 in 3-4 

series, oblong to linear-lanceolate, 1.5-6.0 mm long, 

1.0-1.5 mm wide, apices mostly rounded with minute fringe of hairs, outer surface glabrous; paleae 

trifid with filiform central awn, 5-6 mm long. Rays 

ca. 25-30 in 2 or 3 series, yellow; tube 4.5-5.0 mm 

long, densely pilosulous with fine hairs above; 

limbs mostly near 3 mm long. Disk flowers 2540; 

corollas 4.5-5.0 mm long; tube ca. 2 mm long, with 

a few small hairs at apex; lobes with slender cylindrical hairs on outer surface; resin ducts not present in middle of lobes; anther thecae ca. 1.3 mm 

long, Achenes ca. 1.5 mm long; pappus setae ca. 

30-35, ca. 4 mm long, moderately deciduous, not 

or scarcely broadened below tips. 

ADDITIONAL SPECIMEN SEEN.-COLOSIBIA. CALDAS: Rio 

Quindo above .irmenia, 1300-1500 m. Pennell, Sillip & Hazen 

8694 (US). 

The species is closely related to S. eupatorioides 

but is more than just an extreme form of that species. The corymbose inflorescence with longer pedicels differs from the more thyrsoid form with 

densely corymbose branches in S. eupatorioides. 

The smaller number of ray flowers with longer 

limbs and the larger number of disk flowers provide additional significant distinctions. 

The paniculata group 

3. Schistocarpha croatii Robinson 

FIGURE 9 

Schistocarpha croatii H. Robinson, Phytologia, 29:339, 1975. 

[Type: Chiriqui, Panam;(. Cioat 26411, holotype MO.] 

Stems brownish to reddish, pilose with coarse 

hairs. Petioles 1-4 cm long, winged only in distal 
NUMBER 42 11 

third; lamina ovate, 6-10 cm long and 2.5-5.0 

cm wide, margin coarsely serrate. Inflorescence 

corymbose-paniculate pedicels 3-1 0 mm long, 

densely pilosulous. Heads (Figure 9) ca. 8 mm high; 

involucral bracts 20-30 in ca. 3 series, oblong to 

linear-lanceolate, 2-5 cm long, 1 .O-2.3 mm wide, 

apices mostly obtuse with a minutely toothed edge, 

outer bracts sparsely pilose on outer surface; paleae 

trifid with filiform central awn, 4-5 mm long. Rays 

40-60 in 2 or 3 series, white; tube 2.5-3.0 mm long, 

densely hispidulous; limb 2.0-2.5 mm long. Disk 

flowers 30-70; corollas ca. 5 mm long; tube 1.5-2.0 

mm long, densely hispidulous; lobes with stout 

tapering hairs having 3-6 cells; resin ducts not 

seen in middle of lobes; anther thecae ca. 1.3 mm 

long. Achenes ca. 1.5 mm long; pappus setae 30-35, 

mostly 4.04.5 mm long, moderately deciduous, 

slightly broadened below tips. 

The species is apparently endemic to the Chiriqui region of Panama. 

Chorro, Boquete. Davidson 337 (US). 

ADDITIOXAL SPECIXIEN SEE~.-PANAMA. CHIRIQUi: Bajo 

Most of the distinctions of the species from S. 

eupatorioides given with the original description 

prove unreliable. However, the species is thoroughly distinct in the densely hispid tubes of the 

ray and disk corollas, the color and size of the 

rays, the larger number of disk flowers, and the 

coarse hairs on the lobes of the disk corollas. Closest 

relation is actually to S. paniculata of Costa Rica, 

though the latter differs clearly by the smaller 

number of rays with larger limbs. 

4. Schistocarpha paniculata Klatt 

FIGURES 2, 10 

Schistocarpha paniculata Klatt, Bull. SOC. Roy. Bot. Belg., 

31:210, 1892. [Type: Volcan Irazu, Costa Rica. Pittier 866, 

GH, not seen.] 

Stems brownish to reddish, pilose with coarse 

hairs, with or without long-stalked capitate glands. 

Petioles 2-8 cm long, winged only in distal third; 

lamina ovate to subdeltoid, 6-16 cm long and 

3-13 cm wide, margin closely serrate. Inflorescence 

corymbose-paniculate; pedicels 1-4 cm long, densely 

pilose, with or without stipitate glands. Heads 

(Figure 10) ca. 1.0-1.3 mm high; involucral bracts 

35-40 in ca. 4 series, oblong to lanceolate, 3-6 mm 

long, mostly 1.5-2.3 mm wide, apices obtuse to 

acute, often with densely fringed margin, outer 

surface with few to many hairs or stipitate glands; 

paleae trifid, scarious, 5-6 mm long, awn linear 

with filiform tip. Rays 20-25 in ca. 2 series, white; 

tube 5-6 mm long, densely hispidulous; limb 5-10 

mm long. Disk flowers 40-70; corollas 6-7 mm 

long; tube ca. 3 mm long, densely hispidulous; 

lobes with stout tapering hairs having 3-5 cells 

(Figure 2), resin ducts often present in middle of 

lobes; anther thecae ca. 1.7 mm long. Achenes ca. 

1.8 mm long; pappus setae 30-35, mostly 4.5-5.5 

mm long, moderately deciduous, slightly broadened 

below tips. 

The species is restricted to central Costa Rica. 

