SMITHSONIAN CONTRIBUTIONS TO BOTANY NUMBER 51 

A Revision of the Tribal 

and Subtribal Limits 

of the Heliantheae (Asteraceae) 

Haro Id Rob inson 

SMITHSONIAN INSTITUTION PRESS 

City of Washington 

1981 
ABSTRACT 

Robinson, Harold. A Revision of the Tribal and Subtribal Limits of the 

Heliantheae (Asteraceae). Smithsonian Contributions to Botany, number 5 1, 102 

pages, 210 figures, 1981.-The tribe Heliantheae is expanded to include the 

genera previously placed in the tribe Heienieae and many genera from the 

Senecioneae. Thirty-five subtribes are recognized and described, and the more 

than 265 genera are listed with known, validly described synonyms. A 

theoretical key to subtribes is provided. Various structural and chemical 

characteristics are reviewed, and emphasis is given to resin duct patterns, 

presence of fiber-sheaths in disk corollas, and patterns of striations in achenes. 

The Heliantheae is considered a member of the subfamily Asteroideae in a 

position parallel to and more advanced than the Eupatorieae. 

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_laponlcum Siebold and Zuccarini. 

Library of Congress Cataloging in Publication Data 

Robinson, Harold Ernest, 1932- 

h revision of the tribal and subtribal limits of the Heliantheae (Asteraceae) 

(Smithsonian contributions to botany ; no. 51) 

Biblioqraphy: p. 

Includes Index-. 

1. Compositae-Classification. 2. Botany-Classification. I. Title. 11. Title. Heliantheae 

(Asteiaceae) 111. Series 

QKl.S2747 no. 51 [QK495.C74] 581s 81-607993 [583’.55] AACR2 
Contents 

Introduction ..................................................... 

Historical Considerations .......................................... 

Review of Taxonomic Characteristics ............................. 

Limits and Relationships of the Tribe ............................. 

Key to the Subtribes of the Heliantheae 

A Synopsis of the Subtribes of the Heliantheae ...................... 

Review of the Subtribes of the Heliantheae 

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

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

Subtribe 1 . AMBROSIINAE ....................................... 

Subtribe 2 . ESPELETIINAE ....................................... 

Subtribe 3 . MELAMPODIINAE ................................... 

Subtribe 4 . POLYMNIINAE ..................... .......... 

Subtribe 5 . MILLERIINAE ................................. 

Subtribe 6 . DESMANTHODIINAE ........................ 

Subtribe 7 . CLIBADIINAE ................................. 

Subtribe 8 . PINILLOSINAE ................................ 

Subtribe 9 . GUARDIOLINAE ............................ 

Subtribe 10 . ENHYDRINAE ......................... 

Subtribe 11 . MONTANOINAE ................... .......... 

Subtribe 12 . RUDBECKIINAE .............................. 

Subtribe 13 . ZALUZANIINAE ............................ 

Subtribe 14 . HEPTANTHINAE ............................... 

Subtribe 15 . ECLIPTINAE ...................................... 

Subtribe 16 . HELIANTHINAE ..................................... 

Subtribe 17 . NEUROLAENINAE ............................ 

Subtribe 18 . GALINSOCINAE ..................................... 

Subtribe 19 . DIMERESIINAE ........................................ 

Subtribe 20 . COREOPSIDINAE ................................... 

Subtribe 2 1 . FITCHIINAE 

Subtribe 22 . COULTERELLINAE .................................... 

Subtribe 23 . PECTIDINAE ...................................... 

Subtribe 24 . FLAVERIIKAE ......................................... 

Subtribe 25 . VARILLINAE .......................................... 

Subtribe 26 . CLAPPIINAE .... ................................. 

Subtribe 28 . MADIINAE ............................................ 

Subtribe 29 . HYMENOPAPPINAE ............................. 

Subtribe 30 . LYCAPSINAE ......................................... 

Subtribe 31 . PERITYLINAE ......................................... 

Subtribe 32 . BAERIINAE ............................................ 

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

Subtribe 27 . JAUMEINAE ........................................... 

... 

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iv SMITHSONIAN CONTRIBUTIONS TO BOTANY 

Subtribe 33 . CHAENACTIDINAE ............................... 

Subtribe 34 . GAILLARDIINAE ................................... 

Subtribe 35 . MARSHALLIINAE ................................... 

84 

87 

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Index 96 

Literature Cited ............................................... 90 

...................................................... 
A Revision of the Tribal 

and Subtribal Limits 

of the Heliantheae (Asteraceae) 

Harold Robinson 

Introduction 

The tribe Heliantheae is generally known for 

such members as the sunflowers (Helianthus), the 

ragweeds (Ambrosia), the cockleburs (Xanthium), 

black-eyed Susans (Rudbeckzu), commonly cultivated members of the Coreopsidinae (Coreopsis, 

Cosmos, and Dahlia), and the marigolds (Tagetes). 

More notable to students of the tribe is the diversity reflected in the many obvious taxonomically 

useful characters. Variations in sexuality of flowers including ray flowers that are completely 

sterile, extremes in palea development including 

ornate or greatly expanded forms, and a great 

variety of achene and pappus shapes have resulted in the acceptance of well over 200 traditional genera in the tribe. The obvious diversity 

and the often primitive aspect of members of the 

tribe have caused some authors to assign to it a 

central and ancestral position in the family, and 

in recent decades the group has been subjected to 

more revisions at the generic level than any other 

comparable group of plants. The increasing efforts in the tribe have also provided great 

amounts of chemical and cytological data for 

various members. Nevertheless, until recently, the 

overall understanding of the tribe has rested on 

a treatment that is over a hundred years old. 

Recent studies have shown a contrasting picture of the understanding of various taxonomic 

levels in the Heliantheae. Because of the wealth 

of structural features, the true diversity of the 

tribe has been obvious, and only comparatively 

minor alterations of generic concepts have been 

necessary. However, the adoption of nearly correct generic limits in the Heliantheae has not 

been matched by equally correct ideas regarding 

the limits of the tribe or the relationships between 

the genera. The present effort is directed toward 

this problem. 

The present study has evolved from an initial 

need to publish various results from identification 

and floristic work. As the full taxonomic implications of some structural features have become 

apparent, it has become desirable to provide a 

general revision of the tribe. The revised subtribes 

will be used in floristic treatments of the tribe in 

Ecuador and Peru now in preparation, and they 

provide a refined basis for evaluating cytological 

data that is being fully treated in another publication. The present revision utilizes new characters discussed in recent studies such as striation of 

achene walls (Robinson and Brettell, 1973b), 

resin duct patterns in the flower (Robinson, 

1979b), and fiber sheaths in the corolla (Robinson, 1978a). The study also incorporates improvements from the recent revision of the tribe by 

Stuessy (1977), while avoiding a number of feaHarold Robinson, Department of Botany, National Museum of Natural 

tfistory, Smithsonian Institution, Washinp.ton, D. C. 20560. 
2 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

tures of that revision believed to be in error. The 

limits of the tribe are redefined to include the 

related genera that have been placed in the Helenieae and Senecioneae, and the relationships of 

the Heliantheae to the other tribes are considered. 

Conclusions regarding the relationships are discussed in the light of the ancestral position in the 

family given to the tribe by Cronquist (1955). 

Completion of the present paper has been delayed to take advantage of the published results 

of the Reading Symposium on the Biology and 

Chemistry of the Compositae (Heywood et al., 

editors, 1977), which were not available until the 

spring of 1978. The parts of the symposium that 

are most important for this study are the systematic reviews of the Heliantheae by Stuessy, of 

the Helenieae by Turner and Powell, of the Tageteae by Strother, and of the Senecioneae by 

Nordenstam. Various chemical reviews, such as 

those of the Heliantheae by Swain and Williams 

and of the sesquiterpene lactones by Herz, also 

are important, but data on polyacetylenes are 

taken primarily from the book by Bohlmann et 

al. (1973). Chromosome data have been evaluated in a separate study (Robinson et al., in 

press), but the conclusions are incorporated into 

the present paper. Throughout the study, a primary source has been the continuing microscopic 

examination of floral parts mounted in Hoyer’s 

solution (King and Robinson, 1970). 

The present paper is designed to achieve three 

primary purposes: to examine the origins of some 

previous concepts of the Heliantheae, to describe 

and evaluate characters of potential taxonomic 

value in the tribe, and to align and describe the 

35 subtribes that are recognized in this study. 

thanked for their contributions to this study. The 

drawings of the achene and head structures have 

been prepared by Alice Tangerini of the Department of Botany. The series is based on an earlier 

unpublished series prepared for me in 1972 by 

Rebecca D. Brettell. The microphotographs were 

taken by Victor E. Krantz, staff photographer, 

National Museum of Natural History. The paper 

has benefitted from the work of various authors 

ACKNOWLEDGMENTS.-Many people are to be 

cited as well as from conversations with a number 

of synantherologists, including Jose Cuatrecasas, 

Vicki Funk, R. M. King, Guy Nesom, David 

Rindos, and John Strother. 

Historical Considerations 

The taxonomic history of the Heliantheae revolves around a limited number of interesting 

and influential treatments produced during the 

last 170 years. In almost every case, features of 

these treatments have been retained in recent 

classifications of the tribe, and often this seems to 

be strictly on the basis of tradition. A review can 

be instructive, to the extent that reasons for past 

concepts, whether correct or not, can be recognized. The tribe’s history is discussed here in the 

light of my present concepts. My judgments of 

some works may seem overly kind or harsh independently of the extent to which I consider them 

to be correct; I admit .being influenced by my 

perception of whether various works represent a 

proper use of the information available at the 

time they were produced. In general, those treatments that I consider most flawed in their conclusions are the ones that I find to be seriously flawed 

in their basic approach. 

The tribe Heliantheae was first named by Cassini in 1819. One of the more useful summaries of 

salient features was provided by Cassini in his 

article on the Helianthees in volume 20 of the 

Dictionnaire (1 82 1 : 3 70). 

La calathide est ordinairement radiee, souvent incouronnee. 

quelquefois discoide. Le clinant he est ordinairement squamellifkre, souvent inappendicule. jamais fimbrillif6re. Les 

squames du pkricline sont ordinairement uniseriees ou biseriees,egales ou peu inegales. souvent imbriqueesiq Les feuilles 

sont ordinairement opposees. souvent alternes. souvent trinervees. Les tiges sont ordinairement herbacees. souvent 

liqneuses. Les fleurs sont ordinairement jaunes. souvent 

blanches. quelquefois purpurines. 

The tribe included the subgroups Prototypes, 

Rudbeckiees, Coreopsidees, Heleniees, Tagetinees, and Millerinees. The only major element of 

the present concept that Cassini omitted was the 

Ambrosiees, with its various specializations of 
NUMBER il 

unisexual heads, fused heads, and free anthers, 

which Cassini placed in a separate tribe. 

The work of Cassini presented for the first time 

the primary characters that have been used in 

constructing most subsequent classifications of 

the Asteraceae, including those of Bentham and 

Hooker (1873) and Hoffmann (1890-1894). The 

use of the characters has led to a partial restoration of the tribal concepts of Cassini in the latter 

classifications. However, one essential quality of 

Cassini‘s concept of the Asteraceae was lost by 

most subsequent workers: his recognition of the 

discrete nature of most of the tribes of the family. 

Bentham and those who followed his system seem 

to consider the tribes as mere nodes in a continuum. 

In a diagrammatic representation of the family, 

Cassini (18 16- 18 17) arranged the tribes in a circle, which, in many details, anticipated the presmtly accepted two-parted concept of the Asteraceae (Robinson and Brettell, 1973d; Wagenitz, 

1976; Carlquist, 1976). 

A final consideration regarding the Cassini 

work concerns the sense in which he used the 

term “affinites.” It is possible that Cassini was 

influenced by his contemporary Lamarck, although he would not have been able to openly 

espouse evolution when publishing in the Diction- 

?zaire edited by Cuvier, who was Lamarck’s most 

outspoken opponent. Certainly. the comparative 

phyletic accuracy of Cassini’s work is in clear 

contrast to the classifications of his immediate 

successors who used totally nonevolutionary approaches to classification. 

One can only speculate about what would have 

happened if subsequent authors had built directly 

upon the work of Cassini. Instead, the year of 

Cassini‘s death saw the publication of the appallingly simplistic and superficial study by Lessing 

(1832), which presented a classification that emphasizes the most obvious characters and ignored 

all subtleties. In one case Lessing described the 

new genus Asternma, without benefit of material, 

on the basis of the erroneous suggestion in the 

original description that Monactis dubia H.B.K. 

might be dioecious. The uninsightful nature of 

3 

the work is probably traceable in no small part to 

the extreme youth of the author (1809-1862). 

Unfortunately, Lessing was followed closely in 

the treatment of the Compositae by de Candolle 

(1836-1838). An important aspect of the Lessing 

and de Candolle treatments was the placement 

of most yellow-flowered radiate members of the 

family in a single tribe, Senecioneae. This was 

responsible for the manner in which Bentham 

later treated the Senecioneae as a residuum in his 

classification. 

The Bentham classification of the Compositae 

was presented in Bentham and Hooker in 1873, 

and a separate explanatory article was produced 

by Bentham in the same year. The system represents a full reinvestigation of the characters of the 

family with restoration of most of the essential 

features of Cassini’s classification. The explanatory article even includes a circular representation 

of the tribes similar to that of Cassini. It has been 

argued that Bentham considered the Heliantheae 

the most primitive tribe in the family (Cronquist, 

1955, 1977a), but the idea that the tribe was 

ancestral to others was Cronquist’s alone. 

In his treatment of the Heliantheae, Bentham 

introduced a degree of artificiality. The Heliantheae were rigorously redelimited to exclude 

groups totally lacking paleae and to exclude all 

groups with a well-developed. nonplumose, capillary pappus. The epaleaceous forms with reduced pappus were placed in a separate tribe, 

Helenieae, which was placed adjacent to the 

Heliantheae. The genera with a capillary pappus 

were placed in the more remote and extremely 

heterogeneous Senecioneae. 

General patterns that are preserved in the present treatment can be detected in the Bentham 

organization of the two tribes Heliantheae and 

Helenieae. The Heliantheae are basically paleaceous, connecting through the partly epaleaceous 

Madiinae to the totally epaleaceous Helenieae. 

Within the paleaceous Heliantheae, the first subtribes are epappose, progressing through more 

completely pappose forms and ending with the 

well-developed radial pappus of the Galinsoginae. 

The paleaceous, mostly epappose groups show 
4 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

one further trend from types with separate strictly 

male and female flowers to those with heterogamous heads. These patterns are believed to represent phyletic trends in the tribe. Two subtribes 

in Bentham’s treatment of the Heliantheae that 

were characterized by reduced numbers of flowers 

in the heads are considered here to have been 

artificial. 

The Bentham system was perpetuated essentially intact by Hoffmann in Die naturlichen Pflantenfamilien (1890-1894). 

Small (1 9 1 7- 19 19) proposed a phylogeny of 

the family which patterned the relationships 

closely after Bentham’s system. Senecio was selected as the ancestral genus because it had the 

most species and the widest distribution. Small 

might have drawn some support from the extreme 

diversity of the Senecioneae as delimited by Bentham, but this was not considered. Small’s concepts have no present support. 

The essential points of the more recent phyletic 

system of Cronquist (1955) continue to have some 

support (Conquist, 1977a; Turner, 1977). The 

system virtually enshrines the work of Bentham 

as perceived by Cronquist in what should be 

called the “heliocentric theory of the Compositae.” The Heliantheae are considered the ancestral stock of the family, giving rise directly to all 

the other tribes but one. The family is seen as a 

near continuum, with the exception of the Lactuceae, which are held unnaturally distinct because of their ligulate heads and latex. Almost all 

phyletic stages in the family are presumed to 

survive, with the extant helianthean genus Wyethia closely approximating the ancestor of the 

family. Highly specialized structures such as the 

pappus are presumed to be in their primitive 

form when they have a shape more like a calyx, 

and involucral bracts are presumed primitive 

when they look more leafy. 

The present paper not only disagrees with various conclusions of Cronquist, but also with some 

of his methods of approach. The 1955 paper tried 

to interrelate the tribes of Bentham without a 

proper foundation study of tribal limits. Cronquist did doubt the naturalness of the Helenieae 

as a separate tribe from the Heliantheae, but one 

of the primary examples he cited was the misplaced asterean genus Rigiopappus. Cronquist relied heavily on secondary sources and on studies 

at the floristic level. This superficial approach 

resulted in every morphological convergence 

being misconstrued as a reflection of close relationship. Circular reasoning was employed to justify the ancestral position of the Heliantheae, that 

tribe being said to have primitive character states, 

but the character states being determined as 

primitive primarily by the fact that they are 

found in the Heliantheae. Finally, Cronquist 

completely overlooked the fact that many members of the Eupatorieae share the same “primitive” character states as the Heliantheae, and 

that the former tribe would be more ancestral 

according to his own criteria. 

Carlquist (1957) published a study of the genus 

Fitchia, which used microcharacters in reference 

to some tribal and subtribal relationships of the 

Heliantheae. Fitchia, with its ligulate heads, was 

shown to be a member of the Heliantheae near 

the Coreopsidinae, with no direct relationship to 

the Lactuceae in which it had been placed by 

Bentham. Carlquist showed that other distinctive 

insular genera were also related to the Coreopsidinae, and he made observations on the basic 

dissimilarity of the two genera Lagascea and Coulterella then placed in the subtribe Lagasceinae. 

In recent years, three significant papers on 

relationships within the Heliantheae have been 

published by Stuessy. The first (Stuessy, 1973) 

initiated the much needed dismemberment of the 

subtribe Melampodiinae. The second (Stuessy, 

1976) reduced the subtribe Lagasceinae to synonymy, placing Lagascea in the Helianthinae and 

suggesting a position for Coulterella near Flaveria. 

In the third paper, for the Symposium on the 

Biology and Chemistry of the Compositae, 

Stuessy (1977) presented a complete survey of the 

Heliantheae, including some elements such as the 

Bahiinae and Gaillardiinae from the Helenieae. 

Schistocarpha and Neurolaena, transferred to the 

Heliantheae from the Senecioneae by Robinson 

and Brettell (1973b), were accepted, but other 
NUMBER 31 5 

genera, including many from the Helenieae and 

two from the traditional Heliantheae, were referred to the Senecioneae. Stuessy was not able to 

take advantage of the information in the symposium contribution of Nordenstam (1977), where 

various helianthean genera were rejected from 

the Senecioneae. The Tageteae were not considered by Stuessy and were treated as a separate 

tribe in the symposium (Strother, 1977). Marshallia was considered by Stuessy as possibly Eupatorieae, but the eupatorian Zsocarpha was included, 

apparently because of its well-developed paleae 

and its resemblance to Spilanthes in habit. 

In the Stuessy survey, the name “Helianthinae” was used correctly for the typical element, 

which had been under the name “Verbesininae” 

in the works of Bentham and Hoffmann; however, there was a confusing reintroduction of the 

name “Verbesininae” for a segregate subtribe 

containing many genera, including Lagascea. The 

name “Ecliptinae,” dating from 183 1, was resurrected for a largely natural group of genera that 

previously had been distributed among four different subtribes. Stuessy was able to present an 

entirely natural concept of the Ambrosiinae as a 

result of his 1973 work. The subtribe Engelmanniinae was created, being properly removed from 

the Melampodiinae, but its ties to the Ecliptinae 

were not recognized. The Coreopsidinae was enlarged to contain a number of related genera, but 

a few other unrelated genera were included, and 

the distinctive obcompressed disk achene was not 

mentioned, although it had been noted in older 

literature. The Gaillardiinae was redelimited with 

great accuracy, but was considered close to the 

Helianthinae, possibly because of the superficial 

resemblance between Gaillardia and Rudbeckia. 

The artificiality of some subtribes such as the 

Milleriinae and Neurolaeninae was greatly increased by Stuessy. Among the genera placed in 

the Neurolaeninae was Bebbia, which had been 

traditionally correctly placed in the Galinsoginae. 

Also, the transfer of Spilanthes from the Verbesininae to the Galinsoginae was inexplicable, being 

counter to all recognized evidence regarding achene structure. 

One of the most notable features of the Stuessy 

survey is the great faith placed in chromosome 

numbers, some relationships between genera 

seeming to be based on this characteristic alone. 

A diagram of hypothetical evolutionary relationships suggests three main lines for the tribe, the 

first with X = 15, 16, and 17, the second with 

X = 12, and the third with X = 8 and 9. Such a 

proposal conflicts with the evidence of extensive 

instability in chromosome number in many subtribes of the Heliantheae. 

The symposium treatment of the Helenieae 

(Turner and Powell, 1977) is most notable for the 

complete dissolution of the tribe. A number of 

subtribes are recognized that combine genera 

from the traditional tribes Helenieae and Senecioneae. Unfortunately, the resulting groups are 

placed mostly in the Senecioneae. 

Helianthean elements, including those rejected 

from the Heliantheae and Helenieae by Stuessy 

(1977) and Turner and Powell (1977), are excluded from the Senecioneae in the symposium 

treatment of the latter tribe by Nordenstam 

(1977). Eighteen characters are cited that indicate 

the helianthean rather than senecionean nature 

of the genus Arnica, including phyllotaxy, nectary 

shape, presence of numerous corolla hairs, achene 

wall with a carbonaceous layer, shape of achene 

hairs, pollen structure, epidermal structure and 

UV patterns of the rays, embryo sac with 2 

antipodal cells, presence of apomixis, serological 

data, chromosome number, presence of helenanolides, and the lack of both furanoeremophilines 

and pyrrolizidine alkaloids. Many of the characters apply equally well to other helianthean genera that are to be rejected from the Senecioneae. 

The chemical evidence regarding the tribal 

position of Arnica has been clarified further by 

Robins (1977), especially in regard to pyrrolizidine alkaloids. The presence of a senecionean 

type of pyrrolizidine alkaloid is noted in Doronicum, a genus often placed close to Arnica. The 

chemistry reinforces conclusions based on structural details, that the two genera are not closely 

related, and Doronicum is properly placed in the 

Senecioneae. 
6 SMITHSOXIAX CONTRIBUTIONS TO BOTANY 

As the present study has progressed, a number 

of individual contributions on the genera Lundelliunthus (1978a), Kingianthus (1978b), Podachaenium 

(1978c), Alloispermum (1978d), and Smallanthus 

(1978e) have been published by the author. The 

last four of these involve the recognition of subtribally distinct elements within traditional genera of the tribe. Three papers (Robinson, 19788, 

1979a, 1980a) have validated 16 subtribes, including 15 established as a direct result of the 

study. Information on the distribution of corolla 

fibers in the Ecliptinae was introduced in the 

paper on Lundellianthus (1978a), and various characters including resin ducts have been discussed 

in relation to the limits of the Galinsoginae 

(Robinson, 1979b). 

A separate series of studies has led to the recognition of a new subtribe, Espeletiinae, including a number of new genera (Cuatrecasas, 1976), 

which are adopted in this treatment. 

Review of Taxonomic Characteristics 

Almost any genetically controlled feature of a 

plant is potentially useful in taxonomy at some 

level. Even so, the value of any one character is 

reliably determined only by comparison with a 

phylogeny obtained from maximum correlations 

between all available characteristics. Still, the 

stability of some features might be predicted on 

the basis of the function that they serve. Many 

microstructures are basic to survival and are modified only at great risk. Many parts involved in 

pollination are factors in species isolation and 

vary in the process of speciation. Various secondary metabolites are defensive in nature, as discussed by Cronquist (1977b), and taxonomic significance can vary greatly. General chemical 

types are often of phyletic significance, but individual compounds usually vary with the tastes or 

tolerances of herbivores or parasites. In most 

cases, the absence of a character is found to be 

less significant than variations in a character that 

is present. 

The following survey includes traditional characters used in the classification of the Heliantheae 

(some of which have been overemphasized) and 

other characters that previously have been ignored. Such features are all of potential use in 

generic delimitation and should be included in 

any analytical generic descriptions. Not all are as 

useful above the generic level. 

HABIT.-Cronquist (1955) favored a herbaceous ancestor for the Asteraceae, but since the 

various studies of wood anatomy of the family by 

Carlquist (1966), there seems to be general agreement by all, including Cronquist (1977a), that 

the ancestor was woody. This ancestral condition 

of the family is presumed to extend to most of the 

individual tribes, particularly the Heliantheae. 

The pattern of relationships within the tribe accepted in the present paper in no way conflicts 

with this view, and partial support is provided by 

the evidence that X = 19 is functionally the 

ancestral number for almost all of the extant 

members of the tribe. 

The annual habit occurs widely in the tribe 

and is common in diverse subtribes such as the 

Galinsoginae, Dimeresiinae, Madiinae, and Baeriinae. The same subtribes are characterized by 

chromosome numbers of X = 10 or less. It is my 

present view that both annual habit and low 

chromosome number represent derived conditions in the tribe, being found primarily in three 

separate evolutionary lines, each of which contain 

numerous closely related subtribes showing the 

more common perennial habit and higher chromosome number. 

hYLLoTAxY.-The spatial relationships of 

leaves, especially as they effect the distribution of 

flowers in an inflorescence, may have functional 

importance in a plant. However, the often utilized simple difference between opposite and alternate leaves on vegetative stems does not seem 

to be under strict constraints, and it seems useful 

to keep in mind the simple physiological basis of 

the character. In the meristem, auxin from each 

growing point suppresses the development of new 

growing points in its immediate vicinity. Patterns of distances between the central growing 

point and those leaf primordia already formed in 

the meristem determine whether one or more new 
NUMBER -1 1 7 

primordia will develop during each plastochron. 

Distances in the meristem have no direct relationship to the ultimate width of the stem, as can be 

seen in the mature stems of many alternate-leaved 

Espeletiinae. Near or at the time of inflorescence 

formation, there is a tendency for the meristem 

to become more conical, and the reduction in 

primordium distances may suppress the oppositeleaved condition, even when that condition is 

present in the basal part of the plant. A rare 

reversal of this is seen in Espeletia, where the 

vegetative parts bear alternate leaves, but the 

bracts of the lateral inflorescences are opposite. 

Whatever the basis, the opposite- versus alternate-leaved condition shows some interesting patterns in the family at species, generic, and tribal 

levels. It is notable that two of the three tribes in 

which opposite leaves predominate are the Heliantheae and Eupatorieae. Still, both of these 

have many alternate-leaved species and genera. 

Opposite-leaved exceptions also occur in basically 

alternate-leaved tribes, including Baccharis spicata 

(Lam.) Baillon and B. cotymbosa (R. & P.) Pers. in 

the Astereae, Luciliopsis in the Inuleae, Chuquiraga oppositifolia Don in the Mutisieae, and various 

species in the Senecioneae, including two species 

of Herodotia and most species of Gynoxys. In the 

Vernonieae, a few species of Vernonia such as V. 

stellata (Spreng.) Blake of Brazil and V. sparrei H. 

Robins. and V. harlingii H. Robins. from Ecuador 

have opposite leaves (Robinson, 1976b, 1979c), 

and both opposite and whorled leaves occur in 

the African Bothriocline Oliv. ex Benth. 

To the extent that it can be seen to work, 

phyllotaxy should be used in identification and 

minor phyletic speculation, but the character 

seems very unreliable at higher levels of classification. Phyllotaxy is certainly not sufficiently 

reliable for speculations at the family level, as 

attempted by Cronquist (1955). 

SECONDARY AND TERTIARY HEADs.-The reduction of individual heads and their enclosure in a 

secondary or even tertiary involucre occurs sporadically in the family. A clear example of secondary heads in the Heliantheae is Lagascea 

(Stuessy, 1978). Here, one can see a true involucre 

of semi-paleaceous aspect around each flower, 

and a secondary involucre is formed around the 

cluster. Elsewhere in the family, Echinops has 

flowering structures variously interpreted as secondary or tertiary heads. Expression of genes 

seems to be transferred from one structure to 

another. Furthermore, the physiological machinery by which plants developmentally differentiate 

their parts seems to fail in such complex inflorescences, producing structures that are morphogenetic hybrids. 

INvoLucRE.-Contrary to the generalization of 

Wagenitz (1976), subimbricate or imbricate, 

scalelike involucral bracts occur widely in the 

family in both subfamilies. It is my belief that 

they are the primitive type in the family. The 

various other types, such as 1 -2-seriate interlocking form in the Senecioneae or the herbaceous 

form of many Heliantheae, are endemic to their 

tribes, and they seem specialized. Cronquist 

(1955) holds the herbaceous involucral bract of 

the Heliantheae as primitive because of its greater 

similarity to the vegetative leaf with which it is 

homologous. This ignores the fact that the involucral bracts are bud-scales. a type of structure 

widely produced by higher plants without recourse to intermediate forms. 

Actually, most hypotheses on the origin of the 

asteraceous head do not take into account adequately the budlike nature of the head and the 

probability that it was a specialization associated 

with seasonal flowering. 

PALEAE.-In the Heliantheae, all variations of 

palea development are represented, including the 

most persistent and most foliaceous types in the 

family. The paleae represent the subfloral bracts 

that are theoretically present in all heads, and 

there is a tendency to interpret the most prominent and most foliaceous types as primitive. The 

Heliantheae with the larger paleae, however, are 

among those with more herbaceous involucral 

bracts, and it seems likely that the two homologous structures are both specialized in their leaflike aspects. 

Traditionally the tribe has been restricted to 

those members having paleae, and the epalea- 
8 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

ceous members have been placed in a separate 

tribe, Helenieae. More recently the palea character has been given less support, and Cronquist 

(1955, 1977a) has recommended reduction of the 

Helenieae to synonymy. A few of the epaleaceous 

elements were placed in the Heliantheae by 

Turner and Powell (1977). Most members of the 

present concept of the Heliantheae are paleaceous, and it is probable that this is the primitive 

condition in the tribe. Isolated examples, such as 

Eclipta with its filiform paleae and the epaleaceous Geissopappus group in Calea, evidently are 

reduced from more paleaceous forms. In contrast, 

the paleae in the Madiinae, especially those on 

the disk as in Blepharipappus, the paleae in Hymenopappus newberryi (A. Gray) Johnston, and the 

tendency toward a single row of paleae in Helenium and Gaillardia seem to represent revived structures in subtribes where expression of paleae is 

usually su’ppressed. It is probable that the paleaceous Varilla is related to the epaleaceous Clappiinae, as indicated by Turner and Powell (1977), 

but I interpret the unusual form of paleae in 

Varilla as another example of a reversion from 

epaleaceous ancestors. Such palea variations 

should not always be interpreted as reduction 

series in the Heliantheae or in any other tribe. In 

the Eupatorieae, paleaceous types occur sporadically, and at least some seem definitely to have 

immediate ancestors that were epaleaceous, indicating that caution is needed in interpreting 

the structure. 

The variations in palea development in the 

tribe should not be overemphasized, however. 

The present analysis of the Heliantheae indicates 

that, with the few exceptions, the epaleaceous 

members of the tribe are more closely related to 

each other than to the paleaceous members. This 

suggests more integrity for the “Helenieae” than 

is allowed by Cronquist (1955, 1977a) or Turner 

and Powell (1977). The present study shows, 

nevertheless, that the epalecaeous subtribes are 

best included in the Heliantheae. 

SEXUALITY OF FLowERs.-The Heliantheae 

show great variety in the distribution of partially 

to completely sterile flowers in the heads. The 

basic pattern of all radiate Asteraceae, the female 

ray and the bisexual disk flower, is found in most 

members of the tribe. There are many scattered 

examples of secondary loss of rays, and in a few 

examples such as Tridax, some species have zygomorphic bisexual peripheral flowers (Powell, 

1965). 

Numerous Heliantheae in the paleaceous series 

possess a trait seen in the family only in the 

advanced members of the subfamily Asteroideae 

and a few genera of the remote tribe Mutisieae: 

the suppression of the gynoecium in some flowers. 

In the extreme, whole heads are functionally male 

while other heads are female. The two types of 

heads occur on the same plants in many Ambrosiinae, while Podanthus of the Ecliptinae is dioecious. Petrobium of the Coreopsidinae has been 

credited with dioecy, although this has been questioned (Carlquist, 1957; Stuessy, 1977). A third 

supposedly dioecious genus, Astemma, definitely is 

not (Robinson, 1976a). 

A common specialization in the tribe is the 

functionally male disk flower. The character has 

been a primary basis for the traditional concepts 

of the Milleriinae and Melampodiinae. In such 

cases, the disk style may or may not be defective 

as in Melampodium, where branches and stigmatic 

surfaces are not formed. Such species may have 

a compensating expansion of the ray styles, as in 

Rensonia and Smallanthus. The reliability of the 

character in the Melampodiinae has recently 

been questioned by various authors (Stuessy, 

1973; Turner, 1978a), and various members of 

the subtribe have been transferred to other subtribes which include genera with bisexual disk 

flowers. The present treatment confirms many of 

these observations and makes similar revisions in 

the Milleriinae. 

A nearly unique development in the Heliantheae is the completely sterile ray flower. Both 

stamens and styles are lacking. This specialization 

occurs in all radiate members of the subtribe 

Helianthinae and also occurs in many other genera, such as Encelia, Oyedaea, Balsamorhita, and 

others in the Ecliptinae, Montanoa in the Montanoinae, and all members of the Rudbeckiinae. The 
NUMBER 51 9 

sterile ray seems restricted to the paleaceous members of the tribe, but seems rather randomly 

distributed within that series, being precluded 

only in those members with functionally male 

disk flowers. 

At present it is thought that numerous parallelisms are involved in the distributions of various 

sexual functions in the heads of the Heliantheae. 

In the various combinations, only the obviously 

nonfunctional have been avoided, such as gynoecial suppression in both ray and disk flowers. A 

particularly notable example of variation in a 

closely related group is the occurence of sterile 

rays, functionally male disk flowers, and fertile 

achenes in both ray and disk flowers among 

species presently placed in the genus Wedelia 

(McVaugh, 1972). 

RAY CoRoLLA.-The tip of the limb of the ray 

is basically three-lobed. In most Heliantheae, 

however, the median of the three lobes is reduced 

or even completely lacking. The third lobe is 

more prominent in a number of subtribes such as 

the Gaillardiinae and Coreopsidinae, and it has 

been useful in this study in redelimiting the Galinsoginae and Milleriinae. 

The cells of the ray surface have been studied 

recently by Baagoe (1977a,b, 1978). The length 

and bulge of the cells is often consistent in related 

groups. The shape of the cells of the upper surface 

is usually directly correlated with that of the cells 

on the inner surface of the disk corolla lobes. 

DISK CORoLLA.-Many genera of the Heliantheae have disk corollas containing fibers along 

the veins of the throat. These may occur in sparing amounts or can form complete sheaths. In 

genera such as Borrichia and Rensonia, the fibersheaths extend into the lobes, and Synedrellopsis 

has supplementary strands of fibers between the 

veins of the throat. All the genera showing such 

fibers share carbonized achenes without striations 

and have black anthers. Some of the genera having fiber-sheaths have obvious relatives that lack 

the character, but all the genera having the character are members of the Ecliptinae, and the 

character apparently is unique to that subtribe. 

The genera with fibers seem to form three groups 

within the subtribe. The primary group has already been recognized in large part by Stuessy 

(1977) on the basis of other characters. It centers 

around Wedelia and includes Delilea and Sjmedrellopsis of the traditional Milleriinae, Rensonia and 

Schizoptera from the Melampodiinae, Synedrella 

and Calyptocarpus of the traditional Coreopsidinae, 

and the genera placed by Stuessy in his subtribe 

Engelmanniinae. 

FILAMENTs.-Pubescence on the filaments occurs sporadically in the Heliantheae. The most 

notable examples are Guardiola of the Guardiolinae, Viguiera dentata (Cav.) Spreng. of the Helianthinae, Rojasianthe of the Ecliptinae, Cosmos of the 

Coreopsidinae, and the uniquely glandularhaired Venegasia of the Chaenactidinae. 

ANTHER THEcAE.-Many genera of the Heliantheae have anthers that blacken at maturity due 

to pigments in the outer layer of the thecial wall. 

This blackening sometimes extends to the anther 

appendage, where it may be useful as a species 

character (Heiser, 1957; Robinson, 1979d). Such 

blackening is rare outside of the Heliantheae, but 

it is found in some genera such as Munnozia in the 

Liabeae and in the appendage of Perezia in the 

Mutisieae. 

In the Heliantheae, the black color of the anthers is sometimes poorly developed in subtribes 

where it normally occurs, but there are many 

significant groups in which it is characteristically 

lacking. Pale anthers are a useful key character 

for distinction of the subtribe Ambrosiinae 

(Stuessy, 1977). The pale anthers found throughout most of the epaleaceous subtribes tend to 

support the distinctness of that series within the 

tribe. 

The seemingly black anthers of some genera 

such as Smallanthus and Icthyothere show a more 

reddish color under the microscope, and anthers 

of the Madiinae are often quite reddish. Anthers 

of the anomalous Marshallia tend to be violet. In 

the Neurolaeninae, the Calea group can be delimited by anther thecae with a distincive yellow 

color. In various genera such as Guardiola and 

Schistocarpha the dark anthers may be greenish. 

ENDOTHECIAL CELLS.-A basic feature of the 
10 SMITHSONIAS CONTRIBUTIONS TO BOTANY 

anthers of most angiosperms is the thickenings in 

the walls of the endothecial cells that function in 

the opening of the valves. In some families such 

as the Melastomataceae and in genera such as 

Solanum where the anthers open by pores, the 

thickenings are lacking. In the Asterales, pollen 

release is facilitated by a plunger action of the 

style, and the essential function of the endothecial 

thickenings apparently is diminished, as evidenced by variations and reductions in the patterns. Dormer (1962) has provided a preliminary 

classification of these patterns: the two most important types are "radial," where the thickenings 

are evenly distributed along both the lateral and 

transverse walls, and "polar," where the thickenings are restricted to opposing walls, either the 

lateral or the transverse. 

The radial pattern of thickenings seems characteristic of fully functional angiosperms, and it 

seems primitive in the asteralean line. Since the 

point of origin of the plunger-type style is not 

known, the earliest possible point for reduction of 

endothecial patterns is not known. It is notable 

that the Eupatorieae have only a few minor 

variations on the radial design, while the Heliantheae and most other tribes have primarily variations of the polar patterns. 

The majority of Heliantheae have endothecial 

thickenings polarized on the transverse walls. Radial patterns occur in a few isolated groups in the 

tribe, some species of Melampodium and Dahlia, the 

genus Lasthenia, the subtribe Ambrosiinae, and 

most members of the subtribe Rudbeckiinae. Another variation in the tribe involves polar thickenings on the lateral walls, as in two species of 

Trtdax and the genera Chamaechaenactzs and Oxypappus, all of which apparently represented isolated occurrences. 

A significant trend in the number of thickenings and the shape of the endothecial cells is seen 

in the subtribes once placed in the Helenieae. 

The Peritylinae and Baeriinae have short cells 

with 2-5 thickenings on the transverse walls, 

which reach an extreme in the radial patterns of 

Lasthenia. Genera of the Chaenactidinae such as 

Palafoxia, Florestina, and Schkuhria have short cells 

with single thickenings. Other genera of the subtribe, such as Arnica and its relatives, and the 

genera of the Gaillardiinae have longer elliptical 

cells with 1 or 2 thickenings at the ends. 

ANTHER APPEmAcE.-In the Heliantheae, the 

anther appendages are usually ovate with a constricted base. Such appendages relate the tribe to 

the Anthemideae-Astereae-Inuleae-Senecioneae 

series, which has appendages usually constricted 

at the base and sometimes constricted throughout. The anther appendages of the Heliantheae 

usually have a radiating pattern of cells that 

might reflect an expansion of a basically narrower 

structure. In contrast, the appendages of the Cichorioideae and the tribe Eupatorieae are basically as broad as the thecae and have no radiating 

cell patterns. 

The presence of hairs or glands on the appendage in the Heliantheae is not regarded here as 

primitive, but as a transfer of a character commonly occurring elsewhere in the plant. Such 

anther pubescence is also seen in the Inuleae, in 

Cyathodzne jacquemontii Gagnep. of the Astereae 

(Fayed, 1979), and in the more remote tribe 

Vernonieae. 

STYLE.-Pubescence on the style base occurs in 

only two tribes of the Asteraceae. In the Eupatorieae, the character is widely distributed, and 

there is even one example of hairs on the nectary 

(King and Robinson, 1974). In the Heliantheae, 

hairs are found on the style base in three genera 

of the Chaenactidinae. The presence of such hairs 

supports the general impression of relationship 

between the two tribes. Other tribes apparently 

have completely lost the potential to produce 

such hairs. The form of the pubescence does not 

directly parallel that elsewhere on the plant, and 

its presence does not seem to represent a simple 

transfer of characters. The potential to produce 

hairs on the base of the style is widespread in the 

angiosperms and is regarded here as primitive 

among the advanced families. 

