SMITHSONIAN CONTRIBUTIONS TO BOTANY NUMBER 54 

A Generic Review of 

the Tribe Liabeae (Asteraceae) 

Harold Robinson 

SMITHSONIAN INSTITUTION PRESS 

City of Washington 

1983 
ABSTRACT 

Robinson, Harold. A Generic Review of the Tribe Liabeae (Asteraceae). 

Smithsonian Contributions to Botany, number 54, 69 pages, 16 figures, 1983.-The 

neotropical tribe Liabeae is treated with approximately 157 species aligned in 

15 genera and three subtribes, the Liabinae containing Austroliabum, Cacosmia, 

Chionopappus, Ferreyranthus, Liabellum, Liabum, Microliabum, Oligactis, and Sinclairia; Paranepheliinae containing Paranephelius and Pseudonoseris; and Munnoziinae containing Chrysactinium, Erato, Munnotia, and Philoglossa. The latter 

two subtribes are formally described herein as new. The tribe is placed in the 

subfamily Cichorioideae, and distinctions from the Vernonieae are discussed. 

A phylogeny of the tribe is proposed with probable central Andean origin. 

Various elements of the tribe are traced phyletically and geographically. The 

genera are seen as unusually well isolated from one another compared to 

genera in most of the more successful tribes of the Asteraceae. The apparent 

restriction of reticulating evolution to the early development of the Liabeae is 

suggested as a reason for the comparatively limited success of the tribe. 

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 Cerctdiphyllum japonicum Siebold and Zuccarini. 

Library of Congress Cataloging in Publication Data 

Robinson, Harold Ernest. 1932- 

.4 generic review of the tribe Liabeae (Asteraceae) 

(Smithsonian contributions to botany ; no 54) 

BibliograDhv: D. 

Includis index: 

1. Compositae-Classification. 2. Botany-Classification. I. Title. 11. ‘Title: Liabeae, (Asteraceae) 111. Series. 

QKl.S2747 no. 54 [QK495.C74] 581s [583’.55] 82-10807 
Contents 

Page 

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

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

Tribal Status and Evolutionary Considerations ....................... 3 

Description of the Tribe Liabeae . ............................. 14 

Keys to the Genera of the Liabeae ................................... 16 

Artificial Key to the Genera of the Liabeae ........................ 16 

Technical Key to the Genera of the Liabeae ....................... 18 

The Subtribes and Genera of the Liabeae ................. 

Subtribe LIABINAE ...................................... 

Austroliabum ..................................................... 20 

Cacosmia ...... ................................. 22 

Chionopappus ..................................................... 25 

Ferreyranthus .................................................... 27 

Liabellum ........................................................ 29 

Liabum .................. ........................... 31 

Microliabum .............. ..................... 

Oligactis ......................................................... 37 

Key to the Subgenera of Oligactis ...................... 38 .. Sinclairia ......................................................... 41 

PARANEPHELIINAE, new subtribe .................................... 44 

Paranephelius ..................................................... 45 

Pseudonoseris ..................................................... 47 

MUNNOZIINAE, new subtribe ........................................ 49 

Erato ........................................................... 52 

Munnotia .............. 

57 

Philoglossa ...................................................... 59 

Species Excluded from the Tribe ................................. 62 

Literature Cited ..................................................... 63 

Index ................................................................ 66 

.. ChTsactznium .................................................... 49 

Key to the Subgenera of Munnotia ............................ 

... 

111 

A Generic Review of 

the Tribe Liabeae (Asteraceae) 

Haro Id Rob inson 

Introduction 

The tribe Liabeae consists of approximately 

155-160 species in 15 genera that are restricted 

to the Neotropical Region. Compared with other 

tribes of the family, the Liabeae are inaccessible 

to centers of taxonomy in the temperate and 

colonial areas, having no representatives in either 

eastern North America or in Brazil in eastern 

South America. The most accessible element of 

the tribe during the earlier periods of taxonomic 

study was the small series of Liabum species in the 

Greater Antilles, and this partially explains the 

inadequate attention given to the group in the 

classical treatments of the Asteraceae. The Liabeae Lz’as not recognized as a tribe until the time 

of Rydberg (1927). Until the various papers derived from the work of S.F. Blake (Blake, 1935; 

Cabrera, 1934: Sandwith, 1956), the various elements of the Liabeae were distributed in four 

different tribes, the Senecioneae, Heliantheae, 

Helenieae, and Mutisieae. All aspects of the tribe 

have been in need of revision. The present review 

follows some preliminary efforts published during 

the last 10 years, the two most basic being Robinson and Brettell (1974b) and Robinson (1978a). 

The revised concepts were briefly covered in the 

recent symposium treatment by Nordenstam 

(1977a). 

Species of the Liabeae first appear in the literHarold RobinJon, Department of Bolanj, ‘Vatzonal Museum of ,Ihtural 

History, Smilh ionran Inriitulion, Washing(on, D.C 10360 

ature as members of genera of the Astereae, such 

as Solidago and Amellus. A West Indian species was 

the first to be recognized as representing a distinct 

genus, Liabum, by Adanson (1 763). A number of 

additional generic names were established during 

the next half-century, including Munnozia from 

Peru and various synonyms of Liabum from Central America and the northern Andes. 

The most critical examination of members of 

the tribe before this century was by Cassini (1823, 

1823, 1830). who treated the members as a section 

of the tribe Vernonieae. The genera recognized 

at that time were Liabum Adanson (including 

Andromachia Humboldt, Bonpland, and Kunth 

and Starkea Willdenow), Munnozia Ruiz & Pavon, 

Cacosinia Humboldt, Bonpland, and Kunth, and 

Oligactis Cassini. The characters cited by the various authors or by Cassini distinguished Munnozia 

by its chaffy receptacle, Cacosmia by its lack of a 

pappus, and Oligactis by its few disk flowers and 

double pappus. 

The work of Lessing (1832) treated the Liabeae 

as a subtribe of the Vernonieae. adding one genus, 

Alibum Lessing, and relegating Munnozia to 

“Genera minus cognita.” This disposition was 

followed in part by DeCandolle (1836), but a 

number of mistakes were introduced, especially 

in regard to two new genera. Erato DeCandolle 

was described as a new genus in the Asteroideae 

in the relationship of Psiadia (Baccharis), and Philoglossa DeCandolle was described in the Senecionideae in the relationship of Rudbeckia and Encplia, 

1 
2 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

neither being recognized as relatives of Liabum or 

as relatives of each other. Also, two asterean 

elements were included in the division Liabeae, 

destroying the naturalness of the group that was 

maintained by Cassini and Lessing. 

Paranephelius was described by Poeppig and 

Endlicher in 1843 and was placed near Liabum by 

Weddell (1853- 1837). The latter work also raised 

section Chrysactinium Humboldt, Bonpland, and 

Kunth to generic rank and resurrected Andromachia (including Munnozia). All the elements of the 

Liabeae included in the work of Weddell were 

together in a natural group. 

The work of Bentham and Hooker (1873) established a tradition lasting for over a hundred 

years, but, unfortunately, these authors treated 

the primarily neotropical groups poorly, especially the Liabeae. All elements of the Liabeae 

having a capillary pappus, except the poorly 

known Allendea Llave and Lexarza, were placed 

together in a single genus, Liabum, and transferred 

from the related Vernonieae to a highly artificial 

tribe Senecioneae. In the Senecioneae, Liabum and 

Allendea were placed in a subtribe, Liabeae, with 

the helianthean genera Schistocarpha Lessing and 

Neurolaena R. Brown and the vernonian genus 

Gongrothamnus Steetz. All elements of the Liabeae 

with a different form of pappus were placed in 

different tribes. Philoglossa DeCandolle was 

placed in the subtribe Melampodiinae of the 

Heliantheae, and the new genus Chionopappus 

Bentham, having a plumose pappus, was placed 

in the Mutisieae. Cacosmia Humboldt, Bonpland, 

and Kunth, which had previously been recognized as a relative of Liabum, was placed in the 

Helenieae. The treatment of the Liabeae by Bentham in Bentham and Hooker (1873) must be 

considered totally incompetent, but was accepted 

with few changes along with other parts of the 

general treatment of the Asteraceae by Hoffmann 

in Engler and Prantl (1890-1894), and has been 

perpetuated with little question into the present 

century by such authorities as Cronquist (1955). 

Even as the tradition prevailed in general 

works, progress was being made in isolated studies 

of the Liabeae. In ,Vorth American Flora, Rydberg 

(1927) established the tribal status of the group 

and recognized numerous generic segregates in 

the Mexican and Central American species, most 

notably Sinclairia Rydberg and Liabellum Rydberg. Blake (1935) described the unnamed species 

on which Bentham in Bentham and Hooker 

(1873) had based his genus Chionopappus, and at 

the same time Blake pointed out the true relationship to Liabum. By the time that Cabrera 

(1934) described his Liabellum (= Microliabum Cabrera), he was able to mention the relationships 

of both Cacosmia and Chionopappus to Liabum. 

Sandwith (1956) described a new species of Philoglossa, at which time he cited an unpublished 

manuscript of Blake in which the liaban nature 

of that genus was recognized. 

Recently, this author has been involved in a 

series of studies of the tribe (Robinson, 1976- 

1982; Robinson and Brettell, 1973, 1974; Robinson and Cuatrecasas, 1973): but the major results 

are summarized in two papers. In the first, 

Robinson and Brettell (197413) placed all the 

elements together as a distinct tribe containing 

14 genera. In the second paper, the treatment of 

the tribe in the Flora of Ecuador, Robinson (1978a) 

resurrected one additional genus, Erato, and 

treated the 29 species of the country with illustrations of seven of the eight genera. There remain 

a number of recent additions and refinements in 

the tribe since the first summary paper that were 

not included in the Ecuadorian study, however. 

The present paper provides a more cornplete 

review of the classification and structure of all 15 

genera presently known in the Liabeae and presents evolutionary and geographical information 

not previously discussed. 

ACKNOWLEDGMENTS.-A number of people are 

to be thanked for their contributions to this study. 

The drawings have been prepared by Alice Tangerini of the Department of Botany, and the 

study has benefitted by comments from Jose Cuatrecasas, Vicki Funk, and R.M. King. The assistance of Rebecca D. Brettell in the early phases of 

the work is also greatly appreciated. 
NUMBER 54 3 

Tribal Status and Evolutionary Considerations 

The Liabeae have deeply lobed disk corollas, 

long-spurred anther bases, continuous stigmatic 

surfaces on the inside of the style branches, and 

somewhat irregular distribution of spines on the 

pollen. These features mark the tribe as a member 

of the subfamily Cichoriodeae, a relationship 

shared with the Lactuceae, Vernonieae, Mutisieae, Cardueae. Echinopsideae, Gundelieae, Ereniothamneae, and Arctotideae. Bentham’s placement of the Liabeae in the Senecioneae was based 

on the presence of a capillary pappus and heterogamous heads, but the Senecioneae often have 

short-lobed disk corollas, nonspurred anther 

bases, often paired stigmatic lines on the style 

branches, and regular distribution of spines on 

the pollen, characteristics of the subfamily Asteroideae. In the Cichorioideae, the Liabeae compare most closely to the Vernonieae and some 

Lactuceae by the hairs on the abaxial surfaces of 

the style branches. The Liabeae differ from either 

of these tribes and from most other Cichorioideae 

by their opposite leaves and the radiate heteroga nio u s heads . 

There has beec a tendency by recent authors 

(Nash, 1976: Jansen and Stuessy, 1980) to return 

the Liabeae to the position assigned to them by 

Cassini (1823. 1823, 1830) within the tribe Vernonieae. The position of the two groups together 

in the Cichorioideae is not questioned here, but 

it is not accepted that the two are necessarily each 

other’s closest relatives within the subfamily. A 

marked phyletic gap exists between the two 

groups, and each tribe by its basic characteristics 

falls completely outside of the basic characteristics of the other tribe. One of the character differences, the heterogamous heads, was sufficiently 

marked to cause Bentham’s removal of the Liabeae to the Senecioneae. Equally marked are the 

opposite leaves found in all Liabeae but found in 

only a few specialized Vernonieae. The leaf 

blades of many Liabeae are strongly trinervate, 

a feature never seen in the Vernonieae. The corollas of the Liabeae are usually yellow, whereas 

those of the L’ernonieae are mostly reddish or 

purplish. Only Gongrothamnus Steetz in the Vernonieae is said to have yellowish flowers. The 

Liabeae have a lax form of areolation of endothecia1 cells rarely found in the Vernonieae, and the 

few subquadrate examples of endothecial cells in 

the Liabeae are specialized with details of thickenings different from those of other tribes. The 

Liabeae lack the polygonal arrangement of spines 

or ridges on the pollen that is basic to the Vernonieae (Robinson and Marticorena, in prep.), 

and the style branches are obtuse to rounded at 

the tip, usually not tapering as in the Vernonieae 

(see below). The presence of latex seems basic to 

the Liabeae, a character found in the Lactuceae 

and a few Mutisieae but not found in the Vernonieae. In other chemistry, the Liabeae show no 

compounds that are particularly suggestive of the 

Vernonieae (Robinson et al., 1980). The cytology 

of the Liabeae falls outside of the pattern seen in 

the Vernonieae (Jones, 1977: Robinson et al., in 

prep.), There are no characters that are uniquely 

shared by the two tribes Liabeae and Vernonieae, 

and there are no characters by which the combined tribes could be distinguished from the Lactuceae. It is believed here that combination of the 

two tribes misrepresents the phyletic significance 

of the two groups and ignores extremely marked 

differences exceeding those that distinguish the 

Vernonieae from other tribes such as the Lactuceae. The relationship between the Liabeae and 

Vernonieae does remain useful as a basis for 

phyletic discussion. 

It should be emphasized that the tribe \. ’ernonieae includes more genera and more species and 

is regarded as having more subtribes than the 

Liabeae, and it has a much wider geographical 

distribution, but it does not have markedly 

greater diversity in its basic characteristics (Robinson et al., 1980). In the latter respect the tribes 

are nearly equivalent. 

The tribe Liabeae is similar in size to the 

paleotropical tribes Arctotideae and Calenduleae, 

which occur mostly in the African region. The 

restriction of the Liabeae to the Neotropical Region encourages the idea of relationship to \.ernonieae, the latter being well represented in that 
4 SMITHSONIAK CONTRIBUTIONS TO BOTANY 

area; however, any divergence between the two 

tribes could not have been very recent. The 15 

genera of the Liabeae show a considerable diversity and display some internal phyletic gaps. The 

genera Philoglossa and Erato, which were sufficiently distinct to have been placed in association 

with various members of the Heliantheae by 

various authors (DeCandolle, 1836: Bentham in 

Bentham and Hooker, 1873), are only one part of 

the three subtribes of the Liabeae that are recogniLed herein. Further insights are gained by 

more detailed study of some individual characteristics of the tribe. 

THE ~\NTHERs.-T~~ form of the anther is one 

of the features that marks the Liabeae as a member of the subfamily Cichorioideae. The appendage is not constricted at the base, and the bases 

of the thecae extend well below the point of 

attachment to the collar. Such anther bases present problems in both observation and terminology. In observation, the true length of the extended anther bases is often underestimated, because the upper part of the filament (collar) lies 

in the plane of the anther thecae. On superficial 

examination, insertion seems to be at the base of 

the collar, but the actual point of attachment is 

at the upper end. 