ADDITIO\AL SPECIMENS SEEN. - COSTA RIGA. CARTACO: 

Southern slopes of Volcan Turrialba, ca. 6 kms generally NE 

of San Rafael de Irazu, ca. 8600 ft. King 6826 (US); southern 

slopes of Volcan Turriabla, ca, 12 hm generally h’E of San 

Rafael de Irazu, ca. 8500 ft. King 6837 (US); region of La 

Esperanza, southern slope of Volcan de Irazd. Standley 35354 

(US); Laguna del Reventado, Volcan Iranzu, 2300 m. Pittier 

14089 (US); between Hacienda Central and Finca Quemado, 

about 2800 m. Wi2bur 14341 (DUKE, US). HEREDIA: Vara 

Blanca de Sarapiqui, north slope of Central Cordillera, 

between Poas and Barba volcanoes, 1680 m. Skutch 3399 (US;) 

El Roble, Stork 2003 (US); Volcan Barba above Hacienda 

Sacramento, about 8700 ft. Wilbur G Almeda 17042 (DUKE, 

US). SAN JosE: Route 216, ca. 2-3 kms S of Cascajal, ca. 5000 

ft. King 6785 (US); Las Nubes, about 1500-1900 m. Standley 

38341, 38759 (US); Bords de rio Parrita a Santa Rosa de 

Copey, 1800 m. Tonduz 12194 (US). 

Stipitate glands are present on the stems, pedicels, and involucral bracts of some plants of S. 

paniculata, and Rydberg (1927) distinguishes the 

species on that basis. However, most specimens 

seen from the area completely lack the glands, 

though they differ in no other character. 

The name S. $anicuZata has suffered from some 

confusing misapplications. One specimen from 

Costa Rica (Pittier 13089) bears an annotation by 

B. L. Robinson of some interest: “This is the Costa 

Rican plant which Klatt described as S. paniculata 

in Bull. SOC. Bot. Belg. xxx pt. 1. Klatt later republished the same description and cited a Pringle 

plant as though it were the type, though it was 

very different.” Rydberg (1927) pointed out that 

one of the syntypes in the original paper, Pittier 

3138 of Costa Rica, was actually S. oppositifolia 

(=S. eupatorioides). The situation is confused still 

further by the fact that one other Klatt syntype, 

Pittier 3411 from southern Costa Rica, is actually 

S. wilburii, described below as a new species. 
12 SXIITHSONIAS COSTRIBUTIONS TO BOTANY 

5. Schistocarpha wilburii, new species 

FIGURE 11 

A S. paniculata cui affinis bracteis involucris 

convexioribus in apice valdius deformibus et paleis 

valdius induratis in apice parum latioribus differt. 

Stems brownish to reddish, pilose with coarse 

hairs. Petioles 2-5 cm long, winged only in distal 

third; lamina ovate to subdeltoid, 6-15 cm long 

and 3-11 cm wide, margin closely serrate. Inflorescence corymbose-paniculate; pedicels 5-27 mm long, 

densely pilose. Heads (Figure 11) ca. 1 cm high; 

involucral bracts 35-40 in ca. 4 series, mostly oblong, 2.5-7.0 mm long, mostly 1.5-2.5 mm wide, 

becoming convex and coriaceous below, usually 

with constriction near tip, tips of inner bracts 

acute and densely fringed, tips of other bracts oblong and blunt, darkened and often recurved, 

outer surface of bracts sparsely pilose; paleae trifid, 

indurated to coriaceous, 5-6 mm long, linear awns 

with lanceolate tips. Rays 20-25 in ca. 2 series, 

white; tube 4.0-4.5 mm long, densely hispidulous; 

limb 5-8 mm long. Disk flowers ca. 60-90; corollas 5.0-6.5 mm long; tube 2-3 mm long, densely 

hispidulous; lobes with stout tapering hairs having 

3-6 cells; resin ducts often present in middle of 

lobes; anther thecae ca. 1.7 mm long. Achenes ca. 

1.8 mm long; pappus setae 30-35, mostly 4-5 mm 

long, moderately deciduous, slightly broadened 

below tips. 

TYPE.-COSTA RICA. SAN JosE: About 14 km 

SE of La Asuncion of the Cerro de la Muerte on 

the Interamerican Highway at an elevation of 

about 8500 ft, shrub 2.5 m tall. Rays white; disk 

yellow. 8 Jan 1972. Wilbur, Almeda G Luteyn 

I6097 (holotype DUKE; isotype US). 

PARATYPES.-COSTA RICA. SAS JosE: Vallee 

du General. clairieres, 600 m. Jan 1891. Pittier 

34II (US); cutover montane cloud forest area, 

Cordillera de Talamanca, about 20 km r\' of San 

Isidro de General along Pan American Highway, 

alt. 2800 m. 29 Jan 1965. Williams4 Molina, Williams Q Gibson 28506 (US). 

The species is apparently restricted to the Valle 

del General area of southern Costa Rica. The species is closely related to S. paniculata, but the involucral bracts of well-developed heads present a 

markedly different appearance. Both the involucral 

bracts and the paleae seem to share the general 

effects of a more indurated texture and broader 

tips. 

The pedicellata group 

6. Schistocarpha pedicellata Klatt 

FIGURES 5, 12 

Schistocarpha pedicellata Klatt, Leopldina, 23:146, 1887. 

[Type: Dos Puentes, Veracruz, Mexico. Liebmann 488, holotype C not seen, photo and fragment. US.] 

Neurolaena pedicellata Schultz-Bip. ex Klatt, Leopoldina, 

23.147, 1887. [Nomen nudum in synon)my.] 

Stems reddish to reddish tinged, densely hirsute, 

with many intermixed gland-tipped hairs. Petioles 

1-5 cm long, winged in distal half; lamina ovate, 

6-15 cm long, 3-10 cm wide, margins closely to remotely serrate. Inflorescence laxly to densely corymbose-paniculate; pedicels 4-35 mm long, densely 

hirsute, with many gland-tipped hairs. Heads (Figure 12) 7-9 mm high; involucral bracts 30-40 in 

2 or 3 series, reddish or reddish tinged, oblonglanceolate to lanceolate, 3-6 mm long, 1.0-1.5 mm 

wide, tips short-acute, tips of inner bracts densely 

fringed with pale hairs, margins and outer surface 

with few to many gland-tipped hairs; paleae usually 

trifid with a slender central awn, 4-5 mm long. 