The presence of two distinct stigmatic lines on 

each style branch seems basic in the subfamily 

Asteroideae, whereas a continuous stigmatic inner 

surface is characteristic of the Cichorioideae. 
NUMBER 51 11 

There are fused stigmatic surfaces in the cacalioid 

Senecioneae and in many Heliantheae, but these 

occur in groups showing no other cichorioid features, and these exceptions in the Asteroideae are 

regarded as specialized. Most Heliantheae have 

tWo stigmatic lines, but fusion seems to have 

occurred separately in a number of subtribes in 

the paleaceous series. Fused surfaces are characteristic of the Helianthinae and Zaluzaniinae, 

and they occur in many Ecliptinae and some 

Coreopsidinae. The Galinsoginae, which resemble the Helianthinae in many ways, are sharply 

distinguished by the presence of two stigmatic 

lines. The lines are often very close, and the 

division difficult to see in some Heliantheae with 

two lines, including some of the Galinsoginae. 

RESIN DUCTS OR CArvALs.-Resin or secretory 

ducts occur widely in the Asteraceae. Ducts in 

vegetative tissues have been extensively studied 

by Col ( 1899- 1904). Anderson (1 970) has studied 

secretory canals in floral parts of members of the 

Astereae, and in various reports has emphasized 

the wide distribution of these structures in various 

tribes, even though they are often difficult to 

observe. Actually, ducts are often very obvious 

because they often contain highly colored resin. 

The numerous reddish ducts in the involucral 

bracts and the paleae of many Coreopsidinae are 

sometimes useful in recognition of the subtribe. 

The first significant detailed study of secretory 

ducts in flowers of the Heliantheae seems to have 

been in certain Coreopsidinae and in the closely 

related Fitchiinae (Carlquist, 1957). Ducts in 

some other genera such as Flourensia, Helianthus, 

and W5ethia were also noted in the study. Colored 

ducts seem to reach their greatest development in 

the Coreopsidinae, where they appear in the anther appendges (Robinson and Brettell, 1973a) 

and in the achenes of some genera. Colored ducts 

are not found in the anther appendages or achenes in the other subtribes of the Heliantheae. 

The color of the resin ducts has proven useful 

in the study of the tribe, and the chemical basis 

of the color should be studied more extensively. 

The distribution of some compounds might be 

determinable by simple visual examination. Different types of chemicals might be involved. A 

preparation shows that resin in the style tips of 

Echinacea has some reddish pigment that might 

be derived from the surrounding tissue. The pigment presumably is an anthocyanin. Nevertheless, the ducts have been noted as the site for 

polyacetylene compounds, and it seems these are 

a primary source of the colors. Sorensen (1977) 

indicates that the ducts in some c]ases are colored 

red because of polyacetylenes containing the unstable dithiin ring. The genera listed as having 

such compounds by Bohlmann et al. (1973) have 

shown little or no reddish resin in this study, but 

my observations have been mostly restricted to 

dead material in Hoyer’s solution. 

On the basis of the survey of material in 

Hoyer’s solution, the majority of the subtribes 

lack colored resin. It is particularly rare in the 

epaleaceous series. The lack of colored resin in 

the Ecliptinae of the paleaceous series is notable 

because of the marked color of the resin in some 

of the subtribes with which it previously has been 

confused. The resin is rather reddish in a number 

of subtribes, including the Coreopsidinae and 

Neurolaeninae. The Helianthinae and Galinsoginae (the two subtribes marked by the presence 

of the polyacetylene dehydrofalcarinone) have a 

yellowish resin. The Desmanthodiinae also have 

yellowish resin, but they have not yet been analyzed chemically. 

The relative positions of the ducts in the corollas and styles are also taxonomically significant. 

In the throats of the disk corollas, the ducts and 

veins show three basic patterns. The first pattern 

(Figure 3) is rarest, where one duct is between 

two veins, as in Vartlla. This pattern occurs to a 

lesser extent in such genera as Clappia and Cltbadzum and is slightly developed in Cnxia. The second pattern (Figure 1) is most common, with one 

duct inside of the vein. This can be seen most 

clearly in the Neurolaeninae, where the duct is 

colored, but is also typical of such subtribes as the 

Ecliptinae. The third pattern (Figure 2) has the 

ducts paired, one duct on each side of each vein 

in the disk corolla throat. Such a pattern is seen 

in most Galinsoginae, in many Helianthinae, and 
12 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

FIGLRES I-10.-Resin duct and fiber patterns in flowers of the Heliantheae: 1-3, corollas with 

veins as zigzag lines and with resin ducts as paired lines (1, Calea eilenii H. Robins., with one 

duct overlying each vein in corolla throat; 2, Schzstocarpha bicolor Less., with ducts paired along 

each vein of throat; 3, Varilla mexicana A. Gray, with single duct between paired veins of throat); 

4-7, styles with relative positions of veins and resin ducts (4, Galinsoga type; 5, Tridax-Lagascea 

type: 6, Helianthus type; 7, Heliomeris type); 8-10, corollas of Ecliptinae with patterns of fibers 

(8. Synedreila nodzylora (L.) Gaertn., with sheaths restricted to veins of throat; 9: Rensonia saluadorica 

Blake, with sheaths extending into lobes; 10, Synedrellopsis grzsebachzi Hieron. & 0. Kuntze, with 

fibers dispersed in corolla throat). (Corollas redrawn to appear equal in size.) 
NUMBER 51 13 

in the Desmanthodiinae. When only one of the 

ducts is evident, that duct is located to one side 

of the vein rather than inside. In both the first 

and second patterns, the ducts continue upward 

along the margins of the corolla lobes. In the 

third pattern, the ducts of the corolla lobe are 

inside the veins and are sometimes remote from 

the margins. 

The resin ducts in the style usually are outside 

of the veins, although this is sometimes difficult 

to see. In groups with colored resin, such as the 

Helianthinae, Galinsoginae, Neurolaeninae, and 

Desmanthodiinae, the reliability of the character 

can be tested. In the Helianthinae, the ducts in 

the shaft of the style are outside of the veins and 

extra ducts are sometimes present. In most Helianthinae, the ducts continue outside the veins 

upward into the branches, as shown by Carlquist 

(1957) in Helianthus. The few exceptions include 

Lagascea and Alvordia, where the ducts of the style 

branches are inside the veins. In Tridax balbisioides 

(H.B.K.) A. Gray, T. brachylepis Hemsl., and T. 

tambensis Hieron., the duct arrangement of Lagascea is equalled, being outside in the shaft and 

inside in the branches, but other Galinsoginae 

and all Desmanthodiinae and Neurolaeninae I 

have seen have the ducts inside the veins throughout the style or collateral with the veins. There 

are further examples of discontinuity of the ducts 

at the base of the style branches, especially in 

Heliomeris of the Helianthinae, which is unique in 

that the resin ducts seem to be restricted to the 

branches. Also, the ducts seen in Spilanthes in the 

Ecliptinae are inside the veins in the style 

branches, rather than outside as is normal in 

Acmella and other genera in that subtribe. 

ACHENE WALr.-The surface of the achene 

provides a number of taxonomic characters. The 

“zwillingshaare” have been reviewed by Hess 

(1938), but they have proven of limited use in the 

present study. The hairs are particularly dense 

and long in many Gaillardiinae and in Dyscritothamnus of the Galinsoginae. Such hairs otherwise 

are most notable for their absence (along with the 

pappus) in sporadically occurring “calvous achenes.” A minor achene wall specialization, cortication, is seen in Borrichia, undoubtedly associated 

with the maritime habitat of that genus. A distinctive regular pattern of short epidermal cells is 

found in some genera such as Polymnia and Unxiu. 

The extensive layer of hard fiber development in 

the genus Montanoa contrasts sharply with the 

extensive layer of soft tissue in the genera Smallanthus and Zchthyothere, but all share a tendency 

for elaboration of the wall that is most common 

in the subtribes with paleaceous receptacles and 

functionally male disk flowers. 

As presently viewed, the most significant feature of the achenes is the presence or absence of 

the carbonization or black layer formed by the 

materials known as phytomelanins. The carbonized layer with the accompanying microscopic 

punctate appearance is found only in the tribes 

Eupatorieae and Heliantheae. Phytomelanins are 

present in all the genera brought to the Heliantheae from the Senecioneae except Psathyrotes, but 

they are absent from all true Senecioneae. The 

tribes that lack phytomelanins and micropunctations have cells in the walls of the achene 

containing prominent raphides. In the Heliantheae two subtribes, the Gaillardiinae (including 

the Psilostrophinae) and the Marshalliinae, lack 

phytomelanins and have raphides. The two subtribes are not closely related, and the lack of 

phytomelanins in the Gaillardiinae is evidently a 

derived condition. The Marshalliinae are near or 

possibly in the tribe Inuleae, being out of the 

main line of helianthean development. If phytomelanins are a primitive feature of the family, 

then their absence in the Gaillardiinae, in the 

advanced Asteroideae including Marshallia, and 

in the subfamily Cichorioideae all represent independent losses. 

In the Heliantheae that have carbonized achene walls, there are often strands or interruptions 

in the carbonized layer that are referred to here 

as striations. These are the internal features that 

can be seen macroscopically as fine longitudinal 

streaks on the common sunflower seeds. The striations are not vascularized and they do not seem 

to be structurally related to the supernumerary 

vascular strands that have been mentioned by 
14 SMITHSONIAN COKTRIBUTIONS TO BOTANY 

Carlquist (1957) and Stebbins (1977). Such striations are found in the Heliantheae, but they do 

not occur in the Eupatorieae. Striations seem to 

be basic to the Heliantheae, and their complete 

absence from some groups such as the Clibadiinae, the Ecliptinae, and the Neurolaeninae is 

regarded as a significant specialization. All forms 

of striations may not be equivalent. Rather broad 

bands of cells interrupting the carbonized layer 

occur in Bidens, but these are structurally different 

from striations in other groups. The Coreopsidinae surveyed have otherwise lacked striations. In 

the genera Melampodium and Smallanthus of the 

Melampodiinae, the carbonized areas are 

strongly convex with the striations recessed. In 

other groups such as the Helianthinae and Galinsoginae, where striations have a high reliability, 

the carbonized layer is flat and in the same plane 

as the striations. In some genera such as Acanthospermum, the achenes seem to lack striations, but 

one or two strands of clear cells have been seen. 

In Montanoa, regular recessed striations are present, but they are difficult to see because of the 

dense layer of fibers. The costate structure of the 

achene wall in the subtribes Clappiinae, Flaveriinae, Jaumeinae, and Varillinae represent outward projections of the carbonized layer that 

have no apparent relation to striations. 

CARPOPODIUM.-The basal callus or carpopodium represents the abscission zone at the base of 

the achene. The variations that often are taxonomically useful also are functional. The asymmetric form of the Galinsoginae leaves an isolated 

plate that seems to be pulled loose from the 

receptacle as it dries. The form in the Neurolaeninae is similar, but differs by the firmer cells on 

the inner surface. Both contrast strongly with the 

bilobed form as seen in the Helianthinae, which 

seems to bite itself loose from the receptacle as it 

matures. 

At least some of the asymmetry of the carpopodium in the Heliantheae may be traced to the 

presence of paleae. In Calea, the epaleaceous species (Geissopappus) have more symmetrical carpopodia than the paleaceous species. As a further 

indication of its unique position, Marshallia has a 

symmetrical annuliform carpopodium in spite of 

the presence of stout paleae on the receptacle. 

PAwus.-The morphology of the pappus varies 

greatly in the family, making it useful in identification. But this same variability creates a need 

for caution in using the pappus as a guide to 

relationships. The pappus is widely accepted as a 

remnant of the calyx, and Cronquist (1955) assumed that the more calyx-like, 5-awned, squamose forms were primitive in the family. This 

idea tended to support the concept of a primitive 

Heliantheae, since most genera with a capillary 

pappus had been systematically excluded from 

the tribe as delimited at that time. This idea 

overlooked the highly specialized nature of the 

pappus and presumed the unlikely multiple derivation of a complex capillary pappus from simpler types. 

Further evidence on the primitive pappus form 

is gained from the redelimitation of the Heliantheae, which shows a well-developed capillary 

pappus in members of both the paleaceous and 

epaleaceous series. There is no reason to regard 

the subtribes that are involved (the Galinsoginae, 

Neurolaeninae, and Chaenactidinae) as more specialized than any others in the tribe. In contrast, 

the Gaillardiinae, with the most calyx-like pappus 

of 3 awns, seems clearly specialized in at least the 

achene wall structure and the chemistry. 

At this time, I regard the pappus as the product 

of a “primordium field” around the periphery of 

the upper callus of the achene. I believe the 

primitive form among extant Asteraceae has numerous primordia in one or more series, arranged 

without direct correlation to other structures of 

the achene. The derived forms are reduced, with 

segments limited to positions over the main veins 

of the achene. The well-developed capillary pappus may contain vascularized bristles, as in Helogyne of the Eupatorieae and some Chaenactidinae 

of the Heliantheae. 

The exact equivalence of some pappus structures in the Heliantheae remains in question. 

Pappus awns of many Heliantheae and relatives 

of the Enceliu- Verbesina group in the Ecliptinae 

look superficially similar, but the former are often 
NUMBER il 

dehiscent at the base with a texture more like a 

true pappus. In the Encelia- Verbesina type, the 

awns are structurally continuous with the body 

of the achene and often are connected to wings of 

the achene. Equivalence is cast in doubt further 

by the form in Perymeniopszs of the Ecliptinae, 

where the two clusters of setae at the outer and 

inner limits of the pappus are attached at a level 

below that of the median squamellae (Robinson, 

1978f). 

The calvous-achened condition occurs sporadically in the Asteraceae in individuals of species 

normally having a pappus. The condition is particularly common in the subtribe Heliantheae 

(Robinson and Brettell, 1972b; Robinson, 1977). 

SEED CoA-r.-The surface of the ovule seems to 

be of particular interest in the Heliantheae, being 

ornamented in many members in a way not seen 

in related tribes such as the Eupatorieae. The 

ornamentation of the ovule surface is much in 

need of a survey, and it must be observed with 

care to insure that the correct layer is obtained in 

a comparable stage of maturity. Nevertheless, on 

the basis of slides prepared primarily for other 

purposes, it has been possible to make the following general observations. 

A basic type of ornamentation in the tribe is 

exemplified by Allozspermum in the Galinsoginae 

(Figure 16), which has sinuous lateral walls and 

distinct annulations on the surfaces. Variations 

include elongate, nearly straight-walled, densely 

annulated cells. as in Acanthospermum in the 

Melampodiinae, and more quadrate, irregularly 

lined cells, as in Verbeszna or Eleutheranthera in the 

Ecliptinae (Figure 17). 

The Coreopsidinae seem to be characterized by 

firmer-walled cells having straight sides and no 

annulations (Figure 18). The character is sufficiently marked to support the separation from 

the subtribe of Guardzola, which has elongate, 

densely annulated cells (Figure 15). 

Among the epaleaceous subtribes of the Heliantheae, there are significant differences in the 

seed coat. The Pectidinae have ornamented cells 

with distinct annulations. A few subtribes such as 

the Flaveriinae have cells with only small thick15 

enings along the lateral walls (Figure 19). Ornamentation apparently is lacking in the main series 

of epaleaceous subtribes, including the Hymenopappinae, the Peritylinae, the Baeriinae, the 

Chaenactidinae, and the Gaillardiinae. 

CYToLocx-The chromosome numbers of the 

Heliantheae are reviewed in detail by Robinson 

et al. (in press). An extreme instability is noted in 

much of the tribe, and most subtribes show evidence of aneuploid decrease from X = 19 or 20. 

The authors differ on whether the higher number 

is basic to the whole tribe excepting the Marshallinae, or whether polyploidy occurred independently in different lines of the tribe. Of the 

authors, Robinson and King favor the first interpretation, while Powell favors derivation of a few 

subtribes such as Galinsoginae and Baeriinae directly from the asteroidean base of X = 10. 

The present view maintains the polyploid condition as a unifying specialized character of all 

the Heliantheae of both the paleaceous and epaleaceous series, with subsequent reduction in some 

cases reaching as far as n = 3. 

CHEMISTRY.-A number of relationships are 

beginning to be proposed on the basis of various 

types of chemicals found in the family. Included 

are sesquiterpene lactones, polyacetylenes, flavanoids, cytochrome c amino acid sequences, and 

the Cq photosynthetic pathway. 

Among the sesquiterpenes, the lactones of the 

germacradiene derivatives have proven most taxonomically useful in studies of the Heliantheae. 

Pathways have been suggested and distributions 

of various forms in the family have been summarized in the papers of Herz (1973, 1977). A 

simplified version of the Herz diagram is presented here to show the sesquiterpene lactone 

types that are significant in systematic considerations of the Heliantheae (Figure 21). The simplest types have an undivided 10-carbon atom 

ring. Derived forms include two major groups. 

First are those in which a secondary cross-link 

divides the larger ring equally into two &carbon 

rings. These include the eudesmanolides, the elemanolides, and eventually in a further step to the 

eremophilanolides. The other group includes the 
16 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

FIGURES 11-2O.-Microphotographs of achene structures: 11-14, achene walls (1 1, Melampodium 

microcephalum Less., wall showing striations and initial phytornelanin deposition, X 1 10; 12, 

Acmella sp., wall lacking striations, X 110; 13, Flaueriu linearis Lag., showing part of rib and extra 

knoblike thickenings in phytornelanin, X 260; 14, Pericome cuudutu A. Gray, showing second 

layer of evenly deposited phytornelanin, X 260); 15-20, seed coats (15, Guardiola mexicanu H. & 

B., X 260: 16, Alloispennum carucasanum (H.B.K.) H. Robins., X 260; 17, Eleutheranthera ruderalis 

(Swartz) Sch.-Bip., X 260; 18, Heterospenna pinnatum Cav., X 260; 19, Flaueria linearis Lag., 

X 260; 20, Baileya multzradiata Harv. & Gray, showing cells and raphides, X 260), 
NUMBER 51 17 

I I I 

I m-qpp 

I 

Eremophilanolides 

I y I 

Eudesmanolides OI 

I 

I 

I 

I 

I 

I 

I 

I 

I 

I 

I 

'(1 ) 

Guaianolides 0 

1 

I 

I Helenanolides 0 

FIGURE 21 .-Common forms of sesquiterpene lactones in the Heliantheae showing pathways 

(Simplified from Herz, 1977.) 

sesquiterepene lactones in which the secondary 

link divides the original ring unevenly into 5- 

carbon and 7-carbon rings. These lactones are the 

guaianolides, and by a further step the ambrosanolides and helenanolides. Most tribes of the family 

possess only germacranolides, eudesmanolides, 

guaianolides, and other comparatively simple 

forms and lack the lactones of the groups in which 

further steps are involved. Apparently the eremophilanolides, ambrosanolides, and helenanolides are produced only in the more advanced 

tribes of the subfamily Asteroideae. 

Until recently, the production of eremophilanolides was thought to be uniquely characteristic 

of the Senecioneae (Robins, 1977). More recently 

two occurrences have been reported in the 

Heliantheae, in Dugesza (Bohlmann and Zdero, 

1976) and Lasthenia (Bohlmann and Zdero, 1978). 

Confirmation should be sought for these records. 

These and possibly other scattered occurrences 

should not be taken to reflect any particularly 

close relationship between the two tribes, beyond 

their both being comparatively advanced members of the subfamily Asteroideae. 

The ambrosanolides and helenanolides are derived from the guaianolides by the formation of 

a quaternary carbon at the (2-5 position. Both 

types were once called pseudoguaianolides, and 
H3C- (CH2) 7-CH=CIb (CH2) 7-C0211 oleic acid 

/ 

J 

H3C- (CH2)4-CEC-CH2-CH=CH-(CH 27 ) -COZH 

cis 

crepenynic acid I 

H3C- (CSC) 3-CI+2-CH=CH- (CH2) 7-C02H 

cis 

CI8-triyne acid 

2 B oxidations 

triyne-8,10,12-en-5-ol-l 

I 

H3C-(CEC)3-CHz-CH=CH-(CH2)3-COZH 

\ 

Ii3C- (CH2) 7-C11=CH- (CH2) 5-CH=CH2 

cis 

I 

+ 

H C- (CZC) -CH=CH-CH=CH- (CH2)4-C

3 trans trans 

Centaur X (al

Ic14/ H3C-(CZC)3-(CH=CH)2-(CH2)2-CH20H triyne-d

H3C - (CEC) -CHZ -CH=CH- (CH2) -OH --= 

Melampodium, Sanvitalia, Tridax

Achyrachaena, and Calycadenia 

/\ 

tetrahydropyranes 

Clibadium, Dahlia and 

Ichthyothere 

# 27 

H3C- (CZC) 3-CII=CH-CH-CH- (CH2) 3-0H 

‘0’ 

E ch in ace a 

\ 

H~C- (CGC) 5-~~=~~2 pentaynene 

Thiophenes 
NLMBER il 19 

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20 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

they differ technically by stereoisomerism at the 

C- 10 position. The basic nature of this seemingly 

slight difference is reinforced by the different 

trends seen in lactone formation in the two groups 

(Herz, 1977), as well as by the fact that they 

occur in two widely separated groups of the 

Heliantheae. The stereoisomerism arises at the 

stage where the secondary cross-link is established, and thus it also occurs in the guaianolide 

precursors, but there is no evidence in the literature of any preferential occurrence of guaianolide 

stereoisomers in various Heliantheae. This might 

indicate that it is the elaborating enzyme systems 

that are selective in the guaianolide stereoisomers 

that they can use as a substrate. 

Acetylenic compounds include the phytomelanins of the achene walls that are visible and 

can be surveyed by optical means. Only the 

presence versus absence or the pattern of deposition of the phytomelanins is considered in this 

study. 

A great variety of acetylenic compounds is 

available through chemical analysis, and the 

structure, distribution, and chemical pathways of 

these have been extensively studied and summarized by Bohlmann et al. (1973). These compounds are well represented in the Heliantheae 

by products from a number of different pathways. 

General pathways of polyacetylene formation 

as interpreted from Bohlmann et al. (1973) have 

been diagramed (Figures 22-23). All polyacetylenes are considered to be ultimately derived from 

fatty acids with 18 carbon atoms, passing through 

the sequence: oleic acid-linoleic acid-crepenynic 

acid, the latter being a polyacetylene. A number 

of pathways diverge from CIS crepenymic acid. 

A major series of polyacetylenes has a methyltriyn end-group on the free (nonacidified) end of 

the molecule. With the exception of the erratically 

distributed by-product, Centaur X, that has been 

reported in Dahlia, all methyl-triyn polyacetylenes in the Heliantheae are derived from C14 

triyn-(8, 10, 12)-en-(5)-oI-(l), which is compound 

27 in chapter 2 of Bohlmann et al. (1973). Four 

important subgroups are derived from 27. The 

C13 pentaynene series is widely distributed in the 

family and is the source of distinctive epoxides, 

epoxisulfones, dithiins, and most polythiophenes 

in the Heliantheae. The dithiins of the series are 

in the Ambrosiinae, Melampodium, Milleria, Verbesina, the Baeriinae, and the Chaenactidinae. The 

second group from compound 27 includes tridecatetrayne-diene in Dahlia and Bidens as well as 

thiophenes and dithiins in Rudbeckia. The third 

subgroup from compound 27 is also shortened to 

(213, and it produced a phenol group on the 

initially acidified end of the molecule. The compounds of this pathway found in the Coreopsidinae include some with an additional thiophene 

unit. The fourth subgroup from 27 retains 14 

carbon atoms and produces the “fish poison” 

tetrahydropyrane found in Ichthyothere and the 

leaves and flowers of Dahlia coccinea Cav. An 

obviously related form of tetrahydropyrane with 

a ene-diyn-ene (propenyl) end-group rather than 

a methyl-triyn (propynyl) end-group has been 

reported from Clibadium (Czerson et al., 1979) and 

the roots of Dahlia coccinea. 

A number of series of polyacetylenes lacking 

the methyl-triyn end-group are separately produced from crepenynic acid. One of the simple 

derivations leads to C17 dehydrofalcarinone, 

which is found in a number of tribes of the family. 

In spite of its seeming simplicity, dehydrofalcarinone has a significant distribution in the Heliantheae, being rather characteristic of the subtribes 

Galinsoginae and Helianthinae. There is one 

anomalous record in Iva of the Ambrosiinae. 

A CIS polyacetylene lacking the methyl-triyn 

end-group is known from Dahlia merckii Lehm., 

which is the only example of that rare type of 

polyacetylene in the tribe. 

A major group of polyacetylenes lacking the 

methyl-triyn end-group in the Heliantheae is derived by reduction through c14 compounds to 

various forms with 13 carbon atoms. Among these 

are the ene-tetrayne-ene group with derived thiophenes and a related group that produces a 

phenyl unit at the free end of the molecule. The 

simpler members of the ene-tetrayne-ene series 

occur in Guizotia of the Milleriinae, Tithonia of the 

Helianthinae, Tridax of the Galinsoginae, and 
NUMBER 51 21 

H 3CCH , -CH2 - (CEC) - CH2 - CH= CH - C ?HI 

H,C=CH-CO- (C!C)2-CH2-CH=CH- (CHZ)6-CH3 

dehydrofalcarinone 

Helianthinae, Galinsoginae, Iva 

HjC- (CH=CH),-C~C-CH2-CH=CH-(CHz)6-OR 

H,C-CH=CH-(C~C)z-(CH=CH)z-(CHz)4-CHzOH 

Lasthenia of the Baeriinae, but they are most 

common in the Ecliptinae and the Coreopsidinae. 

The derived types in the series are concentrated 

in the Coreopsidinae, but there is one with a 

phenol unit in Heliopsis of the Ecliptinae and 

some with a thiophene unit in Lasthenia. 

A derivation of the polyacetylene amides suggested by Bohlmann et al. (1973) is included in 

the diagram (Figure 23). The amide unit originates from a reaction of the acidified end of the 

polyacetylene with an isobutyl group from valine 

peptide. In the Heliantheae, the amides are restricted to members of the subtribe Ecliptinae. 

Most polyacetylenes are shortened by loss of carI 

OH 

C3H,-(CZC)Z-CHZ-CH=CH- (CHZ)3-CH-CH2-C02H 

cis 

reacts iiith isobutyl group of 

valine peptide to fom Anides 

C,H,-(CEC),-CH,-CH=CH- (CHZ)4-OH 

- 

bon atoms from the acidified or lower-numbered 

end of the molecule, but the amides seem unusual 

by the losses from the usually stable higher-numbered end. Lengths are variable, with both CE 

and C11 in Acmella and Echinacea. The nonacetylenic amide in Heliopsis of the same subtribe seems 

to have a related derivation as evidenced by the 

double -CH2- unit between the units with unsaturated bonds. 

Greger (1977) has provided an analysis of the 

polyacetylenes for the Anthemideae, and a comparison of the Anthemideae and Heliantheae is 

instructive. Some major pathways seem to differ. 

The C~O series containing dehydromatricaria ester 

:::: Dahlia 

H3CCH,-CHZ- (CX) Z-CHZ-CH=CH- (CHZ) ?-C02H 

Coreopsidinae i 

HjC-CH=CH- (CgC) 2- -CH,-CH=CH- (CHZ) 3-COZR 

H3C-CH=CH- (CEC) 3 -CH=CH-CH=CH2 

CEC -CSC- CH=CH-CH3 

l-phenyl-hepta-1,j-diyn-S-ene 

Coreopsidinae 

jc12/ H3C-(CZC)Z-(CH2)2- (CH=CH),-CONH-CH2-CH-(CH3)Z 

CIz and Cll derivatives in Acmella, 

Coreopsidinae I- J- 

[c131 H3CCH=CH- (CZC),-CH=CH, I trideca-1,ll-diene- 3,s -7,g-tetrayne 

H3CH-CH=CH-CZC 

Bidens, Lasthenta 

FIGURE 23.-Polyacetylenes of the Heliantheae lacking methyltriyne endgroups showing 

suggested pathways. (Based on Bolhmann et al., 1973.) 
SMITHSONIAN CONTRIBUTIONS TO BOTANY 

and the series derived from C13 triyn-(7, 9, 11)- 

en-(4) acid occur in the Anthemideae, but not in 

the Heliantheae. Also in the Anthemideae but 

not in the Heliantheae are numerous furans and 

ringenolethers. Present in the Heliantheae but 

lacking in the Anthemideae are the pentaynene 

series, the tridecatetrayne series, and the ene-tetrayn-ene series. All the latter involve higher 

numbers of triple bonds, those capable of giving 

rise to polythiophenes. As a result, polythiophenes 

are not found in the Anthemideae. 

Furans and ringenolethers are derived in many 

polyacetylene pathways in the Anthemideae, and 

polythiophenes are produced in three interrelated 

pathways in the Heliantheae, examples in which 

the basic pathways are not as important taxonomically as the enzyme sytems that control the 

final elaborations on the molecules. In another 

example, the aromatic polyacetylenes of the 

Heliantheae involve two different basic pathways 

and involve different ends of the molecule, but 

both are almost completely restricted to the Coreopsidinae, where the same enzyme system probably is involved in formation of the phenol unit. 

Three other types of chemical data have been 

mentioned in studies of the Heliantheae. The 

occurrence of aurone and chalcone types of flavanoids has been noted in the Coreopsidinae and 

in Lasthenza of the Baeriinae, and a relationship 

between the two groups has been suggested 

(Bohm, 1977). The two groups also share thiophenes of the ene-tetrayn-ene type. The flavanoid 

anthocyanins are well known for their importance 

as flower pigments. and recently the yellow flavonols have been noted for absorption of UV 

wavelengths in the nectar guide patterns of plants 

such as Rudbeckia and Gaillardia. The similarity in 

UV patterns might have been partial reason for 

the close relationship given the Helianthinae and 

Gaillardiinae by Stuessy (1977). These subtribal 

relationships based on flavanoids are not accepted 

in the present study, and flavanoids do not seem 

particularly useful in studies above the genus 

level. 

Cytochrome c amino acid sequences have been 

studied in two species in the family, Helzanthus 

annuus L. and Guzzotza ab2ssinica Cass., both members of the Heliantheae. The difference of 8 amino 

acids between the two was a partial basis for 

raising the Coreopsidinae to tribal rank by 

Turner and Powell (1977). The conclusion was 

based on the traditional placement of Guizotia in 

the Coreopsidinae, a position that has proven 

erroneous. 

The amino acid sequences of the more variable 

but more easily obtained plastocyanins have been 

analysed for many members of the family, including three genera of the Heliantheae. Guizotia, 

Helzanthus, and Rudbeckia (Boulter et al., 1978). 

Some heterogeneity is seen in each of the helianthean genera, but there is nothing unique to 

members of the tribe. The helianthean genera do 

fall within a general trend shown by members of 

the subfamily Asteroideae. 

The Cq photosynthetic pathway (Kranz syndrome) has been reviewed for the family by Smith 

and Turner (1975), and it has been shown to 

occur in the three helianthean subtribes Coreopsidinae, Flaveriinae, and Pectidinae. The implications of this and other chemical characters 

reported for the Coreopsidinae are considered 

further in the discussion of that subtribe. 

Asteraceae produce inflorescences that obviously 

are able to attract insect pollinators or in some 

cases birds (Leppik, 1977). and some types of 

specializations are known that allow pollinators 

to distinguish individual species (Baagoe, 1977b). 

Still. extreme specializations toward single-pollinator relationships are not known, and most Asteraceae, including most Heliantheae, remain 

very facultative in their pollination mechanisms. 

As a result, studies of specific pollinators furnish 

little insight into the phylogeny of the Asteraceae. 

More significant correlations can be expected 

with the various types of insects that use the 

Asteraceae as hosts. No major surveys have been 

attempted, but many associations have been recorded in various studies and in host indexes. 

One example is the dipteran genus C'rophora, 

whose larvae apparently live almost exclusively 

in achenes of Asteraceae (Steyskal, 1979). Hosts 

IhsECT CORRELATIONS.-MOSt members Of the 
NUMBER iI 

are known for less than half of the species, and 

the full range of hosts probably is not known for 

any. Most of the European species are associated 

with members of the tribes Cardueae and Anthemideae, which are common in that area. Of 

the eight American species having host data, two 

have been recorded from Heliantheae, one from 

Borrichza and one from Smallanthus. 

Another example of the Heliantheae as hosts 

involves the Coleoptera. Two genera of the family 

Bruchidae with an unusual elongate shape, Dahlibruchus and Cosmobruchus, have been described 

from Dahlia and Cosmos respectively (Bridwell, 

1931). Both the host genera are in the subtribe 

Coreopsidinae. Other genera of Bruchidae, for 

which hosts are known, including one from the 

lactucean genus Lapsana, lack the elongate shape. 

Bridwell has suggested, "The form in these interesting species is with little doubt due to the shape 

of the achenes in which they pass their immature 

stage." Bridwell goes on to correlate the more 

flattened shape of Dahlzbruchus with the more 

flattened shape of the Dahlia achenes. It would 

be instructive to discover a species of Bruchidae 

that infests the genus Heterosperma of the same 

subtribe, which has great range of achene shapes. 

Limits and Relationships of the Tribe 

The Heliantheae, as redelimited, consists of 

approximately 3000 species in about 260 genera. 

The tribe is concentrated in North and South 

America, but has a number of pantropical elements and some widely distributed adventives. 

Characteristics that are considered basic or 

primitive in the tribe include the leaves being 

opposite, with alternate-leaved forms derived. 

The leaf blades have palmate as well as pinnate 

venation. The involucre is weakly to strongly 

subimbricate, with the bracts in many of the 

advanced subtribes being herbaceous or dimorphic. Receptacles are basically paleaceous, 

and epaleaceous forms are derived. Both ray and 

disk flowers usually are present. There is a tendency for suppression of gynoecia, which is expressed in sterile rays, functionally male disk 

23 

flowers, or rarely dioecism. Flower pigments are 

mostly yellow. The disk corollas usually have 

short lobes. The anther thecae are often dark or 

reddish, and the bases rarely extend below the 

lower level of the collar. The endothecial cells 

usually have polarized thickenings on the transverse walls, and radial and laterally polarized 

forms are derived. The anther appendages usually 

are ovate and strongly keeled with constricted 

bases, and the abaxial surface may bear glands 

or hairs. The style base has hairs in three genera, 

but it is usually glabrous. The style branches 

usually have paired stigmatic lines. The achenes 

are basically prismatic, with numerous compressed or obcompressed forms derived. The achene walls usually are carbonized and striate, with 

derived nonstriate or rarely noncarbonized forms. 

In most of the paleaceous subtribes having herbaceous involucral bracts, the achenes seem to 

have a distinctive slow maturation, with fully 

mature achenes only in aged heads. The pappus 

is sometimes of the primitive capillary form with 

many bristles, or rarely plumose, but more often 

it is reduced or lacking. The pollen is mostly over 

23 pm in diameter with an echinate surface pattern and a helianthean wall structure (Skvarla et 

al., 1977). 

The tribe is placed in the subfamily Asteroideae on the basis of the short disk corolla lobes, 

the shape of the anther thecae and appendages, 

the basically paired stigmatic lines, the presence 

of suppressed gynoecia, the presence of a carbonized achene wall, and the details of pollen structure. Within the subfamily, the Heliantheae are 

regarded here as parallel to the Eupatorieae, but 

as more specialized than that tribe on the basis of 

the heterogamic heads, the presence of suppressed 

gynoecia, the basically polarized thickenings of 

the endothecial cells, the constricted bases of the 

anther appendages, the striate wall of the achenes, the basically polyploid chromosome numbers, and the more complex polyacetylene and 

sesquiterpene lactone chemistry. The tribe is 

placed closest to the Inuleae, which seems to differ 

primarily by the tails on the anthers, the usually 

flat anther appendage, and the total lack of car- 
24 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

SENECIONEAE ASTEREAE ANTHEMIDEAE 

o enlarged pollen ............. \ ..... 1 { ostyle branch/{ 3 reduced pappus 

pronate ”* a modified pollen wall 

Blennospermatinae 

o style appendages mostly short or truncate 

CALENDULEAE 

G reduced pappus 

.......... G disk styles with short 

Marshal li inae branches 

Gail lardiinae 

.................................................... o loss of carbonization in achene walls t \’ 

HELIANTHEAE ...’ 

z .. o striations in achene wall EUPATOR I EAE 

,.\ o homogomous heads 

G radial thickenings on 

G expanded style appendages 

endothecial cells 

o heterogamous heads 

G tendency for suppressed gynoecia 

G polarized thickenings on 

endothecial cells 

o Anther appendage with basal 

constriction / 

Ancestral Asteroid- with paired stigmatic lines, spinulose pollen, carbonized 

ochene walls, anther appendcges without constricted bases, gynoecia always fertile. 

FIGURE 24.-Schernatic representation of relationships in the subfamily Asteroideae 

bonization in the achene wall. The Anthemideae, 

Astereae, and Senecioneae are considered more 

specialized in structure and chemistry and have 

their closest relationship to the Inuleae. 

The redelimited Heliantheae includes the Ambrosieae Cass., the Coreopsidinae Turner & Powell, the Tageteae Cass., and the Heleniaeae Benth. 

& Hook. f. These are seen as minor specializations 

within the general pattern of helianthean evolution. The primary characters of the segregates are 

often inconstant, and the groups do not seem 

phyletically more significant or isolated than 

most other subtribes of the Heliantheae. 

A number of genera traditionally placed in the 

Senecioneae must be included in the Heliantheae. 

These include Alepidocline, Neurolaena, and Schistocarpha (Robinson and Brettell, 1973b), as well as 

the genera listed by Nordenstam (1977) as members of a provisional “Arniceae”: Arnica, Bartlettia, 

Dimeresia, Haploesthes, Mallotopus, Peucephyllum, 
NUMBER 51 25 

Psathyrotes, Psathyrotopsis, Pseudoclappia, Raillardella, 

and Whitneya. The genus Dyscritothamnus, originally in the Eupatorieae but more recently shown 

to include a radiate species (Paray, 1954), is a 

member of the Heliantheae (Robinson and King, 

1977), and also Lycapsus, which traditionally has 

been placed in or near Alomia H.B.K. in the 

Eupatorieae. 

The redelimitation of the Heliantheae has resulted in removal or confirmation of removal of 

some genera from the tribe. Sheareria Moore of 

China, previously placed in the Milleriinae, is a 

member of the tribe Astereae. It seems related to 

the majority of genera in that tribe having compressed obovate achenes, but is distinct by the 

few-flowered heads and the short, subimbricate 

involucre. The removal of three genera once 

placed in the Heliantheae has been confirmed, 

Microspermum Lag. to the Eupatorieae (Rzedowski, 1970), Olivaea Sch.-Bip. ex Benth. to the 

Astereae (DeJong and Beaman, 1963), and Rigiopappus A. Gray to the Astereae (Robinson and 

Brettell, 1973~). Epallage DC. belongs to the Inuleae (Wild, 1964). In spite of some recent doubts 

(Jeffrey, 1978), reexamination shows that Geissolepzs B. L. Robinson, originally of the Galinsoginae, is a member of the Astereae, although it is 

distinctive and isolated in that tribe (Robinson 

and Brettell, 1972a). 

The impression of phyletic diversity within the 

Heliantheae seems well established. The present 

study tends to confirm that the tribe is the most 

diverse in the family. The measure by which I 

judge is the need for 35 subtribes in the present 

treatment. Other tribes with which I have personal experience and where there is comparability 

are the Eupatorieae with 18 subtribes, the Liabeae with 3 subtribes, the Arctotideae with 2 

subtribes, and the Senecioneae with at most 3 

subtribes. It should be emphasized that the previous impressions of diversity in the Heliantheae 

have been based mostly on the extremely obvious 

nature of many of the characters involved. The 

comparative uniformity and comparative phyletic insignificance previously assigned to the Eupatorieae were the result of the more subtle nature 

of its characters. 

The order of the subtribes in the present treatment is based on a combination of phyletic, 

pragmatic, and traditional considerations. Some 

important general trends seem evident. Most obvious is the isolated position of the Marshalliinae. 

As indicated under that subtribe, Marshalfia is 

outside the main line of evolution in the Heliantheae, and it may ultimately be placed in the 

closely related Inuleae when the limits of that 

tribe are more completely understood. 

Excluding the Marshalliinae, the subtribes of 

the Heliantheae can be arranged in a more or less 

linear series. Combinations of characters place 

the Ambrosiinae at one extreme of the series with 

a transition through various groups with separate 

male and female flowers to members of the tribe 

such as the Helianthinae, the Galinsoginae, and 

the Coreopsidinae. With rare exceptions these are 

all paleaceous. The series continues in the epaleaceous subtribes and reaches its other extreme in 

the Gaillardiinae, which has the most modified 

endothecial patterns in the anthers and which 

lacks carbonization in the achene walls. It is 

notable that the two extremes in the morphological series also represent extremes in the sequiterpene lactone chemistry: the Ambrosiinae with 

ambrosanolides and the Gaillardiinae with helenanolides. This more or less linear arrangement of 

character combinations is seen as a reflection of 

a predominantly two-directional evolutionary diversification from an ancestral stock more like the 

central elements of the series. 