The term for such anthers is also subject to 

confusion. The three terms sometimes used are 

"dorsifixed," "tailed." and "hastate." The first, 

"dorsifixed," is inaccurate, since the collar is attached at the base of the connective and only the 

thecae extend below that point. The term 

"tailed" is undesirable for the Liabeae, since 

tailed anther bases such as those in the Inuleae 

refer to sterile basal extensions. In a few Mutisieae, rarely in the Inuleae and to a slight extent 

in Chionopappus and Ferreyranthus in the Liabeae, 

there is a tail in addition to a fertile part that 

extends below the attachment to the collar, but 

it seems important to have a term that will allow 

distinction between the various forms. The third 

term, "hastate," is also undesirable, since it is also 

used for the very different short type of anther 

base found in many Heliantheae, and also it 

implies a pointed tip, which most Liabeae and 

other Cichorioideae do not have. Because of this 

problem, Jose Cuatrecasas has suggested (pers. 

conim.) adoption of the term "calcarate" or 

"spurred" for the type of anther base that is 

characteristic of the many tribes of the subfamily 

Cichorioideae. In some of the cases mentioned 

above, the anther base would be both calcarate 

and tailed, 

The anther bases in the Liabeae are all calcarate, but there is variation in the ornamentation 

on the base. In the Paranepheliinae and Munnoziinae, the base is essentially plain. In the 

Liabinae, the base of each theca is often fringed 

by projecting cells, and in Chionopuppus and Ferrgranthus, this fringe is borne on a short but 

distinct sterile portion of the theca. In F. zwbascifolius, the short tail is particularly noticeable as a 

fringed coil next to the strongly papillose filament. 

The thecae of the anthers in the subtribe Munnoziinae of the Liabeae show a black coloration 

that is characteristic of almost all members of the 

subtribe. The black color is deposited on the 

outside surface of the valves. although it often 

seems to follow the pattern of the endothecial 

cells. The color is particularly prominent in long 

narrow bands in Erato and Philoglossa. Such coloration is not found elsewhere in the Liabeae but 

is found elsewhere in the Asteraceae, being most 

common in the subfamily Asteroideae in the Heliantheae. It also occurs in a few other isolated 

genera such as Blepharispermum Wight ex DeCandolle of the Inuleae. The occurrence in the 

Munnoziinae is the only example presently 

known in the Cichorioideae. The black color is 

not always well developed in less mature material, 

but it is lacking in the subtribe Munnoziinae in 

only Munnozia hastfolia and its closest relatives, M. 

maronii and M. subuiridis. 

The median endothecial cells show considerable variation in the Liabeae, more than in any 

other group in the family of comparable size. The 

extensive variation appears to fall into a phyletic 

pattern (Figure 1). The variations are primarily 

in the form of the sclerified shields on the inside 

surfaces of the endothecial cells and in the pattern 
NGMBER 34 

M unnozia 

C h rysact in ium 

B 

Philoglossa 

Erato a I 

A 

a 

I Liabum Austroliabum Sinclairia 

iictiy 

I 

I 

Ferreyran th us I 

I I I I1 

Paranephelius 

Pseudonoseris 

I11 

FI(:~,KE I .-Suqgested evolutionary scheme for the Liabeae with illustrations of representative 

enclothecial cells: I. Munnoziinae Lvith anther thecae black, bases of anther thecae unfrinqed. 

and cells of achene \\.all with quadrate raphids: 11. Liabinae with anther thecae pale and bases 

of anther thecae fririged or tailed: 111, Paranepheliinae with anther thecae pale, cells of acherie 

mall with elongate raphids, style branches long. spines of pollen regularl2- disposed. basal 

baculae vestiqial or absent. habit rosulate or subrosulate. (A = stiff hairs with bulbous base\. 2 

1 anqles on achcne; B = spines of pollen regularly disposed; c = cells of anther collar Lvith dense 

aniiiilar rhickeniiigs: 11 = pappus absent: E = anther thecae tailed: F = latex absent; c; = pappus 

brihtles plumose. paleae present: H = cells of achene wall with quadrate raphids. style 1)ranches 

of disk flo\\-ers usually long: I = Argentina and southern Bolivia; J = klexico and Central 

:lmerica: K = with tuberous root: L = heads sessile. bases of anther thecae not fringed.) 

of thickenings or extensions of these shields onto 

the periclinal walls between the endothecial cells. 

In the Munnoziinae, the pattern of the shields 

and their thickenings is different in each of the 

genera. In Chysactinium, thickenings are often solitary at the lou.er ends of short cells. In Munnotia, 

the thickenings vary from solitary at the upper 

and lower ends and strongly arching inwards, to 

niultiple and scattered, or sometimes essentially 

lacking. In Munnorza subgenus Kastnera, the endothecial cells provide the best example of a welldeveloped radial pattern of thickenings in the 

tribe, with a number of thickenings on both 

vertical and transverse walls. In Erato and Philoglossa, where elongate black deposits on the thecae are particularly notable, the thickenings of 

the endothecial cells are lacking from the transverse walls, and the cells form distinct series. In 

Phzloglossa, the individual series of short cells are 

accentuated by the dense thickenings along the 

vertical walls, but in Erato, such thickenings are 

very weak, and the endothecial cells are difficult 
6 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

to see against the background of the black deposits. The Munnoziinae shows a comparatively high 

proportion of its members with the sclerified 

shields filling the width of the endothecial cells 

with thickenings on the vertical walls. These include Erato, Philoglossa, and many atypical members of Munnozia such as the subgenus Kastnera. 

In the remaining two subtribes of the Liabeae, 

the endothecial cells, with few exceptions, have 

sclerified shields distinctly narrower than the endothecial cells. In these, which are similar to the 

endothecium type in typical Munnozia and Chrysactiniurn, the distinct strap-shaped or oval sclerified shields on the inner surface, themselves, superficially look like discontinuous cells. The 

shields are often plain, but in some genera such 

as Chionopappus and Austroliabum, the shields are 

ornamented with lines or pockets of greater or 

lesser thickness. The thickenings on the walls 

between the cells are usually few and are often 

on very long extensions of the shield. Usually only 

one thickening occurs at the end of a cell, but two 

or three closely spaced extensions, in a laciniate 

pattern, are not rare. In Ferreyranthus, Sinclairia, 

Liabellum, and Paranephefius, the endothecial cells 

are notable for having long extensions with nodular thickenings on only one end, these usually 

arching strongly inward. There is a tendency for 

the nodular points in Paranephefius to be more 

common on the lower ends of the cells. In Liabum 

and Ofigactis, the shields are similar to those of 

related genera but more often fill most of the cell 

and usually have single or double thickenings at 

both ends that more closely abut with those of 

adjacent cells. 

In Cacosrnia, which has a habit and leaf surface 

like Ferreyranthus, there are shields in the endothecium that more completely fill the cells, and 

there are scattered thickenings that are often on 

vertical walls. In many thecae of Cacosrnia, there 

are continuous annulations in the endothecial 

cells that are restricted to one end of the cell. The 

form in Cacosrnia is unlike those in other members 

of the subtribe Liabinae, but it approaches the 

forms seen in various members of the Munnoziinae. 

It would be useful to assign a phyletic direction 

to the many patterns of variation in the endothecia1 cells of the Liabeae, but this is not easily 

done. Functional considerations are of little help, 

since the normal function of endothecial thickenings is mostly replaced in the Asteraceae by the 

plunger action of the style. In fact, the broad 

median band of endothecial cells in many Liabeae is probably more important as a weakened 

area in the valve and not capable of providing 

torsion that would help open the valve. The basic 

endothecial cell, as seen in most families of angiosperms, is short with well-developed annulations. Such cells are closely approached in the 

Asteraceae in the Eupatorieae, a tribe showing 

other comparatively unspecialized features within 

the family (Robinson, 1981). The endothecial 

cells are also of a radial form, but weaker and 

completely incapable of function, in most Vernonieae. The latter tribe also shows comparatively unspecialized features, though not as consistently as the Eupatorieae. .4 simple extension 

of the idea of the primitiveness of the radial 

pattern would imply that the broader shields with 

thickenings on the lateral walls in three genera of 

the Munnoziinae were ancestral in the subtribe, 

and Cacosmia would be interpreted as the most 

primitive element in the subtribe Liabinae. 

Actual evidence seems to indicate a different 

evolutionary sequence. The broad endothecial 

shields in the Munnoziinae are of two different 

types and can be interpreted as two separate 

specializations. Those of Erato and Philoglossa are 

not truly radial but are transversely polarized, 

and in Munnozia subgenus Kastnera, the endothecia1 type that most closely approaches the truly 

radial form is in an unquestionably derived element within the genus. 

The alternate form of endothecium, with 

weakly connected shields, presents a different picture with very similar types in Chrysactinium and 

typical Munnozia of the Munnoziinae, in all of the 

Paranepheliinae, and in most of the Liabinae. 

Being farthest from the primitive radial form in 

structure, the weakly connected shields should be 

regarded as specialized in the family. Still, it is a 
NUMBER 54 7 

specialized form that has been attained by such 

diverse members of the Asteraceae as many cacalioid Senecioneae, such subtribes as the Chaenactidinae and Gaillardiinae in the Heliantheae, 

and even Adenoon Dalzell of the Vernonieae. It is 

easiest to assume that the genetic basis for such 

polarized types of endothecial cells was firmly 

established in the Asteraceae at an evolutionary 

level above that of the Eupatorieae but below the 

level of most other tribes such as the Heliantheae 

and Liabeae. Some of the variations as seen in 

Erato and Philoglossa can be interpreted as little 

more than rotations of the axis of polarization in 

shortened endothecial cells. Close examination of 

the endothecial cells in Munnozia subgenus Kastnera shows that they are mostly interpretable as 

having an oblique axis of polarization. Even the 

more completely radial types in some other tribes, 

such as in Dahlia L. and Rudbeckia L. of the 

Heliantheae, would seem to be reversions from 

polarized types, although the mechanism is not 

entirely clear. 

On the basis of present evidence, the endothecia1 form in Cacosmia should be regarded as specialized from more polarized types. Nevertheless, 

the difference in the endothecial type, combined 

with evidence from other characters, would still 

suggest a position for the genus comparatively 

remote from others in the subtribe Liabinae. 

The apparent ease of reversals in endothecial 

cell type in the Asteraceae seems to confirm the 

absence of any significant selective pressures regarding the structure in the family. 

The anther appendage is without notable specializations in the Liabeae except in Oligactis 

subgenus Oligactis. In this subgenus, the ends of 

the cells project from the surface to form distinct 

short papillae. The rest of the Liabeae, including 

Oligactis subgenus Andromachiopsis, have the normal smooth anther appendages. 

THE STYLE.-The structure that has proven to 

be most important in all the classical studies of 

the Asteraceae is the style, and the style remains 

the one most valuable complex of characters for 

the evaluation of relationships of the Liabeae. 

The undivided stigmatic surfaces on the inside of 

the style branches and the hairs on the backs of 

the branches and on the upper shaft immediately 

mark the tribe as a part of what is here recognized 

as the subfamily Cichorioideae sensu Carlquist 

(1976) including the Vernoriioideae Turner ex 

Jansen and Stuessy (1980). The close similarity 

in detail of the styles of the Liabeae and the 

Vernonieae is the principle reason for the combination of the two tribes by various authors 

(Cassini, 1823, 1825, 1830; Turner and Powell, 

1977; Jansen and Stuessy, 1980), and it is worthy 

of a detailed review. 

The style of the Asteraceae is a highly functional structure, and it must be treated as such in 

phyletic speculations. In the family, the basic 

function of receiving pollen is complicated by the 

specialization of the style as a plunger for pushing 

pollen out of the encircling ring of anthers. In 

spite of various short-term evolutionary variations, the basic patterns of stylar evolution in the 

Asteraceae use the style to expel pollen but at the 

same time to prevent automatic self-pollination. 

This is done in three basically different ways. 

The first and least effective method of preventing self-pollination in styles of the Asteraceae 

involves the development of an enlarged sterile 

stylar appendage that pushes the pollen from the 

styles ahead of the emerging stigmatic surfaces. 

This form is most highly developed in the tribe 

Eupatorieae of the subfamily Asteroideae. The 

development of such an appendage may be related to the origin of the sterile band that separates the stigmatic surface into two lines on each 

style branch, a separation that is basic in the 

whole subfamily Asteroideae, excluding some specialized groups. 

The second and most effective method of preventing self-fertilization is the separation of the 

flowers into two types, those with anthers and 

sterile styles serving as plungers and those with 

fertile styles but no anthers. This design is most 

common in the subfamily Asteroideae above the 

level of the Eupatorieae. Included are various 

Heliantheae, Inuleae. Calenduleae. Astereae, Anthemideae, and Senecioneae with partial to complete separation of flower types in the same heads 
8 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

or inflorescences or on separate plants. Such separation is rare in the Cichoroideae but is found in 

an incomplete form in the Arctotideae and in 

incomplete to complete forms in the Mutisieae. 

The third form of protection of the stigmatic 

surface in the Asteraceae is basic to most Cichorioideae. The stigmatic papillae are completely 

enclosed between the young branches as the style 

elongates and pushes through the ring of anthers. 

The surfaces do not become exposed until the 

style is exerted and the branches begin to separate. Such enclosed stigmatic surfaces are comparatively rare in the subfamily Asteroideae, 

where they occur as apparent reversions in specialized groups such as the subtribe Helianthinae 

of the Heliantheae and the cacalioid members of 

the Senecioneae. 

The functional aspects of the style in the Cichorioideae insure. or at least allow for, the basic 

similarity of the style structure in the subfamily. 

The stigmatic surface is continuous and enclosed 

in all members of the subfamily, and the entire 

outer surface is available for pollen-collecting 

hairs or scabrosity extending downward onto the 

shaft. Hairs are restricted to the apex in only a 

few specialized members of the Cichorioideae 

such as the Nassauviinae of the Mutisieae. For 

the rest of the subfamily. variation tends to be 

restricted to two aspects, the sharpness of the 

lower boundary of the scabrous or hairy surface 

and the proportion of the width of the styleplunger mechanism, which is formed bv the hairs 

or scabrae. Thicker styles as in the Arctotideae, 

Cardueae, and many Mutisieae need only short 

scabrae to collect the pollen, whereas thinner 

styles such as those of the Vernonieae, Liabeae, 

and Lactuceae need longer hairs to achieve the 

same function. It is notable that the tribes in 

which the upper style is thickened to serve as a 

plunger also tend to have a sharper lower boundary of the scabrous collecting surface at the point 

where the style becomes narrower. From this 

simple functional point of view, the co-occurrence 

of narrower styles with longer pubescence in the 

Vernonieae, Lactuceae, and Liabeae cannot be 

regarded as such a certain sign of relationship. 

The actual evolutionary history of the style 

form in the Cichorioideae is not certain from 

present evidence. There is no evidence that ancestors of the subfamily ever had a stylar appendage 

or a divided stigmatic surface, and there is no 

value in speculation on that topic, but it would 

be useful to determine whether the ancestral style 

was closer to thai of the Arctotideae with a broad 

tip and short scabrae or to that of the Vernonieae 

with a narrow tip and longer hairs. Both style 

types occur with various head types, homogamous 

or heterogamous, and both are associated with 

various types of pollen. Variations of the St) le in 

the phyletically diverse Mutisieae raise the question of stability of the style form at the tribal 

level. Also, Pseudostzfftzu H. Robinson shows strong 

evidence of a position in or near the L’ernonieae, 

though it has a broad tip on the style. The 

narrower form of style with longer hairs also 

occurs anomalously in the Eremothamneae, 

which has heterogamous heads and a capillary 

pappus but which proves most closely related to 

the Arctotideae as indicated by Leins (1970) and 

Norlindh (1977). The style of Eremothumnzis remains distinct from the other narro\t type4 in the 

subfamily by the bicellular hairs, which are divided by a longitudinal wall. It seems certain that 

both the broad and narrorv style types in the 

Cichorioideae have arisen more than once. For 

these reasons I regard the designation of a primitive style type in the subfamily impossible at this 

time, but the situation is sufficiently complex that 

possible separate origins of narrow style forms in 

the L7ernonieae and Liabeae are worthy of further 

investigation, 

It should be noted that, aside from the narrow 

form, the details of the styles in the Liabeae and 

Vernonieae are basically different. A long tapering and pointed style branch is basic to the 

Vernonieae, with only a few specialized genera 

such as Centrutherum having less-tapered or shortpointed style branches. In the Liabeae, the style 

branches in most genera are comparatively short, 

often less than half as long as the hispidulous part 

of the style shaft, and the tips are narrowly obtuse 

to rounded. Longer style branches occur in the 
NUMBER 34 9 

Liabeae but seem restricted to two groups that 

are considered here as specialized in the tribe, the 

generic pairs Liabum-Oligactis and ParanepheliusPseudonoserzs. Of these, only the latter genus-pair 

has a notable tapering of the branches, and none 

have tips as sharp as in the Vernonieae. Both in 

Liabum and in the ray flowers of Paranephefius, 

there is a tendency for the long style branches to 

be spiralled, a tendency not known in the Vernonieae. 