Rays 15-18 in one series, white?; tube 3 4 mm long, 

minutely hispidulous above; limb 8-12 mm long. 

Disk flowers 40-75; corollas 4.5-6.0 mm long; tube 

1.8-2.5 mm long, minutely hispidulous above; outer 

surface of lobes with numerous sometimes narrow 

1- or 2-celled hairs (Figure 5); without resin ducts 

in middle of lobes; anther thecae 1.5-1.9 mm long. 

Achenes 1.3-1.7 mrn long; pappus setae ca. 35, 

mostly 3.5-4.5 mm long, easily deciduous, slightly 

broadened below the sharp tips. 

The species is restricted to the Orizaba region of 

Mexico. 

ADDITIOA 4~ SPECIMEUS SEE\.-MEXICO. I'ERACRUZ. Eastern slope, Mt. Oriraba, 8000-7500 ft. '\'elson 315 (US); hills 

above Orizdba, 4500 ft. Przngle 6131 (US). 

Schistocal-pha pedicellata is distinctive within the 

Mexican-Guatemalan complexes by the generally 

reddish color and the larger heads. The species is 

the only one in the genus that seems to consistently 
NUMBER 42 13 

bear gland-tipped hairs on the stems, pedicels, and 

involucre. 

The bicolor group 

7. Schistocarpha bicolor Lessing 

FIGURE 13 

Schistocarpha bicolor Less., Linnaea, 6:409, 1831. [Type: 

Mexico. Schiede L+ Deppe, holotype HAL, not seen; isotype 

BU?, not seen.] 

Schistocarpha velutina Rydb., North .American Flora, 34(4): 

304, 1927. [Type: Mirador, Veracruz, Mexico. Liebrnnnn 

490, holotype GH, not seen.] 

Stems greenish yellow to dark brownish, sparsely 

to densely hirsute. Petioles 2-9 cm long, winged in 

at least distal half, often broadly winged to base 

with fused nodal disk; lamina ovate to subdeltoid, 

6-21 cm long and 3-16 cm wide, margins closely 

short-serrate. Inflorescence of numerous corymbose 

panicles on ends of branches; pedicels 2-25 mm 

long, densely pilosulous. Heads (Figure 13) 6-7 mm 

high; involucral bracts 20-25 in ca. 3 series, oblong 

to oblong-lanceolate, 24 mm long, mostly 1.0-1.3 

mm wide, tips rounded to acute, fringed with 

minute hairs, outer surface pilose with few to many 

hairs; paleae trifid with slender central awn, 34 

mm long. Rays 12-15 in one series, white; tube 34 

mm long, hirtellous to minutely pilosulous; limb 

ca. 5 mm long. Disk flowers mostly 35-75; corollas 

4-5 mm long; tube 1.5-2.0 mm long, rather sparsely 

hispidulous above; outer surface of lobes with numerous 2- or 3-celled hairs; with or without resin 

ducts in middle of lobes; anther thecae 1.3-1.5 mm 

long. Achenes 1.1-1.5 mm long; pappus setae ca. 30, 

mostly 3.0-3.5 mm long, somewhat deciduous, 

slightly broadened below the sharp tips. 

The species is restricted to the eastern escarpment 

area of Mexico in the states of Veracruz, Puebla, 

Hidalgo, and San Luis Potosi. 

The species is notable for the unique, broad wing 

that often extends to the base of the petiole and 

onto the node. Unfortunately the wing is not so 

extensive in all specimens, but it always extends at 

least half the length of the petiole. Schistocarpha 

bicolor is the only member of the Mexican-Guatemalan series that characteristically bears a long 

slender awn on the paleae. The awn is most noticeable projecting above the unopened disk flowers in 

the young heads. 

8. Schistocarpha longiligula Rydberg 

FIGURES 3, 4 

Schistocarpha longiligulo Rydb., North American Flora, 34 

(4):305, 1927. [Type: San Xliguel, El Quinchie. Guatemala. 

Heyde & Lux 3383, holotype GH, not seen; isotype US.] 

Stems pale brownish to reddish brown, hirsute. 

Petioles 2-5 cm long, winged only in distal third; 

lamina ovate to subdeltoid, 6-20 cm long, 3-15 cm 

wide, margins closely serrate. Inflorescence broadly 

corymbose-paniculate; pedicels 5-20 mm long, 

densely pilosulous. Heads (Figure 14) 6-7 mm high; 

involucral bracts 20-25 in ca. 3 series, mostly oblong, 1.5-5.0 mm long, 1.0-1.5 mm wide, tips 

rounded to obtuse with fringe of minute hairs, 

outer surface sparsely puberulous distally, usually 

glabrescent; paleae irregularly lacerate, ca. 3.5 mm 

long. Rays ca. 12 in one series, white; tube 3-4 mm 

long, densely hispidulous; limb 4-7 mm long. Disk 

flowers ca. 25-30; corollas 4-5 mm long; tube ca. 

1.5 mm long, sparsely to densely hispidulous; outer 

surface of lobes with sharp, mostly 3- or 4-celled 

hairs (Figure 3); resin ducts usually not present in 

middle of lobes; anther thecae 1.3-1.5 mm long, 

Achenes 1.3-1.6 mm long; pappus setae 3540, 

mostly ca. 3.8 mm long, moderately persistent, 

slightly broadened below the sharp tips. 

The species is restricted to central Guatemala. 

;\DDITIONAL SPECISIEKS SEEN.-GU.ATEJI.ALA. ALTA \'ERArAz: Along the road to San Pedro Cnrcha, ca. 3 kms generally EKE of Coban, ca. 4200 ft. King 7326 (US); Cobin, 

4300 ft. Turckheim 1151 (US), 1350 m. Turckheirn ZI1662 

(US). 1600 m. Turckheim ZI 2131 (US). BAJA \'ERAI'AZ: Jlilio 

Perdido. Lundell Q Contreras 21164 (UTD). 