The helianthean series shows further peculiarities when compared with the supposedly primitive character states in the tribe. High chromosome base numbers and achenes with striations 

are found randomly through all parts of the series, 

and thus are presumed to be primitive in the 

entire series. Probable primitive forms of gynoecia1 suppression, pleiomorphic achene wall striation, and shrubby to arborescent habit are concentrated in the first group of subtribes, but these 

are combined with more herbaceous involucres, 

modified achene shapes, and loss of pappus. In 
26 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

contrast, strongly subimbricate, nonherbaceous 

involucres, prismatic achenes, and radially symmetrical pappus types are found in the Galinsoginae, which is near the middle of the series, but 

these are combined with hermaphroditic disk 

flowers and mostly herbaceous habits. The epaleaceous subtribes are presumably more derived, 

but many members have prismatic achenes and 

a number of the subtribes have members with a 

capillary pappus. Thus, the various parts of the 

series retain different primitive characters, but no 

part shows all of the primitive characters combined. The evidence suggests early divergence 

from extinct ancestral forms. 

The present study has indicated more phyletic 

integrity among the epaleaceous Heliantheae 

than is generally recognized by recent workers. 

With the exception of Geissopappus, which is a 

subgroup of Calea, the various groups of the 

“Helenieae” do not find immediate relatives 

among the conventional Heliantheae, but they 

represent an extension of the typical element of 

the tribe through an intergrading series including 

the Madiinae. This series lacks functionally male 

disk flowers, mostly lacks dark anthers, and seems 

to progressively lose paleae, ornamented seed 

coats, extra thickenings on the endothecial cells, 

and, finally, the carbonized achene wall. Efforts 

to integrate some members of the “Helenieae” 

more closely, such as placing the Gallardiinae 

near the Helianthinae (Stuessy, 1977) or trying 

to link Clappia and Pseudoclappia to members of 

the paleaceous series through Varilla (Turner and 

Powell, 1977) prove erroneous or extremely dubious. In complete contrast, the various genera 

from the Senecioneae seem to integrate well: 

Schistocarpha and Neurolaena into the conventional 

Heliantheae, Huploesthes into the Flaveriinae, 

Pseudoclappia into the Clappiinae, Arnica and its 

relatives into the Chaenactidinae, and Psathyrotes 

into the Gaillardiinae. 

It is worth noting that, in spite of many exceptions, the paleaceous members of the Heliantheae 

are more concentrated in more tropical areas, 

whereas the epaleaceous members are mostly 

temperate or xeric in distribution. 

The helianthean series presented in this study 

agrees in many ways with the traditional treatments of Bentham and Hooker (1873) and Hoffmann (1890- 1894). This is partly the unavoidable 

result of the basic character patterns from which 

all the treatments are derived. Under the circumstances, it has seemed best to leave all elements of 

the Heliantheae as close as possible to their traditional places in the series when there are no 

important reasons to do otherwise. 

Not all problems of subtribal sequence are 

clearly resolved. The Clibadiinae. which shares 

many characters of the Melanipodiinae, is probably close to the Ecliptinae. The Desmanthodiinae, which is superficially similar to the Clibadiinae, has achene-wall and resin-duct characters 

closest to the Helianthinae and Galinsoginae. 

There is a reticulating character pattern between 

the Ecliptinae, Neurolaeninae, Galinsoginae, and 

Helianthinae. Some of these patterns might result 

from basic characters, such as achene-wall striations, being suppressed in various groups. In cases 

of some chemicals or of corolla surfaces. separate 

origins in different parts of the tribe seem likely. 

Nevertheless, some character distributions might 

result from past intergeneric hybridizations. The 

latter situation would conform to situations now 

suspected in some Senecioneae (Robinson and 

Brettell, 1974a) and a few Eupatorieae (Robinson 

and King, 1977). 

A major intent of this study has been the 

redelimitation and recharacterization of the subtribes of the Heliantheae into distinct phyletic 

units. I believe the present system furnishes a 

reliable basis for subtribal assignments of most 

genera and for comparisons of characters. Some 

subtribes, such as the Coreopsidinae, Ecliptinae, 

Galinsoginae, and Helianthinae, which have been 

poorly defined in the past, now show high internal 

consistency in many characters and trends. The 

clear phyletic gaps now seen between most of the 

subtribes are an extra, but not entirely unexpected, result of the study. They indicate that 

even at the subtribal level the ancestral stocks 

have become extinct. 
NUMBER iI 

Key to the Subtribes of the Heliantheae 

27 

The following key is provided to summarize the most obvious differences 

between the various subtribes recognized in this study. An attempt has been 

made to account for some of the exceptional genera by use of parenthetical 

comments. Nevertheless, it should be noted that many genera, including some 

that have all the characters of their subtribe, can be keyed more effectively on 

the basis of their individual characteristics. 

1. Achene walls carbonized, without raphides ....................... ... ..2 

2. Achenes with live or less ribs or unribbed ................................ 3 

Plants with involucres usually herbaceous or coriaceous, usually with 2 or more flowers in 

a head ......................................... .............. 4 

4. Receptacle with disk usually bearing paleae (lacking paleae in some species of Calea 

and Clibadium and in heads with 5 or less flowers): anther thecae often blackened . . 5 

5. Disk achenes prismatic, terete, or compressed, never obcompressed (often with disk 

achenes sterile); leaves usually simple or divided into narrow lobes: anther append6. Disk flowers functionally male, with gynoecia vestigial: veins of disk corolla 

throat never with fiber sheaths ...................... ..... 7 

7. Filaments glabrous; anther thecae not greenish, endothecial cells with 

thickenings on transverse walls or radially arranged ............ 8 

8. Male and female flowers arranged in separate series in the head or in 

separate heads ..................................... 9 

9. Anthers pale, sometimes separate from each other: endothecial 

cells mostly broader than long, with radial thickenings: plants 

mostly specialized for wind pollination ....... AMBROSIINAE 

Anthers often dark, always synantherous; endothecial cells quadrate or longer than wide, usually with polarized thickenings . 10 

10. Leaves alternate, usually closely spirally inserted or 

rosulate ......................................... 11 

11. Small rosulate herbs from Cuba: heads solitary on slender 

erect peduncles; ray achenes with pappus of radially 

11. Coarse rosulate, caulirosulate, or arborescent plants of the 

northern Andes: heads clustered in terminal or lateral 

inflorescences; pappus usually absent (usually with 3 

squamellae in Tamania) .............. ESPELETIINAE 

10. Leaves mostly opposite ........................... 12 

12. Disk corolla with distinct resin ducts paired along veins of 

throat, with yellowish resin . . , , , , , DESMANTHODIINAE 

12. Disk corollas with resin ducts single along veins of throat, 

resin either reddish or colorless .... ...... 13 

13. Achenes with regular, clear, nonrecessed striations; 

ray corollas with expanded, strongly trilobed 

limbs ........................... MILLERIINAE 

3. 

ages with or without glands .................. ......... ... 6 

9. 

arranged squamellae or aristae ..... HEPTANTHINAE 
28 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

13. Achenes either without striations or with irregular or 

recessed striations; limbs or ray corollas mostly bilobed 

or lacking ....................................... 14 

14. Disk corollas without colored resin in ducts; outer 

bracts of involucre usually strongly differentiated 

into spreading foliaceous bracts (lacking in Ichthyothere); achenes usually with grooves, costae, or 

a few partial striations, achenes sometimes enclosed in inner involucral bract ................ 

............................ MELAMPODIINAE 

14. Disk corollas often with reddish resin in ducts; 

outer bracts of involucre not strongly differentiated; achenes without evident costae, grooves, 

or striations, achenes never completely enclosed 

in involucral bract .......................... 15 

15. Anthers pale; limb of ray corollas single and 

expanded or completely lacking; achene glabrous, with regular pattern of isodiametric 

cells on surface ............. POLYMNIINAE 

15. Anther thecae blackened; ray corollas tubular with 3 or 4 radially arranged apical lobes; 

achenes usually pubescent; inflorescence with 

heads often in scorpioid cymes or complex 

glomerules ................... CLIBADIINAE 

Heads with 2 male and 2 female flowers arranged collaterally (small 

repent herbs from the West Indies) ............... PINILLOSINAE 

7. Filaments densely pubescent below collars; anther thecae greenish, 

median endothecial cells strongly noduliferous along vertical walls ...... 

................................................. GUARDIOLINAE 

Disk flowers hermaphroditic (some genera of Ecliptinae have disk flowers 

functionally male, but these usually have distinct fiber sheaths along veins of 

disk corolla throat; also see Unxzu of the Neurolaeninae which has yellowish 

anther thecae) .......................................................... 16 

16. Achenes without striations ........................................... 17 

17. Anther thecae whitish; pappus lacking; carbonization of achenes 

in evenly distributed brownish layer; lamina of leaves rather fleshy 

and deeply lobed (endemic to small Pacific islands off coast of 

Chile) .............................................. LYCAPSINAE 

17. Anther thecae usually blackened, sometimes yellowish; pappus often 

present; carbonization of achenes in unevenly distributed, highly 

porose, black layer ............................................. 18 

18. Disk achenes usually compressed, ray achenes often broadly triangular or obcompressed, achenes sometimes winged; pappus 

usually bilaterally symmetrical or lacking, rarely of radially arranged, short, deciduous setae; veins of disk corollas often with 

8. 

6. 
NUMBER 51 29 

strong fiber sheaths; ducts with colorless resin, ducts in shaft of 

18. Achenes prismatic, never winged; pappus radially arranged or 

lacking; disk corollas without fibers; ducts often with reddish 

.................................... ..... NEUROLAENINAE 

16. Achenes with striations .................... ................. 19 

19. Style branches with continuous stigmatic surface adaxially; achenes of 

disk flowers compressed ..................................... 20 

20. Ray flowers fertile, with style; ducts of flower with colorless resin; 

pappus lacking; basal margins of disk corollas often extended to 

cover tops of achenes, lower throat of disk corollas not 

scabrid ............. ...................... ZALUZANIINAE 

20. Ray flowers sterile or lacking, without styles (vestigial style in 1 

species of Helianthopsis), ray achenes filiform; ducts of flower with 

yellow resin; pappus usually present and bilaterally symmetrical 

(lacking in specimens with calvous achenes) ; basal margins of disk 

corollas not extended, lower throat of disk corollas distinctly 

scabrid ...................................... HELIANTHINAE 

19. Style branches with paired stigmatic lines; achenes of disk flowers 

often terete or prismatic ..................................... 21 

21. Ray flowers sterile .................................... 22 

22. Shrubs or small trees with opposite leaves; paleae becoming 

enlarged with age; ducts with colorless resin; endothecial cells 

with thickenings polarized on transverse walls; achenes 

prismatic .................................. MONTANOINAE 

22. Herbs with alternate or basal leaves; paleae not enlarged with 

age; ducts often with reddish or orange resin; endothecial cells 

with thickenings radially arranged or polarized on vertical 

walls; achenes often compressed; receptacles usually highly 

conical or columnar ...................... RUDBECKIINAE 

21. Ray flowers fertile or lacking ............................ 23 

23. Erect or caespitose terrestrial plants; heads not sessile in the 

axils of leaves on elongate stems; inner surfaces of disk corolla 

lobes papillose ....................................... 24 

24. Heads with 5 or more flowers; ray flowers present, with 

trilobed limb ........................................ 25 

25. Pappus lacking; ducts single along veins of disk corolla 

throat, with colorless or orange resin, ducts in shaft of 

disk style outside of veins; carpopodium symmetrical 

or slightly asymmetrical .............. MILLERIINAE 

25. Pappus usually present, with separate segments radially arranged; ducts paired along veins of disk corolla 

throat, with yellow resin, ducts on shaft of disk style 

inside of veins; carpopodium strongly asymmetrical, 

disk style located outside of veins .............. ECLIPTINAE 

resin, ducts in shaft of disk style located inside of veins ......... 
30 SMITHSONIAN CONTRIBCTIONS TO BOTANY 

outer half remaining as isolated sclerified plate at 

24. Heads with 2 or 3 flowers; ray flowers lacking; pappus a 

fused replicate ring, deciduous as a unit: acaulescent herbs 

from the western United States ...... DIMERESIIKAE 

23. Spreading aquatic plants; heads sessile in axils of leaves on 

elongate stems; inner surfaces of disk corolla lobes 

smooth ............................ ENHYDRINAE 

Disk achenes obcompressed (sterile and vestigial in a few genera with dissected 

leaves); leaves often subcompound with ovate or filiform segments: anther appendages always glabrous, often with duct containing reddish resin ....... 26 

26. Heads heterogamous or discoid, with tubular disk flowers: involucre usually 

with strongly differentiated outer series of herbaceous bracts .COREOPSIDINAE 

26. Heads homogamous with bisexual, 5-lobed, ligulate flowers: involucre with 

coriaceous or woody bracts, without differentiated outer series: plants endemic 

Receptacle usually without paleae on the disk (paleae present among disk flowers in 

some Madiinae and in Hymenopappus newberryi) ; anther thecae usually pale, sometimes 

reddish (blackened in Trichocoryne) ...................................... 2 7 

27. Leaves and involucres spotted with distinct, often elongate secretory pits or 

cavities .................................................... PECTIDINAE 

27. Leaves and involucres without secretory pits or cavities .................. 28 

28. Involucral bracts in single series, each subtending a flower; ray flowers when 

present separated from disk by a single series of paleae (rarely with additional 

paleae on disk); anther thecae often dark reddish; leaves with rather abundant 

intercellular pectic materials (lacking in vernal annuals) ...... MADIIKAE 

28. Involucral bracts usually in more than one series; without a single series of 

paleae inside of ray flowers; anther thecae usually pale, not reddish: leaves 

without pectic materials .................................... 29 

29. Achene walls without striations; leaves distinctly petiolate ....... 30 

30. Achenes usually rather trigonous, shortly to longly stipitate at the 

base; disk corollas with 5 lobes . , . , , , . , , , , . , HYMEKOPAPPIKAE 

30. Achenes slightly to strongly compressed, not stipitate at the base: disk 

................. PERITYLINAE 

3 1. Endothecial cells short with 3-5 thickenings on transverse walls, thickenings sometimes radially arranged; leaves without distinct petiole: 

pappus awns or squamae when present without distinct median costa 

or vascular trace ...................................... BAERIINAE 

31. Endothecial cells short to elongate with 1 or 2 thickenings on transverse 

walls; leaves with or without distinct petioles; pappus awns or squamae 

often with distinct median costa or vascular trace . CHAENACTIDINAE 

Plants with fused, fleshly, slightly winged involucres, with 1 flower in each head; xeromorphic shrubs endemic to southern Baja California , , , , . . , . , , , COULTERELLINAE 

....................................... VARILLINAE 

maturity ........... ... .GALIKSOGIKAE 

5. 

to Polynesia ............................................. FITCHIINAE 

4. 

.. 

corollas usually with 4 lobes 

29. Achene walls with striations; leaves with or without petioles ....... 31 

3. 

2. Achenes with 8-10 ribs; anther appendages glabrous ............................. 32 

32. Receptacle with paleae on the disk 
NUMBER -11 31 

32. Receptacle without paleae, sometimes with hairs .................................. 33 

pappus with numerous capillary bristles in ca. 3 series ................. CLAPPIINAE 

33. Xeromorphic shrubs with alternate leaves, without scarious wings or sheaths at nodes; 

33. Herbs or subshrubs with opposite leaves, with scarious wing or sheath at nodes; pappus 

lacking or with single series of squamae or bristles ............................. 34 

34. Erect herbs or subshrubs with heads grouped in axillary or terminal 

inflorescences .................................................. FLAVERIINAE 

34. Stoloniferous herbs with heads solitary from axils of leaves , , , , . , , , .JAUMEINAE 

I. Achene walls not carbonized, with raphides ............................................ 35 

35. 

35. 

Receptacle without herbaceous paleae, sometimes with spinelike fimbrillae or with a few bracts 

between ray and disk flowers; heads with rays present or absent; corollas usually yellow or 

whitish, with tubes and lobes not long and narrow, throat long: median endothecial cells rather 

elongate ............................................................. GAILLARDIINAE 

Receptacle with herbaceous paleae; heads discoid; corollas lavender or violet, with tubes long 

and narrow, throat very short; all endothecial cells short; endemic to southeastern United 

States 

A Synopsis of the Subtribes of the Heliantheae 

The thirty-five subtribes of the Heliantheae are 

arranged in the following sequence in this treatment. General trends are indicated by the seven 

major groups. The achene wall type and some 

notable features are cited for the individual subI ribes. 

;2. L’sually paleaceous: usually no pappus: disk flowers 

Achenes not striate: often wind-pollinated: male and female flowers often in separate 

heads: anthers pale. often separate. 

2. EwELETiiNAE: Achenes not striate: leaves alternate, 

apically in dense clusters or rosettes; often monopodial, often arborescent. 

Achenes usually striate: ray corolla surface papillose, ray achenes often completely enclosed in modified bract. 

1. POLYMNIIKAE: Achenes not striate: ray corolla surface not papillose, ray achenes not enclosed in 

bract. 

:i. MILLERIINAE: Achenes striate; ray corollas strongly 

t ri lobed, 

6. DESMANTHODIINAE: Achenes with or without striations: rcsin ducts paired in disk corolla, inside 

veins in style. 

7. CLIBADIINAE: .4chenes not striate; ray corollas tubiform: ducts in disk corolla single. 

8. PINILLOSINAE: Achenes not striate; 2 male and ‘2 

female flowers collateral in head. 

male: achenes carbonized. 

1. AMBROSIINAE: 

3, MELAMPODIINAE: 

MARSHALLIINAE 

9. G~IARDIOLINAE: Achenes striate: filaments pubescent below collars; endothecial cells with prominent thickenings polarized on vertical walls. 

B. Paleaceous: usually no pappus; disk flowers hermaphAchenes striate. prismatic; rays fer1 I. MONTANOINAE: Achenes striate, prismatic, rays 

12. R~IDBECKIINAE:chenes striate, mostly comIS. ZALUZANIINAE: Achenes striate. compressed; rays 

C. Usually paleaceous: usually with pappus: disk flowers 

usually hermaphroditic: disk achenes prismatic to 

compressed. carbonized. 

14. HEPTANTHINAE: Xchenes not striate: disk flowers 

roditic: achenes carbonized. 

10. ENHYDRINAE: 

tile: stigmatic lines 2: aquatic herbs. 

sterile: stigmatic lines 2: trees and shrubs. 

pressed: rays sterile: stigmatic lines 2. 

fertile: stigmatic surface single. 

male: short-stemmed rosulate herbs. 

1.5. ECLIPTINAE: Achenes not striate, in disk usually 

compressed: corolla veins often with fiber sheaths; 

style ducts outside veins in shaft. resin not colored. 

Achenes striate. compressed; rays 

sterile: stigmatic surface single; style ducts outside 

veins in shaft, resin yellow. 

17. NEUROLAENINAE: Achenes not striate, prismatic; 

pappus radial: rays fertile: stigmatic lines 2; style 

ducts inside veins in shaft, resin reddish. 

Achenes striate. prismatic; pappus 

radial; rays fertile; stigmatic lines 2: style ducts 

inside veins in shaft. resin yellow. 

19. DIMERESIINAE: Achenes striate, prismatic; 2 or 3 

flowers in head: pappus replicate. deciduous as 

unit. 

16. HELIANTHINAE: 

18. GALISSOGINAE: 
32 SMITHSONIAX CONTRIBUTIONS TO BOTANY 

D. Paleaceous: usually with pappus: disk flowers usually 

hermaphroditic: disk achenes obcompressed, carbonized. not striate. 

20. COREOPSIDINAE: Heads heterogamous, with distinct ray and disk flowers; style branches elongate; 

seed coat not ornamented: no glands on anther 

appendage. 

Heads homogamous. with all flowers 

ligulate and j-lobed: style branches short; seed 

coat not ornamented: no glands on anther appendage. 

E;. Not paleaceous or secondarily paleaceous on disk; disk 

flowers hermaphroditic: achenes carbonized, with or 

bvithout pappus: stigmatic lines 2. 

22. COLILTERELLINAE: Achenes striate, prismatic; heads 

25. PECTIDINAE: Achenes striate, prismatic; leaves and 

involucre with distinct internal oil glands or cavities. 

24. FLAVERIINAE: .4chenes multi-costate; inflorescence terminal or axillary; subshrubs. 

?,?. L'ARILLINAE: Achenes multi-costate; ducts single 

between pairs of veins in corolla throat, resin 

reddish: paleae present. 

26. CLAPPIINAE: Achenes multi-costate; inflorescence diffuse, with few mostly large heads; ducts 

sometimes between paired veins: xeromorphic 

shrubs. 

Achenes multi-costate: heads solitary; 

stoloniferous herbs. 

Achenes not striate: single series of pa2 1, FITCHIINAE: 

with 1 flower. 

27. JAIIMEINAE: 

28. MADIINAE: 

leae inside rays. 

29. HYMENOPAPPINAE: 

50. LYCAPSINAE: 

.Achenes not striate, trigonous. 

Achenes not striate, 4-ribbed; leaves 

spirally inserted. petiolate; disk corollas 4-lobed; 

paleae present, 

3 1, PERITYLINAE: Achenes not striate, compressed; 

leaves opposite. petiolate; disk corollas 4-lobed. 

32. BAERIINAE: .4chenes striate, prismatic to compressed: leaves not petiolate: endothecial cells with 

3-5 thickenings on transverse walls. 

Achenes striate, prismatic to obcompressed; endothecial cells with 1-2 thickenings 

on transverse walls. 

E'. Disk mostly without true paleae; disk flowers hermaphroditic: achenes not carbonized, with raphides in wall; 

pappus radial. 

31. GAILLARDIINAE: Endothecial cells with 1-2 thickG. Paleaceous: disk flowers hermaphroditic; achenes not 

carbonized. with raphides in wall: pappus radial with 

5 squamae. 

53. CHAEXACTIDINAE: 

enings on transverse walls. 

3.5. MARSHALLIINAE: Heads discoid; anthers slightly 

tailed: corollas puberulous, with tubes and lobes 

long and narrow. 

Review of the Subtribes of the Heliantheae 

The following 35 subtribes include many new 

segregates that have been validated separately 

(Robinson, 19788, 1979a, 1980a). A number of 

traditional subtribes also are reduced or altered. 

The most notable examples are the Verbesininae 

and Zinniinae, which are placed in the Ecliptinae; the Bahiinae, which are placed in the 

Chaenactidinae; and the Milleriinae, which are 

completely reinterpreted. 

In the generic listings under the subtribes, the 

accepted genera are marked with an asterisk (*). 

Subtribe 1. AMBROSIINAE 

FIGURES 25-35 

Subtribe AMBROSIINAE Less., Linnaea, 5: 151, 1830. 

Tribe AMBROSIEAE Cass., Journ. Phys.. 88: 191, 1819. 

Family AMBROSIACEAE Link, Handbuch, 1 :8 16, 1829. 

Family PARTHENIACEAE Link,Handbuch, 1:816, 1829. 

Tribe IVEAE Rydb.. N. Amer. F1.. 33(4):3, 1922. 

*Ambrosia L. (including Acanthambrosia Rydb., 

Franseria Cav., Gaertneria Medic., nom./rej ., Hemiambrosia Delpino), * Dicoria Torrey & Gray, * Euphrosyne DC., *Hymenociea Torrey & Gray, *Iva L. 

(including Chorisiva Rydb., Cyciachaena Fres., Leuciva Rydb., Oxytenia Nutt.), *Parthenice Gray, *Parthenium L. (including Villanova Ortega), *Xanthium 

L. (including Acanthoxanthium (DC.) Fourreau) . 

Herbs. Leaves opposite, alternate, or basal. 

Heads with paleae; involucre herbaceous in ca. 2 

series; flowers female or functionally male in same 

or different heads, when separate the female 

heads borne below male heads on same plant and 

often containing 1 flower enclosed in receptacle 

(2 in Xanthium, more in some Ambrosia). Ray 

corollas tubular or with small smooth limbs. Male 

flowers evenly funnelform with indistinct achene; 

lobes short, glanduliferous outside, smooth inside; 

filaments separate or fused to each other; thecae 

pale, usually separate (synantherous in Parthenice 
NUMBER 51 33 

FIGURES 25-35.-Ambrosiinae: 25, Ambrosia trifida L., head in fruiting stage, X 12: 26, 27, 

Xanthium strumarium L. (26, achene, X 6; 27, head in partial section containing pair of achenes. 

X 6); 28, Hymmoclea salsola Torr. & Gray, head in fruiting stage in partial section, X 12; 29. 30. 

Diroria canescens A. Gray, achene, X 12, section and face view; 31, Euphrosyne parthenifolia DC.. 

achene. X 24: 32. 33, Parthenice mollis A. Gray, ray achene with adjacent paleae. X 12. inside 

and outside view: 34, 35, Parthenium hysterophorus L., ray achene with adjacent paleae. X 12. 

inside and outside view. 
34 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

and Parthenzum) : endothecial cells mostly broader 

than long, with radial thickenings, 4-6 thickenings on transverse walls: appendages without 

glands, sometimes apiculate or attenuate. Style 

branches of female flower with 2 stigmatic lines; 

styles of male flower undivided, with tuft of hairs 

at tip. Resin ducts with little or no coloration, 

solitary along veins of male corolla throat. Achenes of female flowers prismatic to obcompressed, 

sometimes ornamented or forming complexes 

with adjacent paleae, usually with many uniseriate hairs near tip; walls of achenes carbonized, 

without striations; cells of seed coat ornamented; 

pappus of a few short spines or lacking. Pollen 

25-30 pm in diameter, often with reduced spinules. 

The Ambrosiinae are most notable among the 

Heliantheae for specializations toward wind pollination. The nonsynantherous condition in the 

more advanced genera of the subtribe is unique 

in the family. The subtribe can be characterized 

by the anther thecae usually lacking black pigment, by the form of the endothecial cells, and 

by the anther appendages being glabrous and 

rather thin. Seven of the genera have been examined for chromosomes and all show a base of 

X = 18. Distinctive sesquiterpenes, ambrosanolides, are recorded from Ambrosia, Hymenoclea, Iva, 

Parthenice, Parthenzum, and Xanthium. 

The relationship of Parthenice and Parthenzum to 

the Ambrosiinae was recognized by Bentham, but 

the two genera were placed technically in the 

adjacent part of the subtribe Melampodiinae in 

the Bentham and Hooker system (1873). Th’ is was 

based primarily on the presence of fused anthers 

and achene-palea complexes in the two genera. 

Achene-palea complexes are also present in the 

Engelmannza group, which likewise was included 

in the traditional Melampodiinae by Bentham 

and Hooker. Stuessy (1973) properly transferred 

Parthenice and Parthenzum to the Ambrosiinae on 

the basis of the translucent anthers, the chromosome numbers, and the chemistry, but he allowed 

for possible relationship to Engelmannza. Such relationship was not mentioned in the revision of 

the tribe by Stuessy (1977), where the Engelmannza 

group was raised to subtribal status. The present 

treatment places the Engelmanniinae in the 

Ecliptinae, where the disk corolla shape, the fiber 

sheaths in the corolla, the black anthers, and the 

endothecial cells are like those of the M’edelia 

series and totally unlike the t\mbrosiinae. The 

achene-palea complexes in the group are evidently of separate origin. 

Many detailed studies of the Ambrosiinae deal 

with specializations associated with wind pollination in the pollen grains or inflorescence form 

(Wodehouse, 1928; Payne, 1963, 1964; Skvarla 

and Larson, 1965; Payne and Skvarla, 1970). The 

pollen has a foot layer thinner than the adjoining 

endexine, has cavus areas, has loosely attached 

exine components on the columellar bases and 

foot layer, has a rather shallow ektexine, and has 

lamellations in both apertural and intercolpal 

areas, all as in other members of the Heliantheae. 

This is in contrast to pollen in the tribe Anthemideae, where specializations toward wind pollination are also found. Within the Ambrosiinae, 

Parthenice, Parthenium, and parts of Iua are fully 

helianthoid in the smaller cavus areas of their 

grains and in the presence of foramina in the 

ektexine. Reports of a different pollen form in 

Parthenice have proven erroneous (Bolick and 

Skvarla, 1976). The more advanced Ambrosiinae, 

including Ambrosia, Dicoria, Euphrosyne, Hymenoclea, 

Xanthium, and typical Iaa, are nonhelianthoid in 

the reduction or absence of foramina. Additional 

specializations in various of the advanced genera 

include a somewhat thickened, less perforate exine, a second layer of columellae, enlarged vesiculate cavus areas, reduction in spinule length, 

and shortening of the colpi. Such pollen grains 

are not found elsewhere in the Heliantheae. 

The subtribe shows little relationship to the 

Melampodiinae as defined in this treatment. The 

nonstriate achenes of the Ambrosiinae suggest 

closer relationship to such subtribes as the Clibadiinae and the Ecliptinae. 

Subtribe 2. ESPELETIINAE 

FIG( RES 36-40 

Subtribe ESPELETIIYU Cuatr . Phhtologia. 35 48, 1976 

*Carramboa Cuatr., *Coespeletza Cuatr , *Espele- 
NUMBER il 35 

tia Mutis ex H. & B., *Espeletiopsis Cuatr., *Libanothamnus Ernst, *Ruilopetia Cuatr., * Tamania 

Cuatr. 

Mostly arborescent or shrubby perennials; 

stems often caulirosula, unbranched or sparingly 

branched (usually dendroid in Libanothamnus). 

Leaves alternate, rosulate, or subrosulate, vaginate at base (tubular in Carramboa and Libanothamnus), usually without distinct petioles. Inflorescence terminal (Libanothamnus, Ruilopetia and Tamania) or axillary. Heads with paleae; involucre 

herbaceous, pluriseriate: flowers female and functionally male in same head. Female flowers in up 

to 6 series, adaxial surface of rays with short 

scarcely mamillose cells. Disk corolla lobes 5, 

short-triangular, hairs of corolla usually blunt or 

clavate at tip (usually sharp in Carramboa); thecae 

slightly darkened, endothecial cells short, with 2 

or 3 thickenings on transverse walls; appendages 

glabrous. Styles of rays with 2 broad stigmatic 

lines: styles of male flowers undivided or shortbranched, truncate, clavate. Resin ducts without 

colorat ion. Achenes of female flowers obovoid, 

somewhat trigonous to tetragonous, glabrous, 

walls carbonized, without striations: cells of seed 

coat irregular, ornamented; pappus lacking (1-3 

squamellae in Tamania). Pollen ca. 30 pn in 

diameter. 

The Espeletiinae are restricted to rather high 

elevations in the northern Andes, where they are 

characteristic of the typical Paramo habitat. The 

center of diversity and probable center of origin 

is in Venezuela, with more recent extension into 

Colombia and northernmost Ecuador. 

Most concepts of the group recognize the single 

genus, Espeletia, based on the familiar caulirosula 

species E. grandglora Humb. & Bonpl. of central 

Colombia. There are consistent chromosome reports of X = 19 for the group (Robinson et al., in 

press). Cuatrecasas (1976). in his reorganization 

of the subtribe, emphasized significant variations 

in habit and correlations with habitat that are 

not evident in the disarticulated fragments usually found on herbarium sheets. 

The genera of the Espeletiinae form a closely 

related series which is, nevertheless, strongly isolated from other subtribes of the Heliantheae. 

The group has been placed traditionally in the 

Melampodiinae and has some similarities to the 

fruticose genus Smallanthus of that subtribe. The 

Espeletiinae are distinct in the alternate leaves 

and the unstriated achenes. They also lack the 

distinct outer series of herbaceous involucral 

bracts that are found in most Melampodiinae. 

The Espeletiinae provides a most striking example of a fundamentally alternate-leaved group 

in the basically opposite-leaved Heliantheae. The 

only example of opposite .leaves in the subtribe, 

the bracts on the axillary inflorescences of Espeletia, represents a highly specialized condition, 

being a unique reversal of the common trend 

toward more alternate leaves in inflorescences 

and occurring in one of the more specialized 

genera of the subtribe. 

Subtribe 3. MELAMPODIINAE 

Subtribe MELAMPODIINAE Lessing. Linnaea. 3: 149, 1830. 

* Acanthospermum Schrank (including Centrospermum H.B.K. non Spreng., Orcya Vell.), *Ichthyothere Mart. (including Latreillea DC., Torrentia 

Vell.), *Lecocarpus Decaisne, *,kfelampodium L. (including Alcina Cav., Dysodium Rich. in Pers., Zarabellia Cass.) , *Smallanthus Mackenzie in Small. 

Herbs, shrubs, or small trees. Leaves opposite. 

Heads solitary or in terminal or axillary inflorescences, with paleae; involucre herbaceous, pluriseriate, usually with expanded outer series; flowers female or functionally male in same head. 

Female flowers in 1-3 series: limbs of corollas 

sometimes reduced, adaxially weakly to strongly 

papillose, apically bilobed. Disk corolla throat 

with veins usually narrow (broadened above in 

Zchthyothere) : lobes 5, with isodiametric, often papillose cells adaxially; thecae of anthers often 

darkened; endothecial cells short with usually 2 

or 3 thickenings along transverse walls (radial 

thickenings in some Melampodium); anther appendages with or without glands. Styles of rays 

with stigmatic lines paired; styles of male flowers 
36 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

FIGURES 36-48.-Espeletiinae-Melampodiinae: 36-39, Espeletia killippzi Cuarr., outer and inner 

ray achenes with sections, X 13; 40, Tamanza chardonti (A. C. Smith) Cuatr., ray achene, X 13: 

4 1-43. Lecocurpus pinnutifidus Decaisne, ray achenes enclosed in bract, X 12 (4 1, section showing 

form of achene);44, 45, Melampodium divaricatum (Rich.) DC., ray achenes enclosed in bract, 

X 12 (45, section showing form of achene); 46, Melampodium appendiculatum B. L. Robins., ray 

achene enclosed in bract, X 12; 47, 48, Acanthospemum hispidum DC., ray achenes enclosed in 

bract, X 12 (48, section showing form of achene). (Figures 36-40 after Cuatrecasas, unpublished.) 
NUMBER 51 37 

FIGURES 49-59.-Melampodiinae-Polymniinae-Neurolaeninae-Milleriinae: 49, Smallanthus uoedaha (L.) Mackenzie, ray achene, X 8; 50, Ichthyothere scandens Blake,' ray achene, X 12: 51, 

Polymnza canadensis L., ray achene, X 12; 52, Unxia camphorata L. f.. ray achene, X 12; 53. 54. 

Millerza quinqueflora L. (53, head at fruiting stage enclosed in involucral bract, X 12; 54, head in 

section showing mature ray achene, X 12); 55, 56, Trigonospermum melampodmdes DC., ray 

achene, cross-section and lateral view, X 17; 57, Sigesbeckia orientalis L.. disk achenes with 

corollas, lateral and inside views, X 12; 58, Rumfordia Jloribunda DC.. ray achene, X 17: 59. 

Guizotza scabru (Vis.) Chiov., ray and disk achenes, X 17. 
38 SMITHSONIAS CONTRIBUTIONS TO BOTANY 

not or scarcely divided at tip, with 2 rudimentary 

stigmatic lines in some Smallanthus. Resin ducts 

without obvious coloration, single along veins of 

disk corolla throat, outside of veins in shaft of 

disk style. Achenes of ray flowers often enclosed 

in subtending involucral bract, slightly compressed, walls carbonized, more or less striate; 

cells of seed coat slightly to strongly ornamented; 

pappus absent. Pollen 25-30 pm in diameter. 

The Melampodiinae has traditionally included 

most members of the Heliantheae with suppressed gynoecia in the disk flowers but lacking 

specializations toward wind pollination. Of the 

taxa included in the subtribe at various times, 

many have been excluded in recent studies, Philoglossa DC. (Sandwith, 1956), Dicranocarpus, Guardiola, Moonia, Parthenice, and Parthenium (Stuessy, 

1973), Baltimora, Clibadium, the Engelmannia group, 

Ichthyothere, Rensonia, and Schizoptera (Stuessy, 

1977), and the Espeletiinae (Cuatrecasas, 1976). 

In the present study, the subtribe has been further 

reduced to a comparatively uniform element in 

which all members show at least some striations 

in the achene, and all but Ichthyothere have an 

expanded outer series of involucral bracts. 

The striations in the achenes of the subtribe 

show variations to an extent that contrasts with 

most other groups of the tribe. In Acanthospermum, 

striations are present only in the form of irregularly placed lines that usually extend only part of 

the length of the achene. The striations of most 

other genera are recessed in grooves, but seem 

otherwise poorly differentiated. Some preparations of Melampodium seem to lack striations, and 

some species of Ichthyothere are much less striate 

that others. One would conclude that the striations of the Melampodiinae are in a more pleiomorphic state than those of most Heliantheae. 

A recent article by Czerson et al. (1979) emphasizes the occurrence of the tetrahydropyrane 

ichthyothereol and the lack of melampolides in 

both Clibadium and Ichthyothere. This is contrasted 

with the condition in Melampodium and Smallanthus 

(sic Polymnia). While this does not overcome the 

objections, based on structure, to relating Ichthyothere closely to Clibadium, it does indicate a greater 

phyletic gap between Ichthyothere and other Melampodiinae. 

In the restricted sense, the Melampodiinae is 

still diverse. The genera Melampodium and Acanthospermum would be regarded as close relatives by 

the traditional character of the inner involucral 

bracts enclosing the ray achenes, but differences 

in shapes and pubescence of disk corollas, different details of the inside of the lobe tips, and the 

basically different achene walls suggest otherwise. 

The distinctions are particularly marked when 

compared to the basic uniformity seen throughout all major elements of Melampodium. All genera 

retained here in the subtribe seem comparatively 

remote from each other. 

Subtribe 4. POLYMNIINAE 

FIGURE 51 

Subtribe POLYMNIIAE H. Robinson. Phytologia. 4 1 :43. 1978 

* Polymnia L. (including Polymniastrum Lam.). 

Shrubs. Leaves opposite. Inflorescence a terminal cyme. Heads with paleae: flowers female 

or functionally male in the same head. Female 

flowers in 1 series; limbs of corollas minute, adaxially nearly smooth, apically unlobed or bilobed. 

Lobes of disk corollas 5, not papillose inside, with 

series of multicellular hairs inside along margin, 

cells of inner surface elongate; thecae of anthers 

pale; endothecial cells short, with 2 or 3 thickenings along transverse walls; anther appendages 

with glands abaxially. Styles of rays with prominent pairs of stigmatic lines; styles of male flowers 

divided at tip, with large coma of hairs abaxially 

ending below tip, without stigmatic surfaces. 

Resin ducts colorless, not obvious, single along 

veins of disk corolla throat, marginal in lobes, 

apparently outside of veins in shaft of disk style. 

Achenes of ray flowers somewhat obcompressed, 

walls carbonized, without striations, with distinct 

thin layer of sclerids, cells of surface subquadrate 

in regular pattern; cells of seed coat mostly elongate, weakly ornamented; pappus absent. Pollen 

25-27 pm in diameter. 

The subtribe contains only two species of Polym- 
NUMBER iI 39 

nza that are endemic to eastern North America. 

The genus is notable for the lack of a differentiated series of outer involucral bracts, the nearly 

smooth upper surface of the rays, the elongate 

cells on the inside surface of the disk corolla lobes, 

the pale anther thecae, the small regular cells on 

the surface of the achenes, and the lack of striations in the achene wall. These furnish a significant contrast to genera of the Melampodiinae 

with which Pobmnia has previously been associated. Smallanthus, a mostly tropical genus often 

included in Pobmnia, remains in the Melampodiinae. 

There is actually no assurance that Polymnza is 

closely related to the Melampodiinae with which 

it shares the lack of pappus and the functionally 

male disk flowers. At the beginning of this study, 

Ckxza was regarded as a member of the Polymniiae, sharing most of the essential characters of 

the subtribe. Examination of two rarer species of 

l'nxia. however, has shown definite relationship 

to the genus Calea and the Neurolaeninae, a group 

in which there is no other example of suppressed 

gynoecia in the disk flowers, and where a pappus 

is usually present (Robinson, 1980b). Polymnza 

does not share the features of the Neurolaeninae 

seen in C'n,tra, but like Chxza, it might have its 

closest relationship to genera having hermaphroditic disk flowers. 

Subtribe 5. MILLERIINAE 

FIGURES 53-59 

Subtribe MILLERIINAE Benth. 8: Hook. f., Gen. PI., 2:190, 

1873. 

*Axiniphyllum Benth., *Guizotia Cass., nom. 

cons. (including Ramtilla DC. ex Wight, Veslingia 

\'is.), *Milleria L., *Rumfordia DC., *Sigesbeckia L. 

(including Limnogenneton Sch.-Bip. ex Walp., Minyranthes Turcz., Sckuhria Moench., nom. rej., non 

Schkuhria Roth, Trimeranthes (Cass.) Cass.), * Trigonospermum Less. 

Herbs with opposite leaves. Heads in terminal 

usually subcymose inflorescences, with paleae. 

Ray flowers female; limbs of corollas papillose 

adaxially, strongly trilobed apically. Disk flowers 

functionally male or hermaphroditic; lobes 

weakly to strongly papillose inside; anther thecae 

blackened; endothecial cells short, with 2 or 3 

thickenings on transverse walls; anther appendages usually without glands (with glands in Guizotia and Milleria). Style branches of rays and 

hermaphroditic disk flowers with paired stigmatic 

lines. Resin ducts without colored resin (distinctly 

colored in Guizotia), ducts solitary along veins of 

disk corolla throat, inside of veins in shaft of disk 

style. Achenes terete to trigonous, with carbonized walls, with distinct striations; cells of seed 

coat irregular or interlocking, with ornate walls; 

pappus absent. Pollen 30-40 pm in diameter. 