The length of the style branches is not subject 

to the same type of functional constraints as other 

aspects of the style discussed above. Still, longer 

branches may be functional by having longer 

exposed stigmatic surfaces and by recurving and 

coiling so as to allow self-pollination. In any case, 

in the Liabeae, the hispidulous portion of the 

style seems to be a comparatively fixed quantity, 

with longer branches generally correlated with a 

shorter hispidulous region on the shaft. The 

shorter form of branch has been considered more 

primitive in the tribe in this study because of its 

occurrence in most of the genera and in the most 

diverse groups such as the Munnoziinae, Sinclairia, and Austroliabum. The longer form of 

branches occur in one case in the obviously specialized higher elevation genus Parunephelius and 

in the other case in the Liabum-Oligactis generic 

pair, where longer branched forms are specialized 

within the group. An interesting intermediate is 

seen in Chionopuppus, which has short branches 

with the papillae of the stigmatic surface continuing below the branches well down into the stylar 

canal. The actual level of separation of the 

branches and development of stigmatic papillae 

seems very plastic in the Liabeae, but that itself 

is a characteristic that seems different from most 

Vernonieae. 

THE PAPPus.-Both the Liabeae and Vernonieae seem to share a basic type of pappus having 

a short, often squamellate outer series and a long 

capillary inner series. Exceptions exist in both 

tribes, such as Paranephelius in the Liabeae and 

Chresta and Eremanthus in the Vernonieae, where 

there is no marked differentiation of an outer 

series. In these cases, the outer series is not simply 

defective but only irregularly shortened and intergrading with the inner pappus. The pappus of 

these latter genera seems to be specialized within 

the respective tribes. As a result of the similar 

pappus, the isolated achenes of the Liabeae and 

Vernonieae are not easily distinguished; however, 

caution should be used in attempting to relate 

the tribes on this basis. There is ample reason to 

believe that a capillary form of pappus is primitive in the Asteraceae (Robinson, 1981), and 

among the tribes having a capillary pappus, a 

short outer series is not rare. A short outer series 

is seen in many Eupatorieae and Astereae and 

even in some Lactuceae (Krigia) and Heliantheae 

(Pectis, Sartwellia, Eutetras) . 

The teeth of the pappus setae in some \'ernonieae and Liabeae share a detail of some interest. 

The character is seen only with special preparation using a clearing agent with an index of 

refraction like that of Hoyer's solution. Pappus 

setae mounted in water show no anomaly in the 

tips of the teeth when examined under the niicroscope, but in Hoyer's solution, under the highpower lens, an area of the tip of the tooth seems 

to disappear because of its index of refraction, 

leaving a denser central core that resembles a 

short mucro. The character appears in 0ligacti.r 

and in all but one West Indian specimen seen of 

Liabum, and it also appears on the tips of the long 

rays of the plumose pappus in Chionopappus. 

The same type of mucro-tip is even more pronounced in a number of species of T'ernonia 

(Robinson, 1980b), but a similar aberration, differing only by a denser and still visible outer wall, 

is seen in the pappus of Eremothamtius of the 

Eremot hamneae. 

RAPHIDS OF THE ACHENE hIALL.-The achenes 

of the Liabeae all have raphids in their walls, as 

is true of all tribes of the subfamily Cichorioideae 

and all but the Eupatorieae and Heliantheae in 

the subfamily Asteroideae. The raphids in the 

Liabeae occur in two distinct forms. elorigate and 

subquadrate, which do not intergrade. Each genus consistently has raphids of one type or the 

otlier. For those with a microscope and cleared 

preparations, the raphids are the most convenient 
10 SMITHSONIAK CONTRIBUTIONS TO BOTANY 

method for an initial separation of the genera 

into two groups. Most of the genera have elongate 

raphids, but the subquadrate form is characteristic of two distinctive groups, the Munnoziinae 

with four genera and the Liabum-Oligactis generic 

pair in the Liabinae. 

Because of the simple nature of the raphids, it 

would be convenient to interpret their two forms 

as representing two basic lines of development in 

the tribe. In addition, the subquadrate form, 

occurring as it does in elongating cells, is not 

quite matched in any of the other tribes observed, 

and it would be natural to assume that it was 

derived within the tribe from a single origin. 

Unfortunately, the pattern is not so simple. The 

two groups with subquadrate raphids strongly 

contrast in most of their other characters and 

seem to represent two extremes within the tribe. 

Genera more intermediate in other characters 

have elongate raphids. Thus, it is necessary to 

assume parallelism or reversions, or even intergeneric hybridizations, to explain the distribution 

of raphid types in the tribe. Since the Munnoziinae seem to arise from the earliest divergence 

in the tribe, it is possible that the subquadrate 

raphids were present in the common ancestral 

type of all existing Liabeae and that the subquadrate raphids in Liabum and Oligactis are a reversion 

or a tenuous relict of this earlier type. 

GEOGRAPHICAL CONSIDERATI0SS.-The tribe 

Liabeae is restricted to the Neotropical Region 

and is further confined within that region to the 

more mountainous parts. In spite of an erroneous 

original claim regarding Munnozia maronii (Andre) 

H. Robinson (Robinson, 1977a), there are no 

members of the tribe presently known from Brazil. Furthermore, there are only a few representatives of Liabum in the Greater Antilles. The tribe 

is primarily found in the western cordilleras of 

the Americas from central Mexico in the north to 

northern Argentina in the south. Only two phyletic lines, Liabum and the Sinclairia-Liabellum generic pair, account for almost all the range extensions northward from South America. The center 

of distribution of the tribe is in the older portion 

of the Andes, with 11 of the 15 genera in Peru 

and eight genera in Ecuador. Such a distribution 

helps explain the fact that the temperate-based 

classical taxonomists failed to understand the true 

nature of the tribe. 

The center of diversity of the tribe may have 

been a factor in the limited distribution compared 

to most other tribes. Groups of Asteraceae originating in middle elevations of the Andes seem to 

be among the least mobile geographically, if evidence from the Eupatorieae is correct (Robinson 

and King, 1977). In the latter tribe. the Andean 

groups tend to remain restricted to the cordilleran 

chain, usually terminating northward in western 

North America and in the south in Argentina. It 

is groups spreading from eastern North America 

and from Brazil in eastern South America that 

tend to reach the Eastern Hemisphere. In this 

regard, it is notable that geological evidence suggests the Andes first formed as offshore islands 

that were isolated until the emergence of the 

surrounding lowlands at the end of the Tertiary 

(Pliocene) (Haffer, 198 1). 

The pattern of distribution of the Liabeae 

within the Neotropical Region is somewhat complex. The pattern can be made clearer by reviewing the subtribal elements one at a time. 

Accepting that the tribe originated in the area 

of Peru or southern Ecuador. the subtribe Paranepheliiinae has extended its range in only one 

direction, upward. The genus Paranephelzus seems 

very specialized for the high elevations where it 

occurs in Peru and Bolivia. The distribution of 

Pseudonoserzs in various parts of Peru is consistent 

with the assumption based on structural features, 

of relationship to Paranephelius. The considerable 

diversity between the three species of Pseudonoserzs, 

the occurrence at somewhat lower elevations, and 

the more generalized form of inflorescence indicate a less specialized, possibly older, or even 

relictual element in the subtribe. On this basis, 

the characters of Pseudonoseris are regarded as a 

better guide to the ancestral form of the tribe. 

The subtribe Munnoziinae is somewhat less 

restricted in its distribution, with two of the genera, Munnozia and Erato, ranging from Costa Rica 

and Columbia in the north to Bolivia in the 
NUMBER 51 11 

south. The center of diversity of the subtribe is 

shared between Ecuador and Peru, with both 

Munnozia and Chrysactinium occurring nearly 

equally in both countries. Erato actually has a 

center of diversity in Ecuador. Only Philoglossa is 

primarily Peruvian, and it has been suggested 

(Robinson and Cuatrecasas, 1973) that the latter 

genus originated in the floristically distinctive 

coastal range of Peru. Four of the five species of 

Philoglossa are restricted to the latter area, while 

one species extends its range from southern Colombia to the interior of Peru with an outlier in 

Bolivia. It seems unlikely that the Munnoziinae 

and Paranepheliinae can be traced to different 

regions of origin on the basis of existing evidence, 

but it is evident that the Munnoziinae have been 

very successful in Ecuador and other areas north 

of the distribution attained by the Paranepheliinae. It is notable that many species of Munnozia 

have individually attained distributions extending from Costa Rica or Colombia in the north to 

Bolivia in the south, exceeding the entire range 

of the Paranepheliinae. 

The Liabinae is the most widely ranging of the 

subtribes and also is the most internally complex. 

Liabum is the most widely distributed genus in the 

tribe and the only representative in the West 

Indies. The genus, nevertheless, seems comparatively uniform in structure, and the distribution 

probably is rather recent in origin. The West 

Indian members include some species diversity, 

but all are closely related and evident derivatives 

of a single introduction. The Mexican and Central American distribution is based on a single 

species, L. bourgeaui, which is closely related to 

two species in the Andes. 

The primary area of speciation of Liabum and 

the closely related Oligactis is in the northern 

Andes, north of the center of diversity of the tribe. 

Nevertheless, Liabum has some of its most divergent forms, such as L. sandemanii and L. steinbachii, 

in Peru and Bolivia. Oligactis has its most specialized elements to the north in Colombia, Venezuela, and Costa Rica, while the less specialized 

subgenus Andromachiopsis is centered in Ecuador. 

Overall, diversity and geography would indicate 

that the Liabum-Oligactis group was among the 

first elements of the tribe in the northernmost 

Andes, perhaps moving into the area as the habitats in the younger northern Andes were first 

becoming available. 

The two woody genera of the tribe, Cacosmia 

and Ferryranthus, present no geographical complications. Both are restricted to Peru and Ecuador with principal diversity in the adjacent parts 

of these countries. These genera share the basic 

style and raphid form with Chionopappus from the 

coastal region of Peru, also within the general 

area where the tribe is concentrated; however, 

the remaining two generic pairs in the subtribe 

represent major geographical extensions. Microliabum and Austroliabum occur in Argentina and adjacent Bolivia, while Sinclairia and Liabellum are 

found in Mexico and Central America. The genera are all sufficiently alike to indicate a related 

complex extending from Mexico through the 

Andes to Argentina, but they are also sufficiently 

distinct to indicate that the dispersal is not of 

recent origin. 

In the case of Austroliabum and :2ilicroliabum, the 

northernmost occurrence is in the Santa Cruz 

region of Bolivia, the same general area in which 

Munnozia, Liabum, and Paranephelius seem to reach 

their southernmost limits. This gives the impression of a continuous distribution for the tribe; 

however, a significant change occurs in the Santa 

Cruz region: the front of the t\ndes changes 

direction from a northwest-southeast direction to 

a north-south direction. The part northwest of 

Santa Cruz, continuing into Peru, faces northeast 

and shelters a more tropical flora. The part south 

of Santa Cruz, continuing into Argentina, faces 

east and has a more temperate flora. It can be 

seen that in geography, as in structure, the Austroliabum-Microliabum group is discontinuous with 

the Liabeae of the neighboring area to the north. 

The Sinclairia-Liabellum generic pair presents a 

different geographical problem, being the only 

group in the Mexican-Central American area 

that gives evidence of any age. The Central American region has been separated from South America until the creation of the Panamanian isthmus 
12 

in the late Pliocene, -3 million years ago. Even 

after that time, less moist, savanna-type floras 

were more predominant in the area (Matthews, 

1980). It is notable that one of the generic pair. 

Liabplluni, is perhaps the most xeric-adapted genus 

in the tribe, being a small herb with a perennial 

tuber. The specializations within the generic pair 

and the overall diversity of the group would 

indicate that Sinclairia and Liabellum belong to an 

element established in the Mexico-Central America area before the climatic shift to a more moist 

flora and probably before the formation of the 

isthmus. The other genera in Central America, 

Munnozia (2 species), Erato (1 species), Oligactis (1 

species), and Liabum (I species), are evidently 

recent immigrants since the climatic changes and, 

with the exception of Liabum, have not reached 

beyond Costa Rica. 

Considering the Sinclairia and Austroliabum 

groups together, both share many structural features, and both share specializations toward less 

moist environments. As such, it is possible to 

suspect that, in spite of their wide separation. 

they represent a related group that made the 

earliest successful migration beyond the area of 

origin of the tribe, both to the north and south. 

The geographical distribution of the Liabeae 

shows an interesting correlation with chromosome 

numbers. According to available records, the genera centered closest to the presumed center of 

origin of the tribe mostly have numbers interpreted as diploid, whereas the genera centered 

farther from the Peruvian center, at least to the 

north, seem basically polyploid. 

PHYLETIC CoNcLustom-The evolutionary divergence within the Liabeae is accentuated by 

the apparent lack of intergeneric hybridization. 

In this way the tribe differs from many of the 

larger more successful tribes where intergeneric 

or even intersubtribal hybridization seems to occur. The hybridization in the latter tribes seems 

to be the source of much of their diversity, and it 

seems to be at least partially responsible for the 

greater success of those tribes. Evidence of intergeneric hybridization has been seen in the Vernonieae (Robinson. unpublished), Eupatorieae 

SMITHSONIAN CONTRIBUTIOKS TO BOTANY 

(King and Robinson, in prep.), Heliantheae (Robinson, 198 1). and Senecioneae (Robinson and 

Brettell. 1974a), and it will probably be found in 

others. It should be noted that many of the 

processes that increase evolutionary success complicate the taxonomy of the groups. As such, the 

lack of intergeneric hybridization in modern Liabeae might help explain the comparative lack of 

success of the tribe. 

The apparent lack of intergeneric hybridization results in comparatively simple distinctions 

of genera and subgenera in the tribe and allows 

for unusually precise keys. Unlike the situation in 

most other tribes of the Asteraceae. it is possible 

to take some advantage of the hierarchical nature 

of the nonreticulating evolutionary patterns. 

Some species appearing to be intermediate between genera have proven to be the result of 

convergence as indicated in the discussion of Liabum. 

The subtribe Munnoziinae is considered the 

most phyletically distinct group of the Liabeae. 

The subtribe consists of four genera that seem to 

be the opposite of the Paranephelius group in a 

number of important characters: the anthers are 

usually black, the style branches are almost always less than half as long as the hispidulous part 

of the style shaft, and the raphids in the achene 

wall are always quadrate. There is also a tendency 

in the group for long-pedunculate heads, and the 

group contains all members of the tribe that 

consistently bear single-headed scapose or longpedunculate inflorescences. The group is readily 

divided into two parts, each containing two genera. AWunnozia and Chysactinium have a tomentose 

pubescence that is characteristic of most of the 

tribe, both have a capillary pappus that sometimes is reduced to as few as five or 10 setae, and 

both have eight to 10 ribs on the achene as found 

in most Liabeae. The other two genera, Erato and 

Philoglossa, lack tomentum on the leaves, having 

instead long stiff hairs with enlarged bases. One 

of the four species of Erato and all five species of 

Phzloglossa have a reduced noncapillary pappus, 

and the angles of the achene are reduced to four 

in Erato and two in Philoglossa. 
NUMBER 54 1 :3 

The second most phyletically distinct group of 

the Liabeae is treated in this study as the subtribe 

Paranepheliinae, with two genera, Paranephelius 

and Pseudono.seri.r. The extreme form of the subtribe. Parunephplius, is a sessile rosulate herb with 

sessile heads. The genus occurs at higher elevations than any other in the tribe, in areas subjected to snow fall. The distinct nature of the 

subtribe is less evident in the related genus Pseudonoseris occurring at lower elevations, having a 

branching inflorescence and a short-caulescent 

habit that is paralleled in other members of the 

tribe such as Liabellum and ,Microliabum. Pseudonoseris differs from Paranephelius by the noncoiled 

style branches of the ray flowers and by the lack 

of high ridges on the receptacle between the 

areolae. The rather large achenes with broadly 

sclerotized ribs tend to relate the two genera, even 

though there is no reason to suppose that those of 

Pseudonoseris have the longer viability that is suspected for Paranephelius. The style branches are 

elongate in both genera, and the raphids in the 

walls of the achenes are also long, but neither 

character is unique to the group, only the combination of the two characters. The best confirmation of relationship of the two genera seems to 

be in the details of the pollen-wall structure, 

neither genus having large baculae such as are 

seen in most other members of the tribe and in 

most members of the subfamily (Robinson and 

Marticorena, in prep.). 