Rydberg distinguished the species from S. seleri 

by longer rays and more linear involucral bracts. 

More recent collections show the same range of 

variation for the rays of both species. The type of 

S. longiligula does have somewhat more pointed 

involucral bracts, but material from Alta Verapaz 

and Baja Verapaz cannot be distinguished on this 

basis from S. seleri. Pubescence of the stem and the 

longer, more septate hairs of the disk corolla lobes 

serve to distinguish S. longiligula, and also the 

pappus setae seem somewhat more persistent. 
14 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

9. Schistocarpha matudae, new species 

FIGURE 15 

A S. bicolore cui affinis differt limbis radiorum 

brevioribus et paleis receptaculorum irregulariter 

laceratis. 

Stems brownish, coarsely and stiffly hirsute. Petioles 1-5 cm long, wing mostly restricted to distal 

half; lamina ovate, 6-15 cm long, 3.0-9.5 cm wide, 

margins closely and sharply serrate. Inflorescence a 

broad corymbose panicle; pedicels 1-10 mm long, 

densely hirtellous. Heads (Figure 15) 6-7 mm high; 

involucral bracts 20-23 in 2 or 3 series, oblong to 

oblong-lanceolate, 2-5 mm long, 1.0-1.3 mm wide, 

obtuse to narrowly acute, tips of inner bracts with 

dense fringe of hairs, outer surface pilose; paleae 

irregularly lacerate, 2.0-2.5 mm long. Rays 12-15 

in one series, white; tube 2.3-3.0 mm long, densely 

hispidulous; limb 1.5-2.0 mm long. Disk flowers 

3040; corollas ca. 4.0-4.5 mm long; tube ca. 2 mm 

long, densely hispidulous; outer surface of lobes 

with numerous small 1-3-celled hairs; resin ducts 

usually present in middle of lobes; anther thecae 

ca. 1.5 mm long. Achenes 1.5 mm long; pappus 

setae ca. 30, mostly 34 mm long, easily deciduous, 

slightly broadened below the sharp tips. 

TYPE.--MEXICO. CHIAPAS: Mt. Ovando. 20 Dec 

1936. hlatzida 709 (holotype US). 

The pubescence and form of the involucral bracts 

and the number of ray and disk flowers indicate 

that the new species is most closely related to S. bicolor of the escarpment areas of Veracruz and Hidalgo. The geographic disjunction, the reduced 

limbs of the rays, and the lack of a central awn on 

the paleae provide ample basis for distinction. 

10. Schistocarpha seleri Rydberg 

FIGURES 6, 16 

Schistocarpha seleri Rydberg, North American Flora, 34(4): 

305, 1927. [Type: zw. San Martin und Ococingo, Chiapas, 

Mexico. Seler 2191, holotype NY; isotype US.] 

Stems yellowish brown, sparsely hirtellous above, 

glabrous or glabrescent below. Petioles 24 cm long, 

winged only in distal third; lamina ovate, 6-15 cm 

long and 3-8 cm wide, margins closely and sharply 

serrate. Inflorescence broadly densely corymbosepaniculate; pedicels 4-20 mm long, densely pilosulous. Heads (Figure 16) 6-7 mm high; involucral 

bracts 20-24 in 2 or 3 series, oblong, 1.5-4.0 mm 

long, 1.0-1.3 mm wide, tips rounded to obtuse with 

fringe of minute hairs, outer surface glabrous or 

nearly glabrous; paleae irregularly lacerate or sometimes with longer central awn, 3-4 mm long. Rays 

12-15 in one series, white; tube 2.5-3.0 mm long, 

sparsely to densely hispidulous; limb mostly 4-7 

mm long. Disk flowers ca. 25-35; corollas 4-5 mm 

long; tube 1.3-2.0 mm long, sparsely to densely hispidulous; outer surface of lobes with small mostly 

1- or 2-celled hairs (Figure 6); resin ducts present in 

middle of lobes; anther thecae 1.3-1.5 mm long. 

Achenes 1.5 mm long; pappus setae ca. 27-30, 

mostly 34 mm long, easily deciduous, broadened 

below the sharp tips. 

The species is restricted to the northeastern escarpment area in Chiapas. 

ADDITIONAL SPECIMENS SEE& .-MEXICO. CHIAPAS: The 

Sukleh of Hohoch'en in the Paraje of Shishintonil. Municipio 

of Tenejapa, 7600 ft. Breedlove 9313 (US); Paraje Balum 

K'anal, Municipio of Tenejapa, 8400 ft. Ton 806 (US). 

In number of involucral bracts and flowers the 

species seems closest to S. bicolor, which occurs in 

the escarpment area farther west in Veracruz. 

Schistocarpha seleri differs by the very restricted 

wing on the petioles, the more sharply serrate 

leaves, the glabrous outer surfaces and shorter tips 

of the involucral bracts, and the usually irregularly 

lacerate paleae. 

The platyphylla group 

11. Schistocarpha chiapensis, new species 

FIGURE 17 

A S. pseudoseleri cui affinis differt caulibus puberulis floribus radii 10 et floribus disci ca. 25. 

Stems reddish brown, minutely puberulous to 

glabrous. Petioles to over 5 cm long, winged in distal 

half; lamina ovate, 6-21 cm long, 3-14 cm wide, 

margins closely short-serrate. Inflorescence broadly 

corymbose-paniculate; pedicels 4-1 5 mm long, 

rather densely puberulous to pilosulous. Heads 

(Figure 17) 67 mm high; involucral bracts 16-20 in 

ca. 3 series, oblong to oblong-lanceolate, 1.54.5 mm 

long, 1.0-1.5 mm wide, obtuse to acute, inner bracts 

densely fringed with hairs, outer surface sparsely 

minutely puberulous; paleae irregularly lacerate, 

3.0-3.5 mm long. Rays 10 in one series, white; tube 
NUMBER 42 15 

3.0-3.5 mm long, densely hispidulous; limb 4-5 mm 

long. Disk flowers ca. 25; corollas 4.5-5.0 mm long; 

tube ca. 1.8 mm long, densely hispidulous; outer 

surface of lobes with numerous short 2- or 3-celled 

hairs; resin ducts present in middle of lobes; anther 

thecae ca. 1.3 mm long. Achenes 1.5 mm long; 

pappus setae ca. 30, mostly 3-4 mm long, easily 

deciduous, slightly broadened below the sharp tips. 