The Milleriinae has been traditionally characterized by heads with few flowers, but that concept is rejected here. Various genera are excluded, 

Sheareria to the tribe Astereae, Delilia and Tetranthus to the Ecliptinae, Zchthyothere to the Melampodiinae, Clibadium, Lantanopsis, and Riencourtia to 

the Clibadiinae, Desmanthodium and Stachycephalum 

to the Desmanthodiinae, and Pinillosa to the Pinillosinae. The genus Milleria itself shows relationships to the group of genera including Rumfordia, 

and the subtribal name is applied to the group in 

this treatment . 

At least part of the present subtribal grouping 

has support from other studies. The presence of 

two highly differentiated series of involucral 

bracts with one of the series strongly glanduliferous has caused Rumfordia and Sigesbeckia to be 

placed together in traditional systems. The studies of McVaugh and Laskowski (1972) and 

Turner (1978a) indicate the relationship of Trigonospermum to Sigesbeckia, and the generic position 

of some species has been disputed. The tie between Rumfordia and Axiniphyllum is mentioned by 

Sanders (1977) and Turner (1978b). 

The position of Guizotia is less certain. The 

genus has been placed in the Coreopsidinae by 

traditional systems and by Stuessy (1977). Baagoe 

(1974) transferred the genus to the Verbesininae, 

but later agreed with Stuessy (Baagoe, 1977a). 

The more definitive concept of the Coreopsidinae 

in this study excludes Guizotia. The placement of 
40 SMITHSONIAN COKTRIBGTIONS TO BOTANY 

Guizotia in the Milleriinae, however, is based 

mostly on the close approximation of technical 

characters and is made in spite of differences in 

habit, the strictly yellow rather than whitish 

flower color, and the colored resin in the corolla 

ducts. The genus is unusual in the Heliantheae 

by its restriction to the African region and its 

uncertain relationship to anything in the Western 

Hemisphere. Sigesbeckia also extends its range into 

the Eastern Hemisphere, but it seems to represent 

a separate, more recent introduction of the typical 

element of the subtribe into that area. 

Guizotia and Hefianthus, in which amino acid 

sequences of the cytochrome c are known, are 

subtribally distinct, but they are clearly in the 

same paleaceous series of the tribe. The differences of 8 amino acids (Boulter et al., 1972) 

indicate considerable instability in the molecule 

in the tribe. All speculations regarding the Coreopsidinae based on Guizotia cytochrome c 

(Turner and Powell, 1977) must be rejected since 

the genus is not a member of that subtribe. 

Various authors such as Stuessy (1977) and 

Turner (1978a) relate the Rumfordia-Sigesbeckia 

series to the Melampodiinae. Relationship to Melampodium is possible, but it seems best to hold 

these genera with strongly 3-lobed rays in a separate subtribe for which the name Milleriinae is 

available. 

Subtribe 6. DESMANTHODIINAE 

FIGURES 60-65 

Subtribe DESMANTHODIINAE H. Robinson, Phytologia, 4 1 :40, 

1978. 

*Desmanthodium Benth., *Stachycephafum Sch.- 

Bip. ex Benth. 

Subshrubs or shrubs with opposite leaves. Inflorescence with scorpioid cymes or syncephala. 

Heads with few flowers, heterogamous, with small 

involucral bracts in 1-2 series, with paleae. Ray 

flowers female; corolla limb lacking or with adaxial papillae and a trilobate tip. Disk flowers functionally male; corolla lobes and sometimes throat 

papillose inside; lobes oblong-ovate, with blunt 

uniseriate-celled hairs outside; thecae of anthers 

pale; endothecial cells with radial thickenings or 

with 3-5 thickenings on transverse walls; anther 

appendage sometimes glanduliferous. Branches 

of ray style with paired stigmatic lines, disk style 

undivided and without stigmatic surface. Resin 

ducts yellowish, paired along veins of disk corolla 

throat, along margin and sometimes midline of 

lobes, ducts inside of veins in disk style. Achenes 

of rays obovate and obcompressed, walls carbonized, with or without striations; cells of seed coat 

not ornamented?; pappus absent. Pollen ca. 25 

pm in diameter. 

Since the habits and the traditional characters 

of Desmanthodium and Stachycephafum are similar, 

their close relationship has never been questioned. 

Nevertheless, a number of basic differences are 

obvious, including the condensed inflorescence, 

the vesicle surrounding the ray flower, the minute 

tubular ray corolla, the usually radial thickenings 

of the endothecial cells, and the distinct striations 

in the achene in Desmanthodium, all in contrast to 

Stachycephafum. The well-developed yellowish resin 

ducts with the pairs along the veins of the corolla 

throat and the papillosity inside the disk corolla 

throats in species of both genera tend to confirm 

the relationship between the genera. The habits 

and the structure of the heads suggest relationship 

to the Clibadiinae, and the Desmanthodiinae are 

positioned accordingly here, but the hairs of the 

corollas are different and the achenes of Desmanthodium are striate. The color and arrangement of 

resin ducts might indicate ties to the Helianthinae 

and Galinsoginae, and the group should be 

checked for dehydrofalcarinone polyacetylenes. 

The exact nature of the vesicle around the ray 

achene in Desmanthodium is not obvious. It is usually referred to as a palea, but it is positioned 

directly inside the subtending involucral bract, 

rather than alternating as it should be. The hairs 

on its surface are like those on the unenclosed 

disk achenes, and it seems the vesicle might be a 

special outer layer of the achene, rather than a 

palea. 

The subtribe is strictly neotropical, centering 

in Central America. Desmanthodium extends into 
NUMBER 51 41 

FIC~TRES 60-79.-Desmanthodiinae-Clibadiinae-Pinillosinae-Guardiolinae-~nhydrinae-Montanoinae-Rudbeckiinae: 60-63, Desmanthodium pefolzatum Benth. (60. disk achene. X 17; 61. 62. 

ray florets in envelope, X 12; 63, ray floret without envelope, X 12): 64. 6.5, Starhjcephalum 

argentinurn Griseb.. ray achenes, lateral and face views, X 12; 66-69. Clibadium surinamtnsc L. (66, 

hair from achene. X 85: 67. disk achene, X 17; 68, 69. ray achenes, lateral and face views, 

X 12); 70-72. Lantnnopsis hispidula Wright ex Griseb. (70, disk achene, X 17; 71. 72. ray achenes. 

lateral and face views, X 12); 73, Pinillosa berteri (Spreng.) Urb., ray achene. X 12; 74. Guardiola 

papplfera P. G. Wilson, ray achene, X 5; 75, Guardiola mexicana H. & B., ray achene. X 12: 76, 

Enhydra sessillis DC., achene, X 17; 77, Montanoa tomentosa DC., disk achene, X 12; 78, KudbecXia 

laciniata L., disk achene, X 12; 79, Ratibida columnijera (Nutt.) Woor. & Standl.. disk achene. 

x 17. 
42 SMITHSONIAK CONTRIBUTIONS TO BOTANY 

northern South America, whereas Stachycephalum 

is disjunct between Mexico and southern South 

America. 

Subtribe 7. CLIBADIINAE 

FICIJRES 66-72 

Subtribe CLIBADIINAE H. Robinson. Phytologia. 41:39. 1978. 

*CLibadium L. (including Ballieria Aubl., Orsinia 

Bertol. ex DC., Osrwaldia Cass., Trixis Sw. non L.), 

* Lantanopsis Wright ex Griseb., * Riencourtia Cass. 

(including Pontesia Vell.). 

Herbs and shrubs with opposite leaves. Inflorescence strongly cymose, often scorpioid. Heads 

heterogamous, with small herbaceous involucral 

bracts in few series; with paleae (lacking on disk 

in some Clibadium). Ray flowers female, up to 26 

in l-many series: corolla tubiform with 3 or 4 

subequal disciform lobes. Disk flowers functionally male, 1-22; corolla without fiber-sheaths in 

throat: lobes slightly to strongly mamillose inside; 

hairs on outer surface of lobes uniseriate-celled, 

with elongate, papillose penultimate cells and 

short, apiculate, smooth apical cell; anther thecae 

and appendages blackened; endothecial cells 

short with 2-4 thickenings on transverse walls; 

appendage sometimes glanduliferous. Branches 

of ray style with paired stigmatic lines, disk style 

undivided and lacking stigmatic surface. Resin 

ducts somewhat reddish, ducts solitary along 

veins of disk corolla throat. Ray achenes obovate, 

subterete or angular to slightly obcompressed, 

often with prominent apical neck or stalk, densely 

pubescent with hairs or glands, walls carbonized, 

without striations; cells of seed coat elongate or 

irregular with ornamented walls; pappus absent. 

Disk achenes with long hairs. Pollen 23-30 pm in 

diameter. 

The subtribe is distributed throughout the 

Neotropical region. Clibadium is widely distributed, but with only one species in eastern Brazil; 

Lantanopsis is West Indian, and Riencourtia is 

mostly Brasilian extending north to Venezuela. 

The group is marked by the strongly cymose 

inflorescence, the disciform ray corollas, the penicillate tuft of distinctive hairs that is often well 

developed on the tips of the disk corollas, and the 

dense pubescence of the achenes of most species. 

A sigmoid twist is seen at the apex of the ray 

achene in Lantanopsis and some Clibadium species. 

The black anthers and nonstriate achene walls 

are reminiscent of the Ecliptinae, but the characteristically male disk flowers with undivided 

styles and the common presence of reddish resin 

justify a separate subtribal status. The lack of a 

pappus and the reduced rays result in a placement here nearer to the Ambrosiinae than the 

Ecliptinae. 

Some other genera have been placed close to 

Clibadium in various treatments. Desmanthodium is 

similar in habit and form of the heads, but has 

differently arranged ducts in the disk corolla with 

yellowish resin, the anthers are pale, and the hairs 

of the corolla lobes are totally different with blunt 

tips and evenly spaced septations. Delilea has 

some resemblance in habit and reduction of the 

male flowers, but the male corolla has well-developed fiber sheaths along the veins of the throat, 

the style is divided, and the sterile achene is 

glabrous. The inflorescence is also actually subumbellate rather than strongly cymose, unlike the 

Clibadiinae. The presence of the fiber sheaths 

indicates that Delilea is a member of the Ecliptinae. 

Ichthyothere was first placed in the same subtribe 

with Clibadz'um by Stuessy (1977), but the two 

genera are juxtaposed in traditional systems and 

generally are considered close relatives. The idea 

is reinforced by the occurrence of the fish poisons 

or tetrahydropyranes in both genera. Nevertheless, Ichthyothere lacks the inflorescence form, disk 

corolla form and pubescence, and achene pubescence characteristic of the Clibadiinae. In addition, the achene is striate and the veins of the disk 

corolla throat are expanded above in unique 

manner in Ichthyothere. The genus is returned to 

the Melampodiinae in the present treatment and 

is not considered a close relative of the Clibadiinae. 

The different basic arrangements of function- 
NUMBER .-)I 43 

ally male and female flowers in the members of 

the Clibadiinae seem worthy of detailed study. 

Subtribe 8. PINILLOSINAE 

Subtribe PiiviLLosmAE H. Robinson, Phytologia, 41:42, 

1978. 

*Ptntllosa Ossa ex DC., *Koehneola Urb.? 

Minute, repent herbs. Leaves opposite, with 

distinct petioles; lamina suborbicular, palmately 

veined. Heads solitary on long peduncles; involucral bracts 4, herbaceous; 2 female flowers and 

2 male flowers borne collaterally, male flowers 

inside of smaller involucral bracts. Corolla of 

female flower obsolete. Corolla of male flower 

with elongate tube and lobes, with very short 

throat; lobes densely and strongly papillose inside; anther thecae blackened; endothecial cells 

subquadrate to broader than long, with radial 

thickenings; anther appendages obsolete. Style 

branches of female flowers large, with paired 

stigmatic lines; style branches of male flowers 

large, with expanded, patelliform tips, densely 

papillose, without stigmatic surface. Resin ducts 

not evident. Achenes of female flowers prismatic, 

I-lobed, walls carbonized, without striations; cells 

of seed coat not ornamented; pappus absent. 

Pollen ca. 25 pm in diameter. 

The subtribe, as represented by Pinillosa of 

Cuba and Hispaniola, is distinct among the 

Heliantheae by a number of characters, including 

the repent habit, the collateral male and female 

flowers of strikingly different forms, the obsolete 

appendage of the anther, the radial thickenings 

of the endothecial cells, the expanded tips of the 

male style branches, and the 4-lobed achenes. 

Koehneola of Cuba, of which only the description 

has been seen, may be related to Pinillosa having 

a similar repent habit, heads with 4 involucral 

bracts and 2 each of 2 types of flowers collaterally 

arranged, deeply lobed disk corollas and an achene without a true pappus. In described features, 

the latter genus differs by the presence of a deeply 

bifid female corolla, less differentiated styles in 

the male flowers, and incompletely suppressed 

gynoecium in the male flowers, and the presence 

of a broad appendage on the anther. 

Pinillosa is one of a group of West Indian genera 

of reduced habit and solitary, long-pedunculate 

heads that traditionally is placed in the Milleriinae. Others of the group are the Cuban genera 

Heptanthus and Tetraperone, which have a rosulate 

habit and have more numerous ray and disk 

flowers in the head; and Tetranthus of Hispaniola 

and the Bahamas, with 4 hermaphroditic flowers 

in the head, simple achene wall structure, and 

fused stigmatic surfaces. The first two are placed 

in a separate subtribe, Heptanthinae, and the last 

is placed in the Ecliptinae. 

The intensely blackened anther thecae and the 

nonstriate achene wall of Pinillosa might indicate 

that it is close to the Ecliptinae. Because of the 

similarities in appearance dnd geography, it is 

possible that Pinillosa is actually closely related to 

Tetranthus, but the many differences in detail 

make this doubtful. 

Subtribe 9. GUARDIOLINAE 

FICCRES 74, 7.5 

Subtribe GUARDIOLINAE H. Robinson, Phytologia. 4 1 :41, 

1978. 

* Guardiola Cerv. ex Humb. & Bonpl. (including 

Tulocarpus Hook. & Arn.). 

Subshrubs with opposite leaves, glabrous or 

subglabrous; leaf blades simple. Inflorescence terminal. Heads often aggregated, heterogamous, 

with involucral bracts subequal in 1-2 series, with 

paleae. Ray flowers female; corolla limb white, 

adaxially papillose. Disk flowers functionally 

male; corolla with elongate tube, deeply 5-lobed; 

lobes on inner surface smooth below and strongly 

papillose above; filaments densely pilose below 

the collar; thecae of anthers blackish green; median endothecial cells strongly noduliferous along 

vertical walls; anther appendage with 1 or 2 large 

glands abaxially. Branches of ray style large, with 

paired stigmatic lines: branches of disk style elongate, sharply and densely papillose on outer sur- 
44 SMITHSONIAN CONTRIBLTIONS TO BOTANY 

face, sometimes with obsolete pairs of stigmatic 

lines on lower inside surface. Resin ducts reddish, 

ducts numerous in involucral bracts; ducts indistinct in throat of disk corolla, often prominent in 

lobes: ducts not evident in styles. Achenes of rays 

slightly obcompressed, strongly indurated, walls 

carbonized, with distinct striations; cells of seed 

coat slightly distorted. with prominent irregular 

thickenings on walls; pappus absent (with crown 

of 5 subquadrate indurate squamae in G. papptfera 

M'ilson). Pollen 30-35 pm in diameter. 

Guardtola usually has been placed in the Melampodiinae because of the suppressed gynoecia 

of the disk flowers. Stuessy (1973) transferred the 

genus to the Coreopsidinae, which it resembles in 

habit and details of the involucre. Initial doubts 

regarding Stuessy's placement derive from the 

presence of glands on the anther appendages in 

Guardzola and the apparent lack of glands on the 

anther appendages of the Coreopsidinae (Robinson and Brettell, 1973a). On further analysis, 

Guardzofa shows distinct striations in the achene 

wall and strongly ornamented cells of the seed 

coat, characters also absent in the Coreopsidinae. 

The scarcely flattened ray achenes further indicate that the divergence between Guardiola and 

the Coreopsidinae occurred before the latter 

gained any of its characteristic features. 

The glands on the anther appendages of Guardzola are distinct and massive, unlike the usual 

biseriate, capitate glands on other Heliantheae. 

In the specimens showing colored resin ducts in 

the disk corolla lobes, the ducts are inside the 

veins and do not join where they terminate at the 

top of the throat. In the species I have sampled, 

there is no variation in the dense pubescence of 

the anther filaments or in the prominent thickenings along the vertical walls of the endothecial 

cells. 

The single genus of the subtribe is restricted to 

Mexico. 

Subtribe 10. ENHYDRINAE 

FIGITRE 76 

Subtribe EXHYDRINAE H. Robinson. Phytologia, 41 :398, 

1979. 

* Enhydra Lour. (including Cryphiospermum P. 

Beauv., Hingtsha Roxb., Meyera Schreb., Sobreyra 

Ruiz & Pav., Sobrya Pers., Tetrads Reinw., Wahlenbergia Schum., non Schrad., non Blumej. 

Aquatic herbs with opposite leaves. Inflorescence of sessile axillary heads. Heads heterogamous, with herbaceous involucral bracts in few 

series, with paleae. Ray flowers female; limbs of 

corollas minute, smooth adaxially. Disk flowers 

hermaphroditic; corolla lobes slightly longer than 

wide, smooth on inner surface, outer surface 

nearly glabrous: thecae of anthers slightly 

darkened, endothecial cells short with single 

thickenings on transverse walls: anther appendages ovate, with glands abaxially. Style branches 

with paired stigmatic lines, with short blunt papillose appendages. Ducts of disk corollas often 

with reddish resin, ducts solitary along veins of 

throat, continued along margins of lobes; ducts 

of style inside the veins. Achenes prismatic or 

terete, short- to long-stipitate at base, walls carbonized, with striations; cells of seed coat inornate; small upper callus present, pappus absent. 

Pollen ca. 27-35 pm in diameter. 

The single genus Enhydra, has a form reminiscent of some Ecliptinae, but the striate subterete 

achenes are completely foreign to that subtribe. 

The reddish resin that sometimes is present in the 

disk corollas and the ducts inside of the veins of 

the style suggest a relationship to the Neurolaeninae, but the striate achenes and the lack of 

pappus favor placement in the tribal scheme 

nearer the Montanoinae. The aquatic specialization of the Enhydrinae seems distinctive. 

Enhydra occurs at lower elevations in tropical 

and subtropical parts of both hemispheres. 

Subtribe 11. MONTANOINAE 

FIGURE 77 

Subtribe MONTA~OINAE H. Robinson, Phytologia. 41:42, 

1978 

*Montanoa Cerv. in Llave & Lex. (including 

Eriocoma H.B.K., non Nutt., Montagnaea DC., 

C'hdea Kunth). 
NUMBER 51 45 

Shrubs and small trees with opposite leaves; 

leaves distinctly petiolate, blades trinervate and 

often lobed. Inflorescence corymbose-paniculate. 

Heads with ray and disk flowers, with small 

herbaceous involucral bracts in 2-3 series; paleae 

present and accrescent with age. Ray flowers 

sterile; limb of corolla long, papillose on upper 

surface. Disk flowers hermaphroditic; corolla 

densely scabrid outside; lobes short- to long-triangular, with stomates on outer surface, papillose 

inside; anther thecae blackened; endothecial cells 

short, with 2 or 3 thickenings on transverse walls; 

anther appendage ovate, with glands and hairs 

abaxially. Branches of style with paired stigmatic 

lines, appendage tapering. Resin ducts without 

evident color. Disk achenes quadrangular obpyramidal, walls carbonized, with striations, surface 

of wall woody; cells of seed coat weakly ornamented; pappus absent. Pollen ca. 25-35 pm in 

diameter. 

The position of the Montanoinae in the present 

subtribal sequence is based primarily on the bisexual disk flowers having uncompressed, epappose achenes. Previous treatments have associated 

Montanoa with various other genera with sterile 

ray flowers, genera placed here in the Helianthinae and Ecliptinae. Rojasianthe of the latter 

subtribe is of interest because it is the only other 

genus in the tribe with paleae markedly enlarged 

with age, but the presence of a pappus and the 

specialized form of disk corolla make close relationship to Montanoa doubtful. Mature achenes of 

Rqasianthe have not been seen, but there seem to 

be no striations in the wall. The achenes in Montanoa are more like those in the Melampodiinae 

and Milleriinae, but the scarcely lobed sterile rays 

and the hermaphroditic disk flowers preclude 

inclusion of the genus in either group. 

The genus Montanoa occurs in Mexico, Central 

America. and northern South America. 

Subtribe 12. RUDBECKIINAE 

FIGURES 78. 79 

Subtribe RUDBECKIINAE H. Robinson. Phytologia, 41:43, 

1978. 

*Dracopis Cass., *Ratibida Raf., *Rudbeckia L. 

(including Heliophthalmum Raf., Lepachys Raf., Obeliscotheca Adans., Peramibus Raf.). 

Plants herbaceous with alternate leaves. Inflorescence with scapose solitary heads or sparsely 

branched. Heads with highly conical or columnar 

receptacles, with ray and disk flowers, involucral 

bracts herbaceous in ca. 2 series, with paleae. Ray 

flowers sterile; limb of corolla prominent, papillose on upper surface. Disk flowers hermaphroditic; lobes short, triangular, papillose on inner 

surface; anther thecae blackened; endothecial 

cells subquadrate, with thickenings radial or on 

vertical walls; anther appendage ovate, with cluster of stout glands abaxially. Branches of style 

with paired stigmatic lines; appendage short to 

long-tapering, with dense brush of long papillae. 

Resin partially reddish; ducts of disk corolla 

throat solitary along veins. Disk achenes compressed to subquadrangular, walls carbonized, 

with striations; cells of seed coat not or slightly 

ornamented; pappus absent or with two aristate 

projections. Pollen ca. 23-27 ym in diameter. 

The Rudbeckiinae is an exclusively North 

American group previously placed in the broadly 

interpreted Verbesininae of Bentham and Hooker 

(1873) and in the more narrowly interpreted 

Helianthinae of Stuessy (1977). Similarities in 

habit and in some details such as sterile rays and 

the shape of the Ratibida achene do suggest a 

relationship to elements from the traditional Verbesininae, which are placed here in the Ecliptinae. The Rudbeckiinae differ from these genera 

and all Ecliptinae by the striate achenes, the 

radial or vertical arrangements of thickenings in 

the endothecial cells, the distinctive cluster of 

stout, cylindrical glands on the anther appendages, and the common occurrence of prominent 

reddish resin in the ducts of the disk corolla 

throat. 

The Rudbeckiinae, as delimited here, seems to 

be an entirely natural group in spite of the more 

truncated style branches and more compressed 

achenes of Ratibzda and the vertically rather than 

radially arranged endothecial thickenings in Dracopis. Echinacea, which is usually regarded as a 
46 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

close relative of Rudbeckia, has nonstriate achenes, 

lacks the cluster of distinctive glands on the anther appendage, has single thickenings on the 

transverse walls of the endothecial cells, and has 

a single, fused stigmatic surface on each style 

branch. Echinacea is evidently not particularly 

close to the Rudbeckiinae, and it conforms in all 

essential details to the present interpretation of 

the Ecliptinae. 

Subtribe 13. ZALUZANIINAE 

FI(:IIRES 80-83 

Subtribe ZALIZANIIYAL H. Robinson, Phytologia, 41:44, 

1978. 

* Chromolepis Benth., *Hybrzdella Cass. (including Chzlzophyllum DC., nom. rej., non Philippi), 

*Zaluranza Pers. (including Chrysophanza Kunth ex 

Less., Ferdinanda Lag.). 

Herbs or shrubs with alternate or basal leaves, 

blades often lobed or dissected. Inflorescence with 

few to many heads in corymbose panicles, or 

heads solitary on peduncles from basal rosette. 

Heads heterogamous; receptacle convex or 

shortly conical; involucre herbaceous in ca. 2 

series: paleae present. Ray flowers female (substerile in Zalutania grayana B. L. Robins. & 

Greenm.); limb of corolla prominent, papillose on 

upper surface, bilobed or obscurely trilobed at 

tip. Disk flowers hermaphroditic; corolla outer 

surface often covered with dense pubescence of 

blunt or gland-tipped hairs, base often with extended margin covering top of achene; lobes 

short, papillosity on inner surfaces rather abruptly demarcated at base: anther thecae blackened; endothecial cells with 2-4 thickenings on 

transverse walls; anther appendages ovate, with 

or without glands abaxially. Style branches with 

single, fused stigmatic surface inside. Resin ducts 

not colored, solitary and sometimes prominent 

along veins of disk corolla throat. Achenes slightly 

compressed, walls carbonized, with striations; 

cells of seed coat not ornamented, without sinuous 

walls; pappus absent. Pollen 25-30 pm in diameter. 

In Stuessy’s revised system of the tribe (1977), 

Zalutanza is placed in the Neurolaeninae, while 

Chromolepis, in spite of its fertile rays, is placed in 

the Helianthinae. Actually, the striate achenes 

and fused stigmatic surfaces of the Zaluzaniinae 

are suggestive of the latter subtribe. Differences 

in the rays, the lack of the distinctive resin ducts, 

and the lack of the characteristic scabrous bases 

of the corolla throats clearly distinguish the Zaluzaniinae from the Helianthinae. Also, the achenes are fundamentally epappose in the Zaluzaniinae, a condition permitting the unique expanded corolla base in Zaluzania, whereas the 

Helianthinae are basically pappose with intermittent calvous forms. 

The fused stigmatic surface of the style 

branches and the fertile rays furnish technical 

distinctions from various other epappose subtribes having hermaphroditic disk flowers. Alternate leaves are consistent in known members of 

the Zaluzaniinae as in the Rudbeckiinae, but the 

character also is found in other subtribes, especially among genera from more temperate areas. 

The subtribe is native to the southwestern 

United States and adjacent Mexico. A South 

American species previously placed in Zaluzania 

is the basis of the genus Kingzanthus of the subtribe 

Ecliptinae (Robinson, 1978b). 

Subtribe 14. HEPTANTHINAE 

E’IGIJRE 84 

Subtribe HEPTANTHINAE H. Robinson, Phytologia, 41:41, 

1978. 

*Heptanthus Griseb., * Tetraperone Urb. 

Plants rosuliform with short erect stems, leaves 

distinctly petiolate. Heads solitary on long peduncles; with ray and disk flowers; involucre herbaceous in ca. 2 series; paleae few or lacking. Ray 

flowers female; limb of corolla papillose above, 

bilobed at tip. Disk flowers functionally male; 

corolla with short lobes, lobes slightly papillose 

on inner surface; anther thecae slightly blackened; endothecial cells short, with thickenings of 

inner cells sometimes radial, inner walls often 

strongly ornamented; anther appendage without 
NUMBER jl 47 

FIGURES 80-90.-Za!uzaniinae-Heptanthinae-Ec!iptinae: 80, 8 1, Chromolepis heterophylla Benth. 

(80. ray achene, X 12: 81, disk achene, X 12); 82, 83, Zalutania angustata (Lag.) Sch.-Bip. (82. 

ray floret, X 12: 83. disk achene, X 12): 84, Heptanthus cochlearfolius Griseb.. ray achene, X 17: 

85. 86. Helzopsis buphthalmozdes (Jacq.) Dunal (85. ray floret with persistent corolla. X 12: 86. 

disk achene. X 17): 87, 88, Zinniagrandzflora Nutt. (87, ray floret with persistent corolla, X 8: 88. 

disk achene. X 12): 89, Sanvitalia aubertii A. Gray, disk achene. X 12: 90, Borrichzafrutescens (L.) 

DC.. disk achene, X 12. 
48 

glands. Style branches of rays with paired stigmatic lines; style of disk flowers distinctly 

branched, densely long-papillose. Resin ducts 

mostly indistinct (reddish and distinct in involucre and paleae of Tetraperone). Ray achenes polygonal or terete, with brownish carbonization, not 

striate, sometimes winged; cells of seed coat rather 

subquadrate, strongly ornamented; pappus of radially disposed squamellae or aristae; disk achenes linear, inornate. Pollen 25-35 pm in diameter. 

The Heptanthinae is regarded here as a natural 

group of two Cuban genera distinct by the small 

erect rosuliform habit, the sterile disk flowers, and 

the prominent, radially disposed pappus of the 

ray achenes. The genera differ in a number of 

features, including the form of the leaf blades, 

which are cordiform with palmate venation in 

Heptanthus and elliptical with subpinnate venation in Tetraperone. Heptanthus apparently lacks 

paleae and has more membranaceous involucral 

bracts. whereas Tetraperone has some thin paleae 

and firmer involucral bracts reminiscent of those 

in the Coreopsidinae. 

The genus Heptanthus is lectotypified here by 

H. cochleariflius Griseb. My concept of the species 

is based on the isotype Wrzght 2819 (US). Material 

of Tetraperone seen has mostly lacked corollas, but 

remnants of one disk corolla were found tangled 

in the bracts of a dried head. 

Subtribe 15. ECLIPTINAE 

FIGI:RES 85- 129 

Subtribe ECLIPTINAE Lessing, Linnaea, 6: 153, 1831. 

Subtribe ZIZSIINAE Benth. & Hook., Gen. PI., 2: 193, 1873. 

Subtribe VERBESININAE Benth. & Hook., Gen. Pl., 2: 193, 

Subtribe ENGELM.ANNIINAE Stuessy, Biol. Chem. Compositae, 

1873. 

645. 1977 [ 19781. 

*Acmella Rich. ex Pers. (including Colobogyne 

Gagnep., Pyrethrum Medic., non Zinn.), *Ancistrophora A. Gray, *Angelphytum Barroso, *Aspilia 

Thou. (including Anomostephium DC., Coronocarpus 

S c h u m . , Dip t e ro t h eca S c h . - B i p . , Gym n o lo m ia 

H.B.K., Harpephora Endl., Wirtgenia Sch.-Bip.), 

* Balsamorhiza Hook., * Baltimora L. (including 

Fougeria Moench., Fougerouxia Cass., Scolospermum 

Less.), * Berlandiera DC., * Blainvillea Cass. (including Eisenmannia Sch.-Bip., nom. nud.), *Borrichia 

Adans. (including Adelmannia Reichb., Diomedea 

Cass., Helicta Cass., Trimetra Moc. & Sesse ex 

DC.) , *Calyptocarpus Less. (including Blainviliea 

subg. Oligogyne (DC.) Baker, Galophthalmum Nees 

& Mart.), *Chrysogonum L. (including Cargiila 

Adans., Diotostephus Cass.), *Delilia Spreng. (including Desmocephalum Hook. €., Elvira Cass., Meratia Cass., non Loisel., Microcoecia Hook. f.), *Dimerostemma Cass. (including Serpaea Gardn.), *Dugesia A. Gray, *Echinacea Moench. (including 

Brauneria Neck. ex Britt., Helichroa Raf.), *Eclipta 

L., nom. cons. (including Abasaloa La Llave & 

Lex. ?, Eupatoriophalacron Mill., Micrelium Forsk., 

Paleista Raf.), *Eleutheranthera Poiteau ex Bosc 

(including Euxenia Chamisso, Fingalia Schrank, 

Kegelia Sch.-Bip., Ogiera Cass.) , *Encelia Adans. 

(including Pallasia L’Herit. ex Ait., non Scop., 

non L. f., non Klotzsch., non Pallassia Houtt.), 

*Enceliopsis A. Nels., *Engelmannia Torr. & Gray, 

*Exomiocarpon Lawalree, * Fenixia Merrill, *Flourensia DC., *Geraea Torr. & Gray, *Helianthella 

Torr. & Gray, *Heliopsis Pers. (including Andrieuxia 

DC., Helepta Raf., Kallias Cass.), *Hoffmanniella 

Schlect., *Kingianthus H. Robins., *Lasianthaea 

DC. (including Telesia Raf.), *Leptocarpha DC., 

*Lindheimera Gray & Engl., *Lipochaeta DC. (including Aphanopappus Endl., Microchaeta Nutt., 

Schizophyllum Nutt., non Fries), * Lundellianthus H. 

Robins., *Macraea Hook. f. (including Trigonopterum Steetz ex Anderss.), *Melanthcra Rohrb. (including Echinocephalum Gardn., Lipotriche R. Br., 

Psathurochaeta DC., Wuerschmittia Sch.-Bip. ex 

Walp.), *Monactis H.B.K. (including Astemma 

Less., Monopholis Blake), *Otopappus Benth. (including Notoptera Urb.), * Oxycarpha Blake, *Oyedaea DC., *Perymeniopsis H. Robins., *Perymenium 

Schrad., *Philactis Schrad., *Phoebanthus Blake, 

*Plagiolophus Greenm., *Podachaenium Benth. ex 

Oerst. (including Dicalymma Lem., Cosmophyllum 

Koch), *Podanthus Lag., nom. cons., non Podanthes 

Haw., *Rensonia Blake, *Rojasianthe Standl. & 

Steyerm., *Salmea DC., nom. cons., non Salmia 
NUMBER 51 49 

Cav. (including Fornicaria Raf., Hopkirkia 

Spreng.), *Salmeopsis Benth., *Sanvitalia Gualt. ex 

Lam. (including Anaitis DC., Lorentea Orteg., non 

Lag.), *Schizoptera Turcz., *Silphium L., *Sphagneticola Hoffm. ?, *Spilanthe.r Jacq., *Steiractinia 

Blake, *SynedreEla Gaertn., nom. cons. (including 

Ucacou Adans.) , *Synedrellopsis Hieron. & Kuntze, 

* Tetranthus Swartz, * Verbesina L. (including 

Achaenopodium Brandeg., Actinomeris Nutt., Altamirania Greenm., Chaenocephalus Griseb., Ditrichum 

Cass., nom. rej., non Hampe, Hamulium Cass., 

Hingstonia Raf., Ochronelis Raf., Phaethusa Gaertn., 

PlatyPteris H.B.K., Ridan Adans., nom. rej., Saubinetia Remy, Tepion Adans., Ximenesia Cav.), * Vigethia Weber, * Wedelia Jacq., nom. cons., non 

Loefl. (including Menotriche Steetz, Niebuhria 

Neck. ex Britt., non Scop., non DC., Pascalia 

Or t eg . , St emmodon t ia Cass . , Thelech itonia Cu at r., 

Trichostemma Cass., non Trichostema L., Trichostephium Cass.), * Wollastonia DC. ex Decaisne, * Wulf- 

$a Neck. ex Cass. (including Chatiakella Cass., 

Chylodia Rich. ex Cass., non Chilodia R. Br., Tilesia 

Meyer), * Wyethia Nutt. (including Alarconia DC., 

Melarhiza Kellogg, nom. provis.), *Zexmenia La 

Llave & Lex., *Zinnia L., nom. cons. (including 

Crassina Scepin, Diplothrix DC., Lepia Hill, nom. 

rej,, Mendezia DC., Sanuitaliopsis Sch.-Bip. ex 

Benth., Tragoceras H.B.K.). 

Annual or perennial herbs, shrubs, or small 

trees. Leaves opposite, alternate or basal, with or 

without distinct petioles. Inflorescence of solitary 

heads on long peduncles or in panicles, axillary 

or terminal. Heads usually heterogamous; involucre herbaceous in 2-4 series, sometimes with 

scarious inner series; paleae present. Ray flowers 

female or sterile, rarely absent; limb of corolla 

papillose above, bilobed or obscurely trilobed at 

tip. Disk flowers hermaphroditic or sometimes 

functionally male; fibrous sheaths often present 

along veins of throat, rarely fibers in lobes or 

between veins; lobes usually short (long in Rojasianthe), slightly to strongly papillose inside, often 

more papillose or with hairlike papillae adaxially 

near margin, lobes often densely scabrous outside, 

sometimes with glands; anther thecae usually 

black (pale in Encelia group and some Verbesina); 

endothecial cells with 1-3 thickenings on transverse walls; appendages ovate, with or without 

glands abaxially. Styles distinctly branched, 

branches rarely with glands abaxially; stigmatic 

surface fused or divided into a pair of lines. Ducts 

of corollas without colored resin (traces of reddish 

resin in Perymenium and Echinacea), ducts solitary 

along veins of disk corolla throat, ducts of style 

shaft, where evident, outside of veins, usually 

outside of veins in branches. Achenes of fertile 

rays usually broadly triangular to obcompressed, 

disk achenes compressed or quadrangular; walls 

carbonized, without striations: cells of seed coat 

slightly irregular, walls not strongly sinuous, usually ornamented on tangential surfaces: pappus 

often with 2 aristae, sometimes with squamellae, 

a circle of short deciduous setae, or lacking. Pollen 

mostly 25-35 pm in diameter. 

The Ecliptinae initially seems rather amorphous, but it is regarded here as natural, being 

comparable phyletically to such subtribes as the 

Ambrosiinae, Galinsoginae, Helianthinae, Pectidinae, and Gaillardiinae. All members have 

blackened, nonstriate achenes and lack well-developed patterns of colored resin in the ducts of 

their corollas. The anthers tend to be strongly 

blackened. Many of the members have fibers 

along the veins of the disk corollas, a feature not 

seen elsewhere in the Heliantheae. Where known, 

the polyacetylenes lack any of the special epoxides 

or ketones found in some other subtribes. Ambrosanolides and helenanolides are lacking. The subtribe contains the only three genera of the tribe 

from which amides have been reported, Acmella, 

Echinacea, and Hehopsis (Bohlmann et al., 1973). 

The only members of the Heliantheae known to 

have glands on the abaxial surface of the style 

branches are also in the Ecliptinae, Podachaenium 

(Robinson, 1978c), Zexmenza mtkanioides (Britt.) 

Blake (Anderson et al., 1979), and species of 

Dimerostemma. In the last case glands occur on 

both the branches and the upper shaft of the 

style. 

The diversity of the group has led to many 

separate subtribes being established. The name 

“Ecliptinae” has priority in the present combined 
SMITHSONIAN CONTRIBUTIONS TO BOTANY 

FIGURES 91- 102.-Ecliptinae: 91, Chrysogonum uirginianum L., ray and disk achenes in bract 

complex, X 12; 92, Engelmannia pinnat+da Torr. & Gray, ray achene in bract complex, X 8: 93, 

Wedelia calycina L. C. Rich., disk achene, X 8; 94, 95, Synedrella nodqlora (L.) Gaertn. (94, ray 

achene, X 12: 95, disk achene, X 12); 96, Rensonia salvadorica Blake, ray achene, X 12: 97, 

Schizoptera peduncularis (Benth.) Blake, ray achene, X 12; 98, 99. S)nedrellopsis grisebachii Hieron. 

& 0. Kuntze, achenes showing variation, X 12; 100, Calyptocarpus uialis Less., ray achene. X 12; 

101, 102, Lundellianthus gualemalensis 0. D. Smith) Rindos (101, ray achene, X 12; 102, disk 

achene, X 12). 
NUMBER 51 51 

concept. Stuessy (1977) accepted a narrower concept which included Wedelia and many related 

genera that had previously been placed in four 

different subtribes. The Stuessy grouping was 

generally defined by a herbaceous habit and carpellate ray florets. The name “Verbesininae,” 

which was used for the typical subtribe of the 

Heliantheae by Bentham and Hooker (1873), was 

retained by Stuessy for a more artificial group 

generally defined as shrubby with neuter ray 

florets. Stuessy also established the subtribe Engelmanniinae for a group of North American 

genera with achene complexes that were once 

placed in the Melampodiinae, and he recognized 

the traditional Zinniinae in which the ray corollas 

are usually fused to the achenes and are persistent 

at maturity. 

The present concept of the Ecliptinae is not 

strictly a combination of Stuessy’s four subtribes. 

Some genera placed in the Ecliptinae by Stuessy, 

such as Enhydra and Garcilassa, and all but one of 

those in his group 1 of the Verbesininae are 

excluded here. Some genera placed elsewhere by 

Stuessy, such as Delilea (Milleriinae) and the Encelia group (Helianthinae), are included here. 

There are significant subgroupings within the 

Ecliptinae. Regarding the typical element, Eclipta 

itself is distinctive by the filiform paleae, but in 

habit and various details of styles and disk achenes it is similar to Wedelia, with which it shares 

the specific flavanoid wedelolactone (Hegnauer, 

1977). In the group related to Eclipta and Wedelia, 

the ray achenes, when fertile, characteristically 

are trigonous to broadly obcompressed, in contrast with the more compressed or quadrangular 

achenes of the disk. The stigmatic lines of the 

group are separated in pairs. As an additional 

character, many of the genera have disk corollas 

of identical, narrowly funnelform shape with 

short, externally scabrid lobes and fibers or fibersheaths along the veins of the throat. This has 

been seen in American species of Aspilia and in 

Berlandiera, Cahptocarpus, Chrysogonum, Delilea, Engelmannia, Exomiocarpon, Fenixia, Hoffmanniella, Lasianthaea, Lindheimera, Lundellianthus, Oyedaea, Renson ia, Sch izop tera, Silph ium, Steiractinia, Sy nedrella, 

Synedrel lopsis, Vigeth ia , Wedelia , Wo 1 last onia , and 

Zexmenia. This extremely natural group includes 

genera placed by Bentham and Hooker (1873) 

and by Hoffman (1890-1894) in the Milleriinae, 

the Melampodiinae, and Verbesininae, and the 

Coreopsidinae. Fiber development reaches its extreme in Rensonia, with broad sheaths along the 

margins of the lobes, and there are many auxillary 

fiber strands in the lower parts of the corolla in 

Synedrellopsis. In the Wedelia group, the paleae 

usually have broad, blunt, and often colored tips, 

a striking contrast to those of the peripherally 

related Eclipta. In accepting the recent resurrection of Lasianthaea (Becker, 1976), it should be 

noted that both it and Zexmenia are in the relationship of Wedelia. Rindos (1980) furnishes significant insights into generic limits in this group. 