The combinations of characteristics in the previous two groups have led to their recognition as 

distinct subtribes in the present treatment. The 

remaining genera of the tribe are treated herein 

in a single subtribe, but, nevertheless, considerable diversity is involved. Many, but not all, 

members are notable for short tails or fringes on 

the bases of the anther thecae, a trait not found 

in the Munnoziinae or Paranepheliinae. 

The subtribe Liabinae can be broken down for 

purposes of discussion into four generic pairs plus 

one other individually distinctive genus. The first 

pair. Liubum-Oliguctis, is distinct from the others 

by the elongate branches of the disk styles and by 

the quadrate raphids of the achene wall. The 

elongate style branches are the only feature the 

group has in common with the Paranepheliinae, 

and they are probably of separate origin. Similarly, the subquadrate raphids of the achene wall 

do not seem to relate directly to those of the 

Munnoziinae. The generic pair apparentlv consistently lacks latex. 

If the differentiated minute mucros on the teeth 

of the pappus are any indication, Liabum and 

Oligactis are probably closest to Chionopuppus, 

which has such points on the rays of the plumose 

pappus segments: however, Chionopuppus otherwise is a very unlikely relative, having a reduced 

number of pappus segments in one series, elongate raphids in the achene wall, shorter branches 

on the disk styles, and the only examples of true 

paleae on the receptacle in the tribe. The reddish 

disk flowers of Chionopappus are also rather distinctive. being unlike any other member of the 

subtribe Liabinae but similar to one species of 

Philoglossa in the more distantly related Munnoziinae. 

The combination of shorter style branches in 

the disk flowers and longer raphids in the achene 

wall, as seen in Chionopuppus, also are seen in the 

remaining three generic pairs in the subtribe. The 

generic pair Sinclairia-Liabellum is notable for its 

distribution in the Mexican and Central American region, for its elongate, somewhat lobed nectaries, and for the only discoid heads in the tribe. 

The two genera, however, show marked differences. the former being larger, often scandent 

plants, and the latter being a small herb with a 

basal tuber. The latter also has the most dissected 

leaves in the tribe. In contrast, the genera Austroliabum and Microliabum are the southernmost element of the tribe having achenes with prominent 

ribs on which the pubescence appears distinctive 

and having a fragile inner pappus series. Microliabum is distinct by its extemely reduced size and a 

unique form of broad inner pappus segments. 

The final pair of genera in the Liabinae, Cacosmiu and Ferreyrunthus, provides the most significant 

insight into the evolution of the subtribe. The 

genera are similar in habit, leaf texture, and 

nodal sheaths. but Cucosmia, as indicated under 
14 SMITHSONAN CONTRIBUTIONS TO BOTANY 

the genus, differs in many details of head structure, achenes, and leaf venation. Most significantly, the endothecial cells of Cacosmia have 

shields showing the cell limits, with scattered 

thickenings and variable annulations, whereas 

those of Ferreyranthus have more specialized strapshaped shields with few thickenings at the upper 

and lower ends. In Cacosmia the anthers are not 

noticeably tailed, and latex has been recorded 

from the stems of some specimens. In Ferryanthus 

the anthers have the most highly developed tails 

in the tribe, and the tests for latex are all negative. 

The general characterization of Ferreyranthus is 

consistent with the broad generic relationship 

in c 1 u ding L ia b um, 0 ligact is, and Ch ionopapp us, 

whereas Cacosmia does not fit this group. Actually, 

Cacosmia shows characteristics such as the endothecial cells that suggest that it diverged from 

the base of the subtribe Liabinae, even before the 

point of divergence of the Paranepheliinae. 

The evolution of the tribe Liabeae remains 

generally simple, but details of the Liabinae as 

represented by Cacosmia, Ferryanthus, Chionopappus, Oligactis, and Liabum indicate some complexity. No simple phyletic scheme can be provided 

for these genera in which many of the characters 

are not in conflict. Too many assumptions of 

parallelism or reversion seem necessary. A solution seems to be the assumption that the past 

primitive Liabinae, unlike modern members of 

the tribe, formed an active plexus of intergeneric 

hybridization. Such a conclusion is not difficult 

to accept, considering the commonness of the 

phenomenon in other, more successful tribes. In 

this way the rather specialized raphids of the 

achene wall in the Munnoziinae and the LiabumOligactis generic pair can be considered of common origin, also the tailed anther bases and lack 

of latex in the typical element of the subtribe 

Liabinae, and the specialized endothecial cells of 

some Liabinae and Munnoziinae. Even the 

woody habits and bullate leaf surfaces of Cacosmia 

and Ferreyranthus might have a remote common 

origin. 

The probable general relationships within the 

Liabeae are summarized by the diagram in Figure 1, where the divergent and hierarchical nature of most tribal elements is evident. The pattern seems unusual in the extent to which most of 

the ultimate units consist of clear generic pairs. 

An arrangement has been adopted here that represents the best correlation of most characters, 

but it is notable that no single characteristic 

completely follows the pattern. 

Description of the Tribe Liabeae 

Tribe \'ERNOMEAE sect. LIABEAE Cassini, Dict. sci. Sat.. 

Tribe VERNONIEAE subtribe LIABISAE Lessing. Linnaea. 

Tribe SENECIONEAE subtribe LIABINAE Benthani in Benthani 

Tribe LIABEAE Rydberg, S. Amer. FI.. 34(4):289. 1927.-H. 

57:338, 1828. 

6:696, 1831. 

and J.D. Hooker. Genera Plantarum, 2:207. 1873. 

Robinson and Brettrell. Phytologia. 25(6):405. 1973. 

Mostly perennial herbs and subshrubs, sometimes scandent; annuals in Microliabum and some 

Munnozia subgenus Kastnera; shrubs or small trees 

in Cacosmia and Ferryanthus. Roots usually not 

specialized, but Liabellum with underground 

tuber, Paranephelius often with prominent fusiform 

lateral roots. Latex usually present, consistently 

lacking only in Liabum, Oligactis, and Ferreyranthus. 

Tomentum usually present on at least undersurfaces of leaves, often on both leaf surfaces, stems, 

and involucre. Erato and Philoglossa with stiff hairs 

having enlarged bases on leaves, stems, and involucre, and without tomentum except sometimes 

on tips of involucral bracts. 

Leaves opposite or congested in rosette, usually 

petiolate or with petioliform base, rarely sessile; 

leaf bases often with stipule-like wing or nodal 

disk, sometimes fused into sheath; leaf blades 

linear to broadly triangular with pinnate or trinervate venation. Inflorescence simple or subcymose, sometimes forming a pyramidal panicle, 

scapose or subscapose from a short leafy stem in 

Chvsactiniunz, some Pseudonoseris, Liabellum, Microliabum, and some Liabum; inflorescence most often 

open with few long peduncles in Munnoziinae. 

Heads never compound or forming complexes. 

Involucre usually subimbricate with bracts in 

many graduated series, subequal with somewhat 
NUMBER 54 15 

foliose outer bracts in some Munnoziinae. Receptacle alveolate, often with projecting crests or 

points, with true paleae in Chionopappus. Ray 

flowers usually present, 3-230, lacking in Liabellum and some Sinclairia. Disk flowers 3-150, bisexual. Corollas usually yellow, ray and disk flowers reddish in Pseudonoseris srysrylowiczii, disk flowers purple in Chionopappus benthamii, Philoglossa 

purpureodisea. and some P. peruuiana, and ray and 

disk flowers whitish or tardily lavender in many 

iVlunnotia jussieui and M. campii. 

Ray flowers in 1 to many series, always fertile 

with no anthers: limb always present, usually 

rather narrow: adaxial cells elongate and nonpapillose: cuticle often smooth, with longitudinal 

ridges in Austroliabum and longitudinal striation 

in IVlunnozia (Baagoe, 1978): achenes not differentiated from those of disk flowers. Disk corollas 

usually narrowly funnelform, throat somewhat to 

distinctly expanded at base in most Munnozinnae 

and some Oligactis; corolla surfaces smooth, the 

cells elongate with sinuous lateral walls: lobes 

elongate, usually linear, usually with stomates 

along margin, lacking stomates in Cacosmia and 

0iigacti.s. and with stomates rare or lacking in 

Liabum; pubescence often of Hieraceum-type with 

long broad bases and small glandular tips: lobe 

tips sometimes with multiseriate spicules or crests. 

Filaments usually smooth in lower part. papillose 

in Ferrp2,ranthu.s r~erbascifblius: anther collar mostly 

with short-oblong cells, \vith walls usually unornamented, \valls ornamented with annular thickenings in Ch~sactinium: thecae usually pale, blackened in all Munnoziinae except A.Lfunnozia hastijolia 

and 2 related species, bases of thecae calcarate, 

extending well below point of insertion on collar. 

bases broad and unfringed in Munnoziinae, more 

or less fringed in many Liabinae, fringed and 

coiled in Ferreyranthus uerbascfolius; endot hecial 

cells often elongate with noncontiguous sclerified 

shields and polarized thickenings on transverse 

walls, short with more lateral thickenings in many 

Munnoziinae and Cacosmia; anther appendage 

thin, longer than wide, not constricted at base, 

not keeled, usually smooth, papillose in Oligactis 

subgenus Oiigactis. Nectary cylindrical, truncate 

in Chionopappus, Cacosmia, and Ferreyranthus, sometimes elongate with upward projecting lobes as in 

Sinclairia. Style base glabrous, usually with evident enlargement; upper style shaft and backs of 

branches covered with hairs: branches with stigmatic papillae over entire inside surface, with 

little or no appendage: style branches of rays 

elongate and sometimes spiralled as in Paranepheiius; branches of disk flowers short (less than 1/3 

to 1/4 as long as hispidulous upper style shaft) 

and blunt in Munnoziinae and such genera as 

Chionopappus, elongate in such genera as Liaburn 

and Ferrqranthus, noticeably tapered only in Pseudonoseris. 

Achenes usually prismatic or subterete with .5- 

10 ribs, 4-ribbed in Erato, compressed with 2 ribs 

in Philoglossa, long-fusiform in Paranephelius; surface with stiff twin-hairs and/or glands, with 

some arachnoid hairs in Paranephelius; cells of 

achene wall with raphids, raphids usually elongate, subquadrate in Liaburn, Oli,gactis, and the 

M u n noz i in ae : carpo pod i u m disc i form. an n u I i - 

form, stopper-shaped, or cylindrical, symmetrical 

or nearly so, usually with small cells in 5-10 

series, sometimes with moderate sized cells in 2- 

5 series. cell walls densely beaded to moderately 

thickened. Achene viability apparently usually 

short, probably longer in Paranppheiius. 

Pappus usually kvith numerous long capillary 

bristles in inner series and with a short outer 

series. inner series of reduced number. and squaniellae of outer series broad in some Munnozia 

subgenus Kastnera and in Austroliabum, latter also 

with inner series easily deciduous, inner series of 

lanielliform segments similar in breadth to outer 

squamellae in Microliabum, pappus of 5 plumose 

bristles in single series in Chionopappus, reduced to 

short deciduous awns in Erato sodiroi and Philoglossa mimuloides, a pair of short awns connected 

by a series of squamellae in P. purpureodisca, totally 

lacking in Cacosmia and 2 Philoglossa species: pappus bristles white in Chyactinium but sordid or 

darker in Munnoria: scabridulae or lateral hairs 

with minute structurally differentiated mucro in 

Liabum, Oligactis, and Chionopappus. 

Pollen 25-50 pm in diameter, spinose, spines 
16 SMITHSOKIAN CONTRIBUTIONS TO BOTANY 

regularly to somewhat irregularly arranged, never 

lophate, large baculae usually present and 

grouped or with a central cavity under each spine, 

cavus variously developed in Paranepheliinae, 

Philoglossa, and Munnotia. 

A few characters provide special problems of 

interpretation. Latex production often seems erratic. Greenhouse specimens of Erato polyrnnioides 

usually show some latex, but the amount varies 

depending on available water and time of day. 

Some weaker tests might not have been recognized as latex in the field. A plant tested at night 

showed no latex. 

The upper surface of the leaf blades often bears 

a coarse pilosity that has frequently been used as 

a taxonomic character. Unfortunately, the character varies within species in many genera such 

as Munnozia and Chrysactinium and varies between 

closely related species in Liabum. 

The receptacle has true paleae in only Chionopappus. In other genera the projections and crests 

in the same positions are unarticulated outgrowths. The latter are particularly prominent in 

Munnotia, being one of the characters originally 

cited by Ruiz and Pavon (1794), but the receptacles of Munnozia subgenus Kastnera completely 

lack crests. The length of the crests or chaff is of 

taxonomic value in Chrysactz'nium but is difficult 

to use, since the heads are often solitary and 

cannot be dissected without destroying a herbarium specimen. 

Many specialized features of the tribe have 

occurred more than once. The pappus has been 

lost in both Philoglossa in the Munnoziinae and 

Cacosmza in the Liabinae. Scandent habits are 

found in two separate genera of the Liabinae, 

Oligactzs of the northern Andes and Sinclairia of 

Mexico and Central America Reduced habits 

with basal leaves and a scapose inflorescence have 

evolved many separate times, in Liabelluin in the 

Sinclazrza relationship, in Microliahum in the Austroliabum relationship, to some extent in Liabum, all 

in the Liabinae, in Chrysactinzum in the Munnoziinae, and in Pseudonoserzs in the Paranepheliinae. 

These and other characters revie\\ed above are 

regarded as evidence of a complex ecolution with 

marked phyletic gaps. This contrasts to the superficial appearance of a continuum that led 

Bentham (1873) to place most members of the 

tribe in a single genus, Liabum. 

Keys to the Genera of the Liabeae 

Two keys to the genera of the tribe are provided. The first key is based on more obvious 

characters and is highly artificial. The second key 

is based on more technical characters and is in 

some parts phyletic. 

Artificial Key to the Genera of the Liabeae 

1. 

I. 

Heads sessile in a basal rosette of leaves; style branches of ray flowers spiralled; plants at higher 

elevations in the central Andes ........................................... Paranephelius 

Heads short- to long-pedunculate, not sessile in basal rosette of leaves, style branches usually 

2. Leaves not tomentose, hairs of leaves and stems stiff, with enlarged bases ............ 3 

3. Small, mostly decumbent herbs: leaf blades trinervate at base: achenes compressed, \vith 

2 ribs; pappus reduced or lacking ............................... Philoglossa 

3. Robust erect herbs or subshrubs; leaf blades with 5-9 veins radiating from base: achenes 

mostly 4-angled; pappus of many awns or bristles .......................... Erato 

... 4 

4. Pappus lacking ........................................................ .Cacosmia 

4. Pappus present .......................................................... 5 

Pappus bristles plumose; receptacle with long narrow strap-shaped paleae; flowers of 

disk red or purple and rays yellow .......................... .Chionopappus 

evenly curved: rarely spiralled 3 .................................................... - 

2. Leaves usually tomentose, at least on undersurface: hairs not stiff with enlarged bases 

5. 
NUMBER 54 17 

5. Pappus setae not plumose; receptacle without paleae, with or without crests or spines; 

disk flowers not reddish and rays yellow .......................................... 6 

6. Inner and outer series of pappus both broadened and flattened; minute plants to 

................................... Microliabum 

6. 