TYPE.-MEXICO. CHIAPAS: Sta. Rita, Mapastepec. Jan 1938. Matuda 2018 (holotype US). 

The species is closely related to S. pseudoseleri, 

and the two occur in closely adjacent regions on different sides of the Mexican-Guatemalan border. 

Since only one specimen of each species is known, 

the value of some characters remains in doubt. The 

leaves of the type of S. chiupensis are larger than 

those seen in the type of S. pseudoseleri, the pedicels 

are generally longer, and the disk corollas differ by 

having resin ducts in the middle of the lobes, but 

these characters are probably subject to variation. 

Experience with related species such as S. platyphylla, however, would indicate that the differences 

in stem pubescence, number of ray flowers, and especially the number of disk flowers, are not so variable. The available stem segment completely lacks 

any of the coarser hairs found in related species. 

The outer surfaces of the involucral bracts are 

minutely puberulous, while those of both S. platyphylla and S. pseudoseleri are puberulous only near 

the tips or are glabrous. 

12. Schistocarpha hondurensis Standley 

& Williams 

FIGURE 18 

Schistocarpha hondurensis Standley & L. 0. Williams, Ceiba, 

3:65, 1952. [Type: Srtn Juancito, Francisco Morazan, Honduras. Williams L- Molina 13780, holotype EAP, not seen; 

isotype US.] 

Stems brownish, sparsely to densely hirsute. 

Petioles 2-7 cm long, winged only in distal half or 

third; lamina ovate to broadly ovate, 6-24 cm long, 

3-17 cm wide, margin closely serrate. Inflorescence 

broadly corymbose-paniculate; pedicels 2-1 5 mm 

long, densely pilosulous. Heads (Figure 18) ca. 8 

mm high; involucral bracts 1620 in ca. 3 series, 

mostly oblong, 1.54.5 mm long, 1.0-1.8 mm wide, 

tips obtuse to short-acute with dense fringe of 

minute hairs, outer surface sparsely puberulous, 

glabrescent; paleae irregularly lacerate, ca. 4 mm 

long. Rays 8 in one series, white; tube 2-4 mm long, 

densely minutely hispidulous; limb 2-5 mm long. 

Disk flowers ca. 20; corollas ca. 5 mm long; tube ca. 

1.5 mm long, scabrid above with few to many short 

hairs; outer surface of throat and lobes evenly scabrid with numerous short mostly 2-celled hairs; resin 

ducts usually present in middle of lobes; anther 

thecae ca. 1.7 mm long, Achenes 1.5-1.7 mm long; 

pappus setae ca. 30, mostly 2.5-3.5 mm long, easily 

deciduous, slightly broadened below apex and 

tapered to a sharp tip. 

The species occurs in El Salvador, Honduras, and 

Nicaragva. 

ADDITIONAL SPECIMEM SEEN. - EL SALVADOR. CHALATE\ANC.O: East slope of Los Esesmiles. Tucker lcM5 (US), SAN 

VICENTE: I’olcan de San I’icente, 1200-1500 m. Standley 21523 

(US). HONDURAS. COMAYAGUA: 10 kms W of Siguatepeque, 

1200 m. Molina 26023 (US). CORTES: entre Buenos Aires y 

Banaderos, Montana San Idalfonso, 1500 m. Molina 11595 

(US), FRANCISCO MORAZAN: La Vuelta del Caite NE de San 

Juancito, Montana La Tigra, 2300 m. Molina 7729 (US); Montana La Tigra, SW de San Juancito, 2000 m. Molina 8818~ 

(US); western slopes of Pena Blanca, San Juancito Mountains, 

1800-2100 m. Morton 7493 (US): San Juancito, 2000 m. 

Williams Q Molzna 13780 (US). SANTA BARBARA: 10 kms W de 

Lago Yojoa, Cerro Santa Barbara, 1500-2000 m. Clewell k 

Hazlett 3896 (Us). NICARAGUA. MAT4CALPA: Between El 

Triunfo and Fuente Pura, NE above Santa Maria de Ostuma, 

1800 in. Molina 20387 (US). 

The original description compared the species 

with S. seleri, but the characters mentioned proved 

unreliable and the species has been reduced to synonymy (Robinson, 1974). On reexamination, the 

species proves thoroughly distinct by the lower number of rays in the head and by the unique, uniformly 

scabrid surface of the disk corollas. 

13. Schistocarpha liebmannii Klatt 

FIGURE 19 

Schistocarpha Ziebmannii Klatt, Leopoldina, 23: 146, 1887. 

[Type: Mexico. Liebniann 42, holotype C, not seen; photo 

and fragment US.] 

Neurolaena liebnzannii Schultz-Bip. ex Klatt, Leopoldina, 

23:146, 1887. [Nomen nudum in synonymy.] 