Baltimora also seems properly placed here (Stuessy, 

1977), even though no fibers have been seen in 

the corollas. The genera of Stuessy’s subtribe 

Engelmanniinae are distinctive in the achene 

complexes in their ray flowers, but they all fall 

within the present series. They have disk corollas 

essentially identical to those of Wedelia in shape, 

pubescence and, except for Dugesza, in presence 

of fiber-sheaths. The broadly margined ray achenes are approached in form in such genera as 

Schizoptera, Synedrella, and Synedrellopsis. 

Dugesia is one of the two genera of the Heliantheae from which eremophiline sesqueterpene lactones have been reported (Bohlmann and Zdero, 

1976). The presence of this class of compounds, 

until recently known only from the Senecioneae, 

must be rechecked. 

The genera Balsomorhiza, Helianthella, and Wyethia are coarse alternate-leaved herbs with sterile 

rays and with fibers usually forming sheaths along 

the veins of the disk corolla throats, but they are 

distinct from the previous group by the fused 

stigmatic surface on the style branches. Although 

lacking fibers and showing prominent resin in the 

style branches, Echinacea also seems to belong in 

this group. 

Borrichia is the remaining genus where fibers 

have been seen in the corolla. The opposite leaves, 

elongate lobes of the disk corollas, and fused 

.. 
52 SMITHSOXIAN CONTRIBUTIONS TO BOTANY 

FIGURES 103-1 16.--Ecliptinae: 103, 104, Zexmenia serrata La Llave (103, disk achene, X 12; 104, 

ray achene, X 12); 105, 106, Laszanlhaeafruticosu (L.) Becker, ray and disk achenes, X 12; 107, 

108. Peymenzopsis ocalifolia (A. Gray) H. Robins., disk achenes, X 12; 109: Steiractinia mollis Blake, 

disk achene, X 12: 110, Melanthera aspera (Jacq.) Small, disk achene, X 12: 11 1, Eleutherantheru 

ruderulis (Swartz) Sch.-Bip., disk achene, X 12; 112, 113, Peiymenium grunde Hemsl. (1 12, disk 

achene, X 12; 113, ray achene, X 12); 114-116, Otopappus verbesinoides Benth. (114, 115, disk 

achenea, X 12; 116, ray achene. X 12). 
NUMBER 51 

stigmatic surfaces seem to preclude inclusion of 

the genus in either of the preceding groups, but 

the distinctive disk corollas and the deciduous ray 

corollas seem to exclude it from the Zinnia relationship where it was placed by Stuessy (1977). 

Borrichia also has distinctive septate hairs on the 

backs of the style branches. 

I am convinced that the traditional Zinniinae, 

even without Borrichia, is an unnatural group. 

The strongly petiolate Phifactis and Heliopsis seem 

close to each other, but not close to Zinnia or 

Sanvitalia. All the Zinniinae share the basic characters of the Ecliptinae, and some genera such as 

Helianthella and some specimens of Zexmenia, not 

in the traditional Zinniinae, seem to have ray 

corollas fused to the achenes. 

A radial capillary pappus in the Heliantheae 

usually is associated with subtribes having prismatic achenes, but four genera of the Ecliptinae 

have short deciduous bristles arranged more or 

less radially: Melanthera, Perymenium, Rojasianthe, 

and Steiractinia. A transitional situation to the 

biaristate condition is seen in Perymeniopsis, which 

has clusters of setae at the outer and inner limits 

of the pappus and has squamellae inserted at a 

slightly higher level between. Various differences 

between genera and the existence of a transitional 

genus indicate that the genera with such a radial 

capillary pappus do not form a natural group. 

Melanthera seems closely related to the epappose 

Wuljja, which also lacks fibers in the disk corollas, 

and is clearly related to the partially pappose 

Wollastonia (Fosberg and Sachet, 1980) of the 

Wedelia group in which corolla fibers occur. Even 

Perymenium and Steiractinia may not be so closely 

related as sometimes thought, since the former 

lacks fibers in the disk corolla which are present 

in the latter. 

The best example of a dioecious member of the 

tribe is Podanthus of Chile, which seems closely 

related to the heterogamous genus Leptocarpha of 

the same area. These genera, along with Macraea 

of the Galapagos and Podachaenium of Mexico and 

Central A4merica, are notable for the slender bases 

of their achenes. 

The small, repent Tetranthus of the West Indies 

53 

has been placed previously in the Milleriinae with 

various other West Indian genera having small 

stature and few-flowered heads. Of the other 

genera, Pinillosa shares the habit and four flowers 

in the heads, but details indicate that the two 

genera are not closely related. The achenes with 

nonstriate walls, the small pappus, the four hermaphroditic flowers each enclosed in one of the 

convolute involucral bracts, and the fused stigmatic surface in Tetranthus all are readily accommodated in the Ecliptinae. 

Among the genera of Ecliptinae with strongly 

compressed disk achenes, both Verbesina and Plagiolophus have broad wings, but the narrownecked achene of the latter suggests that is is 

closer in relationship to Wedelia than Verbesina. 

Genera more closely related to Verbesina include 

Acmella and Salmea, which seem to have fused 

stigmatic surfaces on the style branches, but in 

some more robust specimens there is slight separation of stigmatic lines. ilcmella, which until 

recently has usually been placed in the synonymy 

of Spilanthes, has resin ducts of the style branches 

inside of somewhat broadened veins. In contrast 

Spilanthes urens Jacq., the type species of Spilanthes, 

has the ducts outside the unbroadened veins 

throughout the shaft and branches of the style, as 

in most Ecliptinae. It is notable that the three 

helianthean genera containing amides, Acmella, 

Echinacea, and Heliopsis, while all members of the 

Ecliptinae, do not seem particularly closely related to each other within the subtribe. Only in 

Acmella are the amides acetylenic. 

One group of genera near Verbesina having 

stigmatic surfaces unquestionably fused includes 

Encelia, Enceliopsis, Flourensia, Geraea, and Phoebanthus. The first four of these seem to form a natural 

group, with characteristically pale anther thecae 

and sterile rays. Although the group of four genera is distinctive, every one of its characters is 

duplicated elsewhere among the closely related 

genera of the Ecliptinae. Even pale anther thecae 

are common in some Mexican species of L’erbesz‘na. 

The Encelia group often has been placed close to 

members of the Helianthus group, especially Simsia, and the group was retained in the Helian- 
54 SMITHSOKIAN CONTRIBUTIONS TO BOTANY 

FIGURES 11 7-129.-Ecliptinae: 117, Oxycarpha saedaefolia Blake. achene, X 12: 118, Echznacea 

atrorubens Nutt.. disk achene, X 12: 119, Helianthella unijha (Nutt.) Torr. & Gray. disk achene, 

X 8; 120, hjethia helenioides (DC.) Nutt., disk achene, X 12: 121, 122. Podachaenium eminens (Lag.) 

Sch.-Bip., ray and disk achenes, X 12; 123, Verbesina uzrginica L., disk achene. X 12: 121. 

Podanthus rnitzqui Lindl., achene from female plant, X 12: 125, 126. Macraea laricifolia Ho0k.f.. 

disk achenes, X 12; 127, Spilanthes wens Jacq., disk achene. X 12: 128. Monactis pallatangensis 

(Hieron.) H. Robins., disk achene, X 12; 129, Encelia canescens Lam.. disk achene. X 12. 
NUMBER .jl 55 

thinae in the recent treatment by Stuessy (1977), 

where the Verbesininae were given separate subtribal status. The lack of colored resin in the 

corollas, the nonstriate achenes, the lack of scabrosity on the lower throat of the disk corollas, 

and the apparent lack of dehydrofalcarinone polyacetylenes mark the Encelia group as Ecliptinae 

rather than Helianthinae. 

The genera Monactis and Kingianthus of Ecuador 

and Peru are distinguished from most other Ecliptinae by alternate leaves and prismatic or terete 

achenes. The two genera seem transitional to the 

subt ribe Neurolaeninae, but they differ from the 

latter by the pappus, when present, being asymmetrical, and by the complete lack of colored 

resin. 

No material of Sphagneticola Hoffm. of Rio de 

Janeiro, Brasil, has been seen in this study. Hoffmann (1900) originally assigned the genus to the 

Coreopsidinae, but commented on the ‘yacie Wedeliae.” Neither the description nor a type photograph sh0~7 any basis for placement in the Coreopsidinae, but the habit, the mention of “Antherae nigrae,” and the description of the achenes 

as “uix compressa juniora obscure 2-4-costata, apice 

corona humilisjmbriata coronata, basi in stipitem achaenio proprio dimidio breuiorum contracta ” indicate a 

probable place in the Ecliptinae. 

The Ecliptinae occurs predominantly in the 

Western Hemisphere, where it is well represented 

in both the Nearctic and Neotropical regions. A 

number of separate elements such as Acmella and 

Wollastonia occur naturally in the Eastern Hemisphere. 

Subtribe 16. HELIANTHINAE 

FIGURES 130- 136 

Subtribe HELIANTHINAE Dumort., F1. Belg. Prodr., 71, 1827. 

Subtribe LAGASCEINAE Benth. 8r Hook., Gen. PI., 2:190, 

1873. 

*Agiabampoa Rose ex Hoffm., *Aldama La Llave 

& Lex., *Aluordia Brandeg., *Garcilassa Poepp. & 

Endl., *Helianthopsis H. Robins., *Helianthus L. 

(including Discomela Raf., Echinomeria Nutt., Harpalium Cass., Linsecomia Buckl), *Hehomeris Nutt., 

* Iostephane Benth., *Lagascea Cav., nom. cons. 

(including Calhounia A. Nels., Nocca Cav.), *Pappobolus Blake, * Prionocarpus Blake, * Rhysolepsis 

Blake, *Scalesia Am. (including Zemisne Degener 

& Sherff), *Sclerocarpus Jacq., *Simsia Pers. (including Armania Bertol. ex DC., Barratia Gray & 

Engl.), *Stuessya Turner & Davies, *Syncretocarpus 

Blake, * Tithonia Desf. (including Mirasolia (Sch.- 

Bip.) Benth. & Hook.), * Vzguiera H.B.K. (including Bahiopsis Kellogg, Iiaplocalymma Blake, Hymenostephium Benth., Leighia Cass.). 

Annual or perennial herbs, rarely shrubs. 

Leaves opposite or alternate. Inflorescence with 

heads solitary or sparsely to densely clustered 

(single-flowered heads enclosed in secondary 

heads in Lagascea). Heads usually with ray and 

disk flowers, sometimes discoid; involucre with 

herbaceous bracts in ca. 2 series (more series in 

some Viguiera, fused into single tube in Lagascea); 

paleae usually present (absent in Lagascea), usually strongly conduplicate. Ray flowers sterile; 

limb of corolla with papillose upper surface, tip 

bilobed or obscurely trilobed. Disk flowers hermaphroditic; corolla throat narrowly cylindrical, 

base of throat rounded or bulging with scabrid 

outer surface; lobes usually slightly longer than 

wide, triangular, usually papillose on inner surface; anther thecae black: endothecial cells 

subquadrate or short-oblong, with 1 or 2 thickenings on transverse walls; connective sometimes 

with glands; anther appendage often with glands 

or hairs. Style branches of disk flowers sometimes 

laterally compressed, with single narrow stigmatic 

surface. Ducts with prominent yellowish resin 

(pale resin in Agiabampoa); ducts single or paired 

along veins of disk corolla throat, 2-6 ducts in 

shaft of style outside of veins, usually outside 

veins in style branches but sometimes inside or 

on both sides. Ray achenes sterile, sometimes 

enclosed in involucral bracts (Sclerocarpus, AIdama); disk achenes compressed (terete in Lagascea), walls carbonized, with distinct striations; 

carpopodium of two short, rounded, opposing 

lobes; cells of seed coat not or weakly ornamented; pappus usually of 2 aristae, often with 
56 SMITHSONIAN CONTRIBLTIONS TO BOTANY 

FIGURES 130- 138.-Helianthinae-Neurolaeninae: 130, 13 1, Lagascea moliis Cav. (130, single 

head from cluster showing involucre, X 12; 131. the single achene from the head, X 12): 132, 

Scalesia ufllnis Hook. f., disk achene, X 12; 133, Viguzeru dentutu (Cav.) Spreng.. disk achene, 

X 12: 134. Syncretocarpus sericeus (DC.) Blake, disk achene, X 8: 135. Simsia sanguznea A. Gray, 

disk achene, X 12: 136, Hellanthus annuus L., disk achene, X 12; 137, Neuroiaenu lob& (L.) R. Br., 

disk achene. X 17; 138, Staurochlamys burchellii Baker, disk achene, X 12. 
NCMBER 51 57 

squamellae, sometimes absent. Pollen 27-35 pm 

in diameter. 

The subtribe is almost entirely Nearctic and 

Neotropical in distribution, with the genera centered in Mexico and the United States. A few 

species are widely adventive or cultivated. One 

species of Sclerocarpus apparently is native in Africa. 

The Helianthinae is one of the most concisely 

characterized subtribes in the present treatment. 

All members have sterile rays, single stigmatic 

surfaces on the style branches of the disk flowers, 

ducts outside of the veins in the styles, and striations in the walls of the achenes. Also, the resin in 

the ducts is yellowish in all members except the 

slightly anomalous Agiabampoa. 

The present concept has very little in common 

with the traditional broadly and erratically delimited typical subtribe that has been called variously Helianthinae or Verbesininae. The narrower interpretation of the Helianthinae by 

Stuessy (1977) is an improvement, but remains 

very unnatural, since it includes such genera as 

Balsamorhiza, Encelia, Rudbeckia, and Tetragonotheca, 

while excluding A luordia , Agia bampoa, Lagascea, 

Garcilassa, Scalesia, Aldama, and Sclerocarpus. Of the 

latter series, here placed in the Helianthinae, 

Garcilassa simply never seems to have been critically examined before, and Eliasson (1974) has 

already discussed the relationship of Scalesia of 

the Galapagos. Lagascea and the related Agiabampoa and Alvordia (Stuessy, 1976, 1978) are unusual 

in the subtribe only by specializations toward 

compound heads and by the presence of resin 

ducts inside the veins of the style branches. Aldama 

has recently been resegregated from Sclerocarpus 

(Feddema, 1971), but both share all characters of 

the subtribe Helianthinae and must be rather 

closely related. 

The Encelia group is excluded here, although 

its members closely resemble some genera of the 

Helianthinae such as Simsia. The traditional 

treatments prior to the study by Blake (1913) did 

not separate Sirnsia and Encelia generically. The 

Encelia group has sterile rays, single stigmatic 

surfaces on the style branches, and chromosome 

numbers often n = 17, as in the Helianthinae. 

However, the Encelza group has colorless ducts, 

lacks dehydrofalcarinone, lacks scabrosity on the 

base of the disk corolla throat, and has nonstriate 

achene walls, all of which indicate a position in 

the Ecliptinae. The aristae of the achenes even 

seem different in basic structure, being more of 

an extension of the body of the achene, and not 

a sharply defined or deciduous structure as in the 

Helianthinae. 

The Helianthinae seems to be more closely 

related to the Galinsoginae. The two subtribes 

share similar systems of resin ducts, striate achene 

walls, and polyacetylene chemistry. Both have 

dehydrofalcarinone, which has been reported 

elsewhere in the tribe only in Ioa of the Ambrosiinae (Bohlmann et al., 19731, and both seem to 

lack the thiophenes that are common in other 

subtribes such as the Ecliptinae and Rudbeckiinae. The Helianthinae are readily distinguished 

from the Galinsoginae by the sterile rays, the 

basically compressed achenes and bilateral pappus, and the single stigmatic surface on the style 

branches. In the Helianthinae, the carpopodium 

is a short beaklike pair of lobes that seem to bite 

the achene free at the base as it matures, and 

there is no structure that approaches the specialized, asymmetric, sclerified plate that is characteristic of the Galinsoginae. As a final distinction, 

the resin ducts of the styles usually are completely 

outside of the veins in the Helianthinae and 

usually are completely inside the veins or collateral with them in the Galinsoginae. In rare cases 

such as Lagascea and Tridax, members of both 

subtribes achieve an intermediate condition, with 

ducts outside the veins in the style shaft and 

inside the veins in the branches. 

Most members of the Helianthinae differ from 

the Galinsoginae by having solitary resin ducts 

along the veins of the disk corolla throat. There 

is a tendency for multiple ducts in the style and 

paired ducts in the corolla throat in the Hymenostephium and Haplocalymma parts of Vzguzera. Supplementary resin ducts reach their extreme in 

species like V. sylvatica Klatt of Costa Rica and in 

the related genus Garcilassa. 
58 SMITHSONIAN CONTRIBUTIONS TO BOTASY 

Most members of the subtribe, including the 

genus Viguiera, have a chromosome number of 

X = 17. Recently, the genus Heliomeris has been 

resurrected from the synonymy of Viguiera for a 

series of species having X = 8 (Yates and Heiser, 

1979). Hehomeris of the western United States and 

northern Mexico is scarcely distinct in any other 

way. The exappendiculate style branches differ 

from those of typical Viguiera, but many species 

presently retained in Viguiera have exappendiculate styles. It is therefore significant that Heliomeris 

has a unique pattern of resin ducts in the styles: 

the ducts are prominent in, but restricted to, the 

branches. The termination of the duct is usually 

abrupt, and no ducts are evident in the shaft of 

the style. It is also notable that the chromosome 

number is n = 17, the style is long-appendiculate, 

and the duct is not restricted to the style branches 

in Viguiera porteri (A. Gray) Blake, a species of the 

southeastern United States that was placed in 

Viguiera sect. Heliomeris by Blake (1918). 

Subtribe 17. NEUROLAENINAE 

FIGURES 52, 137-143 

Subtribe NEUROLAENINAE (Rydb.) Stuessy, Turner & Powell, 

Tribe NEUROLAENEAE Rydb., N. Amer. FI., 34(4):303, 1927. 

Biol. Chem. Compositae, 645, 1977 [ 19781. 

*Brasilia Barroso, *Calea L. (including Amphzcalea (DC.) Gardn., Caleacte R. Br., nom. provis., 

Calebrachys Cass., Geissopappus Benth., Leontophthalmum Willd., Meyeria DC., Mocinna Lag.), *Greenmaniella Sharp, *Neurolaena R. Br., *Staurochlamys 

Baker, * Tetrachyron Schlect., * Tyleropappus 

Greenm., *Unxia L. f. (including Greenmania Hieron., Pronacron Cass.) . 

Perennial herbs or shrubs; leaves alternate, opposite, or whorled. Inflorescence usually terminal, 

usually paniculate, sometimes reduced to single 

head. Heads heterogamous or discoid; involucre 

of partially herbaceous to completely coriaceous 

bracts, often in 4 or more series; paleae usually 

present. Ray flowers female; upper surfaces of 

limb with oblong scarcely mamillose cells, usually 

distinctly trilobed at tip (reduced and unlobed in 

some Unxia). Disk flowers usually hermaphroditic, functionally male in Cizxia; corolla lobes 

short (in some Neurolaena) or long (in Calea 

group and some Neurolaena), cells of inner surface 

oblong and scarcely bulging: anther thecae black 

(in Neurolaena and Staurochlamys) or yellow (in 

Brasilia, Calea, Tyleropappus and C'nxia) ; endothecia1 cells with 1-3 thickenings on transverse walls; 

anther appendage ovate, with or without glands 

abaxially. Branches ofstyles with paired stigmatic 

lines. Ducts of corollas usually with prominent 

reddish resin (lacking in typical Neurolaena) , ducts 

solitary along veins of disk corolla throat, along 

margins in lobes, ducts in shaft of style inside of 

veins. Achenes prismatic, walls carbonized, without striations; cells of seed coat weakly ornamented; pappus radially symmetrical, usually of 

numerous bristles or squamellae (lacking in 

Unxia). Pollen 25-35 pm in diameter (to 55 pm 

in Calea belemii H. Robins.). 

Traditionally Neurolaena has been placed with 

the genus Schistocarpha in the tribe Senecioneae, 

apparently being excluded from the Heliantheae 

because of its well-developed pappus of numerous 

capillary bristles. The paleae in the two genera 

are in conflict with the epaleaceous condition of 

the true Senecioneae and are not homologous to 

the form of chaff found in the often associated 

Liabeae. Such paleae were undoubtedly a factor 

in the unexplained creation of a separate tribe by 

Rydberg (1927). In transferring the two genera 

to the Heliantheae, Robinson and Brettell 

(1973b) first noted basic differences in the achene 

wall that could indicate that Neurolaena and Schistocarpha were not each other's closest relatives. In 

formally adopting the subtribe in the Heliantheae, Stuessy (1977) included an assortment of 

genera in addition to Neurolaena and Schistocarpha, 

genera as diverse as Bebbia, Clappia, Dyscritothamnus, Greenmaniella, Varilla, and Zalurania. The 

group parallelled in part the artificial grouping 

included at the same time by Turner and Powell 

(1977) in their Varillinae. Though many similar 

genera are included by Stuessy, Tridax is left in 

the Galinsoginae, perhaps because of its more 

herbaceous habit. 
NUMBER il 59 

FIGURES 139- 149.-Neurolaeninae-Galinsoginae: 139, Calea g’amaicensis L., disk achene, X 12; 

140. Calea (Geissopappus) caleoides (DC.) H. Robins., disk achene, X 12: 141, Brasilia sickii 

Barroso. achene, X 12; 142, 143, Tetrachyron orizabensis Sch.-Bip. ex Klatt. (142, ray achene. 

X 12: 143, disk achene, X 12); 144, Tetragonotheca texana Gray X Engelm.. achene. X 12: 145, 

Galinsoga quadriradiata R. & P., disk achene, X 17: 146, Jaegeria hirta (Lag.) Less., disk achene. 

X 24: 147. Schistocarpha eupatorioides (Fenzl) 0. Kuntze, disk achene, X 17: 148, 149, Alloispermum 

caracasanum (H.B.K.) H. Robins (148, ray achene, X 8; 149, disk achene. X 12). 
60 SMITHSONAN CONTRIBUTIONS TO BOTANY 

Of the genera included by Stuessy (1977), only 

.Veurolaena and Greenmaniella are retained here. 

Most confusion seems to have involved genera 

placed here in the Galinsoginae such as Schistocarpha, Bebbia, Dyscritothamnus, and Alloispermum, the 

last resurrected from the synonymy of Calea. Both 

subtribes share prismatic achenes, a radially arranged pappus, colored resin, a position of the 

resin duct inside the veins in the shaft of the style, 

and usually a subimbricate involucre. The Neurolaeninae differ by the resin being reddish, the 

ducts being solitary along the veins of the disk 

corolla throat, the ducts being marginal in the 

corolla lobes, by the nearly smooth adaxial surfaces of the corollas, by the generally longer lobes 

of the disk corollas, and by the achenes lacking 

striations. The two subtribes share a similar asymmetric form of carpopodium on the achene, but 

in the Neurolaeninae, the inner cushion has 

firmer cells that don’t collapse at maturity. 

Chemically the Neurolaeninae lack the dehydrofalcarinone polyacetylenes that are characteristic 

of the Galinsoginae, but have thymol derivatives 

in both *Veurolaena and Calea. Thymol derivatives 

are not presently known from any other paleaceous subtribe in the Heliantheae (Robinson et 

al., 1978). 

The achene wall structure and chemistry of the 

Neurolaeninae are closer to the Ecliptinae than 

the Galinsoginae, but the prismatic achenes, radial pappus, and frequent reddish resin are signifciant distinctions. The Neurolaeninae are not 

so obviously a natural subtribe as the Galinsoginae. Staurochlamys is particularly distinct in its 

lack of a subimbricate involucre. Neurolaena also 

is distinct from the Calea group by its capillary 

pappus, dark anther thecae, alternate leaves, and 

the inconsistent presence of reddish resin. 

In the course of this study, the genus Calea has 

been redelimited. Allotspermum has been resurrected from synonymy and placed in the Galinsoginae (Robinson, 1978d), and C. skutchii Blake 

has been transferred to Podachaenium in the Ecliptinae (Robinson, 1978~). 

Tetrachyron is held separate from Calea, but it is 

similar to that genus in the position of the style 

ducts, the prismatic achenes without striations, 

and the radial (4-parted) pappus. Characters that 

make placement in the Neurolaeninae uncertain 

are the lack of complete differentiation of the 

inner surface of the carpopodium and the sometimes double ducts along the veins of the disk 

corolla throat. The genus has been monographed 

by Wussow and Urbatsch (1979). 

Some membes of the Calea group show reduction in paleae or suppression of disk flower gynoecia. Epaleaceous species of Calea were treated 

as a separate genus Geissopappus by Bentham and 

placed in the tribe Helenieae in Bentham and 

Hooker (1873). The genus Unxia also has paleae 

of reduced size, but it is most notable for the 

functionally male disk flowers. The styles of the 

disk flowers are completely sterile, lacking either 

branches or stigmatic papillae. The reductions in 

the two genera are totally independent of similar 

reductions elsewhere in the Heliantheae. 

The subtribe is restricted to the Neotropical 

region. 

Subtribe 18. GALINSOCINAE 

FIGURES 144- 152 

Subtribe GALINSOCINAE Benth. & Hook., Gen. PI., 2:198, 

1873. 

* Alepidocline Blake, *Alloispermum Willd. (including Allocarpus H.B.K., Calydermos Lag.), 

*Aphanactis Wedd., * Bebbia Greene, *Cuchumatanea 

Seid. & Beaman, *Cymophora B. L. Robins., *Dyscritothamnus B. L. Robins., *Faxonia Brandeg., 

* Galinsoga Ruiz & Pavon (including Aduentina 

Raf., Stemmatella Wedd. ex Benth. & Hook., Stenocarpha Blake, Vargasia DC., non Bertol. ex 

Spreng., Vasargia Steud., Vigolina Poir., Wiborgia 

Roth, nom. rej., non Thunb.), *Jaegeria H.B.K. 

(including Aganippea Sesse & Moc. ex DC., Heliogenes Benth., Macella Koch), *Otetza La Llave, 

*Sabazia Cass. (including Bariasa Steud., Tricarpha 

Longpre) , *Schistocarpha Less. (including Neilreichia Fenzl, Zycona 0. Kuntze), *Selloa H.B.K., 

nom. cons., non Spreng. (including Feaea Spreng., 

Feaella Blake), * Tetragonotheca L. (including Bikera 
NUMBER 51 61 

Adans., Gonotheca Raf,, Halea Torr. & Gray, Tetragonosperma Scheele), * Tridax L. (including Balbisia 

Willd., nom. rej., non Cav., Bartolina Adans., Carphostephium cass., Mandonia Wedd., Ptilostephium 

H.B.K., Sogalgina Cass.). 

Annual or perennial herbs, occasionally shrubs. 

Leaves usually opposite, sometimes in basal rosette. Heads one to few on short to long peduncles, 

or in corymbose usually terminal panicles. Heads 

usually heterogamous; involucre usually subimbricate, bracts in 2-5 series, coriaceous or sometimes herbaceous; paleae present (rare in Alepidocline), sometimes deeply trifid (in Faxonia, some 

Sabazia, some Schistocarpha). Ray flowers female; 

limb of corolla papillose on upper surface, usually 

white, usually trilobed at tip. Disk flowers hermaphroditic; lobes of corolla short-triangular, 

mamillose or papillose on inner surface; anther 

thecae rather pale to slightly darkened; endothecia1 cells subquadrate or oblong, with 1-3 thickenings on transverse walls (thickenings on vertical 

walls in a few Tridax); anther appendages ovate, 

with or without glands. Style branches with stigmatic lines paired. Ducts with yellowish resin, 

ducts usually paired along veins of disk corolla 

throat, on inner side of veins in lobes and often 

remote from margin, ducts from adjacent lobes 

not fused at sinus; usually with two ducts in style 

inside of veins (4 ducts collateral to 4 veins in 

style shaft of Oteiza, Schistocarpha, and some Tridax; 

4 ducts outside of 2 veins in Tridax brachylepis 

Hemsl.). Achenes prismatic, walls carbonized, 

with striations: carpopodium asymmetrical, with 

hardened plate on outer side, with cushion of soft, 

collapsing cells on inner side; cells of seed coat 

irregular with sinuous walls, often strongly ornamented; apex of achene with distinct callus usually bearing radially disposed bristles or squamellae (plumose bristles in Bebbia and Tridax), 

pappus sometimes lacking (lacking on ray achenes and present in disk in Alloispermum, Faxonia, 

and some Sabazia). Pollen 23-35 pm in diameter 

(smaller and apparently nonfunctional in Faxonia). 

The subtribe was established by Bentham and 

Hooker (1873) for members of the Heliantheae 

having a terete or prismatic achene bearing numerous pappus elements. The listing of genera 

included some extraneous genera such as Balduina, 

Dubautia, and Marshallia, while close relatives of 

Galinsoga such as Jaegeria and Sabazia were excluded. The only member of the Neurolaeninae 

included was Calea, but the genus Neurolaena itself 

was excluded from the tribe. An improved concept is evident in the recent publications of 

Stuessy (1977) and Canne (1978), but the definition remains imprecise. In his key to subtribes, 

Stuessy emphasizes the conical receptacle and 

usually translucent anthers, characters that are, 

at best, unreliable. Steussy includes Spilanthes of 

the Ecliptinae and Zsocarpha of the tribe Eupatorieae, but excludes Bebbia and Schistocarpha which 

are placed in his extremely artificial subtribe 

Neurolaeninae. Most recently, natural limits have 

been recognized between Galinsoginae and the 

Neurolaeninae (Robinson, 1979b), and Schistocarpha has been transferred to the former, while the 

typical element of Calea has been transferred to 

the Neurolaeninae. 

The Galinsoginae is regarded here as a totally 

natural group of Neotropical genera which is 

represented in the Eastern Hemisphere only by 

adventive species of Galinsoga and Tridax. The 

subtribe shares many characters with the Neurolaeninae, but it seems most closely related to the 

Helianthinae with the characters discussed under 

that subtribe. The extensive development of ducts 

in the flowers of both subtribes with yellow resin 

and dehydrofalcarinone polyacetylenes is regarded as particularly significant. 

The prismatic achenes, radially symmetrical 

pappus, and often subimbricate involucre of the 

Galinsoginae conform with states held in this 

study to be primitive in the tribe. In contrast, the 

reduced, often annual habit of the plants seems 

more derived. The low chromosome numbers in 

the group, where base numbers over X = 10 are 

lacking, may be a reflection of the trend toward 

the short-lived perennial or annual habit, but it 

could be a relict of the X = 10 that is believed to 

be basic for the subfamily Asteroideae (Robinson, 

et al., in press). The distinctive, shrubby Dyscri- 
62 

SMMITHSONIAN COKTRIBUTIONS TO BOTANY 

PICURES 150- 156.-Galinsoginae-Dimeresiinae-Coreopsidinae: 150, Bebbia juncea (Benth.) 

Greene, achene, X 12; 15 1, Tridax procumbens L., disk achene, X 12; 152, Dyscritothamnusfilifolius 

B. L. Robins.. achene, X 8; 153, Dimeresia howellti A. Gray., achene, X 12; 154, 155. Coreopsis 

lancedata L., disk achenes, X 12 (154, inner surface; 155, outer surface): 156, Bidens pzlosa L., 

disk achene. X 7. 
NUMBER il 63 

tothamnus with its achenes densely covered with 

“zwillingshaare” is the most divergent element in 

the subtribe, and may represent a more ancestral 

habit A chromosome report for the latter would 

be very instructive. 

Limited observations of the surfaces of the 

ovule indicate potentially useful variations in the 

subtribe. The cells are irregularly shaped and 

annulately ornamented in Schzstocarpha, in at least 

one Trzdax, and to a lesser extent in Sabazia lzebmannzz Klatt. The cells in the equivalent layer in 

Allozspermum and Sabatia acoma (Blake) Longpre 

have nearly straight walls and no annulate ornamentation. The cells of Selloa are irregular in 

shape and mostly inornate. In Galinsoga jilformzs 

Hems1 , the cells are elongate with closely sinuous 

walls, and the cells seen in Jaegerza are elongate 

with nonsinuous inornate walls. 

Through the kindness of Drs. Ornduff and 

Strother at the University of California at Berkeley, it has been possible to make a microscopic 

examination of the sparing material of Faxonza 

puszlla Brandeg., an endemic of southern Baja 

California which is known only from the type 

specimen The immature but well-formed achenes lack carbonization, but they show the correlated punctations in a striated pattern characteristic of the Galinsoginae. The carpopodium of 

the ray achenes, the double or even triple resin 

ducts along the veins of the disk corolla throat, 

and the ducts inside the veins of the style seem to 

confirm this placement. The trifid paleae of the 

receptacle and the pappus restricted to the disk 

achenes find their equal in various other genera 

of the subtribe. The ray corollas of Faxonza are not 

tubular as represented by Brandegee (1894), but 

are mostly reduced to a filiform limb. The leaves 

are long-petiolate and mostly narrow-bladed, as 

originally described. 

Some unique features of Faxonza tend to indicate an apomictic condition. No stigmatic papillae have been seen on either the ray or disk 

styles, and the anthers are very reduced and 

unopened with thin-walled, mostly slightly distorted pollen. The apparently nonfunctional pollen grains are unique in their size, usually ca. 15 

pm in diameter, although sometimes varying to 

ca. 20 pm. Grains smaller than 23 pm have been 

noted otherwise in the Asteraceae only in the 

tribe Eupatorieae. The seemingly defective nature of Faxonia might suggest a hybrid origin, but 

sources for such a hybrid are not obvious and 

recent origin is not likely. Faxonia seems to be an 

extremely reduced ephemeral. I suspect a greatly 

reduced chromosome number with some loss of 

essential genetic material. The genus must have 

persisted for a considerable time strictly through 

apomixis. 

Subtribe 19. DIMERESIINAE 

FIGURE 153 

Subtribe DIMERESIINA~ H. Robinson. Phytologia. 11 40. 

1978. 

* Dzmeresza A. Gray (including Eremznula 

Greene) . 

Annual herbs with leaves in sessile rosette, 

petioles scarcely distinct. Heads aggregated in a 

sessile cluster, discoid: involucre of 2 or 3 herbaceous bracts; paleae absent. Flowers 2 or 3 per 

head, disciform, hermaphroditic: corolla reddish; 

throat narrowly cylindrical, sparsely glanduliferous outside; lobes 5, ovate, strongly papillose 

inside, glabrous outside; anther thecae pale: median endothecial cells elongate, elliptical, with 

single thickenings at ends; anther appendages 

ovate, with glands abaxially. Base of style with 

distinct node; style branches with paired stigmatic lines, with papillae abaxially nearly to base. 

Resin ducts not evident, without colored resin. 

Achenes terete, walls carbonized and appearing 

castaneous, with striations represented by regular 

grooves; carpopodium indistinct; cells of seed coat 

with strongly sinuous walls, slightly ornamented 

on surfaces; pappus deciduous as a unit, segments 

united at base, basal sheath of pappus folded 

back on itself, segments densely fringed above 

with hairlike cells. Pollen ca. 30 pm in diameter. 

The subtribe contains a single species, Dzmeresza 

howellii A. Gray, from the western United States. 

The plants are easily distinguished by the few- 
64 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

flowered heads aggregated into a sessile cluster in 

a basal rosette of leaves. The basally fused pappus, which falls as a unit, also is distinctive. 

Gray (1886) described the anthers as “caudatosagittatae” at the base and placed the genus in 

the Inuleae, where it remained in the treatment 

by Hoffmann (1890-1894). Actually, the anthers 

are scarcely longer at the base than many other 

members of the Heliantheae, and the tips are not 

caudate. The carbonized and striate achenes confirm the helianthean nature of the genus. 

The lack of paleae is correlated with the reduced number of flowers in the head, and its does 

not seem to reflect a position near the epaleaceous 

series of subtribes in the Heliantheae. The achene 

walls of Dimeresia lack the knoblike thickenings in 

the carbonization that are present in most epaleaceous subtribes. The combination of achene wall 

structure, strongly papillose corolla lobes, and 

pappus indicates a position near the Helianthinae 

and Galinsoginae, and the terete achenes, paired 

stigmatic lines, and reduced annual habit are 

most like the latter. The lack of distinct resin 

ducts and the obsolete carpopodium furnish a 

technical basis for distinction of the Dimeresiinae, 

but numerous other subtle details such as the 

difference in endothecial cells and texture of the 

pappus suggest greater phyletic distance from the 

Galinsoginae and possible independent development of the reduced habit. 

Subtribe 20. COREOPSIDINAE 

FIGL~RES 154-16 1 

Subtribe COREOPSIDINAE Less., Linnaea, 5: 153, 1830. 

Family COREOPSIDACEAE Link. Handbuch, 1 :768, 1829. 

Subtribe PETROBIINAE Benth. & Hook., Gen. PI., 2: 192, 1873. 

Tribe COREOPSIDEAE Turner & Powell. Biol. Chem. Compositae, 724. 1977 [1978]. 

* Bidens L. (including Campylotheca Cass., Delucia 

DC., Diodonta Nutt., Kerneria Moench.), *Chrysanthellurn Rich. (including Adenospermum Hook. & 

Arn., Chrysanthellina Cass., Collaea Spreng., non 

DC., Neuractis Cass., Sebastiania Bertol.), *Coreocarpus Benth. (including Acoma Benth.), *Coreopsis L. 

(including Agarista DC., non Don, Anacis Schrank, 

Calliopsis Reic h b . , Chrysornelea Tausc h, Chrysostemma Less., Coreopsoides Moench., Diplosastera 

Tausch, Electra DC., non Panzer, Epilepsis Benth., 

Leachia Cass., Selleophytum Urb.) , *Cosmos Cav. 

(including Adenolepsis Less.), * Cyathomone Blake, 

*Dahlia Cav. (including Georgina Willd.), *Dicranocarpus A. Gray, *Ericentrodea Blake & Sherff, *Eryngiophyllum Greenm., *Glossocardia Cass., *Glossogyne Cass. (including Diodontium F. Muell., Gynactis Cass.), * Goldmanella Greenm.? (including 

Goldmania Greenm., non Rose ex Micheli), *Guerreroia Merr., * Henricksonia Turner, * Heterosperma 

Cav. (including Microdonta Nutt.), *Hidalgos La 

Llave & Lex., *Isostigma Less., *Megalodonta 

Greene, *Microlecane Sch.-Bip., *Moonia Arn. (including Pentalepsis F. Muell.), *Narualina Cass. 

(including Needhamia Cass., non Scop.), *Oparanthus Sherff, * Petrobium R. Br. (including Laxmannia 

Forst. & Forst., nom. rej., non R. Br.), *Thefesperma Less. (including Cosmidium Nutt.). 

Annual and perennial herbs, shrubs, and small 

trees. Leaves usually opposite, usually with dissected blades (alternate with asymmetrical, simple blades in Goldmanella). Heads usually in lax, 

corymbose or cymose panicles often solitary on 

long peduncles, not or rarely congested. Heads 

usually heterogamous, gynoecia sometimes suppressed in ray flowers or in disk flowers, plants 

tending to be dioecious in Petrobium; involucre 

usually with distinct outer series differing in structure and color: paleae present, scarious, not conduplicate. Flowers usually yellow, sometimes purple, reddish, or whitish. Rays usually present and 

female, sometimes sterile or lacking; limb of corollas papillose on upper surface, often prominently trilobed at apex. Disk flowers usually hermaphroditic, sometimes functionally male (Dicranocarpus, Hidalgoa, Moonia) : corolla lobes 

slightly to distinctly longer than wide, cells of 

inner surface short, smooth to mamillose or papillose, outer surface often with short, sharp hairs 

or spicules; filaments usually smooth, lower part 

hirsute in Cosmos; anther thecae usually slightly 

to strongly darkened; endothecial cells short, usually with 2-4 thickenings on transverse walls (ra- 
NUMBER .il 65 

dially thickened in Dahlia); anther appendage 

usually ovate (very short and truncate in Hzdalgoa), without glands, often with central duct containing colored resin. Styles usually branched, 

disk styles unbranched in some genera (such as 

Dicranocarpus and Hidalgoa) , branches of fertile 

styles with stigmatic lines usually paired, sometimes with lines indistinct or fused. Ducts commonly with reddish resin; ducts usually prominent in inner involucral bracts, often present in 

paleae and achenes; ducts of disk corolla throat 

usually paired along veins, usually outside of 

veins in styles (apparently inside in unbranched 

disk style of Dicranocarpus); isolated pocket of 

colored resin often prominent in anther appendage. Achenes of both ray and disk flowers obcompressed, sometimes winged on lateral margins 

(Coreopsis, Heterosperma), achenes when all fertile 

sometimes dimorphic or heteromorphic (Heterosperma, some Bidens) walls carbonized, basically 

without striations (broad, uncarbonized bands 

present in some Bidens), carbonization showing 

additional pattern of knoblike thickenings; cells 

of seed coat not ornamented, with straight somewhat thickened walls; pappus usually present, 

often of a pair of lobes or aristae lateral to the 

corolla, rarely polysetose (Ericentrodea) or bearing 

4 paleaceous squamellae (on disk achenes of Henrzcksonia). aristate forms often strongly retrorsely 

barbellate (many Bidens). Pollen 25-35 pm in 

diameter. 