Anther thecae usually blackish; style branches of disk flower less than 1/3 as 

long as hispidulous upper part of shaft ................................... 8 

8. Pappus white; cells of anther collar with walls obscured by annular 

thickenings; scapose plants with leaves rosulate or on short stems; latex 

Pappus sordid or colored; cells of anther collar not obscured by annular 

thickenings; plants usually with elongate leafy stems; latex present .............................................. ........ Munnozia 

Anther thecae pale; style branches of disk flowers often 1/2 as long as 

................................ 9 

9. Inflorescence with all bracts or with all but the basal bracts and branches 

subopposite or alternate; receptacle scarcely alveolate, without hairs, 

chaff, or projections ............................................. 10 

10. Plants with short or rosulate leafy stems; leaf venation essentially 

10. Plants with prominent erect leafy stems, primary leaves strongly 

trinervate; pollen not caveate ...................... Austroliabum 

Inflorescence with all or at least primary bracts and branching opposite: 

................ 11 

11. Leaves with pinnate venation; achenes bearing both glands and 

setulae ........................................................ 12 

12. Scandent shrubs; leaf bases not stipulate or with only an 

adjacent lobe on node; raphids in achene walls quadrate; tips 

of cells of pappus teeth differentiated in thin-walled 

mucro ............................................. Oligactis 

Shrubs and small trees; leaf bases with stipules fused into small 

sheath: raphids in achene walls elongate; tips of cells of pappus 

teeth not differentiated ....................... Ferreyran th us 

Leaves trinervate; achenes with setulae, with or without glands. . 13 

13. Small, few-headed plants with underground tuber; leaves usually deeply palmately or subpinnately lobed: ray flowers 

lac king ........................................... Lia bellum 

Small to large herbs or vines, without underground tuber; ray 

flowers present or absent ................................. 14 

14. Leaf bases sagittate ........ Munnozia hastifolia group 

14. Leaf bases cuneate or rounded ...................... 15 

Petiole bases and nodes without wings or lobes, 

leaves sometimes in whorls of 3 or more; stems 

usually not white-tomentose, mostly terete; latex 

present; inflorescence in thyrsoid or corymbose panicle; ray flowers sometimes lacking; achenes with 

10 cm high ............... 

At least the inner pappus s m; larger plants over 10 cm high ............ 7 

7. 

absent ? ................................................ Chrysactinium 

8. 

7. 

hispidulous upper part of shaft or longer 

pinnate; pollen caveate ............................ Pseudonoseris 

9. 

receptacle with minute hairs or chaff or projections 

12. 

1 1. 

13. 

15. 
18 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

elongate raphids in walls; tips of teeth of pappus 

setae not differentiated; pollen grains 35-50 pm in 

diameter Sin clairia 

Petiole bases or nodes with wings or lobes, leaves 

always opposite; stems usually white-tomentose or 

flocculose pubescent; latex absent; inflorescence a 

cymose or subumbellate panicle; ray flowers always 

present; achenes with subquadrate raphids in walls; 

tips of teeth of pappus setae with thin-walled mucro; 

pollen grains 25-35 pm in diameter Liabum 

15. 

Technical Key to the Genera of the Liabeae 

1. Cells of achene wall with subquadrate raphids; anthers with or without black thecae .......... 2 

2. Anther thecae pale, with bases bearing teeth or projecting cells; pappus with tips of projecting 

cells bearing a minute thin-walled mucro; lobes of disk corollas without evident stomata; shaft 

of disk style with hispidulous part less than twice as long as style branches; plants without 

latex ...................................................................................... 3 

3. Small to large herbs, rarely subscandent; leaves slightly to strongly trinervate; inflorescence 

cymose or subcymose; achenes with only setulae and no glands; style with hairs often 

scarcely extending onto upper part of shaft .................................... Liabum 

Scandent shrubs; leaves pinnately veined; inflorescence a corymbose or thyrsoid panicle; 

achene with glands and setulae; style with hairs distinctly extending onto segment of 

upper shaft ................................................................... Oligactis 

Anther thecae usually blackish, with bases not bearing teeth or projecting cells; pappus with 

tips of projecting cells not specialized; lobes of disk corollas with stomata; shaft of disk style 

with hispidulous upper part usually twice as long as branches or longer; plants often with 

latex ............................................................................. 

4. Leaves not tomentose, hairs of leaves and stems stiff, with enlarged bases . . , . 

5. 

5. 

3. 

2. 

Small mostly decumbent herbs; leaf blades trinervate at base; achenes compressed 

Robust erect herbs or subshrubs; leaf blades with 5-9 vein 

achenes mostly 4-angled; pappus of many awns or setae ...................... Erato 

................... 6 

Pappus white; cells of anther collar with walls obscured by annular thickenings; 

scapose plants with leaves rosulate or on short stems; inflorescence of single heads on 

very long unbranched peduncles; disk corolla lobes with nearly sessile glands scattered 

over outer surface, without longer glands or hairs; latex absent? .... Chrysactiniurn 

Pappus sordid or colored; cells of anther collar not obscured by annular thickenings; 

plants small to robust herbs with distinct leafy stems, usually with branching inflorescence; disk corolla lobes usually with elongate hairs or glands near tips; latex 

............................................................. Munnozia 

Branches of disk style filiform, longer than the hirtellous upper part of the shaft; tube of disk 

corolla longer than the limb; plant small with leaves congested on short stems or in rosette; 

leaf venation pinnate or weakly trinervate .............................................. 8 

with 2 ribs; pappus reduced or lacking ...................... 

4. Leaves usually tomentose below; hairs not stiff with enlarged bases 

6. 

6. 

1. Cells of ach elongate raphids; anthers with thecae never black ... 7 

7. 
NUMBER 54 19 

8. Heads sessile in basal rosette of leaves; style branches of ray flowers spirally contorted; 

pappus outer series of irregularly shorter setae, not sharply differentiated; plants with or 

without latex 

Heads raised in unbranched or alternately branched inflorescence above a short but 

distinct leafy stem; style branches of ray flowers not spirally contorted; pappus with outer 

series of distinct short squamellae; plants with latex ............... ... Pseudonoseris 

Branches of disk style narrowly oblong to linear, shorter than the hirtell upper part of the 

shaft; tube of disk corolla shorter than limb; plant small to large, never a sessile rosette; leaf 

venation pinnate to strongly trinervate ............................................ 

9. Pappus of 8-10 plumose setae; heads with yellow rays and purple disk flowers; receptacle 

with long slender paleae ........................................... ... Chionopappus 

9. Pappus without plumose setae, sometimes absent; heads with all flow yellow; receptacle 

without paleae, sometimes with short crests or projections ........................... 10 

10. Woody shrubs or small trees of the northern Andes; leaves usually with bullate upper 

surfaces, with petiole bases of leaf pair fused into a sheath; involucral bracts closely 

and regularly multiseriate . , . , , . ............................................ 11 

1 1. Pappus lacking; achenes glabrous, 4-5-angled; leaves trinervate with strongly 

ascending secondary veins; heads cylindrical with - 10 flowers ...... Cacosmia 

11. Pappus present, with capillary setae; achenes setuliferous or glanduliferous, 10- 

ribbed; leaves pinnately veined; heads campanulate with 20-35 flowers 

Herbs or vines; leaves without obvious bullate upper surfaces, with bases of petioles 

rarely forming a sheath; involucral bracts usually in few or irregular series . . , . 12 

12. Plants of Argentina and Bolivia; inner pappus easily deciduous; pubescence of 

achene usually different on ribs; ray flowers always present; branching of 

inflorescence mostly or entirely alternate ................................... 13 

13. Minute herbs of higher elevations; inner pappus of broad laminate 

13. Coarse herbs of medium elevations; inner pappus of capillary 

elements , . ...... .............................. Austroliabum 

Plants of Mexico and Central America; inner pappus moderately persistent; 

pubescence of achene not different along ribs; ray flowers often lacking; branching of inflorescence often opposite ............... .................... 14 

14. Leaves sessile or winged to base, usually palmately lobed; small herbs with 

basal tuber; style with broad often nodular base .............. Liabellum 

14. Leaves petiolate, without wings or stipules, not lobed; plants often large or 

scandent, without basal tuber; style with narrow base ......... .Sinclairia 

. . , . ................................................ Paranephelius 

8. 

.. ....................................................... Ferreyran th us 

10. 

segments ...................................... .......... Microliabum 

12. 

The Subtribes and Genera of the Liabeae nephelius and Pseudonoseris; and subtribe Munnoziinae containing Chrysactinium, Erato, Munnotia, 

The present and Philoglossa. treatment recognizes three subtribes and 15 genera in the Liabeae, which are 

arranged as follows: subtribe Liabinae containing 

Austroliabum, Cacosmia, Chionopappus, Ferreyranthus, 

Liabellum, Liabum, Microliabum, Oligactis, and Sin- Annuals, perennial herbs, vines, shrubs, or 

clairia; subtribe Paranepheliinae containing Para- small trees; leaves mostly ovate to linear, rarely 

Subtribe LIABINAE 
20 SMITHSONIAS CONTRIBUTIONS TO BOTANY 

with angulate margins, pinnately or trinervately 

veined. Heads in cvmose, corymbose, or subumbellate panicles: disk corollas with bases of throats 

usually not abruptly expanded: anthers pale, thecae often with lobes or teeth at base: shields of 

endothecial cells usually not contiguous, usually 

with thickenings polarized on transverse walls; 

style branches short to elongate: raphids of achene wall quadrate to elongate. Pollen grains with 

spines somewhat irregularly disposed, spines with 

small to large columellae that are hollow or in 

groups. 

The subtribe contains the majority of the genera and species of the Liabeae and has the widest 

geographical distribution, reaching from Mexico 

and the West Indies in the north to Argentina in 

the south. The nine genera are as follows. 

A us trolia b urn 

FlGL'RE 2 

Au~/ro/iabutn H. Robinson and R.D. Bretrell. Phytologia, 

28(1):48, 1974. [Type-species: Liabum candidum Grisebach.] 

Perennial herbs or subshrubs, with milky sap. 

Leaves opposite, becoming alternate in inflorescence, distinctly petiolate, petioles winged or stipitate at the base: blade triangular to lanceolate, 

trinervate at the base. margin serrate to dentate, 

Lvhite-tomentose below and sometimes on both 

surfaces. Inflorescence mostly cymose, alternately 

branched at least above, pedicels often elongate. 

Heads broadly campanulate; involucral bracts 

50-75 in -3-4 series, unequal to subequal. narrowly oblong to lanceolate, inner bracts narrowly 

attenuate. often densely covered with long-stipitate glands on outer surface, sometimes Lvhitetonientose: receptacle essentially glabrous. Flowers yellow. Ray flowers 20-30 in 1-2 series, corolla 

with tube slender, hirsute, limbs linear with few, 

sparse, nonglandular, biseriate hairs: style 

branches elongate, not spiralled. Disk flo\vers 30- 

173, corolla narrowly funnelform, not abruptly 

expanded at base of throat, tube narrow, tube 

and most of throat sparsely hirsute: lobes with 

stomates along margins, mostly glabrous, with 

some hairs distally: filament smooth or papillose. 

walls of cells of anther collars not strongly annulated: anther thecae pale, long and narrowly 

tapering at base with a few projecting cells, 

scarcely tailed, median endothecial cells oblong. 

thickened shields noncontiguous, oval or linear, 

irregularly ornate, attenuate and uncinate with 

1-3 points at lower end, appendages long-ovate, 

1'12-2 times as long as wide, smooth, cells shortly 

oblong with truncate to somewhat oblique endwalls: nectary short, not or scarcely lobed: often 

obscure: style base with large, distinct, abrupt 

node: style branches slender, less than 1/2 as long 

as the hispidulous portion of the shaft. Achenes 

prismatic, obovate, distinctly narrowed below, 

strongly costate with - 10 ribs. densely setiferous 

on the ribs and often also in the furrows; cells of 

the achene walls with elongate raphids: carpopodium narrow rim of -3-5 rows of rather small 

subquadrate cells, cell walls slightly thickened, 

porose; pappus with 20-30 rather easily deciduous inner setae in 1 series, teeth of setae simple, 

shortly and stiffly spreading; pappus outer series 

of slightly to distinctly squamiform small to large 

elements. Pollen grains 35-45 pm in diameter, 

spines unevenly disposed, with distinct internal 

colu mellae . 

Austroliabum and the related .Wicroliabum together form the southernmost element of the 

tribe. The former genus occurs along the eastern 

slopes of the Andes in the area from central 

Bolivia into northern Argentina. The northern 

limit is near Santa Cruz, where the escarpment 

of the Andes turns westward and tends to shelter 

a more tropical flora. '4s such, the northernmost 

limit of Austroliabum scarcely reaches the southernmost limits of the large Andean genera Liabum 

and Munnozia. 

The genus Austroliabum differs from both Liabum 

and Munnozia by the elongate raphids in the cells 

of the achene wall. The elongate raphid form is 

shared with other genera of the Liabeae such as 

Chionopappus, Cacosmia, Ferreyranthus, Pseudonoseris, 

and Parunephelius of the Andes and the distant 

Sznclairzu of Central America. Austroliabum also has 
NUMBER 54 

E'IGI.RE i'.-,4us/rolzabum cundidmi (Grisebach) Robinson & Brettell: A. habit. X !?: B. head. post 

anthesis. X 4: c, ray corolla, X 8: D. disk corolla, X 8: E. disk style. X 8: F. bases of anther 

thecae. X 35: G. median endothecial cells: X 400: H, achene, X 8. 

'I 
22 SMITHSONIAN CONTRIBUTIOKS TO BOTANY 

a distinctive receptacle, which is prominently convex in older heads and has a simple alveolate 

pattern. The ribs on the achene are very prominent, and in some species the twin-hairs are restricted to the ribs. Even in the species where the 

hairs are not restricted, the prominence of the 

ribs causes the appearance of unequal distribution. The outer pappus elements of all species are 

highly differentiated, their greater breadth correlating with the somewhat reduced number of 

setae in the inner pappus. The setae of the inner 

pappus are more easily deciduous than those in 

most other members of the tribe. 

The leaves of ilustroliaburn show great variation. 

The blades are strongly deltoid and trinervate in 

A. pofymnioides but narrowly lanceolate and pinnately veined in A. eremophilum. The pubescence 

includes long-stalked glands in A. pobmnioides but 

is densely white-tomentose without glands in A. 

candidurn. The petioles are often strongly winged, 

and either the auricles'of the wing or stipule-like 

lobes seem to be present in all species. 

The genus contains the following three species. 

ilu.\/ro/zaburn candzdurn (Grisebach) H. Robinson and Brettell. 

Phytolog-ia. 28( 1):49. 1971. Argentina. 

Z,iclhm randidrim Grisebach. S!.mbolae. 203. 1879. 

Licthum uuriruidutn (;risebach. Symholae. 203. 1879. 

Z,icihrr,n ,\rhiTi.ho\iiin Blake. ,J, \Vash. Acad. Sci.. 17:293. 

1927. 

Lici/norr randiduin .\ubrirrhoruin (Blake) Cabrera. Bol. Soc. 

Argent. Bot.. 2:95. 1917. 

.I/i.s/io/iuhwi ~rernophilum (Cahrera) €1. Rohinson and Brettell. 

Ph>.toloqia. 28( l):19. 1974. .~rqentina. 

Lici/~i~/, Prrmrp'iilum Cahrera. Bol. Soc. Argent. Bot.. 2:%, 

l9+i. 