Stems dark brownish to slightly reddish, glabrous 

to slightly puberulous. Petioles 14 cm long, distinctly winged only in distal y3-’/3; lamina ovate to 

elliptic, 620 cm long, 3-9 cm wide, glabrous above, 

slightly puberulous on veins below, pale below, mar- 
16 ShIITHSOSIAN COSTRIBUTIOSS TO BOTANY 

gins minutely serrulate to remotely serrate. Inflorescence lax with numerous corymbose panicles; 

pedicels 2-13 mm long, sparsely to densely puberulous. Heads (Figure 19) ca. 7 mm high; involucral 

bracts 16-20 in ca. 3 series, mostly oblong, 1.5-4.0 

mm long, 1.0-1.5 mm wide, tips rounded with fringe 

of minute hairs, outer surface glabrous; paleae irregularly lacerate, 2-3 mm long. Rays 8 in one series, 

white; tube 3.5-4.0 mm long, glabrous; limb ca. 4 

mm long. Disk flowers ca. 12; corollas 4.04.5 mm 

long; tube ca. 1.5 mm long, glabrous or nearly glabrous; lobes with small 2- or 3-celled hairs; weak 

resin ducts sometimes present in middle of lobes; 

anther thecae ca. 1.5 mm long. Achenes ca. 1.5 mm 

long; pappus setae ca. 22-25, mostly 2.5-3.0 mm 

long, easily deciduous, not broadened below tips. 

The species is restricted to the eastern escarpment 

area of Veracruz in Mexico. 

ADDITIONAL SPECIMENS SEEN. - MEXICO. VERACRUZ: along 

the road to Coscomatepec, ca. 7 km SW of Coscomatepec, ca. 

4400 ft. King 6487 (US); Zacuapain. Purpus 7753, 10662 (US). 

The ovate leaf blades and the nearly glabrous surfaces of the stems and leaves make the species the 

most distinctive vegetatively in the genus. Only S. 

pseudoseleri has a smaller number of disk flowers 

in the head. 

14. Schistocarpha platyphylla Greenman 

FIGURE 20 

Schistocarpha platyphylla Greenm., Publ. Field. Columb. 

Sfus., Bot., 2:274, 1907. [Type: Quezaltenango, Guatemala. 

Kellerman 5295, holotype F, not seen; isotype US.] 

Schistocarpha kelleimanii Rydb., North American Flora, 

34(4):306, 1927. [Type: Sacatepequez, Guatemala. Kellerman 7293, holotype NY, isotype US.] 

Stems brownish to slightly reddish, rather sparsely 

hirtellous, usually glabrescent. Petioles 2-9 cm long, 

sometimes winged nearly to base, wings wider above; 

lamina ovate to broadly ovate, 6-25 cm long and 

3-22 cm wide, margins closely serrulate. Inflorescence broadly corymbose-paniculate; pedicels 2-17 

mm long, densely pilosulous. Heads (Figure 20) 6-8 

mm high; involucral bracts 16-20 in ca. 3 series, 

oblong to oblong-lanceolate, 2-6 mm long, 1.0-1.5 

mm wide, tips of inner bracts often acute with 

dense fringe of hairs, outer surface glabrous or 

slightly puberulous near tip; paleae irregularly lacerate, 2.5-3.0 mm long. Rays ca. 10 or rarely 12 in 

one series, while; tube 2.5-3.5 mm long, densely 

hispidulous; limb lacking or vestigial and linear to 

2 mm long. Disk flowers 15-25; corollas 4.5-6.0 mm 

long; tube 1.5-2.5 mm long, densely hispidulous; 

outer surface of throat with sparse short hairs, with 

short mostly 2- or 3-celled hairs denser on lobes; 

with or without resin ducts in middle of lobes; 

anther thecae 1.5-1.7 mm long. Achenes ca. 1.5 

mm long; pappus setae ca. 30, 3-5 mm long, easily 

deciduous, slightly broadened below the sharp tips. 

The species is represented by numerous collections from southern Guatemala and also occurs in 

the following areas: 

ADDITIONAL SPECIMENS EXAMINED. - EL SALVADOR. SANTA 

ANA: Slope of A)eco, N\V flank of Volcan de Santa Ana, ca. 

1710 m. Tucker 1280 (US). SONSONATE: Forest Finca La Cumbre, 1600 m. Molina & MontaZuo 21737 (US). MEXICO. 

CHIAPAS: Volcan Tacana north, 2100 m. Matuda 2951 (US). 

OAXACA: L’icinity of Concordia, Distr. Pachutla, 600 m. 

Makrinius 584 (US); vicinity of Concordia, Montecristo, 1100 

m. Makrinius 661 (US); vicinity of Cafetal Concordia, 400-650 

m. Morton 6 nlakrinius 2552 (US). 

Most specimens from Guatemala, including the 

types of both S. platyphylla and S. kellermanii, completely lack limbs on the rays. The specimens occuring disjunct in Oaxaca are most notable for the 

consistent presence of small, linear, sometimes 

deeply cleft limbs on the rays. The Oaxaca specimens also seem more inclined to have 12 rays in 

the head. The latter specimens do not seem otherwise distinct, and such patterns of disjunction have 

been seen in many other plants such as some species 

of Ageratina (King & Robinson, 1978). One of the 

specimens from El Salvador (MoZina (i. Montaluo 

21629) differs from all other species that were seen 

by the presence of a long, slender, central awn on 

each palea. 

15. Schistocarpha pseudoseleri, new species 

FIGURE 21 

A S. seleri differt bracteis involucris paucioribus 

acutis, floribus radii ca. 8, floribus disci 8-10. 

Stems brownish, sparsely coarsely hirsute, sometimes glabrescent. Petioles 2-3 cm long, winged 

only in distal half; lamina ovate, 6-12 cm long, 3-7 

cm wide, margins closely serrulate. Inflorescence of 

many dense corymbose panicles at ends of branches; 

pedicels 1-9 mm long, densely pilosulous. Heads 

(Figure 21) 6-7 mm high; involucral bracts 15-18 
NUMBER 42 17 

in ca. 3 series, oblong to oblong-lanceolate, 1.5-5.0 

mm long, 1.0-1.4 mm wide, obtuse to acute, tips of 

inner bracts more acute and densely fringed with 

hairs, outer surface glabrous; paleae irregularly 

lacerate, 2.5-3.0 mm long. Rays usually 8 in one 

series, white; tube 3.0-3.5 mm long, densely hispidulous; limb 3.5-5.0 mm long. Disk flowers 8-10; 

corollas ca. 5 mm long; tube 2.0-2.5 mm long, 

densely hispidulous; outer surface of lobes with 

numerous small 1- or 2-celled hairs; resin ducts 

weak or lacking in middle of lobes; anther thecae 

ca. 1.3 mm long. Achenes 1.5 mm long; pappus 

setae ca. 30, mostly ca. 4 mm long, easily deciduous, 

slightly broadened below the sharp tips. 