The Coreopsidinae and the Fitchiinae form a 

natural group that is widely distributed pantropically. No other element of the tribe is as well 

represented in the Eastern Hemisphere. The 

number of genera on isolated islands suggests that 

the wide distribution is correlated with superiority of long-range dispersal mechanisms; probably 

it is not indicative of greater age for the group. 

The Coreopsidinae and Fitchiinae can be distinguished as a group from all other subtribes of 

the Heliantheae by the obcompressed achenes of 

both the ray and disk flowers. Some problems are 

possible in placing genera with sterile disk achenes, but some other characters are available. The 

habit with opposed dissected leaves, the numerous prominent resin ducts in the inner involucral 

bracts, and the differentiated outer involucre usually provide a distinctive appearance. Ducts in 

the anther appendages and achenes with prominent reddish resin are not known in any other 

subtribe. The frequent presence of a reddish duct 

and complete lack of glands on the anther appendages of the Coreopsidinae has been noted by 

Robinson and Brettell (1973a). The stigmatic 

surface in the subtribe seems ancestrally separated into a pair of lines though these are not 

distinct in some of the members. The carbonization in the achene wall is basically nonstriate, 

although a specialized form of broad banding has 

been seen in Bidens, which may or may not be 

homologous to striations seen in other subtribes 

of the Heliantheae. 

The Coreopsidinae traditionally have contained a large core of closely related genera, but 

the actual limits have been unnatural because of 

some extraneous genera that have been included 

as well as some relatives that have been excluded. 

Some improvements are seen in the recent survey 

by Stuessy (1977); the changes in that study can 

be evaluated as follows. An extraneous element 

that includes Cabptocarpus and Synedrella. having 

obcompressed ray achenes but prismatic disk achenes, is correctly transferred to the Ecliptinae by 

Stuessy. Guizotia, which was transferred to the 

Verbesininae by Baago e (1974). was returned to 

the Coreopsidinae by Stuessy. The terete, striate 

achenes, the ornamented seed coats, and the glanduliferous anther appendages all dictate against 

inclusion of Guizotia, and that genus is placed in 

the Milleriinae in this study. Of the genera included in the Coreopsidinae for the first time by 

Steussy (1973, 1977), Dicranocarpus of the western 

United States and Mexico, Moonia of the Indian 

Ocean region: and Petrobium of St. Helena are 

unquestionably related, but Jaumea, Guardiola, 

Staurochlamys, and Venegasia are not. Jaumea is 

placed here in the Jaumeinae and Venegasia in the 

Chaenactidinae. Guardiola differs from the Coreopsidinae by the strongly annulate, elongate cells 

of the seed coat, the striate achenes, and the 

distinctive form of large gland at the base of the 
66 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

FIGIJRES 157~166.-Coreopsidinae-Fitchiinae-Coulterellinae-Pectidinae: 157, 158, Heterosperma 

pinnatum Cav.. achenes, X 12 (157, central achene; 133. outer achene): 159. 160, Dahlia coccinea 

Cav. (159, disk achene. X 6; 160, disk achene with palea, X 6): 161. Hidalgoa ternata La Llave 

& Lex., ray achene. X 8; 162, Fitchia nutans Hook. f., achene. X 6: 163, 164, Coulterella capztata 

L'asey & Rose (163, head with fused involucre and flower, X 12: 164, the single achene from the 

head. X 12): 165, 166. Tagetes subulata Cerv. ex La Llave 8r Lex., achenes. X 8. 
NUMBER il 67 

anther appendages. Guardiola has nodular thickenings on the lateral walls of the subquadrate 

median endothecial cells, a precise pattern not 

duplicated elsewhere in the tribe. Guardiola is 

placed here in a separate subtribe Guardiolinae. 

Staurochlamys is rejected from the Coreopsidinae 

because of the glands on the anther appendages, 

the prismatic rather than obcompressed achenes, 

and the totally dissimilar differentiated outer involucral bracts. The genus is placed in the 

Neurolaeninae in this study. 

Certain features of the monotypic Goldmanella 

of Central America have caused both Stuessy and 

myself to place the genus in the Coreopsidinae. 

There are numerous resin ducts in the involucral 

bracts, and the achenes are slightly broadened. 

Still, the alternate leaves with broad, simple, 

asymmetrical blades and the subimbricate involucre make the genus thoroughly anomalous in 

the subtribe. 

Turner and Powell (1977) have proposed a 

tribe Coreopsideae containing various elements 

included in the Coreopsidinae by Stuessy as well 

as Fitchia and elements such as Jaumea from the 

Jaumeinae. It was a second instance where these 

authors proposed a group including both paleaceous and epaleaceous genera. Their treatment is 

not supported by the evidence in this study. Part 

of the difficulty arises from the failure of Turner 

and Powell to either describe their tribe or to 

indicate any of its points of difference from the 

Heliantheae or even the Senecioneae. One of 

their reasons for recognizing a separate tribe apparently was the differences of eight amino acids 

in the sequences of the cytochrome c of Helianthus 

and Guizotia. This is not really relevant, since 

Guizotia is not a member of the Coreopsidinae. 

No amino acid sequences of either cytochrome 

c or plastocyanin have been determined for any 

member of the Coreopsidinae as presently delimited, but a number of other types of chemical 

data are available. 

In reference to the flower color of the Asteraceae, Cronquist (1955) stated “that the yellow 

pigments are partly plastid-borne carotinoids and 

partly water-soluble anthoxanthins chemically 

allied to the anthocyanins, and that several yellow 

pigments including both carotinoids and anthoxanthins may be present in a single flower.” The 

particular group of anthoxanthins of interest in 

the 1955 paper were the anthochlors or aurone 

and chalcone flavonoids. According to Cronquist , anthochlors had been found in nearly 200 species 

of the Coreopsidinae, and except for several species of Baeria (Lasthenia) they were not known 

elsewhere in the family. Cronquist cited Gertz 

(1938), Geissman (1941a,b, 1942), and Geissman 

and Heaton (1943, 1944). The presence of anthochlors in other families seemed possible. Regarding the anthochlors, Cronquist said, “Within 

the Compositae, their very wide distribution in 

and usual restriction to the Coreopsidinae furnish 

further evidence of the naturalness of that subtribe.” 

Much refinement has occurred in the knowledge of anthochlors since 1955. Three papers in 

the Symposium on the Biology and Chemistry of 

the Compositae mentioned the compounds (Harborne, 1977; Heywood et al., 1977; and Bohm, 

1977), and an extensive study has been published 

by Crawford and Stuessy (1981). Both types of 

anthochlors prove to occur in both subfamilies, 

with the phloroglucinol-based anthochlors in the 

Cardueae (Carthamus), Eupatorieae (Chromokaena) 

and Inuleae (Gnaphalium, Helichrysum) and resorcinol-based anthochlors in the Lactuceae (Cabcoseris, Malacothrix, Pyrrhopappus) and the 

Heliantheae. In the Heliantheae, anthochlors occur in two epaleaceous genera, Lasthenia and Syntrichopappus, and in four genera of the Helianthinae, Helianthus, Simsia, Tithonia and Viguiera, in 

addition to various genera that have been placed 

in the Coreopsidinae. In the Crawford and 

Stuessy (1981) study, all 25 genera tested that are 

placed in the Coreopsidinae by both Stuessy 

(1977) and the present treatment proved to have 

anthochlors. It is notable that two genera not 

recognized here as members of the subtribe, Guardiola and Venegasia, lack anthochlors. The genera 

Staurochlamys and Guizotia, included in the Coreopsidinae by Stuessy (1977) but placed in the Neurolaeninae and Milleriinae respectively in the 
SMITHSONIAK CONTRIBUTIONS TO BOTANY 

present treatment, apparently have anthochlors. 

The mutual occurrence of anthochlors in the 

Coreopsidinae and in Lasthenia is paralleled by 

the apparent restriction of thiophene forms of the 

ene-tetrayne-ene polyacetylenes to these same two 

groups in the Heliantheae. For this reason Bohm 

(1977) safely concluded: ‘‘. . . on the basis of anthochlor and acetylene chemistry Lasthenia would 

be judged to lie closer to Coreopsidinae than to 

Senecioneae . . . .” There seems to be no basis, 

however, for Harborne’s (1977) comment, “Recent work . . . has indicated that Lasthenia and 

Syntrichopappus (Helenieae) could well be accommodated on morphological grounds in the Coreopsidinae (Heliantheae) . . . .” 

The Coreopsidinae seem to have a greater diversity of polyacetylenes than any other subtribe 

of the Heliantheae. Involved are both of the 

major pentaynene and ene-tetrayne-ene pathways with various derived thiophenes. Aromatic 

endgroups are present in the acetylenes of many 

Coreopsidinae, formed on either end of the molecule. Such aromatic acetylenes are found elsewhere in the tribe only in Heliopsis of the Ecliptinae. Unique aromatic thiophenes are found in 

Coreopsis and Bidens. Rarer types of polyacetylenes 

found in Dahlia include a tetrahydropyrane like 

that in Zchthyothere and the only examples in the 

Heliantheae of CIS acetylenes lacking methyltriyne endgroups. 

The Kranz syndrome or C4-phostosynthetic 

pathway provides more efficient use of ambient 

COz at higher temperature and higher light intensity than the common CS pathway, and it 

seems to confer advantages to some plants in hot 

or exposed environments. Occurrence of the character is somewhat erratic among higher plants. 

The pathway was orginally discovered in sugar 

cane, and it is found in various other grasses 

including part but not all of the genus Panicum. 

In the chenopodiaceous genus Atriplex, a species 

with the Cq pathway forms hybrids with a species 

lacking the trait. The distribution of the syndrome in the Asteraceae has been reviewed by 

Smith and Turner (1975) using isotope ratios of 

COZ from combustion of fragments of herbarium 

material. Tests of various Astereae and Heliantheae revealed Cq ratios in five genera of the 

Coreopsidinae, Chrysanthellum, Eryngiophyllum, 

Glossocardia, Glossogyne, and Isostigma, four of 12 

species tested of Flaveria in the Flaveriinae, and 

in one genus, Pectis, of the Pectidinae. The lack of 

any special relationships between the three subtribes, and the numerous members of all three 

subtribes that lack the Kranz syndrome, indicate 

that the syndrome is independently 

each group. Even within the groups 

character is probably monophyletic, 

be reversions or complications from 

tion. 

derived in 

where the 

there may 

hybridizaSubtribe 21. FITCHIINAE 

FICI:RE 162 

Subtribe FITCHIINAE Carlquist. Univ. Calif. Publ. Bot.. 29:5, 

1957 

*Fztchia Hook. f. 

Shrubs or trees, glabrous except in leaf axils. 

Leaves opposite, with distinct petioles; blades 

simple, entire or with one to several teeth. Heads 

campanulate, terminal, solitary or rarely in 

groups of two or three; peduncle recurved; heads 

homogamous; involucre multiseriate, subimbricate, bracts coriaceous to woody: paleae present, 

not conduplicate. Flowers hermaphroditic; corollas ligulate with 5 narrow lobes: adaxial surface 

of lobes mostly smooth, tips with laciniate crests 

outside, each lobe with 3 or more veins; anther 

thecae pale: endothecial cells short, with 2 or 3 

thickenings on transverse walls; anther appendages lanceolate, nearly flat, w’ithout glands. Style 

branches short, with continuous stigmatic surface. 

Ducts or secretory canals numerous in involucre, 

paleae, and flowers, resin not obviously colored; 

ducts of achene numerous, accompanying and 

slightly interior to veins, usually not reaching 

base or tip of achene, short ducts often present in 

awns (absent in F. speciosa Cheeseman); ducts of 

flowers usually paired along and slightly outside 

of veins below sinuses, usually single along marginal veins of lobes; ducts present in anther con- 
NUMBER 51 69 

nectives; incomplete ducts of style positioned outside of veins. Achenes broadly obcompressed, 

walls carbonized, without striations, numerous 

subsidiary veins present, style veins continuous 

with median veins of inner and outer surface; 

cells of seed coat not ornamented; pappus of two 

awns borne laterally, vein usually present in awn; 

pubescence and teeth of body and awns of achene 

antrorse. Pollen 40-70 pm in diameter. 

The subtribe contains the single distinctive 

Polynesian genus Fitchia. The genus is unquestionably related to the Coreopsidinae with which 

it shares the characteristically obcompressed achenes. In both structure and geography, the Coreopsidinae seem a likely ancestor to the Fitchiinae. 

The specialized homogamous ligulate heads of 

Fitchia resemble those of the Lactuceae (Cichorieae), and the genus was placed in that tribe by 

Bentham and Hooker (1873) and Hoffman 

(1890-1894). Drake del Castillo (1898) did not 

agree, placing the genus in the Mutisieae. Stebbins (1933) rejected the genus from the Lactuceae 

and suggested a position in the Mutisieae or 

Heliantheae. The proper relationship and distinct 

subtribal status of Fitchia were established by 

Carlquist (1957) in his detailed taxonomic and 

anatomical study of the genus. 

Subtribe 22. COULTERELLINAE 

FIGURES 163. 164 

Subtribe COULTERELLINAE H. Robinson. Phytologia, 4 1 :40, 

1978. 

*CoultereLla Vasey & Rose. 

Brittle, succulent shrubs, with glabrous stems 

and leaves, lower leaves opposite. Heads aggregated in corymbose to subumbellate terminal 

inflorescences; involucre fused into utriculiform, 

low-winged cyathium, fleshy, becoming corky at 

maturity; flowers 1. rarely 2, paleae absent. Flowers discoid, hermaphroditic; corollas greenish-yellow, 10-nerved: throat short; lobes elongate, 

smooth, with strong marginal and median nerves; 

anther thecae pale: endothecial cells oblong, with 

1 or 2 thickenings on transverse walls; anther 

appendages minutely glanduliferous. Style 

branches papillose on complete abaxial surface, 

stigmatic lines paired. Ducts not distinct, without 

obvious colored resin. Achenes prismatic, walls 

carbonized, with distinct striations; cells of seed 

coat elongate, with lateral walls slightly ornamented; pappus absent. Pollen ca. 40 pm in 

diameter. 

The single genus Coulterella contains one species, C. capitata Vasey 8r Rose, growing mostly on 

beaches around La Paz Bay and on nearby islands in southern Baja California. As represented 

in the colored photograph in the paper by Moran 

(1970), the branches of Coulterella have a crassulaceous aspect. Because of the single-flowered 

heads, the genus was originally placed close to 

Lagascea in the subtribe Lagasceinae. The general 

similarity of Coulterella to the Flaveriinae and 

rejection from the proximity of Lagascea was first 

reported by Stuessy (1976), and additional support for such a treatment has been obtained from 

chemical evidence (Robinson et al., 1978). Simple 

thiophenes are found in Coulterella, and polythiophenes are present in the Flaveriinae and Pectidinae. Thiophenes are not known in Lagascea or 

other members of the subtribe Helianthinae to 

which it is related. Also, Lagascea and other 

Helianthinae contain dehydrofalcarinone, which 

is not found in Coulterella or the related subtribes 

Flaveriinae and Pectidinae. 

In spite of the similarity of Coulterella to groups 

such as the Flaveriinae, a separate subtribal status 

is required. The achenes of Coulterella lack the 

numerous costae and the extra knoblike thickenings in the carbonization that are present in the 

Flaveriinae and Clappiinae. The corollas differ 

by their short throats and long lobes. Carlquist 

(1957) noted that the median and lateral bundles 

of the lobes all terminate freely in the lobe tips, 

and that all 10 veins continue separately into the 

achenes instead of being joined at the top of the 

achenes as in other genera. Carlquist also found 

that secretory ducts or canals such as are found 

in Lagascea were lacking. 

Of the related subtribes, the Pectidinae, which 
70 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

has noncostate, striate achenes, seems most similar to Coulterella. It is notable that the Pectidinae 

also have examples of fused involucres. The involucres of Coulterella might be interpreted to have 

a slight glandular development at the top, but 

the distinct type of gland characteristic of the 

Pectidinae is lacking. The specialized floral anatomy of Coulterella is not found in the Pectidinae. 

Subtribe 23. PECTIDINAE 

FICIJRES 165- 167 

Subtribe PECTIDINAE Less., Linnaea, 5: 134, 1830. 

Tribe TACETEAE Cass., Journ. Phys., 88: 162, 1819. 

Subtribe TACETINAE Less., Linnaea, 63253: 1831. 

*Adenopappus Benth., *Adennphyllum Pers. (including Clomanocoma Cass., Lebetina Cass., Schlechtendalia Willd., nom. rej., non Less.), *Chrysactinia A. Gray, *Dyssodia Cav. (including Boebera 

Willd., Boeberastrum (A. Gray) Rydb., Rosilla Less., 

Syncephalantha Bart 1 .), * Gymnolaena (DC.) Rydb., 

*Harnackia Urb., *Hydropectis Rydb., *Hymenatherum Cass. (including Aciphyllaea (DC.) A. Gray, 

Comaclinium Scheidw. & Planch., Dysodiopsis (A. 

Gray) Rydb., Gnaphalopsis DC., Lowellia A. Gray., 

Thymophylla Lag.), * Lescaillea Griseb., * Leucactinia 

Rydb., *NicolLetia A. Gray, *Pectis L. (including 

Cheilodiscus Triana, Chthonia Cass., Cryptopetalon 

Cass., Helioreos Raf., Lorentea Lag., non Orteg., 

Pectidium Less., Pectidopsis DC., Seala Adans., Stammarium Willd. ex DC. in syn.), *Porophyllum Guett. 

(including Kleinia Jacq., non Mill.), *Schizotrichia 

Benth., *Strotheria Turner (including Graciela 

Rzed.), * Tagetes L. (including Diglossus Cass., Enalcida Cass., Solenotheca Nutt.), * Urbinella Greenm., 

* Vilobia Stroth. 

'4nnual or perennial herbs, subshrubs, or 

shrubs, with opposite or alternate leaves. 

Inflorescence usually diffuse, with heads solitary 

and peduncled or aggregated (forming compound 

heads in two species of Dyssodia). Leaves and 

involucres with prominent, often swollen, pellucid 

glandular spots or cavities. Heads usually heterogamous; involucre uniseriate or with a few subtending bractlets, bracts free or fused; paleae 

absent. Rays when present female; corolla limb 

papillose on upper surface. Disk flowers hermaphroditic (achenes of disk sterile in 1 species of 

Pectis); corollas usually glabrous or with minutely 

clavate small hairs, distinctly capitate hairs present in Pectis; lobes usually short, inner surface 

usually papillose, papillae evenly distributed or 

concentrated near margins, papillae sometimes 

hairlike, fringe of hairlike papillae also sometimes 

present inside of veins in corolla throat; anther 

thecae pale; median endothecial cells short to 

laxly elongate, end walls with 0-2 thickenings; 

anther appendage without glands. Style branches 

with paired stigmatic lines. Ducts without or with 

small amounts of colored resin; ducts along veins 

of corolla throat single; ducts in shaft of style 

outside of veins. Achenes cylindrical or prismatic, 

walls carbonized, with striations; cells of seed coat 

weakly ornamented; pappus in 0-3 series, of various setae, squamellae, or laciniate fused elements, sometimes lacking. Pollen 25-40 pm in 

diameter. 

The subtribe is widely distributed in the Neotropical region from Mexico southward to Argentina. Two species of Tagetes are widely cultivated 

as ornamentals. 

The Pectidinae have been readily distinguished 

in the past because of their general aspect and 

their characteristic secretory cavities. The cavities 

produce numerous monoterpenes which seem to 

discourage insect attack. Sesquiterpene lactones, 

which are common anti-herbivore compounds in 

other groups of Heliantheae, seem to be absent in 

the Pectidinae. Because the subtribe seems rather 

distinctive, it sometimes has been given tribal 

rank (Strother, 1977). The actual differences of 

the group are minor, however, compared to those 

of some other subtribes in the Heliantheae. 

The Pectidinae contains two subgroups, each 

recognized at the subtribal level by Strother 

(1977). The first group, containingonly the genus 

Pectis, is defined by Strother as having leaves 

always opposite and simple with bristly cilia at 

the base, having ray florets borne on the bases of 

the subtending bracts rather than on the receptacle, and having anther appendages and the 
NUMBER ii 71 

FICURES 167-177.-Pectidinae-Flaveriinae-Clappiinae-Jaumeinae-~'aril~inae-Madiinae: 167. 

Pecfis sessilij7ora (DC.) Sch.-Bip., achene, X 12; 168. 169, Flaceria trinerLia (Spreng.) C. Mohr 

(168, achene. X 17: 169, top of ray achene with corolla, X 17): 170. Sarlie~ellia mexicana .A. Gra).. 

disk achene. X 17: 171, Haploesthes greggzi A. Gray, disk achene. X 17: 172. Chrilla mexicana A. 

Gray, achene, X 17: 173, Clappia suaedaefolia A. Gray, achene, X 12: 174. Pseuduclappia nrenaria 

Rydb.. achene, X 8; 175, Jaumea carnusa (Less.) A. Gray.. achene, X 12: 176. Blepharipappus scabpr 

Hook., achene. X 12: 177. Madia glumerata Hook., ray achene, X 12. 
72 SMITHSONIAN COSTRIBUTIONS TO BOTANY 

style branches short. The remaining genera, the 

Tagetinae, are distinguished by Strother as having leaves opposite, alternate, or both, mostly 

pinnately parted, with simple blades rarely having bristly cilia at the base, having all florets 

being borne on the receptacle, and having the 

anther appendages lance-ovate and the style 

branches elongate. The carpopodium in Pectis is 

nearly symmetrical, while those of other genera 

are distinctly asymmetrical. BaagGe (1977a) reports a smoother senecionoid ligule surface in 

Pectis compared to a more papillose helianthoid 

type in Tagetes, Akolletia, and Schizotrichia. Both 

types occur in Chrysactinia, however. The Cq photosynthetic pathway is characteristic of Pectis, 

where Smith and Turner (1975) have examined 

20 species, and apparently is restricted to that 

genus: twelve genera of the Tagetinae, Adenophyllum, Chrysactinia, Dyssodia, Gymnolaena, Hymenatherum? Leucactinia, .Vicolletia, Porophyllum, Schizotrichia, Strotheria, Tagetes, and Urbinella were tested 

by Smith and Turner with negative results. 

Colored resin has been seen in small amounts 

in the disk corollas of Adenopappus and Nicolletia. 

Papillosity of the inner surface of the disk corolla 

lobes varies. Limited sampling indicates that 

there is none in Pectis, Gymnolaena, and Schizotrichia 

eupatorioides Benth. Papillosity covers the complete 

inner surface in Tagetes, Chrysactinia, Hymenatherum, 

Leucactinia, and Porophyllum. Papillae are restricted 

to the edges of the lobes in various genera including iVicolletia and in the Peruvian species Dyssodia 

lopez-mirandae Cabrera which Strother (1969) 

would place in Schizotrichia. 

The present study recognizes Vilobia Strother 

(1968) and the segregates Adenophyllum and Hymenatherum from Dyssodia at the generic level. In 

reversing a previous judgment (Robinson, 1973), 

Vilobia is distinguished from Tagetes on the basis 

of a laciniate pappus-type not equalled among 

the numerous variations found in the latter genus. 

Nevertheless, the two genera are almost certainly 

very closely related. The segregates of Dyssodza 

were not given generic rank by Strother (1977), 

but they were considered sufficiently distinct to 

be treated individually. Strother's schematic diagram of relationships among the "Tageteae" 

clearly suggests that the subgenera of Dyssodia are 

not each other's closest relatives. 

Keil (1978) has reported sterile achenes in the 

disk flowers of Pectis linfolia L. var. hirtella Blake, 

representing a further isolated occurrence of this 

character in the Heliantheae. 

Subtribe 24. FLAVERIINAE 

FIGURES 168- 1 7 1 

Subtribe FLAVERIINAE Less., Synopsis. 235. 1832. 

*Flawria Juss. (including Brotera Spreng. in 

Schrad., non Cav., Dilepis Suess. 8: Merxm., 

Neuenbergia Willd., Vermfuga Ruiz 8: Pav.), *Haploesthes A. Gray, *Sartze)ellia A. Gray. 

Herbs or subshrubs, glabrous or slightly puberulous. Leaves opposite, united by narrow scarious wing across node. Inflorescence terminal in 

laxly corymbose panicle or in dense cyme, or 

axillary. Heads heterogamous; involucre with 

bracts thinly to thickly herbaceous, in ca. 2 series, 

subequal: paleae absent. Ray flowers female; 

limb of corollas mostly smooth. Disk flowers hermaphroditic; corolla lobes short, mostly smooth, 

with cells often enlarged, outer surface usually 

glabrous (puberulous with moniliform uniseriate 

hairs below in Flaveria trineru" (Spreng.) C. 

Mohr); anther thecae pale; endothecial cells 

short, with 2-4 thickenings on transverse walls; 

anther appendage oblong, without glands. Style 

branches short, truncate, with paired stigmatic 

lines. Ducts usually without obvious colored resin 

(some reddish resin seen in Flaueria linearis Lag.) : 

ducts of disk corolla throat solitary along veins, 

ducts of lobes marginal. Ducts possibly present 

outside of veins in style shaft, ducts present inside 

veins in style branches. Achenes prismatic, with 

numerous ribs, walls carbonized with regular pattern of extra thickenings often nearly outlining 

cells of adjacent layer, striations lacking: carpopodium distinct with many rows of small cells, 

asymmetrical to nearly symmetrical; cells of seed 

coat elongate, with regular pattern of alternating 

thickenings restricted to lateral walls; pappus 
NUMBER 51 73 

absent in Flaveria (a small crown or squamellae in 

2 species), of bristle-tipped squamellae in Sartwellia, of capillary bristles in Haploesthes. Pollen 25- 

27 pm in diameter. 

The subtribe is widely distributed in the drier 

and more open habitats in tropical and subtropical America. One species of Flaveria occurs in 

Australia, and two other species have been introduced into Asia and Africa. 

The present concept of the Flaveriinae follows 

that of Turner and Johnston (1961) and Turner 

and Powell (1957). The close relationship of the 

three genera seems to be confirmed by the structure of the many-ribbed or multicostate achenes, 

and by the cellular details of the seed coat. The 

variation in the pappus in the subtribe has resulted in the traditional placement of the genera 

in two different tribes, Flaveria and Sartwellia in 

the Helenieae and Haploesthes in the Senecioneae. 

Regarding pappus variation in the subtribe, it is 

notable that an obsolete pappus occurs in 2 species of Flaveria (Powell, 1978). 

The polyacetylenes of the Flaveriinae include 

a number of highly derived forms of the pentaynene series. Tri-thiophenes are present and are 

known elsewhere in the tribe in the Pectidinae, 

one Gaillardia and in Eclipta. Sulfones also occur 

in Flaveria and are found elsewhere in Acanthospermum of the Melampodiinae, Guizotia of the Milleriinae, Eclipta, and Gaillardia. The sulfones of 

Flaveria are not epoxidized. 

Subtribe 25. VARILLINAE 

FIGURE 172 

Subtribe L.ARILLINAE Turner & Powell ex H. Robinson, 

Phytologia. 41 :44, 1978 [Subtribe L’ARILLINAE Turner & 

Powell. Biol. Chem. Compositae, 719, 1977 [ 19781, nom. 

nud. without type genus]. 

* Varilla A. Gray. 

Shrubs; plants very brittle when dry; stems 

sometimes with pale, carnose, glabrous surface; 

leaves opposite or alternate. Inflorescence terminal on branches, with heads solitary on long 

peduncle or clustered in corymbose panicles. 

Heads discoid; involucral bracts in ca. 2 series, 

with prominent median longitudinal reddish 

duct; paleae present, similar to involucral bracts. 

Ray flowers lacking. Disk flowers hermaphroditic; corolla with numerous glands or short hairs 

on lower part; throat cylindrical; lobes narrowly 

triangular, cells of inner surface elongate with 

closely sinuous lateral walls; anther thecae pale; 

endothecial cells short, with 1-3 thickenings on 

transverse walls; anther appendage without 

glands. Style branches with paired stigmatic lines. 

Ducts of involucral bracts, paleae, and corolla 

throats with reddish resin; ducts of corolla throats 

single between pairs of veins; ducts of style colorless, apparently present outside of veins at least 

in shaft. Achenes prismatic, with numerous costae, walls carbonized with slight striations, without regular pattern of knoblike thickenings; carpopodium obsolete, hidden within circle of costal 

lobes; cells of seed coat not ornamented; pappus 

absent or formed from short partially carbonized 

projections of achene body. Pollen 30-35 pm in 

diameter. 

The subtribe contains the one genus, Varilla, 

with two species in the southwestern United 

States and northern Mexico. The two species 

differ in many characteristics. In V. mexicana A. 

Gray, the inflorescence is corymbose, the leaves 

often are opposite, the stems are greenish and 

become brown with age, the tubular throats of 

the corollas are more abruptly expanded at the 

base, the hairs of the corolla base have glandular 

tips, the pollen is smaller with smaller spines, and 

the achenes bear both trichomes and the peculiar 

pappus-like outgrowths at the apex. In V. texana 

A. Gray, the heads are solitary, the leaves are 

rigorously alternate, the stem surface becomes 

thickened and whitish, the corolla throats are less 

abrupt at the base, the hairs are not glandular, 

the pollen is distinctly larger and has larger 

spines, and the achenes are glabrous. In spite of 

the differences, the anatomy of the achene wall 

with its costae and basal lobes, the cellular detail 

of the corolla lobes, the duct between the pairs of 

veins in the corolla throat, and the anatomy of 

the style are the same. The uniformity of the 
I4 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

latter characters between two species that are at 

least subgenerically distinct demonstrates the 

comparative reliability of these characters. These 

are a factor in the exclusion of all other genera 

from the present concept of the subtribe. 

In the first proposal of subtribe Varillinae 

(Turner and Powell, 1977), five genera were mentioned, including three with paleae, one with 

prominent hairs on the receptacle, and one with 

glabrous receptacles. Varilla was thought to represent an ancestral “prototype” from which others 

might have evolved by reduction. Present observations show that two of the paleaceous genera, 

Bebbia and Dyscritothamnus, are unquestionably 

members of the Galinsoginae and are not closely 

related to Varilla or the epaleaceous genera. In 

contrast, between the paleaceous Varilla, the setiferous Clappia, and the totally epaleaceous Pseudoclappia, there are some correlations in corolla 

lobe structure, costate achenes, and, to some extent, resin ducts. Still, there are significant differences, as indicated under the subtribe Clappiinae, 

where the latter two genera are placed in this 

study. It remains probable that the paleaceous 

Varilla is most closely related to the epaleaceous 

Clappia and Pseudoclappia, but I suspect the paleae 

of Varilla are derived from epaleaceous ancestors 

and that all three genera are properly placed in 

the epaleaceous series of the Heliantheae. 

The pattern of a prominent resin duct between 

two veins is one of the three possible basic types, 

but is the least common type in the tribe. This 

type finds its best expression in the reddish ducts 

in the corolla throats of the genus Varilla. The 

type is not unique to Varilla, however, and has 

been seen in less well-developed forms in some 

Clibadiinae, some Rudbeckiinae, and in both the 

corollas and styles of the genus Clappia. 

Subtribe 26. CLAPPIINAE 

Subtribe CLAPPIINAE H. Robinson, Phytologia, 41:39, 1978. 

*Clappia A. Gray, *Pseudoclappia Rydb. 

Shrubs, stems becoming rather pale and carnose; leaves alternate, with narrow sessile blades. 

Inflorescence diffuse, with long-pedunculate 

heads. Heads heterogamous; involucre with 

bracts in 2-4 series, broad and strongly subimbricate in Clappia; herbaceous to subcarnose; paleae lacking, receptacle setiferous in Clappia. Ray 

flowers female; limb of corollas nearly smooth 

above with elongate cells. Disk flowers numerous, 

hermaphroditic; corolla with a narrowly cylindrical throat; lobes narrowly triangular, elongate, 

smooth on inner surface with elongate cells, lateral walls of cells sinuous; anther thecae pale; 

endothecial cells oblong, with 1 or 2 thickenings 

on transverse walls; anther appendages without 

glands. Branches of disk styles with papillae distally on outer surface, with paired stigmatic lines. 

Ducts with some reddish resin in Clappia, without 

reddish resin in Pseudoclappia, involucral bracts of 

Clappia with numerous longitudinal dark ducts; 

ducts of disk corolla throat solitary along veins, 

partially between paired veins in Clappia; reddish 

ducts in style branches of Clappia between paired 

veins. Achenes prismatic, costate, walls carbonized, indistinctly striate, with regular pattern of 

additional carbonized knoblike thickenings; carpopodium distinct; cells of seed coat elongate, 

with annulately ornamented walls; pappus of 

numerous capillary bristles in ca. 3 series. Pollen 

35-40 pm in diameter. 

The two genera of the subtribe occur in Texas 

and the immediately adjacent areas of Mexico 

and New Mexico. Clappia is found in the Chihuahuan Desert region, and Pseudoclappia is in the 

Tamaulipan Desert. 

The genus Clappia has usually been placed in 

the Helenieae in or near the Pectidinae because 

of the dark resin lines in the leaves and involucres 

(Bentham and Hooker, 1873; Rydberg, 1923; 

Smith and Turner, 1975). Hoffmann (1890-1894) 

placed the genus in the same tribe, but in the 

Jaumeinae. Although recognizing the general resemblance, Rydberg (1923) placed his genus Pseudoclappia in the Senecioneae. A more reasonable 

treatment was that of Turner and Powell (1977), 

where both genera were placed together in a 

broadly circumscribed Varillinae. Some of the 
NUMBER 51 75 

artificial aspects and nomenclatorial problems of 

the Varillinae as originally proposed by Turner 

and Powell are mentioned in this treatment under 

that subtribe, but a comparison of the Clappiinae 

with the genus Varilla is still necessary. 

In the geographical distribution, xeric nature, 

shrubby habit, pale stems with alternate leaves, 

and brittle condition of the dried plants, the 

Clappiinae closely resemble at least one of the 

species of Varilla. Anatomical study shows additional similarities in the costate achenes, the elongate, nonpapillose cells of the disk corolla lobes, 

and the tendency for a reddish resin duct between 

paired veins in the disk corolla throat. A reddish 

duct is also present between paired veins in the 

style branches of Clappia. In spite of such similarities, the basic difference in the lack of paleae in 

the Clappiinae is reenforced by many other characters. The achenes of the Clappiinae, which have 

a well-developed carpopodial shield on the outer 

side, the pattern of knoblike thickenings in the 

carbonization of the achene wall that lacks evident micropunctations, the presence, where 

known, of ornamentation on the seed coat, the 

narrower anther collars with more evenly thickened walls, and the short bases of the anther 

thecae all differ from Varilla. The Clappiinae also 

have a prominent, multiseriate, capillary pappus, 

whereas a true pappus is completely lacking in 

Varilla. There is ample reason to treat the Clappiinae and Varilla in separate subtribes. Nevertheless, as mentioned under the Varillinae, a 

relationship seems probable. In such a grouping, 

the epaleaceous condition would seem to be more 

primitive, and the paleae of Varilla would be a 

structure resurrected in the form of the homologous involucral bracts. 

It seems likely that the Clappiinae are also 

closely related to the Jaumeinae and Flaveriinae, 

which have costate achenes. The superficial resemblance of the resiniferous lines of Cluppiu to 

the secretory cavities of the Pectidinae is misleading, and the latter subtribe is more remote in 

relationship. 

Subtribe 27. JAUMEINAE 

FIGURE 175 

Subtribe JAUMEINAE Benth. Br Hook.. Gen. PI.. 2: 199. 1873. 

*Jaumea Pers. (including Coinogyne Less.). 

Perennial herbs with creeping branched rootstocks, glabrous; leaves opposite, narrow, sessile, 

fleshy, fused at base into short, distinct sheath. 

Heads solitary, terminal, sometimes appearing 

axillary, on distinct peduncles. Heads with or 

without rays: involucre of fleshy bracts in ca. 3 

series; paleae lacking. Rays when present female; 

limb of corolla smooth on upper surface, tip with 

2 or 3 short narrow lobes. Disk flowers hermaphroditic; corolla throats narrowly cvlindrical: lobes 

short, smooth; anther thecae pale; endothecial 

cells short oblong or subquadrate. with 2 or 3 

thickenings on transverse walls: anther appendages ovate, without glands. Style branches with 

rounded tips, with stigmatic lines joining at top. 

Ducts with prominent reddish resin; ducts solitary along veins of disk corolla throat, ducts of 

style branches inside veins. .4chenes prismatic, 

multi-costate, glabrous, walls carbonized, without 

striations; carpopodium short and broad, sometimes with spicules in upper part; cells of seed 

coat not ornamented; pappus of short narrow 

squamellae or absent, without obvious median 

veins. Pollen ca. 45 pm in diameter. 

The subtribe is limited here to two species, 

Jaumea carnosa (Less.) A. Gray of western North 

America, having ray flowers and lacking a pappus, and J. linearfolia (Juss.) DC. of southern 

South America, which lacks ray flowers and has 

a short pappus. Other species that have been 

placed in Jaumea are treated here in the genera 

Espejoa and Hypericophyllum in the subtribe Chaenactidinae. The latter genera differ by the longer 

disk corolla lobes with pubescent surfaces, the 

single thickenings on the transverse walls of the 

endothecial cells, the lack of reddish resin in the 

corollas, the unribbed, usually pubescent achenes, 

and the tendency for strong central veins in the 

pappus elements. 

Various genera have been placed in the Jau- 
/b SMITHSONIAN CONTRIBUTIONS TO BOTANY 

meinae in the past, including Oliuaea Sch.-Bip. of 

the Astereae (DeJong and Beaman, 1963) and 

Cacosmia H.B.K. of the Liabeae (Robinson and 

Brettell, 1974b). Helianthean genera include 

Ciappia (treated here in the Clappiinae), Arnicastrum and Venegasia (treated here in the Chaenactidinae), and Geissopappus, which is included in 

Caiea of the Neurolaeninae (Robinson, 1975). 

Turner and Powell (1977) include the Juameinae in their new tribe Coreopsideae, but the 

typical members of the latter group are characterized by paleaceous receptacles and obcompressed achenes. The closest relationship of the 

Jaumeinae is probably to such subtribes as the 

Clappiinae and Flaveriinae, which also have 

multi-costate achenes. 

Subtribe 28. MADIINAE 

FICCRES 176- 18 1 

Subtribe MADIINAE Benth. Sr Hook., Gen. PI.. 2: 198, 1873. 

Tribe MADIEAE Jepson. FI. West. Middle Calif., 486. 1901. 

Subtribe RAILLARDELLINAE Rydb.. N. Amer. F1.. 34(4):318, 

1927. 

*Achyrachaena Schauer (including Lepidostephanus Bartl.), *Adenothamnus Keck, *Argyroxiphium 

DC., * Biepharipappus Hook.f., * Blepharizonia 

Greene, *Calycadenia DC. (including Osmadenia 

Nutt.), *Dubautia Gaud., *Hemizonia DC. (including Centromadia Greene, Hartmannia DC.), * Holocarpha (DC.) Greenm., *Hoiozonia Greene, *Lagophyiia Nutt., *Layia Hook. & Arn. ex DC., nom. 

cons., non Hook. & Arn., 1833 (including Calliachyris Torr. & Gray, Caiiichroa Fisch. & Mey., 

Caiiigiossa Hook. & Arn., Eriopappus Arn. in Lindl., 

Oxyura DC., Toiiatia Endl.), *Madia Molina (including Amida Nutt., Anisocarpus Nutt., Harpaecarpus Nutt., Madaria DC., Madariopsis Nutt., Madoreila Nutt.), *Raillardella A. Gray in Benth. & 

Hook., * Wiikesia A. Gray. 

Annual or perennial herbs, or rarely shrubs as 

in Dubautia, rosuliform or caulorosula in 

Argyroxiphium. Leaves opposite at least in early 

stages, often becoming subopposite or alternate, 

sometimes bearing specialized, multicellular, capitate glands. Inflorescence terminal, usually cymose or with cymose branches, sometimes elongate or virgate with some acropetal development, 

rarely with solitary heads. Heads usually heterogamous, rays lacking in Dubautia, Raillardella, Wilkesia, and Layia discoidea Keck; involucral bracts 

in a single series except in Hemizona angusttfoiia 

DC., each bract usually enfolding a ray flower; 

when rays lacking, a single series of bracts surround disk; paleae usually in single series subtending outer disk flowers, arranged in a circle 

and more or less connate (except in Achyrachaena): 

interior of disk usually epaleaceous (with inner 

paleae in Biepharipappus, typical Hemizonia, Hoiocarpha, and 2 species of Layia); involucral and 

receptacular bracts with strands of fibers usually 

separate from and alternating with veins. Ray 

flowers when present female: corolla limb papillose on upper surface, strongly trilobed at tip. 