.lu.\/ro/iccbu~n /iu/pwioidr.t (1I.E. Fries) 11. Robinson and Brettell. Ph!.rologia. 28( 1): 49. 19i4, Arqenrina. Bolivia. 

1,inhirni poll,mrioidit R.E. Fries. Xrki\. Rot.. .?I( l3):21. 1906 

Licihirrii viu/qdiIjb/iirirl ~liischlrr. Enql. Bot. Jahrb. 50. 

.lui/~o/ru/iuni inrc/,qrdr!/~//ciori (hluschlet) €1. Robinson and 

Beihl.. 3:85. l9l,3. 

Brrrtell. Phytologia. L'8( 1): 19. 197-4. 

Cabrera (1947) has provided a key by which 

the three species can be distinguished. 

Cacosmia 

Fl(;17Rk. 3 

Coco\rriio €Iuniholdt. Bonplaiid. and Kiinth. Nov. Gen. et 

Sp.. ed. fol. -4:E7. 1818. 17'!ye-~pecies: C'crcosmia iu,qosn 

Hiimholdt. Bonpland. and Kunth: oriqinally inonotypic.] 

Clairzilleu DeCandolle. Prodr.. 5:636. 1836. /Type-species: 

Clarrz rllea quinquenewia DeCandolle.] 

Erect shrubs, sometimes densely branching. 

Stems terete, densely pubescent, with or without 

milky sap. Leaves opposite, bases fused into a 

sheath, with petioles or subsessile; blades oblongovate to narrowly lanceolate, base broadly to 

narrowly rounded, margins serrulate to subentire, 

minutely bullate above, densely pale-tomentose 

below, trinervate or quinquenervate from near 

base with strongly ascending veins. Inflorescence 

terminal on leafy branches, densely corymbose or 

with densely corymbose branches, branching opposite: pedicels short, densely tomentose to puberulous. Heads cylindrical; 20-25 involucral 

bracts in 5-6 series mostly in 5 ranks, ovate to 

narrowly elliptical, tips narrowly rounded to 

acute, inner more pointed bracts not in ranks, 

outer surface glabrous or obscurely pubescent; 

receptacle slightly convex. alveolate, glabrous. 

Flowers yellowish, rarely whitish. Ray flowers 5 

in a single series; corolla narrow: limbs short, 

elliptical, mostly glanduliferous on outer surface, 

minutely trilobed at tip, style branches slender, 

not spiralled. Disk flowers 5-6; corollas narrowly 

funnelform, with glands on outer surface, slightly 

to densely hirsute, tubes short and slightly narrowed, gradually expanding at base of throat; 

lobes without stomates; filaments smooth; anther 

collar with few to many bulging cells, annulations 

of cell walls weak; thecae pale. untailed but minutely digitate at base; endothecial cells shortly 

oblong, with scattered nodular thickenings on 

transverse or vertical walls, with few annular 

thickenings sometimes restricted to one end of 

cell: appendages oblong-ovate, - 1Y2 times as long 

as wide, smooth, with narrowly oblong cells having truncate to oblique end walls; nectary short 

and unlobed; style base with a distinct large 

node: style branches of disk flowers linear, shortly 

to rather narrowly pointed, 1/3-2/3 as long as 

the hispidulous portion of the shaft. Achenes 

prismatic, outer sometimes subtriquetrous, inner 

with 4-5 angles, faintly costate, 8- 10-veined, glabrous. walls of achene with elongate raphids: 

carpopodium indistinct, born on abruptly nar- 
NUMBER 54 

FICL.RE Y.--Cuco~mzu rugmu Humboldt. Bonpland. and Kunth: A, habit. X lh: B. head with 1 ray 

removed, X 10: c. ray corolla. X 10: D. disk corolla, X 10: E. bases of anther thecae, X 35: t.. 

median endothecial cells. X 400; G. disk achene with style. X 10. 
24 SMITHSONIAS CONTRIBUTIONS TO BOTANY 

rowed base of achene: pappus lacking. Pollen 

grains -30 pm in diameter, with spines rather 

unevenly dispersed, with distinct internal columellae grouped under spines. 

Because of the lack of a pappus, Cacosmia, 

during its long history, has been kept separate 

from Liabum. Nevertheless, shortly after the description by Humboldt, Bonpland, and Kunth 

(1818), the genus was recognized as a relative of 

Liabum by Cassini (1823, 1830). The genus remained with Liabum in the treatments of Lessing 

(1832) and DeCandolle (1836). Cacosmia was 

placed in the new tribe Helenieae by Bentham 

and Hooker (1873), remote from Liabum, which 

was placed in the Senecioneae. Unfortunately, 

the Bentham and Hooker treatment during the 

last 100 years has become endowed with the force 

of tradition, and the separation between Cacosmia 

and other Liabeae has become widely accepted. 

Still, isolated authorities such as Cabrera (1934) 

recognized the relationship between Cucosmia and 

Ll'abunz, and the pollens were compared by 

Skvarla and Turner (1966). The two genera were 

more formally associated under the tribal concept 

of the Liabeae by Robinson and Brettell (1973a, 

1974b). The relationship has been recognized in 

all recent treatments (Nordenstam, 1977a; 

Robinson, 1978a). 

During most of the studies by this author, the 

genus Cucosmia has been considered a close relative of Ferryanthus of the same subtribe. The two 

genera share a more lvoody habit, a sheath at the 

node formed of the fused paired leaf bases, and 

bullate upper leaf surfaces. Some differences such 

as the lack of pappus in Cacosmia seem obvious 

but kvould not necessarily bar close relationship. 

The consistently pinnate venation of Ferryanthus 

contrasts totally with the nearly longitudinal 3-5 

veins in Cacosmia, but some variation between 

pinnate and trinervate venation is seen within 

related groups elsewhere in the tribe. e.g.. LiabumO1icqactis. The differences in involucre form and 

achene shape do present more serious considerations, and they call the closeness of relationship 

into question. It is the form of the thickenings in 

the endothecial cells, however, that indicates that 

the similarities between the two genera are probably spurious. An overview of the tribe shows that 

the endothecial cells are a conservative character 

of considerable phyletic reliability, yet those of 

Cacosmia are comparatively specialized, whereas 

those of FerrTranthus are comparatively primitive. 

No workable phylogeny can be constructed using 

this character without separating the two genera. 

The present view would derive Cucosmia from 

near the base of the subtribe Liabinae above the 

point of divergence from the Munnoziinae but 

before the point of origin of the Paranepheliinae. 

In retrospect, a number of other characteristics 

seem significant, such as the minutely digitate 

but untailed bases on the anther thecae and the 

documented presence of latex in at least some 

specimens of Cacosmia. It makes more phyletic 

sense to be able to exclude Cacosmia from the 

typical series in the subtribe Liabinae of Liabum, 

Oligactis, Ferryanthus, and Chiompappus, which 

will have sonic tail formation on the base of the 

anther thecae and which have consistently shown 

no latex in those for which reports are available. 

The involucre of Cucosmia is one of the most 

distinctive features of the genus. Both the cylindrical shape and the ranking of the median series 

of bracts are unique in the tribe. 

Cacosmia has usually been regarded as containing one species, and the subdivisions recognized 

by Hieronymus (1901) were at the varietal level. 

Two new species were recognized recently in 

rapid succession by Robinson (1976a) and Nordenstam (1977b). All the species occur in southern 

Ecuador near the Peruvian border, and two of 

the species are nearly to completely restricted to 

that area. 

The three presently recognized species are as 

fo 1 1 0 w s . 

Carosmza harlirigii Nordensrani. Bot. Notiser. 130:283. 1977. 

C,iimmia hierorymi H. Robinson. Phytologia. 31( 1) :16. 1976. 

C,hcosmiu riigoha var, arachnoidea Hieronymus. Engl. Bot. 

Chcosmia rugosu Humboldt. Bonpland. and Kunth. Nov. Gen. 

C/uiwiiIta quinqutneri iu DeCandolle. Prodr.. 5:636. 1836. 

Ecuador. 

Ecuador. 

Jahrb.. 28:619. 1901. 

et Sp.. ed. fol.. 1:228, 1818. Ecuador. Peru. 
c,'uco\rniu qmriquener;~za (DeCandolle) Steudel. Nonien. Bot.. 

Ci(co,\mIu rugotu \.ar. nii'm Hieronynius. Engl. Bot. Jahrb.. 

Crm\ rn 10 I ugu \a va r . X/ngi 1 €3 . Robinson . P h >- t ol og i a, 

ed. 2. 1::377. 1840. 

28 : 6 1 9. 190 1 . 

.13( l):&8. 1976. 

Keys to the three species and to the three 

varieties of Cacosniia rugosa have been provided by 

Nordenstam (1977b) and Robinson (1978a). 

Ch ion opapp u s 

t'l(;l.Rk 4 

C,'//io,io//c~/~~rc,\ Bent ham in Benrham and Hooker f.. Gen. 

Plant.. .3:485% 1873. Ilectotype: Chionopuppus benthumti 

Blake: genus originally described without a named specie.] 

Erect shrubs, moderately branched. Stems terete with thinly evanescent arachnoid tomentum, 

latex not noted. Leaves opposite, bases of petiole 

pair fused to form short sheath, petioles very 

short: blade ovate, base broadly rounded to subtruncate, margins serrate to serrulate. nearly 

smooth above, pale-tomentose below, strongly trinervate from near base. Inflorescence with heads 

in groups of 3 at tips of branches, cymose; pedicels 

4-50 mm long, 1 or usually 2 small foliose bracts 

at base of head. Heads broadly campanulate; 50- 

55 involucral bracts in -5 unequal graduated 

series, ovate to narrowly lanceolate, tips acute to 

shortly acuminate, outer bracts with distinct herbaceous tips. outer surface with evanescent arachnoid pubescence: receptacle convex, with long, 

narrow, strap-shaped paleae. Rays yellow, -40 in 

1-2 series: base of corolla extremely narrow; limbs 

linear. minutely bilobed at apex, glabrous; style 

branches elongate, not spiralled. Disk flowers red, 

corollas narrowly funnelform; tubes extremely 

narrow, gradually expanding at base of throat, 

outer surface glabrous, lobes with stomates near 

margins: filament smooth, cells of anther collar 

without strong annular thickenings on walls; thecae pale. shortly tailed and minutely digitate at 

base; median endothecial cells oblong, with strapshaped to oval, somewhat spirally ornamented 

sclerified shields. often with attenuate uncinate 

tips at lower end: appendages smooth, oblong to 

oval, cells mostly elongate with truncate to 

oblique end walls: nectary short and unlobed: 

style base with distinct node: style branches of 

disk flowers less than 1/2 as long as the hispidulous upper portion of the shaft: stigmatic papillae 

extending far down into the stplar canal. Achenes 

prismatic, with 8-10 ribs, with minute usually 

appressed setulae, walls of achene with elongate 

raphids; carpopodium indistinct, born on abruptly narrowed base of achene: pappus of 8-10 

long, plumose, persistent setae, tips of lateral hairs 

appearing minutely mucronate in Hoyer's solution, outer series of pappus lacking. Pollen grains 

35-37 pm in diameter, with spines rather unevenly dispersed, with distinct internal columellae 

grouped under spines. 

Chionopuppus was originally described by Bentham in Bentham and Hooker (1873) and was 

placed in the Mutisieae in spite of its opposite 

leaves that were nearly unique in that tribe. The 

plumose pappus was evidently a factor in the 

disposition, since plumose setae of a different 

detailed structure are found in the Mutisieae. 

The genus was stated to include a single species 

from Peru, but the species was not named. Blake, 

in 1935, was the first to name the species on 

which Bentham had based his genus over 50 years 

before, and at the same time Blake indicated the 

proper position of Chiompappus as a relative of 

Liubum. Subsequent mentions of the genus have 

followed the disposition of Blake (Cabrera, 1954; 

Robinson and Brettell, 1974b). 

Chiompappus is unique in the Liabeae in the 

form of its pappus, both in the plumose condition 

and in the reduced number of setae in a single 

series. The lateral hairs of the pappus segments 

have the type of complex, internally differentiated tips that are seen on the pappus teeth in 

Liabum and Oligactis. Other peculiarities of Chionopuppus include the reddish-purple flowers of the 

disk, a character once thought to be unique in 

the tribe, but recently the character has been 

found in a species of Philoglossa. The style has 

shorter branches as in most genera of the tribe, 

but material cleared and mounted on microscope 
26 SMITHSOKIAN CONTRIBUTIOKS TO BOTANY 

FICLRE 4.--ChiunupappuJ berrtharnii Blake: A, habit. X k: B. head X 2lh: c. ray corolla. X 10; D. 

disk corolla, X 10; E, disk style, X 10; F. ba5e5 of anther thecae, X 35; c,, achene, X 10: H, achene 

wall showing setulae and raphids, X 200. 
NUMBER 54 27 

slides shows that the stigmatic papillae do not 

stop at the base of the branches but continue well 

down into the stylar canal. The style branches 

differ from the longer forms in Liabum and Oligactis by only the fusion that persists above the 

lower level of the stigmatic papillae. The achene 

has a reduced number of ribs or angles, and its 

base is abruptly constricted to a nearly obsolete 

carpopodium, both characters reminiscent of Cacosniia. The short but distinct tails on the anthers 

are more reminiscent of Liabum, Oligactis, and 

Ferrey ran th 11s. 

The thickenings of the endothecial cells with 

extensions on only the lower ends are most like 

those of the Sincfairia group in Mexico and Central America and the Austroliabum group in Argentina and southern Bolivia. 

The genus contains only the one known species, 

which is restricted to the coastal region of Peru. 

C%ior2ol,n/,/,iti brri//iariiii Blake. Journ. Wash. Acad. Sci., 

2.5: $92. 1935. Peru. 

The species is treated in detail by Blake (1935). 

Ferreyran thus 

FIOIJRE 5 

Frrryj,ran/h~is H. Robinson and Brerrell, Phytologia. 28( 1):50. 

197-1. [Type-species: Atidrornnchia z,erbascij-ilia Humboldt. 

Bonpland. and Kunth.] 

Shrubs or small trees, moderately branched. 

Stems terete to slightly hexagonal, without apparent milky sap. Leaves opposite, bases of petiole 

pairs fused into distinct sheath; petioles distinct, 

narrowly winged; blade ovate to elliptical, base 

usually rounded, margin serrate to subentire, usually rugose with minutely bullate upper surface, 

pale tomentose below, pinnately veined. Inflorescence densely corymbose paniculate, branching mostly opposite: pedicels short, whitish-tomentose. Heads broadly campanulate: 45-55 

strongly graduated involucral bracts in -5 series, 

ovate to lanceolate, tips acute to short acuminate. 

outer surface with evanescent arachnoid pubescence: receptacle with distinct short chaff. Flowers yellou.. Rays 8-12 in single series: base of 

corolla extremely narrow; limbs short, mostly 

glabrous; style branches not spiralled. Disk flowers 12-25; corollas narrowly funnelform, tubes 

slightly narrowed, gradually expanding at base of 

throat, outer surface with glands, with few hairs 

sometimes on tips of lobes: lobes with stomates 

near margins: filaments smooth to densely papillose, cells of anther collar with few or no annulations on walls; thecae pale, short-tailed and 

strongly fringed at base; median endothecial cells 

oval to oblong, usually with short protruding 

single nodular thickenings at upper and lower 

ends of sclerified shields; appendages oblongovate, 1Y2-2 times as long as wide, smooth with 

elongate cells having mostly oblique end walls: 

nectary short and unlobed: style base with a 

distinct node: style branches of disk flowers elongate, -10-12 times as long as wide. Achenes 

prismatic, - 10-ribbed, variously setiferous and 

glanduliferous. cells of achene wall with elongate 

raphids: carpopodium stopper-shaped, cells 

rather small, in 6- 10 series, with thickened walls: 

pappus with inner series of 10-15 persistent setae, 

tips slightly to distinctly broadened, pointed, 

teeth simple, with outer series of 10-15 short 

narrowly squamiform setae. Pollen grains 25-40 

pm in diameter, with spines rather unevenly dispersed. with distinct internal columellae grouped 

under spines. 