TYPE.-GUATEMALA. SAN MARCOS: Near 

Aldea Fraternidad, between San Rafael Pie de la 

Cuesta and Palo Gordo, west facing slope of the 

Sierra Madre Mountains, wet mountain forest, 

1800-2400 m. 10-18 Dec 1963. Rays white, herb 4 

meters tall. Williams, Molina ri7. Williams 26256 

(holotype US). 

The species has furnished at least part of the 

basis for reports of S. seleri from Guatemala. Differences are evident in the smaller numbers of involucral bracts, ray flowers, and disk flowers in the 

heads. Such a reduced number of flower parts is 

approached elsewhere in the genus only in S. liebmannii of central hlexico, but the latter differs by 

its distinctive nearly glabrous remotely serrate and 

abaxially paler leaves. The lack or near lack of 

resin ducts in the middle of the disk corolla lobes 

may be of some significance. 

16. Schistocarpha sinforosii Cuatrecasas 

FIGURES 4, 22 

Schistocarpha sinforosii Cuatr., Trab. Mus. Cienc. Nat. 

Madrid, Bat., 29:43, 1935. [Type: Tolima, Colombia. 

Cuatrecasas 2879, holotype MA, not seen; photo US.] 

Liabum eupatorioides Muschler, Bot. Jahrb. 50, Beibl., 3:83, 

1913. [Type: Cajamarca, Peru. Weberbauer 4084, holotype 

B, destroyed; photo and fragment US; not Schistocarpha 

eupatorioides (Fenzl) 0. Kuntze.] 

Stems brownish yellow to slightly reddish, hirsute 

to villous. Petioles 2-9 cm long, winged in distal 

part, wing sometimes extending narrowly to base; 

lamina ovate to broadly ovate, 6-22 cm long, 3-18 

cm wide, margin closely serrate. Inflorescence 

corymbose-paniculate, often widely spreading; pedicels 3-20 mm long, densely villosulous or hirtellous. 

Heads (Figure 22) 7-10 mm high; involucral bracts 

16-20 in 2-3 series, mostly oblong to oblong-lanceolate, 3-7 mm long, 1-2 mm wide, tips often sharply 

acute and densely fringed, outer surface glabrous to 

sparsely pilose; paleae irregularly lacerate, 3-4 mm 

long. Rays usually 8-10 rarely 12 in one series, 

white; tube 4-6 mm long, glabrous to densely 

hirtellous; limb 6-11 mm long. Disk flowers mostly 

20-30; corollas 6-7 mm long; tube 2.0-2.5 mm long, 

with few to many small hairs above; lobes scabrid 

with short tapering 2- or 3-celled hairs (Figure 4); 

resin ducts usually present in middle of lobes; 

anther thecae ca. 1.7-2.3 mm long. Achenes 1.5-2.0 

mm long; pappus setae ca. 30, mostly 4.0-5.5 mm 

long, moderately deciduous, slightly broadened below apex and tapered to sharp tips. 

The species is represented by many collections 

from Colombia and occurs as a disjunct in the 

northern half of Peru, but no collections have been 

seen from intervening Ecuador. 

The species has the reduced numbers of involucral bracts and ray flowers that indicate relation to the S. platyphylla group of Central America 

and hlexico. Schistocarpha sinforosii is most distinct from other species of the group by the longer 

limbs of the ray flowers and the generally larger 

size of the heads. There is some variation in the 

species. Specimens with more pointed inner involucral bracts, glabrous or sparsely pubescent tubes 

of the ray flowers, and hirtellous pedicels are restricted to Colombia, but specimens with less 

pointed bracts, densely pubescent tubes of the rays, 

and villosulous pedicels occur in both Colombia 

and Peru. A single Peruvian collection (Amazonia: 

Chachapoyas. Hutchison ri7. Wright 4923, US) represents an extreme form with the undersurfaces of 

the leaves tending to be reddish and the involucral 

bracts being arranged in 3 or 4 markedly unequal 

series. 

Excluded Species 

Schistocarpha steyermarkiana H. Robinson, Phytologia, 29:248, 1974. [Guatemala. = Oteiza ruacophila (J. Donn. Smith) Fay, Phytologia, 31:16, 

1975.1 

Schistocarpha triangularis Rusby, Bull. New York 

Bot. Gard., 4:392, 1907. [Bolivia. Bang 2477. Isotypes at US. = Senecio alternifolius (Schultz-Bip.) 
18 >.MITHSOSIAN CONTRIBUTIONS TO BOTANY 

Greenm., Ann. Mkouri Bot. Gard., 10:76, 1923. been found and the original description mentions 

Also known as Gynoxys repanda Wedd., Chlor. triangular leaves and multiseriate involucral 

And., 1:77, 1856. The holotype at NY has not bracts that are not present in the isotypes.] 

FIGURE 23.--Map showing Schistocarpha species having more 

than 20 ray flowers in the heads. (Solid line = S. paniculata; 

stipples = S. wilburii; square E S. croatii; solid dots = S. 

margaritensis; arrows = direction of dispersal of S. stripes = S. longiligula; solid dot = S. matudae.) 

eupatorioides from presumed area of origin.) 

FIGURE 24.-Map showing Schistocarpha species having 12- 

18 ray flowers and 20-40 involucral bracts in heads. (Square 

= S. pedicellata; solid line = S. bicolor; stipples = S. seleri; 

FIGURE 25.-Map showing Schistocarpha species having 8-12 

ray flowers and 1G20 involucral bracts in heads. (Star = S. 