Disk flowers hermaphroditic (functionally male 

in at least some disk flowers of Hemizonia sect. 

Centromadia and sect. Deinandra, Holocarpha, and 4 

species of Madia) ; corolla lobes short and papillose 

on inner surface, with few helianthean hairs; 

anther thecae usually darkened; endothecial cells 

short, usually with 1 or 2 thickenings on transverse walls: anther appendages without glands, 

with cell walls often elaborately thickened. Style 

branches with paired stigmatic lines: styles of disk 

flowers usually with a short triangular or acuminate appendage, lower abaxial surfaces of 

branches usually smooth (numerous hairs continuing onto shaft in Biepharipapfus) : styles in functionally male flowers often reduced, often without 

stigmatic papillae. Pectic intercellular deposits in 

leaves and sometimes in bracts (except in vernal 

annuals Achyrachaena and Layia). Duct system usually reduced, present in stems and leaf-bases in 

Achyrachaena, Adenothamnus, Wiikesia, a few species 

of Layia, Madia, and Raillardella, present in corollas 

and styles of Achyrachaena, Lagophylia, some species 

of Dubautia; ducts sometimes solitary along veins 

of corolla throat, ducts with reddish resin prominently paired along veins of corolla throat and 

inside veins of style branches in Achyrachaena, 

traces of 4 ducts seem evident outside of veins in 
NUMBER jl 77 

FIGURES 178- 191 .-Madiinae-Hymenopappinae-Lycapsinae-Peritylinae: 178. 179. Lajia glandulosa (Hook.) Hook. & Am. (178, disk achene, X 12; 179, ray achene. X 12): 180. Wilkesia 

gymnoxiphium A. Gray, achene, X 12; 181, Dubautia laxa Hook, & Am.. achene with corolla, 

X 12: 182, Hymenopappusjilijolius Hook., achene with corolla, X 8: 183, Loxothysanus pedunculatus 

Rydb., achene, X 17; 184. 185, Villanova titicacensis Mey. & Walp. (184. achene. X 17: 183. 

achene with corolla: X 17); 186, 187, Galeanapratensis (H.B.K.) Rydb., achenes, outer and inner 

view, X 13: 188, Lycapsus tenuifolius Philippi, ray achene. X 17: 189, Penole emotyi Torrev. 

achene, X 17: 190, Amauria rotundijolia Benth., achene, X 17: 191, Eutetras palmeri A. Gray. 

achene. X 12. 
78 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

style shaft of Holocarpha. Achenes prismatic, 

curved or sometimes slightly compressed, walls 

carbonized, usually with striations but obscured 

by dense cell layers in Hemizonia and Holocarpha, 

carbonization unbroken but slightly grooved in 

Calycadenia; where seen, cells of seed coat without 

ornate thickenings, with strongly sinuous walls in 

Holocarpha, with pustulate inner surface in Achyrachaena and Layia; pappus of various bristles, 

awns, or squamellae, or lacking, usually radially 

arranged. Pollen 30-35 pm in diameter. 

Members of the Madiinae are generally known 

as tarweeds, evidently because of the common 

occurrence of abundant pectic material in the 

leaves. The subtribe has a distinctive distribution, 

with most genera concentrated along the Pacific 

coast of the United States and Baja California. A 

few genera occur in the Hawaiian Islands, and 

one or two species of Madia occur as a disjunct 

element in Chile. 

The subtribe has been reviewed recently in an 

anatomical study by Carlquist (1959), and that 

study contains citations of various other papers 

of that author dealing with members of the group. 

With few exceptions, the group can be recognized 

by the single series of subequal involucral bracts 

enfolding the ray florets and the single series of 

usually partially connate paleae surrounding the 

disk. Carlquist (1959) cited a few developmental 

trends within the subtribe, the development of 

distinct fiber strands in the involucral bracts and 

paleae, the loss of secretory canals or resin ducts, 

and the presence of capitate trichomes with biseriate stalks which are modified into more complex structures in some genera. The achene walls 

of the Madiinae seem to be basically striate, and 

the cells of the seed coat basically inornate, but 

interesting variations occur in both structures. 

Traditionally, the subtribe has been placed 

among the paleaceous Heliantheae, and the 

darkened anther thecae might seem to support 

such a position. No close relatives are evident 

among the paleaceous Heliantheae, however, and 

in many respects the Madiinae resemble members 

of the epaleaceous series. Under the microscope, 

as preserved in Hoyer’s solution, the pigment of 

the anther thecae of the Madiinae appears reddish. Except for reddish pigments in a few genera 

such as Smallanthus and Zchthyothere of the Melampodiinae, the pigments of the most genera of the 

paleaceous subtribes appear blackish or greenish 

under the microscope. In this study, the Madiinae 

are regarded as remote from other dark-anthered 

members of the tribe, and are aligned among the 

basically epaleaceous subtribes of the Heliantheae. 

Stuessy (1977) suggested that the reduced annual herb Faxonia, of Baja California, might belong to the Madiinae. The genus has been examined and has been placed in the subtribe 

Galinsoginae in this study. 

Subtribe 29. HYMENOPAPPINAE 

FIGURES 182-187 

Subtribe HYMENOPAPPINAE Rydb., N. Am. Fl., 34( 1):43, 

1914. 

*Galeana La Llave & Lex. (including Chlamysperma Less.) Hymenopappus L’H6r. (including 

Rothia Lam., nom. rej., non Schreb., nec Rothia 

Pers., nom. cons.), *Loxothysanus B. L. Robins., 

* Trichocoryne Blake?, * Villanova Lag., nom. cons., 

non Ortega (including Vasquezia Phil.). 

Annual, biennial, or perennial herbs or shrubs. 

Leaves opposite or alternate, sometimes in basal 

rosettes. Inflorescence subscapose, or terminal on 

leafy branches, cymose. Heads heterogamous or 

discoid; involucre of subequal herbaceous bracts 

in 2 or 3 series, mostly ovate; paleae lacking 

(except in Hymenopappus newberryi (A. Gray) Johnston). Ray flowers when present female; corolla 

limbs papillose on upper surface. Disk flowers 

hermaphroditic; corolla glanduliferous on outer 

surface, throat campanulate; lobes 5, papillose on 

inner surface; anther thecae pale; endothecial 

cells short, with usually 2 or 3 thickenings on 

transverse walls; anther appendages ovate, with 

or without glands. Style branches with paired 

stigmatic lines, papillose only near tip abaxially, 

short-pointed or truncate at tip. Without evident 

colored resin; ducts of style, if present, outside of 
NUMBER .5l 79 

veins. Achenes rather trigonous or quadrangular, 

with shortly to longly stipitate base, surfaces 

winged or tuberculate in Galeana, walls carbonized, without striations, carbonization with additional pattern of knoblike thickenings; cells of 

seed coat elongate, with ornate thickenings restricted to lateral walls or lacking; pappus lacking 

or of radially arranged squamellae, squamellae 

with central cells forming distinct costa. Pollen 

25-35 pm in diameter. 

The subtribe is concentrated in the region of 

Mexico and the southwestern United States with 

one genus, Villanova, having some species endemic 

in western South America. 

The subtribe forms part of a series of five 

(Hymenopappinae, Peritylinae, Baeriinae, 

Chaenactidinae, and Gaillardiinae), which I consider unquestionably closely related. These form 

the core of what was recognized by Bentham and 

Hooker (1873) and Hoffmann (1890-1894) as the 

tribe Helenieae. The Hymenopappinae differ 

from the others of the series by the distinctly 

carbonized achene walls lacking striations. The 

achenes of the Hymenopappinae also tend to be 

distinctive in the trigonous or quadrangular 

shape and the stipitate bases. In both Hymenopappus and Loxothysanus, where a pappus is present, 

the squamellae have a similar differentiated costal region. 

The subtribe contains two elements of very 

different habits. Hymenopappus contains biennial 

or perennial often subscapose plants with alternate leaves forming a basal rosette. The remaining annual genera Galeana and Villanova, and the 

shrubby Loxothysanus, branch above the base and 

have opposite leaves in at least the lower part of 

the plant. 

The genus Trichocoryne was described by S F. 

Blake in 1924 with a single species, T. connata 

Blake. Relationship to Galeana was suggested, but 

the more numerous bracts and flowers of the 

heads were noted. The genus appears to have 

been overlooked by more recent summaries of the 

epaleaceous Heliantheae. Examination of the 

type specimen from Durango, Mexico, leaves the 

position of the genus in question, but the nonstriate, generally trigonous achenes would favor 

placement near the Hymenopappinae. The four 

small plants on the sheet seem more like annuals 

than perennials as described by Blake. The leaves 

are linear and opposite, and are fused into a short 

sheath at the base. The pedunculate heads are 

solitary on the tips of the stems and branches. 

There are 10 broadly rounded involucral bracts 

having subhyaline margins. The receptacle was 

described as convex or low-conical, but it is very 

highly conical. Paleae are lacking. The 5-7 rays 

were described as sterile with “inane” achenes, 

but they have styles and are only lagging in 

development. The corollas are 5-lobed and are 

covered with short, clavate, uniseriate hairs. The 

base of the achene is not stipitate, and the cells of 

the seed coat are slightly distorted with no ornamentation on the walls. The plant is, by all 

indications, a member of the epaleaceous series, 

but the anther thecae are distinctly blackened. 

The thickenings of the endothecial cells are numerous and radially or subradially disposed. 

Subtribe 30. LYCAPSINAE 

FIGURE 188 

Subtribe LYCAPXNAE H. Robinson, I’hytologia. 46: 120. 1980. 

* Lycapsus Phil. 

Small perennial herbs. Leaves closely spirally 

inserted; petioles long and slender from short 

broadened bases, blades fleshy, irregulary divided 

into a few narrow lobes. Inflorescence axillary, 

with single heads on long slender peduncles. 

Heads heterogamous; involucre with slightly herbaceous narrow bracts irregularly disposed in ca. 

2 series; paleae present. Ray flowers present, female; limb of corolla broad, papillose on upper 

surface, tip broadly trilobed. Disk flowers hermaphroditic: corollas 4-lobed: inner surface of 

lobes papillose, outer surface with glands and a 

few short, 2-3-celled, uniseriate hairs; anther thecae pale; endothecial cells with 2 or 3 thickenings 

on transverse walls: anther appendage without 

glands. Style base distinctly nodulose, branches 

with paired stigmatic lines, tips short acute, tri- 
80 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

angular, papillose on abaxial surface of appendage. Ducts not evident, without colored resin. 

Achenes prismatic, fusiform with ca. 4 costae, 

walls carbonized in an even and continuous coating, without striations, without additional pattern 

of knoblike thickenings, marked with only cell 

outlines and micropunctations; cells of seed coat 

not seen. Pappus absent. Pollen ca. 27 pm in 

diameter. 

The subtribe Lycapsinae consists of a single 

species, Lycapsus tenuifolius Phil., that is geographically isolated, being endemic to the remote, 

rather inaccessible islands of San Ambrosio and 

San Felix off the coast of Chile. 

The genus Lycapsus is initially distinguishable 

by the closely spirally inserted leaves with deeply 

pinnately dissected, fleshy laminae. On the basis 

of inadequate information, the genus was reduced 

to synonymy under Alomia H.B.K. in the tribe 

Eupatorieae by Reiche (1901), and the species 

was treated as a member of that genus by B. L. 

Robinson (1913). The genus was resurrected and 

removed from the Eupatorieae by Johnston 

(1935) who noted the radiate condition of the 

heads. According to Johnston, “The plant has 

fertile pistillate marginal florets with a 3-toothed 

ligule about once and a half the length of the 

tube. The tubular inner florets appear to be 

hermaphroditic and sterile. The style branches 

are linear, flattened and abruptly contracted into 

a short triangular apex. The receptacle bears 

conspicuous slender scales which seem to separate 

the marginal florets from the inner ones. Except 

for the bracteate receptacle the plant is very 

suggestive of some of the helenioids or even certain asterioids.” The material seen in this study 

differs from the Johnston description only by the 

presence of distinct paleae among as well as 

around the disk flowers and by the more evident 

fertility of the disk flowers. The plant is clearly a 

member of the Heliantheae as defined here. 

The genus is sufficiently distinctive to be placed 

in a separate subtribe. In fact, the closest relationship within the tribe is only tentatively resolved. 

As noted by Johnston (1935), the genus is suggestive of the helenioids, although it has paleae. 

Present observations find distinctly petiolate 

leaves, 4-lobed corollas, and nonstriate achenes 

with an even layer of carbonization, all as in the 

subtribe Peritylinae. The costae and carpopodial 

structure of the achenes are matched by members 

of the latter subtribe, especially Eutetras. It is 

suggested here that the Lycapsinae are close to 

the latter opposite-leaved, epaleaceous subtribe, 

and that the paleaceous receptacle and less regular involucre represent a reversion from an epaleaceous condition, a situation equivalent to that 

seen in Varilla. 

Geographically, the Lycapsinae are isolated 

from the primary ranges of potentially related 

subtribes such as the Peritylinae in western North 

America. Nevertheless, the disjunct occurrences 

of Madia of the Madiinae and of Perityle emoryi 

Torr. of the Peritylinae in northern Chile are two 

examples of separate, more recent introductions 

of such subtribes into areas near to San Ambrosio 

and San Felix. 

Subtribe 31. PERITYLINAE 

FIGURES 189-191 

Subtribe PERITYLINAE Rydb., N. Amer. FI., 34(1): 11, 1914. 

Subtribe AMAURIINAE Rydb.. N. Amer. Fl., 34(1):29. 1914. 

*Amauria Benth., *Correllia Powell, *Eutetras A. 

Gray, *Pericome A. Gray, *Perityle Benth. (including Closia Remy, Laphamia A. Gray, Leptopharynx 

Rydb., Monothrix Torrey, Nesothamnus Rydb., Pappothrix (A. Gray) Rydb.). 

Annual or perennial herbs or subshrubs, rarely 

low shrubs. Leaves usually opposite, at least near 

base of plant; petioles distinct. Inflorescence 

rather laxly cymose or with densely subcorymbose 

branches, heads distinctly pedunculate. Heads 

heterogamous or sometimes discoid; involucre 

with numerous narrow herbaceous bracts, mostly 

uniseriate; paleae lacking. Ray flowers when present female; limb of corolla papillose on upper 

surface, tip usually distinctly trilobed. Disk flowers hermaphroditic; corollas usually 4-lobed; inner surface of lobes weakly to strongly papillose, 

outer surface often with peglike hairs; anther 
NUMBER 51 81 

thecae pale; endothecial cells with 1-3 thickenings on transverse walls; anther appendage without glands. Style branches with paired stigmatic 

lines, tips acute, abaxial surface with many hairs 

on at least the distal half. Ducts without colored 

resin: ducts of style outside of veins. Achenes 

slightly to strongly compressed, usually with 2 or 

4 costae, walls carbonized, usually in even, continuous coating without striations, often without 

additional pattern of knoblike thickenings and 

having only cell margins and micropunctations 

evident; cells of seed coat not ornamented, sometimes with sinuous or zigzig walls; pappus of 

bristles or scales, usually arranged bilaterally, 

pappus absent in Amauria and some species of 

Perityle. Pollen 23-27 pm in diameter. 

The subtribe occurs in the southwestern United 

States and northern Mexico, with one species, 

Perityle emoryi Torrey, having a disjunct range in 

North America, Peru, and Chile. 

The taxonomic history has been reviewed recently in a survey of the subtribe by Powell and 

Turner (1974). Those authors state, “The unifying characters of the Peritylinae are radially flattened achenes with conspicuous to inconspicuous 

callous margins: a pappus of short laciniate squamellae and/or rather stout short-ciliate bristles; 

one or two equal to subequal series of involucral 

bracts which partially enclose the peripheral achenes; narrow slightly flattened, subulate-filiform 

style branches; and four-lobed disk corollas.” The 

Powell and Turner survey, nevertheless, retained 

three genera in the subtribe, Eatonella, Oxypappus, 

and Hulsea, which have 5-lobed disk corollas. 

Also, Hulsea has a 2-3-seriate involucre. The present treatment excludes these three genera, which 

differ from the natural element of the subtribe by 

their striate achenes, their sessile leaves, and their 

chromosome numbers not based on X = 18. 

Eatonella and Oxypappus with X = 10 seem to be 

members of the subtribe Baeriinae, and Hulsea 

with X = 19 is a member of the Chaenactidinae. 

Amauria, Correllia, and Perityle have a distinctive 

form of achene wall carbonization with an even 

layer of deposition interrupted only by wellspaced micropunctations. This seems to be a specialization within the subtribe, although similar 

achene walls occur in some species of Lasthenia in 

the related subtribe Baeriinae. The remaining 

two genera of the Peritylinae, Pericome and Eutetras, have a more conventional form of achene 

wall with a granular structure and an additional 

regular pattern of knoblike thickenings. In Pericome, the conventional wall structure seems to 

underlie a layer of the Perityle-type, but in Eutetras, 

the Perityle-type layer is completely lacking. 

The endothecial, cells of all but one genus of 

the subtribe are characterized by 2 or 3 thickenings on the transverse walls. In Pericome the endothecial cells usually have single thickenings on 

the transverse walls. 

Subtribe 32. BAERIINAE 

FIGURES 192- 196 

Subtribe BAERIINAE Benth. & Hook., Gen. PI., 2:200, 1873. 

Subtribe ERIOPHYLLINAE Rydb., N. Amer. F1.. 34(2):81, 

1915. 

* Amblyopappus Hook. & Am. (including Aromia 

Nutt., Infantea Remy in Gay), *Antheropeas Rydb., 

* Baeriopsis Howell, * Eatonella A. Gray, * Eriophyllum Lag., *Lasthenia Cass (including Baeria Fisch. 

& Mey., Burrielia DC., Crockeria Greene ex A. 

Gray, Dichaeta Nutt., Hologmne Bartl., Ptilomeris 

Nutt., Rancagua Poepp. & Endl., Xantho Remy), 

*Lembertia Greene, *Monolopia DC., *Oxypappus 

Benth., *Pseudobahia (A. Gray) Rydb. 

Annual or perennial herbs; leaves opposite or 

sometimes alternate, crowded on short stems in 

Baeriopsis; petioles lacking or indistinct. Inflorescence terminal, with heads solitary on erect peduncles, laxly cymose, or in subcorymbose clusters. Heads usually heterogamous, sometimes discoid; campanulate to cylindrical: involucral 

bracts herbaceous, in ca. 2 series, ovate to orbicular, narrow in Oxypappus; paleae lacking. Ray 

flowers when present female; corolla limb papillose on upper surface, weakly to strongly trilobed 

at tip. Disk flowers hermaphroditic; corollas usually 5-lobed; inner surface of lobes papillose, outer 

surface often with few to many short, flattened, 
82 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

biseriate, nonglandular hairs, glabrous in Oxypappus; anther thecae pale; endothecial cells short, 

usually with 2-5 thickenings on transverse walls 

(with radial thickenings in Lasthenia; cells elongate with thickenings on lateral walls in Oxyfappus); anther appendages with or without glands. 

Style branches with paired stigmatic lines, tips 

subtruncate or shortly apiculate. Ducts without 

colored resin; ducts of style shaft apparently outside of veins, veins of style branches near outer 

surface and ducts possibly absent, ducts apparently present inside of veins in branches and 

upper shaft of Oxypappus style. Achenes prismatic 

to compressed, walls carbonized, with striations, 

usually with additional knoblike thickenings, 

with a smooth layer of carbonization in some 

species of Lasthenia; cells of seed coat not ornamented; pappus of awns or squamellae or lacking, 

awns and squamellae without sharply demarcated costal region. Pollen mostly 25-30 pm in 

diameter. 

Members of the subtribe are concentrated in 

the western United States and northern Mexico. 

One species, Amblyopappus pusillus Hook. & Am., 

has a disjunct distribution extending to Peru and 

Chile in South America. 

Within the related series of epaleaceous subtribes, the Baeriinae differ by the lack of distinct 

petioles on the leaves, by the carbonized achenes 

with distinct striations, and by the usually short 

endothecial cells with 2-5 thickenings on the 

transverse walls. The short clavate hairs on the 

disk corolla lobes of a number of the genera 

always seem to be biseriate rather than monoseriate as in some of the other subtribes in the 

series. Most genera of the Baeriinae are further 

distinguished from the related Peritylinae by the 

broader involucral bracts. 

Oxypappus is evidently related to this subtribe 

by the form of the leaves and the achenes. However, the genus has a number of anomalous features: an involucre with narrow bracts of the 

Perityle-type, elongate endothecial cells with thickenings on the vertical walls, and resin ducts of the 

style inside the veins in the upper part of the 

shaft. 

Various members of the subtribe that have 

been examined show weakly to strongly developed conical receptacles. Also, a number of members, including Eatonella, Monolopia, and parts of 

Lasthenia, have compressed achenes. The achene 

of Monolopia is of particular interest. The shape of 

the carpopodium indicates that the achene ordinarily lies against the conical receptacle in a 

tangential position, but the veins of the style are 

continuous with the edges of the achene, rather 

than with the flattened surfaces, showing that the 

achene is anatomically compressed rather than 

obcompressed. 

The genus Lasthenia is here considered a natural 

genus which can be distinguished anatomically 

from others of the subtribe by the more radial 

pattern of thickenings on the endothecial cells. 

The genus is also of interest chemically. It is one 

of the two elements in the Heliantheae that has 

long been known to contain anthochlor pigments 

(Bohm et al., 1974), and Bohm (1977) mentions 

that Lasthenia, on the basis of five species in two 

sections, differs from other “Helenieae” by having 

polyacetylenes based on the ene-tetrayne-ene series rather than the pentaynene series. The monothiophenes in Lasthenia of the ene-tetrayne-ene 

series are particularly notable. It seems to be 

coincidence that both types of chemicals, the 

anthochlors and the ene-tetrayne-ene thiophenes, 

are most prominent elsewhere in the tribe in the 

subtribe Coreopsidinae. Contrary to the suggestion of Harcombe (1977), there is no other reason 

to associate Lasthenia with the Coreopsidinae. 

Previously unnoticed in Lasthenia is the internal 

structure of the achene wall, which varies between 

two types. Lasthenia chrysostoma (Fisch. & Mey.) 

Greene, L. coronaria (Nutt.) Ornduff, L. leptalea (A. 

Gray) Ornduff, L. minor (DC.) Ornduff, and L. 

plaQcarpha (A. Gray) Greene have walls with 

evenly deposited phytomelanin interrupted only 

by micropunctations and striations. Lasthenia chrysantha (Greene ex A. Gray) Greene, L. glaberrima 

DC., L. glabrata Lindl., and L. microglossa (DC.) 

Greene have the phytomelanin more irregular 

and granular, with an additional pattern of knoblike thickenings. The wall type in the first series 
NUMBER 51 83 

FIGURES 192-200.-Baeriinae-Chaenactidinae: 192, 193, Baertopsis quadalupensu J. T. Howell. 

disk flowers with achenes, X 12; 194, Lasthenta chrysostoma (Fisch. & Mey.) Greene, disk achene. 

X 17; 195. Oxypappus seemannii (Sch.-Bip.) Blake, disk achene. X 17: 196, Eatonella congdonii A. 

Gray, achene, X 12; 197, Espejoa mexicana DC., achene, X 8: 198, Palafoxta sphacelata (Kutt. ex 

Torr.) Cory, disk achene, X 8; 199, Bahia absinthtfolta Benth., disk achene with corolla. X 12: 

200, Hypertcophy llum multicaule Hutchins., achene, X 6. 
84 SMITHSOSIAN CONTRIBUTIONS TO BOTANY 

is the same as that seen without striations in the 

typical members of the subtribe Peritylinae. Examination of a series of L. microglossa indicates the 

character is stable within a species. Attempts to 

correlate with the groups of Ornduff (1966) show 

the first type of wall in sections Baeria, Platycarpha, 

Ptilomeris, and one species of Burrielia. The second 

wall type is in sections Hologymne, Lasthenia, and 

another species of Burrielia. Only in section Burrielia has hybridization been reported by Ornduff 

between species shown here to have different wall 

types, and those hybrids were sterile. 

The variation in the achene wall in Lasthenia 

can be compared also with the results of the 

detailed study of the flavonoid pigments in the 

genus (Bohm et al., 1974). The first type of achene 

wall, the Perityle-type, is found in groups with the 

most highly developed anthochlor chemistry as 

well as those with the nonanthochlor flavonoid 

patuletin. The second wall type is found in species 

which have less complex anthochlors or species 

with no specialized flavonoids. In the flavonoid 

study, the species with anthochlors are regarded 

as primitive within the genus, and the species 

without anthochlors are considered derived. As 

such, the Perityle-type of achene wall would also 

be primitive within the genus. It is notable that 

neither the Ornduff sections, the flavonoid types, 

nor the achene wall types correlate completely. It 

is notable, also, that anthochlors and Perityle-type 

achene walls are not found in the other genera of 

the Baeriinae, and they almost certainly are not 

primitive in the subtribe. Just as the anthochlors 

do not seem to reflect close relation to the Coreopsidinae, the evenly deposited phytomelanin of 

the achene wall does not seem to reflect a direct 

relationship of Lasthenia to Perityle. 

Subtribe 33. CHAENACTIDINAE 

FIGURES 197-204 

Suhtribe CHAENACTIDINAE Rydb., N. Amer. Fl., 34(1):63, 

Suhtribe BAHIINAE Rydb., N. Amer. FI., 34(1):31, 1914. 

Suhtribe PALAFOXIINAE Rydb.. N. Amer. Fl., 34(1):58, 1914. 

1914. 

* Achyropappus H.B.K., *Arnica L., * Arnicastrum 

Greenm., * Bahia Lag. (including Amauriopsis 

Rydb., Picradeniopszs Rydb., Stylesia Nutt., Virletia 

Sch.-Bip. ex Benth. & Hook. in syn.), *Bartlettia 

A. Gray, *Chamactis DC. (including Acarphaea 

Harv. & Gray ex Gray, Acicarphaea Walp., Macrocarphus Nutt.), *Chamaechaenactis Rydb., *Espejoa 

DC., *Florestina Cass., *Hulsea Torrey & Gray, 

*Hymenothrix A. Gray (including Trichymenia 

Rydb.), *Hypericophyllum Steetz, *Jamesianthus 

Sherff & Blake, *Mallotopus Franch. & Sav., *Orochaenactis Coville, *Palafoxia Lag. (including Loxaxeta Raf., Othake Raf., Paleolaria Cass., Polypteris 

Nutt.), *Peucephyllum A. Gray, *Platyschkuhria 

Rydb., *Psathyrotopsis Rydb. (including Pseudobartlettia Rydb.), *Schkuhria Roth., nom. cons., non 

Sckuhria Moench (including Cephalobembix Rydb., 

Chamaestephanum Willd., Hopkirkia DC., non 

Spreng., Mieria La Llave in La Llave & Lex., 

Tetracarphum Moench) , *Syntrichopappus A. Gray, 

* Thymopsis Benth. in Benth. & Hook., nom. cons., 

non Jaubert & Spach (= Neothymopsis Britt. & 

Millsp.), * Venegasia DC., * Whitneya A. Gray. 

Annual, biennial, or perennial herbs (a shrub 

in Peucephyllum). Leaves opposite, alternate, or 

basal, with or without distinct petioles. Inflorescence of many heads or of single heads on erect 

peduncles, sometimes subscapose to laxly cymose 

or clustered at tips of short leafy branches. Heads 

usually heterogamous, sometimes discoid; involucre of 2 or 3 series of herbaceous bracts, bracts 

broadly to narrowly ovate or lanceolate, usually 

subequal; paleae lacking. Ray flowers when present female; corolla limbs usually papillose on 

upper surface, nearly smooth in Bahia. Disk flowers hermaphroditic (functionally male in Neothymopsis polyantha Urb.?); corollas usually with 5 

lobes; inner surfaces of lobes smooth or papillose; 

anther thecae pale: endothecial cells short or long, 

with 1 or 2 thickenings on transverse or end walls, 

thickenings only on vertical walls in Chamaechaenactis; anther appendage with or without glands. 

Style base usually glabrous as in other subtribes, 

but with hairs in Arnica, Chaenactis, and Chamaechaenactis; style branches with paired stigmatic 

lines. Without colored resin; shaft of style with 

ducts outside of veins. Achenes usually terete or 
FIGURES 20 1-2 10.-Chaenactidinae-Gaillardiinae-Marshalliinae: 20 1, Peucephyllum schottii A. 

Gray, achene, X 8; 202, Chaenactis douglasii Hook. 8z Am., achene, X 8; 203, Thymopsis thymoides 

Griseb., achene, X 17; 204, Arnica cordifilia Hook., disk achene. X 8; 205, Psilostrophe tagetina 

(Nutt.) Greene, ray floret with persistent corolla, X 12: 206, Baileya multiradiata Haw. Br Gray, 

ray floret with persistent corolla, X 12; 207, Trichoptilium incisum (A. Gray) A. Gray, achene, X 

12; 208, Psathyrotes annua (Nutt.) A. Gray, achene, X 12; 209, Helenium autumnale L., disk achene, 

X 17; 210. Marshallia graminlfolia (Walt.) Small, achene, X 17. 
86 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

prismatic, obcompressed in Bartlettia, walls carbonized, with or without additional pattern of 

knoblike thickenings, with striations; cells of seed 

coat usually not ornamented, with weakly annulated walls in Psathyrotopsis; pappus of bristles, 

awns, or squamellae, in l-many series, with or 

without differentiated central costa, sometimes 

with long vascular trace. Pollen 27-40 pm in 

diameter. 

The subtribe is predominantly North American with most genera in the western United 

States, Mexico, or Central America. A few species 

of Bahia and Schkuhria occur in western South 

America. The poorly understood and perhaps 

improperly assigned genus Thymopsis is endemic 

to the West Indies. Mallotopus is in eastern Asia, 

and Arnica extends from North America into Europe. A number of species of Hypericophyllum occur 

in Africa. 

The genera of the Chaenactidinae can be divided roughly into two groups on the basis of the 

endothecial cells of the anthers. The series of 

genera that includes Bahia, Achyropappus, Espejoa, 

Florestina, Hynienoth rix, Hypericophy llum, Pa la foxia, 

Plapschkuhria, and Schkuhria has mostly short endothecial cells in regular rows with well-defined 

transverse walls. The group shows some general 

trends toward smooth surfaces on the ray corollas 

and more strongly nerved pappus elements. The 

pappus elements often contain vascular traces, 

the capillary pappus in Hypericophyllum is sometimes vascularlized to 2/3 its length, and Palafoxia 

has awns with veins more than half the length. 

The achenes of the group usually show a regular 

pattern of knoblike thickenings in the carbonization at full maturity. 

The typical group of the Chaenactidinae, 

which includes Arnica, Arnicastrum, Chaenactis, Chamaechaenactis, Hulsea,Jamesianthus, Mallotopus, Orochaenactis, Peucephyllum, Psathyrotopsis, and Whitneya, has elongate, often elliptical endothecial cells 

that are not in distinct rows and are pointed at 

the ends. The pappus in the group is mostly more 

delicate, with little or no central costa and no 

vascular trace. Arnica, Chaenactis, and Chamaechaenactis have hairs on the base of the style, a character 

found elsewhere in the family only in the tribe 

Eupatorieae. Genera in the group have carbonization of the achene wall lacking any additional 

pattern of knoblike thickenings. 

Two subtribes, the Bahiinae and Chaenactidinae, could be recognized on the basis of the 

foregoing alignment, but a few genera included 

in the present concept do not fall easily into either 

of the two groups. One of these singularly distinctive genera is Venegasia, which is unique in the 

stalked glands on the anther filaments. Another 

genus, Bartlettia, has the only truly obcompressed 

disk achenes in the tribe outside of the Coreopsidinae-Fitchiinae group. Thymopsis, a reduced 

West Indizn genus, is placed in the subtribe with 

reservations, but it has striate prismatic achenes, 

distinctly petiolate leaves, and usually single 

thickenings on the endothecial cells. 

The typical element of the Chaenactidinae 

seems closely related to the next subtribe, Gaillardiinae, as indicated by the reports of helenanolide sesquiterpene lactones in Arnica (Evstratova 

et al., 1971; Poplawski et al., 1971). The carbonized versus uncarbonized achene wall is the character by which the two subtribes are distinguished 

here. This character receives its most severe test 

in the genera Psathyrotes and Psathyrotopsis. These 

two genera share a strikingly distinctive pappus 

form, and they have been treated as parts of a 

single genus in the monograph by Strother and 

Pilz (1975), but they recognized cytological, geographical, and some structural differences between the two elements. The two genera were 

listed separately by Nordenstam (1977). The present study shows that the genus Psathyrotopsis of 

the Chihuahuan region has corollas glanduliferous throughout, corolla lobes with biseriate bilobed hairs on the outer surface and evenly distributed papillae on the inner surface, anther 

appendages that are ovate and glanduliferous, 

style branches that are haired abaxially to the 

base, achene walls that are carbonized, and cells 

of the seed coat that are ornamented with annulate thickenings. Psathyrotes of the Sonoran and 

Great Basin regions has corollas that are glabrous 

below, corolla lobes with uniseriate hairs outside 
NUMBER .>I 87 

and long papillae forming a marginal fringe inside, anther appendages that are spathulate and 

without glands, style branches that are haired 

only at the tips, achene walls that are not carbonized, and cells of the seed coat that are not 

ornamented. The two genera may yet prove 

closely related, but no character except the pappus has been found to suggest this. For the present, the two are placed in separate subtribes. 

Psathyrotopsis is a member of the Chaenactidinae, 

whereas typical Psathyrotes is placed in the Gaillardiinae close to Trichoptilium. 

The African Welswitschiella Hoffm. has not 

been seen in this study, but the conventional 

placement is near Hypericophyllum. 

Subtribe 34. GAILLARDIINAE 

FIGURES 205-209 

Subtribe GAILLARDIINAE Less., Linnaea, 6:516, 1831. 

Subtribe HELENIINAE Less., Synopsis Gen. Comp., 237, 1832. 

Tribe HELENIEAE Benth. & Hook., Gen. PI.. 2: 199, 1873. 

Subtribe RIDDELLIINAE Hoffm.. Nat. Pflanzenfam. IV, 5: 

253. 1891. 

Family HELENIACEAE Bessey, Ann. Missouri Bot. Gard., 2: 

163, 1915. 

Subtribe TETRANEURINAE Rydb., N. Amer. FI., 34(2): 100, 

1915. 

Subtribe PSILOSTROPHINAE Turner & Powell, Biol. Chem. 

Cornpositae, 723, 1977 [ 19781, based on RIDDELLIINAE 

Hoffrn. 

*Actinospermum Elliott, *Arnblyolepis DC., *Baileya Haw. & Gray ex Torrey, *Balduina Nutt., nom. 

cons. (Mnesiteon Raf.), *GailLardia Foug. (including Agassizia Gray & Engelm., Calonnea Buchoz, 

Cercostyfos Less., Guentheria Spreng.), *Hefeniurn L. 

(including Actinea Juss., Brassavofa Adans. nom. 

rej., non R. Br., nom. cons., Cephalophora Cav., 

Dugaldia Cass., Espeletiopsis Sch.-Bip. ex Benth. 

& Hook. in syn., Craemia Hook., Hecubaea DC., 

Heleniastrum Fabricius, Leptophora Raf., Leptopoda 

Nutt., Mesodetra Raf., Oxylepis Benth., Tetrodus 

Cass.), * Hyrnenoxys Cass. (including Actinella Juss. 

ex Nutt. nom. rej., non Pers. nec Lewis nom. 

cons., Phileozera Buck1 . , Picradenia Hook., Rydbergia 

Greene), *Macdougalia A. Heller, * Platyleima (A. 

Gray) Cockerell, *Plurnrnera A. Gray, *Psathyrotes 

(Nutt.) A. Gray, *Psilostrophe DC. (including Riddellia Nutt.), * Tetraneuris Greene (including Actinella Pers. nom. rej., non Lewis), *Trichoptilium A. 

Gray. 

Annual, biennial, or perennial herbs or small 

shrubs: caulescent or rarely acaulescent; leaves 

alternate or basal, petiole usually not well delimited, sometimes lacking. Inflorescence often scapose or subscapose with one or a few heads, more 

rarely with heads in densely corymbose clusters. 

Heads usually heterogamous, sometimes discoid; 

involucre with herbaceous bracts in ca. 2 or 3 

series, usually subequal; receptacle plane or convex to subglobose, usually alveolate without 

paleae, sometimes with a few bracts between rays 

and disk, more or less fimbrillate with short-conical to elongate spinelike fimbrillae in Gaillardia, 

or with spinose crests in Balduina. Ray flowers 

when present usually female, sterile in many species of Gaillardia and some Helenium; corolla limbs 

often with elongate rugose cells adaxially, usually 

strongly trilobed at tip, rays papery and remaining attached to achene in Baileya and Psilostrophe. 

Disk flowers usually hermaphroditic, functionally 

male in Plummera; corollas with inner surfaces of 

lobes somewhat papillose, often with glands on 

outer surface; anther thecae pale; median endothecial cells more elongate, often elliptical, 

with 1 or 2 thickenings on end walls; anther 

appendage without glands. Branches of fertile 

styles with paired stigmatic lines. Colored resin 

not evident. Achenes prismatic, with wall not 

carbonized, cells of wall with distinct raphides; 

cells of seed coat not ornamented, ovules containing distinct raphides; pappus with various shaped 

awns or squamellae, with 5 paleae dissected above 

into numerous bristles in Tric-hoptilium, with numerous coarse to fine bristles in 1-4 series in 

Psathyrotes. Pollen 30-40 pm in diameter. 

The subtribe is widely distributed in the Western Hemisphere with a concentration in the drier 

parts of western North America. Some species of 

Heleniurn, Gaillardia, and Hymenoxys occur in South 

America in the region from Peru southward. A 

few species are widely cultivated and adventive. 

This typical element of the Bentham tribe 
SMITHSONIAN CONTRIBUTIONS TO BOTANY 

Helenieae is now widely accepted as an integral 

part of the Heliantheae (Stuessy, 1977; Turner 

and Powell, 1977). Stuessy (1977) expanded the 

traditional concept of the subtribe to include such 

genera as Balduina, which had been placed in the 

Galinsoginae, and Psilostrophe and Baileya, which 

had been placed in a separate subtribe Riddelliinae by Hoffman (1890-1894). The Stuessy concept of the Gaillardiinae correlates closely with 

the distribution of the character of noncarbonized 

achene walls, a character not used by Stuessy in 

deriving his concept. 

In treating the same genera, Turner and Powell 

(1977) differ from Stuessy (1977) in a number of 

details, including their retention of a separate 

subtribe Psilostrophinae (Riddelliinae). The latter group is traditionally distinguished by the 

persistent, papery ray corollas that fall with the 

achenes, but the two genera also differ by the 

achene walls having closely spaced, fibrous costae. 

The discoid genera Psathyrotes and Trichoptilium 

were not included in the Gaillardiinae by Stuessy 

(1977). Both genera have broad cells covering the 

surface of the achene with dark granular inclusions in the lumina, unlike other genera of the 

subtribe. The pappus of Trichoptilium is a set of 

five squamellae, each dissected distally into a 

series of numerous bristles. Psathyrotes has a pappus with bristles of similar structure which are 

borne separate to the base and are borne in more 

than one series. The two forms are undoubtedly 

related, although the pattern of bristle attachment is totally different. The Psathyrotes pappus is 

remarkably similar to that seen in the genus 

Psathyrotopsis in the Chaenactidinae. Other characters indicate the genera, one with carbonized 

achene walls and the other without, are not so 

closely related as once believed. Nevertheless, it is 

possible that the tendency for reduced carbonization seen in the typical Chaenactidinae has 

given rise to two separate lines that totally lack 

carbonization in the achene walls. 

The specialized sesquiterpene lactones, the helenanolides, are notable for their near restriction 

to the subtribe Gaillardiinae. They have been 

reported in the genera Baileya, Balduina, Gaillardia, 

Helenium, Hymenoxys, and Tetraneuris (Herz, 1973), 

a distribution closely supporting the concept of 

the subtribe by Stuessy (1977). Helenanolides also 

have been reported from Arnica of the closely 

related subtribe Chaenactidinae and from two 

genera, Geigeria Griessel. and Inula L.. of the 

closely related tribe Inuleae (Herz, 1973). 

The distinctive polyacetylenes, the epoxvsulfones, seem to be restricted to the genera Gaillardia 

and Helenium (Bohlmann et al., 1973). Other less 

specialized sulfones also are present, but such 

nonepoxidized forms occur elsewhere in the 

Heliantheae, as indicated under the Flaveriinae. 

There is no support from this study for the 

suggestion by Stuessy (1977) that the Gaillardiinae are closely related to the Helianthinae. The 

Stuessy suggestion was based on such characters 

as the large solitary heads, the alternate or basal 

leaves, and the tendency toward elongate receptacles found in the Gaillardiinae and various 

elements that he included in the Helianthinae. 

Rudbeckia and Gaillardia are particularly similar in 

superficial appearance, but all other evidence 

dictates against close relationship. 