Ferreyranthus has a pappus with a capillary inner 

series and a short outer series, the type of pappus 

that is found in Liabum and most other genera of 

the tribe. For this reason, the position of the 

species of Ferreyranthus in Liabum was not questioned until the tribal revision of Robinson and 

Brettell (1974b). The genus is most easily distinguished from Lrabum by the shrubby or arborescent habit, but also it differs by the shorter style 

branches of the disk flower, the mixed glands and 

setulae of the achenes, the presence of elongate 

raphids in the cells of the achene walls, and the 

presence of stomates on the disk corolla lobes. 

Ferreyranthus remains in the relationship of Lrabum 

by the apparent lack of latex in the stems and by 

the presence of well-developed tails on the anther 

bases, but the teeth of the pappus setae do not 
2% SMITHSONIAN CONTRIBCTIONS TO BOTANY 

FIGVRE .i.-Ferre)mn/htc cerbusczjuliiiJ (Humboldt. Bonpland, and Kunth) Robinson Cyr Brettell: 

A, habit, X %; B, head. X 392; c. disk corolla. X 18; D, upper filament and bases of anther 

thecae, X 35; E. disk style, X 18; F, achene. X 15. 
NUMBER 54 29 

have the minute mucro-tips that are seen in 

Liabum, Oligactis, and Chionopappus. The shrubby 

habit, bullate leaf surfaces, and nodal sheaths of 

the petiole bases might reflect a close relationship 

between Ferreyranthus and Cacosmia, but a number 

of characters, including the form of the sclerified 

shields in the endothecial cells, have led to a 

rejection of that idea in this treatment. The latter 

points are discussed under Cacosmia. 

The genus Ferreyranthus is notable for some of 

the largest members of the tribe, with specimens 

of F. verbascz3folius and F. excelsus being described 

as small trees. The anthers of Ferreyranthus have 

the most highly developed tails in the tribe, with 

a fringe of long projecting cells near the tip. The 

common F. zlerbasciflius is the only member of the 

tribe with a hirtellus anther filament, such projecting cells apparently being absent from even 

the closely related F. rugosus. Four of the species 

of Ferreyranthu.r have bullate leaves, but the upper 

leaf surface in F. excelsus and its relatives is smooth 

or nearly so. 

Ferreyranthus was once common on slopes along 

roads in southern Ecuador, but according to R.M. 

King in his most recent visit, this and other woody 

plants have mostly disappeared in recent years, 

apparently the victims of a mobile, mechanized 

pulping operation. It will be interesting to see if 

there is any capacity for recovery. 

The genus Ferreyranthus is mostly restricted to 

Peru, where all seven species occur. The most 

common species, F. verbasczfolius, also occurs in 

southern Ecuador. 

Ferreyranlhus e.ucrl~ui (Poeppig and Endlicher) H. Robinson 

Andtutnachia rwcel.\a Poeppig and Endlicher, No\,. Gen. & 

Liahurn excel~um (Poeppig and Endlicher) Blake, Journ, 

Ferrql'tatithus frutirosus (Muschler) H. Robinson, Phytologia, 

Liabum JiultcoJutn Muschler, Engl. Bot. Jahrb. 50, Beibl.. 

Oligaclis J'rulirosa (Muschler) H. Robinson and Brettell, 

Ferrqymn//ius rarnunii H. Robinson. Phytologia, 5 l(3): 170: 

arid Brettell. Phvtologia. 28( 1):5 1, 1974. Peru. 

Sp.. 3:44, 1843. 

Wash. .4cad. Sci.. 17:293, 1927. 

,51(3):169, 1982. Peru. 

3:81. 1913. 

Phytologia. 28(1):38, 1971. 

1982. Peru 

Fetreyranlhus rugosus (Ferreyra) H. Robinson and Brettell. 

Phytologia. 28(1):51% 1974. Peru. 

Lzahuin rugosurn Ferreyra. Publ. Mus. .'Javier Prado" Bot.. 

ser. B., 20:s. 1965. 

Fme)ranthus z'aginans (Muschler) H. Robinson arid Brettell. 

Phytologia. 28(1):51, 1974. Peru. 

Liahurn r'aginanj Muschler, Engl. ISot. ,Jahrb. 50, Beibl.. 

3:79. 1913. 

Ferreyranthus r~erhasctfolius (Humboldt. Bonpland, and Kiinth) 

H. Robinson and Brettell. Phytoloyia. 28(1):>1, 1974. 

Ecuador, Peru. 

z4ndromachia r'erbasr~jolia Humboldt. Bonpland. and Kunt h. 

Nov. Gen. et Sp.. ed folio, 1:79, 1818. 

Liabutn wthasc$~liurn (Humboldt, Bonpland. and Kunth) 

Lessing, Linnaea. 6: 700, 183 1, 

Liahurn salr~iifolturn Hieronynius, Engl. Bot. Jahrb., 28:622. 

1901. 

Liabutn pseudnsali'i2S0l2utn Hieronyn~us. Engl. Hot. Jahrb., 

36:502. 1905. 

F~rreyrunthus pseududLi@ius (Hieronynius) H. Robinson 

and Brettell, Phytologia. 28(1):5 1, 1971. 

Ferrqyranthus vernnniozdes (Muschler) H. Robinson and Brettell, 

Phytologia, 28(1):51. 1974. Peru. 

Liabum lowrii Cabrera, Bol. Soc. Argent. Bot.. 10:29. 1962. 

F~rt~pznlhus /or'urii (Cabrera) H. Robinson and Brettell. 

Phytologia. 28(1):51. 1974. 

There is no key to all the species, but the 

distinctions of some are mentioned by Robinson 

(1978a), and those with smooth upper leaf surfaces are discussed by Robinson ( 1982). 

Lia bellum 

FI(;IIRE 6 

Liabellutn Rvdberg, North ,%mer. Flora. 31(1):294. 1927. 

/Type-species: L~nhurn palmeri '4. Gray.] 

Low, unbranched or rarely branched, perennial herbs from a tuberous root. Stems short, with 

short internodes, terete, with dense white arachnoid tomentum, with milky sap? Leaves opposite, 

sessile, narrowly to broadly perfoliate, base sometimes constricted and petioliform, margins usually lobed to deeply lobed, sometimes bipinnatifid, rarely simply serrulate, rather smooth and 

pilose above, densely white-tomentose below, 

strongly trinervate from near basal 1/3 or 1/4. 

Inflorescence thyrsoid to scapose, branching opposite below or alternate throughout: pedicels 

elongate, 3- 10 cm long, white-tomentose, with or 
30 SMITHSONIAN CONTRIBUTIONS TO BOTANY 
NUMBER 54 31 

without purple long-stipitate glands. Heads 

broadly campanulate; 20-40 involucral bracts in 

-4 series, unequal, narrowly ovate to lanceolate, 

tips narrowly obtuse to acuminate, outer surface 

with whitish arachnoid tomentum, with or without purple long-stipitate glands; receptacle plane 

to slightly convex, slightly alveolate, with or without short hairs. Rays lacking. Disk flowers yellow, 

25-35; corollas narrowly funnelform; tubes narrow, moderately expanding at base of throat, 

outer surface puberulous to hirsute; lobes with 

stornates near the margins, often with stipitate 

glands at tip; filament smooth, cells of anther 

collar without obvious annular thickenings on 

walls; thecae pale, scarcely crenulate at base; 

median endothecial cells oblong, with oval to 

strap-shaped sclerified shields bearing spiral ornamentation and 1 to many nodes usually at 

lower ends; appendages oblong-ovate, 1 Y2 times 

as long as wide, smooth, elongate cells with truncate to oblique walls; nectary nearly twice as high 

as wide, with irregularly crenulate margin: style 

base broadened above nectary, but node indistinct until shaft withers; style branches 1/4-2/3 

as long as the hispidulous upper portion of the 

shaft. Achenes up to 3 mm long, prismatic, with 

- 10 ribs, densely long-setuliferous, cells of achene 

wall with elongate raphids; carpopodium a narrow rim with rather small subquadrate cells in 4- 

8 series, walls slightly thickened, densely beaded; 

pappus of 40-50 persistent setae, up to 7-13 mm 

long, tips not broadened, teeth simple, with outer 

series of 20-40 short usually narrowly squamiform 

setae. Pollen grains mostly 40-45 pm in diameter, 

with spines rather unevenly dispersed, with a few 

large distinct internal columellae grouped under 

spines. 

Liabrllum is closely related to Sinclairia, with 

which it forms a northern element of the tribe 

centered in Mexico. The species of Liabellum are 

mostly restricted to western Mexico in the area of 

Jalisco and Sinaloa. In the original description of 

the genus, Rydberg (1927) mentioned the herbaceous habit and tuberous roots, which are the 

most notable distinctions from Sinclairia. Rydberg 

also emphasized the acute involucral bracts in 

Liabellum, but such bracts are also found in species 

placed here in a broadened concept of Sinclairia. 

The leaf dissection cited by Rydberg is true of all 

members of Liabellum except the recently described L. gentvli, but all species of Liabellum have 

leaves that are sessile or lobed from the base. In 

Sinclairia, all the species have unwinged petioles 

and lack lobes at the nodes. 

All species of Liabellum and some species of 

Sinclairia lack rays, being the only discoid members of the tribe Liabeae. The condition is unquestionably derived in the tribe. 

Liabellum has an unenlarged style base or a base 

that appears broader only after the style shaft has 

withered, a condition close to that of Sinclairia; 

however, the nectary is not as elongate as in 

Sincla iria. 

The genus contains the following four species 

from southwestern Mexico. 

LinbPllurn angus/i~simuin (A. Gray) Rydherg. North Arner. 

Liabiim angustissirnuin .A. Gray. Proc. Amer. .-\cad.. 22:432, 

Liabellum cerisinum (B.L. Robinson) Rydberg. North rimer. 

Liaburn cerwium B.L. Robinson, Proc. Amer. Acad.. 29:3 17. 

Liabelluin gentryi H. Robinson. PhytoloSia. 4 1 (1):46. 1978. 

Liabellurn palmeri (A. Gray) Rydherg. North Anier. Flora. 

Liaburn palnieri A. Gray. Proc. Anier. .-\cad.. 22:432, 1887. 

Flora, 34(4):295. 1927. Mexico. 

1887. 

Flora, 34(4):'294, 1927. Mexico. 

1894. 

Mexico. 

34(4):295, 19'27. Mexico. 

A key to the three species with dissected leaves 

is provided by Rydberg (1927). 

Liabum 

FILLIR~ 7 

Liabum Adanson, Fam., 2: 13 1, 1763. [ Ixcrotype: Liaburn brownei Cassini (= L. iimbtllatum (L.) Schultz-Bipontinus).] 

Starkea Willdenow, Sp. PI.. 3:22 16, 1803. [Type-species: 

Amellus umbellatus L.: originally monotypic.] 

Andromachla Humboldt and Bonpland, PI. Aequin.. 2: 104, 

1809. [Type-species: .Indromachin igniarin Humboldt. Bonpland, and Kunth.] 

Allendea Llave and Lexarza. Nov. \'eg. Descr.. 1:lO. 1824. 

[Type-species: .4llendea lunc~olata L.lave arid Lexarza (= 

Liaburn bourgeaui Hieronymus) .] 
32 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

L’iuiania \Yilldenow ex Leasing, Linnaea, 4:3 18, 1829. [Typespecies: I’it3zania bicoior Willdenow ex Lessing (= Liabum 

mPia.rtornoidP.t (Humholdt, Bonpland, and Kunth) LessiW.1 

Terrestrial perennial herbs or subshrubs, sparingly branched, rarely subscandent. Stems 

scarcely to strongly hexagonal, thinly to densely 

tomentose, rarely glabrous, apparently without 

milky sap. Leaves opposite, bases of petioles connected across the node by a wing which is often 

expanded into a nodal disk; petioles narrowly to 

broadly winged; blade broadly ovate to narrowly 

elliptical or lanceolate, base acute to slightly cordate, never angulate, margins remotely to closely 

serrulate or denticulate, upper surface nearly flat 

or with slightly prominulous veins, densely tomentose below, trinervate from near base (pinnate in L. diehlii). Inflorescence cvmose, often 

subumbellate, terminal or partly axillary; pedicels less than 5 cm long, covered with white 

arachnoid tomentum. Heads broadly campanulate; 50-150 involucral bracts in -5 unequal 

series, mostly lanceolate to linear, acute to acuminate, outer surface usually with evanescent 

arachnoid pubescence; receptacle with bristly 

chaff or high ridges. Flowers yellow. Rays 20- 

120, usually in 2 series; corolla with tube narrow, 

nearly glabrous, or puberulous above, limb short, 

mostly glabrous, minutely bilobed at tip; style 

branches not spiralled. Disk flowers 10-85; corollas narrowly funnelform, glabrous or with upper 

tube and lower throat puberulous, throat not 

abruptly expanded at base, lobes with stomates 

rare or lacking, tips smooth or densely spiculiferous, sometimes with slender arachnoid hairs; filament smooth; cells of anther collar with walls 

not strongly annulated; thecae pale, digitate at 

base, median endothecial cells oblong to elliptical 

with 1-2 thickenings on transverse walls; anther 

appendages oblong-ovate, cells rather elongate 

with truncate or oblique end walls; nectary short, 

not or scarcely lobed; style base with small to 

large node; style branches of disk flowers elongate, 

filiform, often coiled, hispidulous part usually not 

or scarcely extending onto shaft (extending onto 

shaft in L. solidagineum), tips narrowly obtuse. 

Achenes prismatic, 10-ribbed, setuliferous with 

straight white hairs (also with glands in L. sandemanzi), cells of achene wall containing small quadrate raphids; carpopodium distinct, stoppershaped or annuliform, the small subquadrate cells 

in 4-8 series, walls of cells with large, dense, 

beaded thickenings; pappus with 17-40 capillary 

inner setae, with teeth of setae appearing mucronate-tipped in Hoyer’s solution, with an outer 

series of short narrow squamellae or setae. Pollen 

grains 25-35 pm in diameter, with spines rather 

unevenly dispersed, with distinct internal columellae in groups under spines, columellae sometimes very narrow and numerous. 

The pollen of the Liabum-Oligactis generic pair 

is 23-35 pm in diameter, which is smaller than 

that in many other genera of the tribe. The lack 

of stomates on the lobes of the disk corolla is also 

an unusual condition in the tribe, stomates being 

present in all other genera except Cacosmia. 

The Andean members of Liabum were placed 

in a separate genus, Andromachia, by Humboldt 

and Bonpland (1809), and the Mexican species 

was the basis of Allendea Llave Sr Lexarza described in 1824. Both names seem to owe their 

existence to purely geographical considerations, 

since the species involved possess no basic differences from the typical Liabum of the West Indies, 

except perhaps a tendency to be more strongly 

trinervate. The groups are not recognized here 

even as subgenera. 

A more complex situation is found in the generic segregate Oligactis Cassini, which was resurrected by Robinson and Brettell (1974b). The 

genus Liabum was distinguished from Oligactis in 

the latter work by the mostly nonscandent habit, 

the trinervate rather than pinnate leaf venation, 

the subumbellate rather than corymbose or thyrsoid inflorescence, and the achenes without 

glands. Since 1974, the styles in Liabum have been 

seen to differ by the usually short zone of the 

upper shaft bearing hairs. The latter character 

finds its greatest exceptions in L. igniarium of 

Colombia and Ecuador and in L. solidagineum of 

Peru and Bolivia. Exceptions to the achene pubescence and leaf venation were not known at the 
NUMBER 54 

FIC~~RE 7.--Lraburn utnbellututn (L.) Schultz-Bipontinus: A, habit, X ’%: B. head, X 3%: C. disk 

corolla. X 10; D, disk style. X 10: E, bases of anther thecae, X 35; F. achene. X 9. 