Iiebmannii; stipples = S. platyphylla; solid dot = S. chiapensis; square = S. pseudoseleri; solid line = S. hondurensis; 

arrow = presumed source of the South American S. sinforosii.) 
Literature Cited 

Bentham, G., and J. D. Hooker 

1873. Ordo LXXXVIII: Compositae. Genera Plantarurn, 

2: 163-533, 536-537. 

Candolle, A. P. de 

1836. Ordo CII: Compositae. Prodromus Systematis h'aturalis Regni T'egetabilis, 534-685. 

Fay, J. J. 

1975. Sew Comliinations in Perynzenium and Oteiza 

(.kteraceae-Heliantheae). Phytologia, 31:16-17. 

1976. Oteiza. In D. L. Sash and L. 0. TVilliams, Compositae. Flora of Guatetnala, 24(12):256-277. 

Fenzl, E. 

1850. Nova Quaedam Genera et Species Plantarum Vascularium. Denk~chriften der Kaiserlichen Akademie 

der M'issenschafteu, 1:253-264, plate 30. 

Hoffmann, 0. 

1894. Compositae. In A. Engler and K. Prantl, Die 

na turl ich en PPanzeizfarn ilien, 4(5): 87-394. 

King, R. RI., and H. Robinson 

Studies in the Eupatorieae (;\steraceae), CLXVIII: 

.-idditions to the Genus Ageratina. Phytologia, 38: 

1978. 

323-355. 

La Llave, P. de 

1832. 

Lessing, F. 

Sobre Dos Suevos Generos de L'egetales (Gomezia, 

Oteiza). ,Tlesico, Registro Trirnestre, 1:39-41. 

1831. Synanthereae Rich. In D. Schlechtendal and AD. 

Chamisso, Plantarum Mexicanarum a cel. viris 

Schiede et Deppe collectarum, recensio brevis. Linnaea, 6:397-411. 

Robinson, H. 

1974. Studies in the Heliantheae (.\steraceae), 111: A 

Sew Species of Schistocarplia. Phytologia, 29:247- 

250. 

1'375. Studies in the Heliantheae (Asteraceae), IV: A 

Sew Species of Schistocarplza from Panama. Phytologia, 29:339-342. 

Tribal Revisions in the .isteraceae, IV: The Relationships of Sezirolaenn, Srliistocarphn, and Alepidocline. Phytologia, 25:439-445. 

Robinson, H., and R. D. Brettell 

1973. 

Rydberg, P. A. 

1927. Tribe 14: Neurolaeneae. Sorth American Flora, 

34(4): 303-308. 

Stuessy, T. F. 

1977 [1978]. Chapter 23: Heliantheae-Systematic Review. 

In t'. H. Heyood, J. B. Harhorne, and B. L. 

Turner, The Biology and Chemistry of the Cornpositae, pages 621-671. 

Sew Species and Combinations in Snbazia (Heliantheae, Galinsoginae). Wrighiia, 5(8):302-305. 

Turner, B. L. 

1976. 

19 
Ageratina species, 16 

Alepidocline, 2 

Alloispermum, 2 

Anthemideae, 1 

Aphanactis, 2 

Bebbia, 2 

Brasilia, 2 

Calea, 2 

Cuchumatanea, 2 

Cymophora, 2 

“Eupatorium,” 1 

Galinsoga, 2, 3 

Galinsoginae, 2 

Gynoxys repanda, 18 

Jaegeria, 2 

Helenieae, 1 

Heliantheae, 1, 2 

Liabuin eupatorioides, 17 

Neilreichia, 8 

eupatorioides, 1, 8, 9 

Neurochlaena, 1 

h’eurolaena, 1, 2, 3 

lindeni, 9 

intermedia, 2 

liebmannii, 15 

macrophylla, 2 

pedicellata, 12 

Neurolaeneae, 1 

Neurolaeninae, 2 

Liabeae, 1 

Oteiza, 2, 3 

acuminata, 3 

raucophila, 3, 17 

Perymenium, 3 

discolor, 1 

Sabazia, 2 

Schistwarpha, 1, 2, 3, 4, 8 

bicolor, 1, 3, 4, 5, 8, 9, 13, 14 

bicolor group, 4, 5, 13 

chiapensis, 5, 9, 14, 15 

croatii, 3, 4, 9, 10 

eupatorioides, 3, 4, 8, 9, 10, 11 

eupatorioides group, 4, 9 

hoflrnanii, 10 

hondurensis, 4, 5, 9, 15 

kellerinanii, 3, 16 

liebmannii, 4, 5, 9, 15, 17 

lindeni, 9 

longiligula, 4, 5, 9, 13 

margaritensis, 3, 4, 8, 9, 10 

matudae, 3, 5, 9, 14 

oppositifolia, 10, 11 

paniculata, 3, 4, 9,11, 12 

paniculata group, 3, 4, 8,10, 11, 12 

pedicellata, 3, 4, 9, 12 

pedicellata group, 4, 12 

platyphylla, 3, 4, 5, 9, 15, 16 

platyphylla group, 3, 4, 5,14 17 

pseudoseleri, 4, 5, 9, 14, 15, 16 

seleri, 4, 5, 9, 13, 14, 15, 16, 17 

sinforosii, 3, 5, 8, 9, 17 

steyerinarkiana, 17 

triangularis, 17 

uelutina, 3, 13 

wilburii, 4, 9, 11, 12 

Selloa, 2 

Senecio alternifolius, 17 

Senecioneae, 1, 2 

Senecioninae, 1 

Senecionoideae, 1 

Staurochlamys, 2 

Tetragonotheca, 2 

Tridax, 2 

Tyleropappus, 2 

Zycona, 8 

oppositifolia, 8, 9 

20