The name Gaillardiinae has priority at the 

subtribal level over the name Heleniinae. Until 

recently, however, I have believed the name 

Heleniinae would take priority when used within 

a segregate tribe Helenieae. This was apparently 

true under older versions of the International Code 

of Botanical Nomenclature as reflected by Solbrig 

(1963), but by present versions of the Code (Stafleu et al., 1978), such priority for subtribal names 

based on the type genus of the tribe occurs only 

in the subtribe containing the type genus of the 

family, i.e., the subtribe Asterinae in the tribe 

Astereae. It should be noted that names of subtribes such as the Helianthinae could be displaced 

if older names are found. 

Subtribe 35. MARSHALLIINAE 

FIGURE 210 

Subtribe MARSHALLIINAE H. Robinson, Phytologia, 4 1 :42. 

1978. 
NUMBER 31 89 

*Marshalha Schreb. nom. cons., non J. F. Gmel 

(including Persoonia Michx. nom. rej., non J. E. 

Smith, nom. cons. nec Willd., Phyteumopsis Juss. 

ex Poir., Trattenikia Pers. non Trattinickia Willd.). 

Perennial herbs; leaves alternate or basal. Inflorescence scapose or subscapose, with 1-few 

heads. Heads discoid; involucral bracts herbaceous, subequal, in 2-3 series; receptacle convex 

or conical, with rigid linear paleae. All flowers 

disciform, hermaphroditic; corollas lavender or 

becoming violet; outer surface with numerous 

uniseriate hairs, often with interspersed glands; 

basal tube long and slender, throat very short; 

lobes linear, papillose on inner surface, with expanded cushion or firm papillae on outer surface 

at tip; anther thecae slightly to strongly violet; 

endothecial cells short, with 2 or 3 thickenings on 

transverse walls; anther appendages ovate, 

strongly carinate, without glands. Style branches 

with paired stigmatic lines, style tip subtruncate 

or with a short blunt appendage, outer and lateral 

surfaces distally with numerous, long, blunt papillae. Ducts and resin not evident. Achenes turbinate, with 3 main costae and 5 weaker intercostal veins, with setae and glands, walls not 

carbonized, with laxly quadrate or oblong cells, 

raphides minute; carpopodia symmetrical, annuliform, with cells wider than high; seed coat 

with cells of one of the layers quadrate, not 

ornamented; pappus of 3 lanceolate or triangularovate, acute squamellae, with outer surface scabrid. Pollen ca. 40 pm in diameter. 

The single genus of the subtribe, Marshallia, is 

endemic to the eastern United States. 

The Marshalliinae are remote from other subtribes of the Heliantheae. Uncarbonized walls of 

the achene are found in only one other subtribe, 

the Gaillardiinae, but otherwise there is no evidence of close relationship. The Gaillardiinae and 

the subtribes to which it is related are all basically 

epaleaceous, whereas the Marshalliinae have 

prominent rigid paleae of a distinctive form. In 

the Heliantheae, the presence of paleae seems to 

be correlated with some asymmetry of the carpopodium, but in Marshallta the carpopodium is 

expanded and symmetrical with larger cells than 

in other members of the tribe. 

There have been recent suggestions that Marshallza belongs in the tribe Eupatorieae (Stuessy, 

1977; Turner and Powell, 1977), but the keeled 

anther appendage with a constricted base, and 

the polarized thickenings of the endothecial cells 

are totally foreign to that tribe. The uncarbonized 

achenes, which are unusual in the Heliantheae, 

are even more unusual in the Eupatorieae, and 

the latter tribe never has well-developed raphides. 

The relationship of Marshallza seems to be reflected better in the recent discovery of prenyl 

flavonoids in M. grandglora Bead. & Baynt. (Bohlmann et al., 1979). These unusual flavonoids are 

known elsewhere in the family only in Flourensia 

and Xanthzum of the Heliantheae and in Helichrysum of the Inuleae. These are the two tribes with 

which Marshallta shares structural features. In the 

keeled anther appendages, colored anther thecae, 

and the pappus of five lanceolate squamellae, 

Marshallza seems closest to the Heliantheae, and 

it is retained in the tribe in the present study. 

Nevertheless, the lack of carbonized walls in the 

achene is characteristic of the Inuleae, and short 

to medium-length tails can be seen on the bases 

of anthers in some species of Marshallia. Careful 

redefinition of the limits between the two tribes 

may eventually show that the Marshalliinae can 

be best accommodated in the Inuleae. 

Because of the isolated position of Marshallza 

compared to other members of the Heliantheae, 

the genus with one chromosome report of n = 9 

(Jones, 1970) cannot be assumed to share the 

polyploid ancestry suggested for other members 

of the tribe (Robinson et al., in press). 
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1979. 
Index 

(Synonyms and page numbers of main entries of taxa of Heliantheae in italics) 

.I hU.\U/(JU. 48 

Acanthambrosia, 32 

Acanthosperrnurn. 14. 15. 35. 38. 73 

.4canthoxanthium, 32 

Acarphaea, 84 

;lchaenopodium, 49 

Achyrachaena. 76. 78 

Achyropappus. 84, 86 

Acicarphaea, 84 

‘4ciph-J lluea, 70 

Acmella. 13, 16 (fig. 12). 21. 48, 49. 53, 

.5 5 

Aroma, 64 

Arlinea, 87 

Actinella. 87 

Actinella Juss. ex Nutt., 87 

Actinella Pers.. 87 

Actinomeris, 49 

Actinospermum. 87 

Adelmannia, 48 

Adenolepsis, 64 

Xdenopappus. 70, 72 

.Adenophyllum. 70. 72 

.‘I denospri mum. 64 

Adenothamnus. 76 

.4dwntina, 60 

.4gariippea, 60 

Agarisra, 64 

A,gar?sta. 64 

.-l,gar sir la, 8 7 

Agiabampoa. 55, 57 

Alarronia. 49 

‘4 lcina, 3.7 

Xldama. 55, 57 

hlepidocline. 24, 60. 61 

Allocarpus, 60 

Alloisperrnum, 6. 15, 60, 61, 63 

hispidum. 36 (figs. 47. 48) 

caracasanum, 16 (fig. 16), 59, (figs. 

148, 149) 

Alomia. 25, 80 

‘4Itamirania, 49 

‘41vordia. 13. 55. 57 

Arnauria. 80. 81 

Amauriinae, 80 

rotundifolia, 77 (fig. 190) 

Amauriopsis, 84 

Amblyolepis. 87 

.4mblyopappus, 81 

pusillus, 82 

Ambrosia. 1, 32. 34 

trifida. 33 (fig. 25) 

Ambrosiaceae, 32 

Amhrusiear, 24. 32 

Xnibroii&es, 2 

Ambrosiinae. 5, 8, 9, 10, 20. 25, 27. 31. 

,4mtda, 76 

A tnph icu lea. 58 

Anacis, 64 

;lnai/is. 4.9 

Ancistrophora, 48 

Andrieuxia, 48 

Angelphyturn. 48 

,4nisucarpus. 76 

Anomostephium, 48 

Anthernideae. 10. 21. 22. 23, 24, 34 

Antheropeas, 81 

Aphanactis. 60 

Aphanopappus, 48 

Arctotideae. 25 

Argyroxiphiurn. 76 

;Irmania, 55 

Arnica, 5, 10, 24, 26, 84, 86, 88 

Arnicastrum, 76, 84, 86 

.4romia, 81 

Aspilia. 48. 51 

Aslemma, 3. 8. 48 

Asterales, 10 

.4stereae, 7. 10, 11, 24, 25. 39, 68. 76. 

88 

Asterinae. 88 

Asteroideae, 8. 10, 11, 13, 17, 22, 23, 

61 

Atriplex. 68 

Axiniphyllum. 39 

32. 34. 42. 49, 57 

cordifolia, 85 (fig. 204) 

Baccharis corymbosa, 7 

Baeria, 67. 81. 84 

spicata, 7 

Baeriinae. 6, 10, 15. 20. 21. 22. 30, 32. 

Baeriopsis. 81 

Bahia. 84, 86 

Bahiinae, 4. 32, 84. 86 

Bahiopsis, 55 

Baileya. 87. 88 

206) 

79. 81. 82. 84 

guadalupensis. 83 (figs. 192. 193) 

absinthifolia. 83 (fig. 199) 

niultiradiata. 16 (fig. 20). 85 (fig. 

Ballirria, 42 

Balbisia. 61 

Balbisia, 61 

Balduina. 61. 87. 88 

Balsamorhiza, 8. 48. 51. 57 

Baltirnora, 38. 48. 51 

Barratio, 5j 

Bartlettia. 24, 84. 86 

Bart o iina, 6 I 

Batiasa, 60 

Bebbia. 5. 58. 60, 61, 74 

juncea. 62 (fig. 150) 

Berlandiera. 48, 5 1 

Bidens. 14, 20. 64. 65. 68 

pilosa. 62 (fig. 156) 

Bikera. 60 

Blainvillea. 48 

Blaincillea subg. Oligoune. 48 

Blepharipappus. 8. 76 

scaber. 71 (fig. 176) 

Blepharizonia. 76 

Boebera, 70 

Boeberastrum, 70 

Borrichia. 9. 13, 23, 48. 51, 53 

Bothriocline. 7 

Brasilia, 58 

Brassavola, 87 

Brassavola, 87 

Brauneria, 48 

Brotera, 72 

Brotera, 72 

Bruchidae. 23 

Burrielia, 81, 84 

frutescens. 47 (fig. 90) 

sickii, 59 (fig. 141) 

96 
NUMBER 51 97 

Cacosmia, 76 

Calea, 8, 9, 14, 26, 27, 39, 58, 60, 61, 

76 

belemii, 58 

caleoides, 59 (fig. 140) 

eitenii. 12 (fig. 1) 

janiaicensis, 59 (fig. 139) 

skutchii. 60 

Caleacte, 58 

Calebrachys, 58 

Calhounia, 55 

Callzachyris, 76 

Callichroa, 76 

Calliglossa, 76 

Calliopszs, 64 

Calonnea, 87 

Calycadenia, 76, 78 

Calycoseris. 67 

Calydermos, 60 

Calyptocarpus, 9, 48. 51, 65 

Campklotheca, 64 

Cardueae, 23. 67 

Cargi 1 la, 48 

Carphojtephium, 61 

Carramboa. 34. 35 

Carthamus. 67 

Centromadia, 76 

Centrospermum. 35 

Centrospermurn, 35 

Cephalobembix, 84 

Cephalophora, 87 

Cercostplos, 87 

Chaenactidinae, 9, 10, 14, 15, 20, 26, 

30, 32. 65, 75, 76, 79, 81, 84, 86, 

87. 88 

vialis. 50 (fig. 100) 

Chaenactis, 84, 86 

Chaenocephalus, 49 

Chamaechaenactis. 10: 84, 86 

Chamaestfphanum, 84 

Chatiakella, 49 

Cheilodiscus, 70 

Chiliophyllum. 46 

Chiliophyllum, 46 

Chilodia, 49 

Chlampsperma, 78 

Charisma, 32 

Chromolaena, 67 

Chromolepis, 46 

Chrysactinia, 70. 72 

Chyanthellzna, 64 

Chrysanthellum, 64. 68 

Chrysogonum, 48, 51 

douglasii, 85 (fig. 202) 

heterophylla, 47 (figs. 80, 81) 

virginianum, 50 (fig. 91) 

Chrysomelea, 64 

Chrysophania, 46 

Chrysostemma, 64 

Chthonia, 70 

Chuquiraga opposirifolia, 7 

Chylodia, 49 

Cichorieae, 69 

Cichorioideae, 10, 13 

Clappia, 11, 26, 58, 74, 75, 76 

suaedaefolia, 71 (fig. 173) 

Clappiinae, 8, 14, 26, 31, 32, 69, 74, 

Clibadiinae, 14, 26, 28, 31, 34, 39, 40, 

Clibadium. 11, 20, 27, 38, 39, 42 

surinamense, 41 (figs. 66-69) 

Clomanocoma, 70 

Closia, 80 

Coespeletia, 34 

Coinogvne, 75 

Collaea. 64 

Collaea, 64 

Colubogpne, 48 

Comaclinzum, 70 

Coreocarpus, 64 

Coreopsidaceae, 64 

Coreopsideae, 64, 67. 76 

Coreopsidees. 2 

Coreopsidinae. 1, 4, 5, 8, 9, 11, 14, 15, 

75. 76 

42. 43, 74 

20. 21. 22, 23, 24, 25, 26. 30. 32, 

39. 40. 44. 48, 51, 55, 64, 65, 67, 

68, 69: 82, 84, 86 

Coreopsoides, 64 

Coreopsis, 1. 64. 65, 68 

Coronocarpus, 48 

Correllia, 80, 81 

Cosmzdium, 64 

Cosmobruchus, 23 

Cosmophyllum, 48 

Cosmos, 1, 9, 23, 64 

Coulterella. 4, 69. 70 

Coulterellinae, 30, 32, 69 

Crassina, 49 

Crockeria, 81 

Cvphiospermum, 44 

Cr).;btopetalun, 70 

Cuchumatanea, 60 

Cyathodine jaquemontii, 10 

Cyathomone, 64 

Cyclachaena, 32 

Cymophora, 60 

lanceolata, 62 (figs. 154, 155) 

capitata, 66 (figs. 163, 164), 69 

Dahlia, 1, 10, 20, 23, 64, 65, 68 

coccinea, 20, 66 (figs. 159, 160) 

merckii, 20 

Dahlibruchus, 23 

Deinandra, 76 

Delilea. 9. 39, 42, 48. 51 

Delucin, 64 

Desmanthodiinae, 11. 13. 26, 27, 31. 

39, 40 

Desmanthodium, 39, 40, 42 

Desmocephalum, 48 

Dicahmma, 48 

Dichaeta, 81 

Dicoria, 32, 34 

Dicranocarpus, 38. 64. 65 

Diglossus, 70 

Dilepis, 72 

Dimeresia, 24, 6.?, 64 

howellii, 62 (fig. l53), 63 

Dimeresiinae. 6. 30, 31. 63. 64 

Dimerostemma. 48, 49 

Diodonta, 64 

Diodonlium, 64 

Diomedea, 48 

Diotostephus, 48 

Diplosastera, 61 

Diplothrix, 49 

Dipterotheca, 48 

Discomela, 55 

Ditrichum. 49 

Ditrichum, 49 

Doronicum, 5 

Dracopis. 45 

Dubautia. 61. 76 

Dugaldza, 8 7 

Dugesia. 17. 48, 5 1 

Dyscritothamnus, 13. 25, 58. 60. 61, 74 

I)ysodiopJiJ, 70 

Dysodium, 35 

Dyssodia, 70. 72 

perfoliatum, 41 (figs. 60-63) 

canescens. 33 (figs. 29. 30) 

laxa. 87 (fig. 181) 

filifolius. 62 (fig. 152) 

lopez-mirandae. 72 

Eatonella, 81. 82 

Echinacea, 11, 21, 45. 46, 48. 49, 51, 

congdonii, 83 (fig 196) 

53 

atrorubens, 54 (fig 118) 

Echinocephalum, 48 

Echinomerza, 55 

Echinops, 7 

Eclipta, 8, 48, 51, 73 

Ecliprinae, 5, 6, 8, 9, 11, 13, 14, 15, 21, 

26, 28, 29, 31, 32, 34. 39, 42, 43, 
SMITHSONIAN CONTRIBCTIONS TO BOTANY 

44. 45. 46, 48, 49, 51, 53, 55, 57, 

60. 61, 65, 68 

Eisenmannia, 48 

Electra, 64 

Electra, 64 

Eleutheranthera, 15, 48 

Elrsrra, 48 

Enalcida, 70 

Encelia, 8, 14. 15. 48, 49, 51, 53, 55, 57 

canescens, 54 (fig. 129) 

Enceliopsis, 48, 53 

Engelmannia, 34. 38, 48, 51 

pinnatifida, 50 (fig. 92) 

Engelmanniinae, 5, 9. 34, 48. 51 

Enhydra. 44, 51 

sessilis, 41 (fig. 76) 

Enhydrinae, 30, 31, 44 

Epallage, 25 

Epileprs, 64 

Ereminula, 63 

Ericentrodea, 64, 65 

Eriocoma, 44 

Eriocoma, 44 

Eriopappus, 76 

Eriophyllinae, 81 

Eriophyllum, 81 

Eryngiophyllum, 64, 68 

Espejoa, 75, 84, 86 

mexicana, 83 (fig. 197) 

Espeletia, 7, 34, 35 

grandiflora, 35 

killippii. 36 (figs. 36-39) 

ruderalis, 16 (fig. 17), 52 (fig. 11 1) 

Espeletiinae, 6, 27, 31, 34, 35, 38 

Espeletiopsis, 35 

Espeleliopsis, 87 

Eupatorieae, 4, 5, 7, 8, 10, 13, 14, 15, 

Eupatoriophalacron, 48 

Euphrosyne, 32, 34 

Eutetras, 80; 81 

Euxenia. 48 

Exomiocarpon, 48, 5 1 

23. 25, 26, 61, 63. 67, 80, 86, 89 

parthenifolia, 33 (fig. 3 1) 

palmeri, 77 (fig. 191) 

Faxonia, 60, 61, 63, 78 

Feaea, 60 

Feaella, 60 

Fenixia, 48, 51 

Ferdinanda, 46 

Fingalta, 48 

Fitchia, 4, 67, 68, 69 

pusilla, 63 

nutans, 66 (fig. 162) 

speciosa, 68 

Fitchiinae, 4, 11. 30, 32, 65, 68, 69, 86 

Flaveria, 4, 68, 72, 73 

linearis, 16 (figs. 13, 19), 72 

trinervia, 71 (figs. 168, 169), 72 

Flaveriinae, 14, 15, 22, 26, 31, 32, 68, 

69, 72. 73, 75, 76, 88 

Florestina. 10, 84, 86 

Flourensia, 11, 48, 53, 89 

Fornicaria, 49 

Fougeria, 48 

Fougerouxia, 48 

Franseria, 32 

Gaertneria, 32 

Gaillardia, 5, 8, 22, 73, 87, 88 

Gaillardiinae. 4, 5! 9, 10, 13, 14, 15, 22, 

25, 26, 3 1 32, 49, 79, 86, 87, 88: 89 

Galeana. 78. 79 

pratensis, 77 (figs. 186, 187) 

Galinsoga. 60, 61 

filiformis, 63 

quadriradiata, 59 (fig. 145) 

Galinsoginae. 3, 5, 6, 9, 11. 13, 14, 15, 

20, 25, 26, 30, 31, 40, 49, 57, 58, 

60. 61, 63, 64, 74, 78, 88 

Galophthalmum. 48 

Garcilassa, 51, 55, 57 

Geigeria, 88 

Geissolepis, 25 

Geissopappus, 8, 14, 26, 58. 60, 76 

Georgina. 64 

Geraea, 48, 53 

Glossocardia. 64, 68 

Glossogyne, 64, 68 

Gnaphalium. 67 

Gnaphalopsis, 70 

Goldmanella, 64, 67 

Goldmania, 64 

Goldmania, 64 

Gonotheca, 61 

Graciela, 70 

Graemia, 8 7 

Greenmania, 58 

Greenmaniella, 58, 60 

Guardiola, 9, 15, 38, 43, 44, 65, 67 

mexicana. 16 (fig. 15), 41 (fig. 75) 

papifera, 41 (fig. 74), 44 

Guardiolinae, 9, 28, 3 1, 43, 67 

Guen lheria, 8 7 

Guerreroia, 64 

Guizotia, 20, 22, 39, 40, 65, 67, 73 

abyssinica, 22 

scabra, 37 (fig. 59) 

Gymnolaena, 70, 72 

Gymnolomia, 48 

Gynactis, 64 

Gynoxys, 7 

Halea, 61 

Hamulium, 49 

Haplocalymma, 55, 57 

Haploesthes, 24, 26, 72, 73 

Harnackia. 70 

Harpaecarpus, 76 

Harpa lium. 55 

Harpephora, 48 

Hartmannia, 76 

Hecubaea, 87 

Heleniaceae. 87 

Heleniastrum, 87 

Helenieae, 2. 3, 4, 5, 8, 10, 24, 26, 60, 68, 

Hdldnides, 2 

Heleniinae, 87, 88 

Helenium, 8, 87. 88 

Heiepta, 48 

Hdlianthkes, 2 

Helianthella, 48, 51, 53 

uniflora, 54 (fig. 119) 

Helianthinae. 4, 5, 8, 9, 11, 13, 14, 20, 

22, 25, 26, 29, 31, 40, 45, 46, 49, 

51, 53, 55, 57, 61, 64, 69, 88 

greggii, 71 (fig. 171) 

73, 74, 79, 82, 87, 88 

autumnale, 85 (fig. 209) 

Helianthopsis, 29, 55 

Helianthus, 1, 11, 13, 22, 40, 53, 55, 67 

Helichroa, 48 

Helichrysum, 67, 89 

Helicta, 48 

Heliogenes. 60 

Heliomeris, 13, 55, 58 

Heliophthalmum, 45 

Heliopsis, 21, 48, 49. 53, 68 

Helioreos, 70 

Helogyne, 14 

Hemiambrosia, 32 

Hemizona 76, 78 

angustifolia, 76 

Henricksonia, 64, 65 

Heptanthinae, 27, 31, 43, 46, 48 

Heptanthus, 43, 46, 48 

cochlearifolius, 47 (fig. 84), 48 

Herodotia, 7 

Heterosperma, 23, 64, 65 

annuus, 22, 56 (fig. 136) 

buphthalmoides, 47 (figs. 85, 86) 

pinnatum, 16 (fig. 18), 66 (figs. 157, 

158) 

Hidalgoa, 64, 65 

Hingst on ia , 49 

ternata. 66 (fig. 161) 
NITMBER 51 99 

Hinglsha. 44 

Hoffmanniella. 48. 5 1 

Holocarpha. 76, 78 

Holo,gmne. 81. 84 

Holozonia, 76 

Hopkirkia DC.. 84 

Hopkirkia Spreng.. 49. 84 

Hulsea. 8 1. 84, 86 

Hybridella, 46 

Hy-dropectis. 70 

Hymenatherum. 70, 72 

Hymenoclea, 32, 34 

salsola. 33 (fig. 28) 

Hymenopappinae. 1.5, 30, 32. 78, 79 

Hynienopappus. 78. 79 

filifolius. 77 (fig. 182) 

newberryi, 8. 30, 78 

Hymenostephium, 55. 57 

Hynienothrix. 84. 86 

Hymenoxys. 87, 88 

Hypericophyllum. 75. 84. 86, 87 

multicaule. 83 (fig. 200) 

Ichthyothere. 9, 13. 20, 28, 35, 39, 42, 

68, 78 

In fantea. 81 

Inula. 88 

Inuleae, 7. 10, 13. 23. 24. 25, 64, 67, 88, 

Iaotephane, 55 

Isocarpha. 5, 61 

Iso5tigma, 64, 68 

Iva. 20. 32, 34, 57 

ILeae. 72 

scandens. 37 (fig 50) 

89 

Jaegeria. 60, 61, 63 

hirta, 59 (fig. 146) 

Jamesianthus, 84, 86 

Jaumea, 65. 67. 75 

carnosa. 71 (fig. 175), 75 

linearifolia, 75 

Jaumeinae. 14, 31, 32, 65, 67, 74, 75, 76 

Kallias, 48 

Kegelia. 48 

Kernerza, 64 

Kingianthus, 6, 46, 48, 55 

Kleinia, 70 

Kletnia, 70 

Koehneola. 43 

Lactuceae, 4, 67, 69 

Lagascea, 4. 5, 7, 13, 55, 57, 69 

Lagasceinae, 4. 55, 69 

mollis, 56 (figs. 130, 131) 

Lagophvlla. 76 

Lantanopsis, 39, 42 

Laphamia. 80 

Lapsana. 23 

Lasianthaea. 48. 51 

Lasthenia, 10. 17, 21. 22. 67, 68, 81, 82, 

hispidula. 41 (figs. 70-72) 

fruticosa, 52 (figs. 105. 106) 

84 

chrysantha. 82 

chrysostoma. 82. 83 (fig. 194) 

coronaria, 82 

glaberrima. 82 

glabrata, 82 

leptalea, 82 

microglossa, 82. 84 

minor. 82 

platycarpha. 82 

Latreillea, 35 

Laxmannia. 64 

Laxmannia. 64 

Layia. 76. 78 

discoidea. 76 

glandulosa. 77 (figs. 178, 179) 

Leachia, 64 

Lebetina, 70 

Lecocarpus, 35 

Lezghia. 55 

Lembertia. 81 

Leontophthalmum, 58 

Lepachys, 45 

Lepia, 49 

Lepzdostephanus. 76 

Leptocarpha, 48, 53 

Leptophapx. 80 

Leptophora. 87 

Leplopoda. 8 7 

Lescaillea, 70 

Leucactinia. 70, 72 

LeuciiNa, 32 

Liabeae. 9, 25. 58, 76 

Libanothamnus, 35 

Limnogenneton, 39 

Lindheimera, 48, 51 

Linsecomia, 55 

Lipochaeta. 48 

Lipotriche, 48 

Lorentea Lag., 49, 70 

Lorentea Orteg., 49, 70 

Lowellia, 70 

Loxexeta , 8 4 

Loxothysanus, 78, 79 

Luciliopsis, 7 

Lundellianthus, 6, 48, 51 

pinnatifidus, 36 (figs. 41-43) 

pedunculatus, 77 (fig. 183) 

guatemalensis, 50 (figs. 101, 102) 

L\capsinae. 28. 32. 79, 80 

Lycapsus. 25% 79. 80 

tenuifolius, 77 (fiq 188). 80 

Macdougalia. 87 

,ifacella. 60 

Macraea. 48, 53 

Marrocarphus. 84 

..Madaria, 76 

,.Madariopsis. 76 

Madia, 76. 78, 80 

,l.ladieae. 76 

Madiinae. 3. 6. 8. 9. 26. 30. 32. 76, 78. 

80 

.2.ladorella, 76 

Malacothrix. 67 

Mallotopus. 24. 84, 86 

,Mandonia, 61 

Marshallia. 5, 9, 13. 14, 25, GI. 89 

laricifolia. 54 (figs. 125. 126) 

glomerata, 71 (fig. 177) 

graminifolia. 85 (fig. 2 10) 

grandiflora. 89 

Marshallinae. 13, 15, 2.5, 31. 32. 88. 89 

Megalodonta. 64 

Melampodiinae. 4. 5. 8. 9, 14. 15, 26, 

28. 31, 34. .?5. 38. 39, 40, 42. 44. 

45. 51. 73. 78 

Melampodium. 8. 10, 14, 20. 35% 38. 40 

appendiculatum. 36 (fig. 46) 

divaricatum. 36 (figs. 44, 45) 

microcephaluni, 16 (fig. 11) 

aspera. 52 (fig. 110) 

Melanthera, 48. 33 

..Melarhira. 49 

.Ilelastomataceae. 10 

Menderin, 49 

Menotriche. 49 

Meratia. 48 

Meratia. 48 

Mesodetra, 87 

,l.leyera, 44 

Meyeria, 58 

Micrelium, 48 

Microchaeta. 48 

Microcoecia. 48 

Microdonta, 64 

Microlecane, 64 

Microspermum, 25 

Mierza. 84 

Milleria, 20. 39 

Milleriinae. 5, 8. 9, 20, 2,5, 27, 29, 31, 

32. 39. 40, 43. 45, 51, 53, 65. 67, 73 

Mill&ndes. 2 

Minyranthes, 39 

quinqueflora. 37 (figs. 53. 54) 
100 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

Mirasolia ij 

.MneJ ileon. 87 

Llorinna. 38 

.Monactis. 48, 55 

dubia. 3 

pallatangensis. 54 (fig 128) 

Monolopia. 81. 82 

.L!unophuliJ 48 

Monothrix. 80 

.Montagnaea. 44 

Montanoa, 8. 13. 14, 44, 45 

tonientosa, 41 (fig 77) 

hlontanoinae. 8, 19. 3 1, 44. 45 

hloonia. 38. 64. 65 

hlunnoLia. 9 

hlutisieae. 7. 8. 9, 69 

Narialina. 64 

Needhamia. 64 

.Veedhamia 64 

,Veilreichia. 60 

.\eothimopsis. 84 

polbantha. 84 

.Vesothamnus. 80 

.2euenbergia. 72 

.Veuractis. 64 

Seurolaena. 4% 24. 26. 58, 60. 61 

.Veurolaeneae, 58 

Neurolaeninae. 5. 9. 11, 13. 14. 26, 28, 

29. 31. 39. 14. 46, 55% 58. 60, 61, 

67. 76 

Nicolletia, 70. 72 

Kiebuhria. 49 

.!iebuhria. 49 

.Vocca. i.5 

.Votoptera, 48 

lobata, 56 (fig 137) 

Obeiisrothera. 4.7 

Orhronelis. 49 

Ogiera. 48 

Oliguone subg , Bluiniillea. 48 

Olicaea. 25, 76 

Oparanthus. 64 

Orcpa. 75 

Orochaenactis. 84. 86 

Orsinia. 41 

Ostnadenia, 76 

Osaaldia. 42 

OteiLa. 60. 61 

OthaXe, 84 

Otopappus, 48 

Ox\ carpha. 48 

cerbesinoides, 52 (figs 114-1 16) 

saedaefolia 54 (fig 11 7) 

Oxpiepis. 87 

Oxcpappus. 10, 81. 82 

Oxtlenia. 32 

Oxlura. 76 

O\edaea, 8, 48. 51 

seemannii. 83 (fig 195) 

Palafoxia, 10. 84. 86 

PalaJoxiinae. 84 

Puleista. 48 

Paleolaria, 84 

Pallasia. 48 

Pallasia. 48 

Pallassia. 48 

Panicum, 68 

Pappobolus. 55 

Pappothrix. 80 

Partheniaceae, 32 

Parthenice. 32. 34. 38 

Parthenium. 32. 34, 38 

Puscaiia, 49 

Pectidinae. 15. 22. 30. 32, 49. 68. 69. 70, 

Pertidium. 70 

Peclidopsis. 70 

Pectis. 68, 70. 72 

sphacelata. 83 (fig. 198) 

mollis, 33 (figs. 32. 33) 

hysterophorus. 33 (figs. 34. 35) 

73. 74. 75 

linifolia var. hirtella, 72 

seasiliflora, 71 (fig. 167) 

Pen ta lep i s . 64 

Peramibus. 45 

Perezia. 9 

Pericome. 80. 81 

caudata. 16 (fig. 14) 

Perityle. 80, 81. 82, 84 

emoryi. 77 (fig. 189): 80. 81 

Peritylinae. 10. 15. 30, 32, 79. 80, 81, 

Persoonia. 89 

Persoon ia , 89 

Perymeniopsis, 15, 48, 53 

Perymenium, 48, 49. 53 

Petrobiinae. 64 

Petrobium, 8, 64, 65 

Peucephyllum, 24, 84, 86 

schottii, 85 (fig. 201) 

Phaethusa, 49 

Philactii. 48. 53 

Phiieorera. 87 

Philoglossa. 38 

Phoebanthus, 48, 53 

82. 81 

ovalifolia. 52 (figs. 107: 108) 

grande. 52 (figs. 112. 113) 

l’@teumupJis. 89 

l’irraden ia. 8 7 

t’irroderiicipsi~. 84 

Pinillosa. 39. 43. 53 

Pinillosinae. 28. 31. 39. 4.3 

Plagiolophus. 48. 53 

Plar).carpha. 84 

Plaryleima. 87 

Pia ( yp teri s . 49 

Platyschkuhria. 84. 86 

Plummera. 87 

Podachaenium, 6. 48. 49. 53. 60 

eminens, .54 (figs. 121, 122) 

Podanthes. 48 

Podanthus, 8. 48, 53 

mitiqui. 54 (fig. 124) 

Polymnia. 13. 38. 39 

canadensis. 37 (fig. 51) 

Polptnriias~rum. 38 

Polyrnniinae. 28. 31, 38. 39 

Polrpteris. 84 

I’nntesia. 42 

Porophyllum. 70. 72 

Prionocarpus. 5.7 

l’ronacron. 58 

Prototypes. 2 

l’sathurorhaeta. 48 

Psathyrotes. 13. 25. 26. 86, 87, 88 

Psathyrotopsis, 25, 84, 86. 87. 88 

Pseudobahia. 81 

Pseudobartiettia. 84 

Pseudoclappia. 25, 26. 74 

arenaria. 71 (fig. 174) 

Psilostrophe. 87. 88 

tagetina, 85 (fig. 205) 

I”srioJ/ruphinae. 13, 87. 88 

PtilomeriJ, 81. 84 

Ptilostephium. 61 

Pyrethrum. 48 

l!yrethrum, 48 

Pyrrhopappus, 67 

berterii. 41 (fig. 73) 

annua. 8.5 (fig. 205) 

Ralllardella. 25. 76 

Kailiardeilinae, 76 

Ram ti lla. 3 9 

Kancaguu, 81 

Ratibida. 45 

Renmnia. 8. 9 38. 48, 51 

Rhvsolepis, 5.7 

Ridan. 49 

Kiddellin. 87 

columnifera, 41 (fig 79) 

salvadorica. 12 (fig 9). 50 (fig 96) 
NUMBER 31 101 

Riddelliinae. 87. 88 

Riencourtia. 39. 42 

Rigiopappus. 4, 25 

Rojasianthe, 9. 45. 48, 49. 53 

Rosilla. 70 

Rothia. 78 

Hothia, 78 

Rudbeckia. 1, 5. 20. 22, 45, 46. 57. 88 

Rudbeckiees. 2 

Rudbeckiinae. 8, 10. 29, 31. 45. 46, 57, 

RuilopeLia, 35 

Runifordia. 79. 40 

Rydbergia. 87 

laciniata. 41 (fig 78) 

74 

floribunda, 37 (fig 58) 

Sabazia. 60. 61 

acoma, 63 

liehmannii, 63 

Salmea. 48. 53 

Salmeopsis. 49 

Salmia, 49 

Sanvitalia. 49, 53 

Sani icaiiopsis. 49 

Sartwellia, 72. 73 

Saubinetta. 4.9 

Scalesia. 55. 57 

Schistocarpha, 4. 9. 24. 26. 58. 60. 61. 

auberrii, 47 (fig. 89) 

niexicana, 71 (fig. 170) 

affinis. 56 (fig. 132) 

63 

bicolor. 12 (fig. 2) 

eupatorioides 59 (fig. 147) 

Schizophylluni, 48 

Schtsophyllum. 48 

Schizoptera. 9. 38. 4.9. 51 

peduncularis, 50 (fig. 97) 

Schizotrichia. 70. 72 

eupatorioides. 72 

Sckuhriu. .79> 84 

Schkuhria, 10. 39, 84. 86 

Schlechtendalia. 70 

Schlechtendalia. 70 

Sclerocarpus. 55. 57 

.Sculospermum. 48 

Seala. 70 

SebaJtiania, 64 

Selloa. 60. 63 

Selleophylum, 64 

Senecio. 4 

Senecioneae. 2, 3, 4, 5. 7. 10. 11. 13, 17. 

Serpaea. 48 

24, 2~5? 26, 51, 58. 67. 68, 73. 74 

Sheareria, 25, 39 

Sigesbeckia, 39, 40 

Silphium. 45. 51 

Simsia. 53, 55. 57, 67 

Smallanthus, 6, 8, 9, 13, 14, 23, 35, 38, 

orientalis, 37 (fig. 57) 

sanguinea. 56 (fig. 135) 

39. 78 

Sobreyra, 44 

Sobrya, 44 

Sogalgina, 61 

Solanum, 10 

Solenotheca, 70 

Sphagneticola, 49. 55 

Spilanthes, 5, 13, 49, 53, 61 

urens, 53: 54 (fig. 127) 

Stachycephalum, 39, 40. 42 

argentinum, 41 (figs. 64, 65) 

Stammarium, 70 

Staurochlamys. 58, 60. 65% 67 

burchellii. 56 (fig. 138) 

Steiractinia. 45, 51, 53 

mollis. 52 (fig. 109) 

Stemmatella. 60 

Stemmodontza, 49 

Stenocarpha. 60 

Strotheria. 70, 72 

Stuessya, 55 

Splesia, 84 

Sjncephalantha, 70 

Syncretocarpus, 55 

Synedrella, 9, 49. 51, 65 

Synedrellopsis. 9, 49, 5 1 

uvedalia, 37 (fig. 49) 

sericeus. 56 (fig. 134) 

nodiflora, 12 (fig. 8), 50 (figs. 94. 95) 

grisebachii. 12 (fig. lo), 50 (figs. 98, 

99) 

Syntrichopappus, 67, 68, 84 

Tageteae. 2, 5, 24, 70. 72 

Tagetes, 1, 70, 72 

Tagetinae. 70. 72 

Tagerinees, 2 

Tamania, 27, 35 

Telesia. 48 

Tepion, 49 

Tetracarphum, 84 

Terrachyron, 58, 60 

Tetragonosperma, 61 

Tetragonotheca. 57, 60 

subulata. 66 (figs. 165, 166) 

chardonii, 36 (fig. 40) 

orizabensis. 59 (figs. 142. 143) 

texana, 59 (fig. 144) 

Tetraneurinae, 87 

Tetraneuris, 87, 88 

Tetranthus. 39, 43, 49. 53 

Tetrads, 44 

'Tetraperone, 43, 46. 48 

Tetrodus, 87 

Thelechitonia. 49 

Thelesperma, 64 

Thy mop hy 1 la, 70 

Thymopsis. 84. 86 

Tilesia, 49 

Tithonia, 20. 55. 67 

Tollatia, 76 

Torrent ia, 35 

Tragoceras, 49 

Trattentkia. 89 

Trattinickia. 89 

Tricarpha, 60 

Trichocoryne. 30. 78. 79 

Trichoprilium. 87, 88 

Trichosrema. 49 

Trichostemma. 49 

Trichoslephium, 457 

Trichymenia. 84 

Tridax. 8, 10. 12 (fig. 5). 20. 57. 58. 61, 

thymoides, 85 (fig. '703) 

connata. 30 

incisum, 85 (fig. 207) 

63 

balbisioides. 13 

brachylepis, 13, 61 

procumbens. 62 (fig. 15 1) 

tambensis, 13 

Trigonopter'rm. 48 

Trigonospermurn, 39 

Trimeranthes, 39 

Trimetra. 48 

Trixis. 42 

Trixis. 42 

Tulocarpus. 43 

Tyleropappus, 58 

melampodioides. 37 (figs. 55, 56) 

1 kacou, 49 

13dra. 44 

Cnxia. 11. 13. 28, 39. 58. 60 

camphorata, 37 (fig 52) 

Lrbinella. 70. 72 

Lrophora. 22 

\'argaiia. 60 

Chrgasia. 60 

arilla. 8. 11, 26 58, 73. 74. 75. 80 

mexicana, 12 (fig 3). 71. (fig 172), 

73 
SMITHSONlAh CONTRIBUTIONS TO BOTANY 

texana 73 

\ arillinae, 14% 30. 32, 58 73, 74, 75 

Vasargta, 60 

Ihsquezia, 78 

\ enegasia, 9. 65, 67, 76, 84, 86 

Lerbesina. 14, 15, 20, 49 53 

\irginica, 54 (fig 123) 

TkrbeJininae, 5. 32, 39, 45, 48 49, 51. 55, 

Vermifuga, 72 

\ ernonia, 7 

harlingii, 7 

,parrei, 7 

arellata. 7 

Vernonieae, 7, 10 

l’eslingm, 39 

\ igethia 49, 51 

I’igoiina. 60 

\ iguiera 5.5, 57. 58. 67 

57. 65 

dentata, 9. 56 (fig 133) 

porteri. 58 

sylvatica, 57 

\’illanova, 78, 79 

titicacensis, 77 (figs. 184, 185) 

Villanova, 32, 78 

Vilobia, 70, 72 

Cirletia. 84 

Wahlenbergia, 44 

Wahlenbergia, 44 

h‘edelia. 8, 34, 49, 51, 53 

calycina, 50 (fig. 93) 

Welswitschiella, 87 

M‘hitneya, 25, 84. 86 

M’iborgia. 60 

Wiborgin, 60 

Wilkesia, 76 

M.’I rtgenia, 48 

Wollastonia, 49, 51. 53, 55 

CZ’uerJchmittia. 48 

W‘ulffia. 49, 53 

gymnoxiphium, 77 (fig. 180) 

Wyethia, 4, 1 I, 49. 5 1 

helenioides. 54 (fig. 120) 

Xanthium, 1, 32, 34, 89 

Xantho. 81 

Ximenesia, 49 

strumarium. 33 (figs. 26, 27) 

Zaluzania, 46, 58 

angustata, 47 (figs. 82, 83) 

grayana, 46 

Zaluzaniinae, 1 I, 29, 31, 46 

Zarabellia. 35 

ZemiJnP, 55 

Zexmenia, 49, 51, 53 

mikanioides, 49 

serrata, 52 (figs. 103, 104) 

grandiflora, 47 (figs. 87. 88) 

Zinnia: 49, 53 

Zinniinae, 32, 48, 51, 53 

Zycona. 60