33 
34 SMITHSOKIAN CONTRIBUTIONS TO BOTANY 

time the genus Oligactis was resurrected, but problems are encountered in species described more 

recently by this author. In L. sandemanii of central 

Peru, the achenes bear glands, and in L. diehlii 

from the Department of Cuzco in Peru, the leaves 

have pinnate venation. The last two examples 

cited are regarded here as parallel occurrences of 

the characters in specialized species of Liabum 

from areas where the genus Oligactis does not 

occur. The longer pubescent area on the style 

shaft correlated with somewhat shorter branches 

in I,. igniarium and L. solidagineum remains most 

likely to represent a character with common origin in Oligactis, and it is regarded here as the 

pleisiomorphic condition in Liabum. As such, Oligactis is still regarded as the genus most closely 

related to Liabum but is not regarded as being an 

immediate relative. The phyletic gap between the 

two genera is no less broad than the gap between 

the species L. solidagineum and OIigactis subgenus 

Andromachiopsis. 

The wide geographical distribution of Liabum 

seems to be mostly of recent origin. The entire 

Mexican and Central American range is due to 

one species, L. bourgeaui, which is very closely 

related to L. asclepiadeum of Venezuela and Colombia and L. ferryrii of Peru. As mentioned 

above, the West Indian element in Jamaica, Hispaniola, and Cuba is an interrelated complex 

that is obviously derived from a single introduction. The only real diversity in the genus occurs 

in the Andes, where most of the species are concentrated in Colombia and Ecuador. The related 

genus Oligactis is nearly confined to the same 

latter area and shows most of its diversity in 

Colombia. It is assumed that the basal stock of 

the generic pair originated in this more northern 

part of the Andes. 

The time of origin of the Liabum-Oligactis generic pair may be of interest if the geographic 

area of origin is Colombia. The northernmost 

Andes are the most recently elevated part of the 

range. Both the diversity within the generic pair 

and the phyletic gap between them and other 

genera would indicate that a considerable time 

period is involved in the evolution of the group. 

The group may antedate the general availability 

of habitats in the Colombian Andes. It is possible 

that the earliest members of the Liabum-Oligactis 

line occurred in Colombia in an outlying area 

such as the Nevada de Santa Marta before suitable habitats were available elsewhere in that 

region, but it is notable that the likely primitive 

types of Liabum such as L. igniariunz are not now 

found on Santa Marta or north of southern Colombia. It should also be noted that the lower 

elevation species of Liabum, L. acunzinatum and L. 

amplexicaule, by all indications, are specialized in 

the genus and have recently extended their range 

from Colombia or Ecuador in the north to Peru 

and Bolivia in the south along the Amazonian 

fringe of the Andes. 

The genus contains the following 37 species 

from Mexico, Central America, the West Indies, 

and the Andes of South America. 

Liabum arurninatum Rusby. Descr. S. Xmer. PI.. 161, 1920. 

Liabum amplexicaule Poeppig and Endlicher. Nov. Gen. et Sp.. 

Liabum ulei Hieronymus, Verh. Bot. Ver. Brand., 68:206, 

Liabum asclepiadeum Schultz-Bipontinus, Linnaea, 20:52 1; 

Liabm caliense Hieronymus, Engl. Bot. Jahrb., 28:623. 

Liabum suhurnbellatum Rusby. Deacr. S. Amer. PI., 159, 1920. 

Liabum barahvnense Urban, Arkiv. Bot. (Stockh.). 23A(1 1):85. 

1931. Dominican Republic. 

Liabum barclajae H. Robinson. Phytologia. 34(3) :286. 1976. 

Ecuador. 

Liubzirn hourgeaui Hieronymus in C'le, Verh. Bot. \'er. Prov. 

Brand., 48:208. 1907. Mexico, Central America. 

Allendfa lunreolata Llave and Lexarza, Nov. \.eg. Descr.. 

1: 10. 1824 [not L. /anceo/alum Schultz-Bipontinus]. 

Lzabum crispurn Schultz-Bipontinus. Jour. Bot.. 1 :236. 1863. 

Cuba. 

Liabuni cubense Schultz-Bipontinus, Jour. Bot., 1:236. 1863. 

Cuba. 

Liabum dirhlzi H. Robinson, Phytologia. 46(2):99. 1980. Peru. 

Liabum eggerszi Hieronymus. Engl. Bot. Jahrb.. 28:624. 190 I. 

Liabum erioraulon Poeppig and Endlicher. Nov. Gen. et Sp.. 

Liabumjalratum Rusby, Descr. S. Amer. PI.. 161. 1920. CoEcuador, Peru. Bolivia. 

3:43, 1843. Ecuador. Peru. 

1907 [ 19061. 

1817. Colombia. \.enezuela. 

1901. 

Ecuador, Peru. 

3:43. pl. 249, 1843. Peru, Bolivia. 

lombia. 
NUMBER 54 35 

Liahum Jerrp~rii H. Robinson. Phytologia. 31(3):287, 1976. 

L2ahum ,/liiribundrm Lessing. Linnaea. 6: 702. 183 1. Ecuador. 

Liahutn grandfloruni (Humboldt, Bonpland. and Kunth) Lessilndrornachia grand2flora Humboldt. Bonpland. and Kunth, 

Licihurn wdvrbaucri Muschler. Engl. Bor. Jahrb. 50. Beibl, 

Liahrrni amp1esan.r Blake. Joiirn LVash. .&cad. Sci.. 17:292. 

Liaburn /,piuriutn (FIumboldt. Bonpland, and Kunth) Lessing, 

ilndtornachia i~qniarru Humboldt. Bonpland, and Kunth. 

Liahntn bonplnrrdii Cassini. Dict. Sci. Nat.. 26:206, 18213. 

I,iabutri luhrnannii Hieronymus. Engl. Bot. Jahrh., 19:61. 

1894. 

Liahurn hinpii H. Robinson. Phytologia, 34(3):288, 1976. Ecuador. 

Liabuin inacbridei H Robinson. Phytologia. 34(3):290, 1976. 

Peru. 

Liaburn tnelas/otnoides (Humholdt. Bonpland. and Kunth) 

Lessinq. Linnaea. 6:(i99, 183 1. Colombia. 

Andrum/rhia rnelastotnuide.~ Humholdt . Bonpland. and 

Kunth, Nov. Gen. et Sp.. ed folio, 4:79. 1818. 

Liahuvi nigro-pilu.sinn Hieronymus. Engl. Bot. Jahrh.. 29:59, 

1900. Ecuador. 

Liahun7 tiudrcoulc €1. Robinson. Phyrologia, 35( 1):37. 1976. 

Peru 

Linbirtn oblarrrcolaturri Urhan and Ekman. Arkiv. Bot. 

(Stockh.). 2Sh( 11):89. 1931. Dominican Republic. 

I.inhun~ ozat~Ji~liirni Urban. hrkiv. Bot. (Stockh.), '13A(1 l):86. 

Liubiirn poiteuicr (Cassini) Urban. Arkiv. Bot. (Stockh.). 

Aiidrorncrchra poitecm/ Cassini. B~ill. SOC. Philoni.. 1817: 184. 

Liabutn poiycephaliciri Urban, Arkiv. Bot. (Stockh.). 23'4 

Liahutii \(iIfpn.sc Domke in Diels, Bibl. Bot.. 116: 168. 1937. 

Liabuni .\andrmanir €1, Robinson. Phytologia. 35(6):488. 1977. 

Liabrini ~elleunum Urban. Arkiv. Bot. (Stockh.). 23A(1 1):86. 

Liabum \olidugitieurn (Humboldt, Bonpland. and Kunth) Less- 

.Indiomnr/ua .\olidaginm Humholdt. Bonpland, and Kunth, 

Liabuni jiuli~otorncntosurri 0, Kunrze, Rev. Gen.. 3(2): 163. 

Peru. 

Peru. 

ing. Linnaea. 6:701. 1831. Ecuador. 

Nov. (;en. et Sp.. ed folio. 4:77. 1818. 

3:78. 1913 

19'7. 

Linnaea. 6:698. 183 1. Colomhia. Ecuador. 

Nov. (;en er Sp.. ed folio. 4:77. 1818. 

193 1, Dominican Republic. 

.\(I 1):87. 1931. Haiti. 

1817. 

(I 1):88. 1931. Haiti. 

Colomhia. Ecuador. 

Peru. 

1931. Haiti. 

ing. Linnaea. 6: 700. 183 1. Peru. Bolivia. 

Kov. <;en. et Sp.. ed. folio. 1:78. 1818. 

1898. 

Liabum acutfolzutn Cuatrecasas, Collect, Bot.. Barcinone. 

Liaburn steinbachzi H. Robinson, Phytologia. 3.5(6):489. 1977. 

Liabum slipulatum Rushy. Descr. S. Amer. PI.. 160, 1920. 

Liabum subacaule Rydberg, North hier. Flora. 34(4) :290. 

Liabum domingense Rydberg. North Anier. Flora. 34(4):29 1, 

Liabum triariae H. Robinson. Phy-tologia. 34(3):29l. 1976. 

Lzaburn umbrllatum (L.) Schultz-Bipontinus, Jour. Bot.. 1 :236. 

3:299. 19.73. 

Bolivia. 

Colombia. Ecuador. 

1927. Haiti, Dominican Republic. 

1927. 

Colombia. 

1863. Jamaica. 

A4rnellus umbe1Ialu.s L.. Syst. Nat. ed.. 10: 122.5, 17.X. 

Starkea umbellala (L.) LVilldenow. Sp. PI 

Liaburn bruwwz Cassini. Dict. Sci. Sat.. 26:203. 1823. 

Liabum cargasii H. Robinson, Phytologia. 34(3):29'2, 1976. 

Liabum wright~i Grisebach, Mem Xmer. .-\cad.. n. ser.. 8:5 1.5. 

Liaburn longipes Urban. Fedde. Rep. Sp. Nov.. 26 115, 1929. 

Liabum wurdackii Ferreyra, Publ. hlus. Hist. Nat. "Javier 

Peru. 

1862. Cuba. 

Prado" Bot.. ser. B.. 20:2. 1965. Peru. 

Rydberg (1927) has provided a partial treatment of the species north of South America, but 

there have been considerable changes since that 

time. The Ecuadorian species have been treated 

by Robinson (1978a), and some additional distinctions of species are mentioned by Robinson 

(1976~). 

Microlia bum 

FIGI.RE 8 

Mirroliabum Cabrera, Bol. Soc. .L\rgenr. Bot.. 5:21 I, 19.55. 

Liabellum Cahrera. Not. hlus. La Plata. 17(84):76. 1934 [not 

Liabellum Rydberg, 19271. [Type-species: Lidellurn hurnile 

Cahrera.] 

Angelianthus H. Robinson and Brettell. Phytologia, 28(1):48. 

1974. 

Annual herbs to 10 cm tall, unbranched or 

sparsely branched; stems covered with appressed 

white tomentum and hirsute with coarse minutely 

gland-tipped purple hairs; internodes short to 

-I cm long. Leaves opposite, sometimes congested at base of plant: bases of leaf pairs fused 

into short sheath, without distinct petioles: lamina elliptical to subrhomboid, faintly trinervate, 
36 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

FIG~.RE 8.--,Mirrolzaburn humile (Cahrera) Cahrera: 4. habit. X 1: B. head. X 3’12: c. ray corolla. 

X 13; D. disk corolla, X 16; E. disk style. X 16; F, anther, X 16; c, achene. X 16: H. elenient of 

inner pappus, X 35. 

entire to subdentate, acute at tip, green and 

smooth above, glabrous to coarsely pilose, whitetomentose below. Heads single on slender peduncles or scapes, broadly campanulate. Involucral 

bracts -10-20 in -2 series, subequal, ovate to 

broadly lanceolate, apex acute, hirsute with purple glandular hairs on outer surface; receptacle 

flat to slightly convex, foveolate, without chaff. 
NUMBER 34 37 

Flowers yellow. Ray flowers 10-18; tubes hirsute; 

limbs narrowly elliptical, with tinge of red, with 

few to many short hairs on outer surface: style 

branches elongate, not spiralled. Disk flowers 

- 15; corollas glabrous below, tubes expanding 

gradually into throat; lobes with large stomates 

near margin, with numerous short glandular hairs 

on outer surface and with cluster of stout nonglandular hairs near tip: anther collars with annular thickenings on walls; thecae pale, not digitate at base; median endothecial cells oblong, 

sclerified shield mostly with weak irregular coiled 

ornamentation, often with long uncinate extension on lower end; appendages smooth, with 

shortly oblong to subquadrate cells; style base 

with prominent node, branches short, -0.5 mm 

long. Achenes prismatic, obovate, with mostly 8 

shallow costae with long setae, intercostal areas 

glabrous, cells of achene wall with elongate raphids; carpopodium with narrow rim of small, 

thin-walled, firm, subquadrate cells in -2-3 rows; 

pappus distinctly biseriate, short outer squamae 

positioned over furrows of achene, inner easily 

deciduous, broadly winged by fused spreading 

lateral cells, lanceolate to oblanceolate, apices of 

cells simple. Pollen grains -35 pm in diameter, 

with spines unevenly dispersed, with distinct internal columellae. 

Microliabum was described by Cabrera (1954, as 

Liabellum) during a period in the history of the 

tribe when genera were based exclusively on pappus characters. In the case of Microliabum, the 

distinction was the broad lamellate form of the 

inner pappus segments. a form resembling the 

outer pappus segments in width. Though segments of the outer series are sometimes broad in 

other genera of the Liabeae, the inner segments, 

when present, are always capillary. The pappus 

difference, the comparatively short anther bases, 

and the small size of the plants provide ample 

distinction for the species, which is specialized for 

higher elevations in the southern Andes. Nevertheless, critical examination indicates close relationship to the more robust species from lower 

elevations in the same area of the southern Andes, 

which until recently were placed in Liabum. Both 

Microliaburn and the related species have the same 

prominent costae on the achene, a prominent 

short outer pappus, a very deciduous inner pappus, elongate raphids in the cells of the achene 

wall, and an identical form of thickenings in the 

short endothecial cells. In spite of the similarity, 

the separation seems best, and a separate generic 

name, Austroliabum, has been provided for the 

related group. 

The genus Microliabum was originally known 

from only Jujuy in northern Argentina but since 

has been collected farther south in the Province 

of Salta. It is likely to be found farther north, 

also, in adjacent southern Bolivia. 

The original illustration of the genus by Cabrera (1954) shows a plant with strictly rosulate 

leaves and strap-shaped inner pappus segments 

without a stipitate base. Material seen from the 

Province of Salta has a slight elongation to the 

vegetative stem, and the segrnents of the inner 

pappus are oblanceolate with a long narrow base 

(see Figure 8). It is possible that more than one 

species is involved, but this cannot be confirmed, 

since the herbarium at Tucurnan has refused to 

send the type specimen from Jujuy on loan and 

has not replied to requests for information on 

details of the plant. Except for the kindness of Dr. 

Zardini at the Universidad Nacional de La Plata, 

who arranged for the loan of a fragment of some 

Salta material, the genus would have been completely unavailable for this study. It is hoped that 

further material will be collected of this inadequately known genus. 

The genus is monotypic with the species as 

fOllO\VS. 

.Mirroliaburn hurnile (Cabrera) Cabrera. Bol. Soc. Argent. Bot ., 

Liahrllurn humiie Cabrera. Sot. Mus. La I'lata. 1 7(84): 78. 

,4ngelzunfhus huiniiis (Cabrera) H. Robinson and Brettell, 

5:21 1. 195.5. Argentina. 

1954. 

Phyrologia 28(1):48, 1971. 

Oligac tis 

FIC;r:RE 9 

OlzgactiJ (Humboldt, Bonpland. and Kunth) Cassini. Dicr. 

Sci. Nat., 36: 16. 1825. 

.Indroinuchia sect. OiigartiJ Humboldt. Bonpland. and Kunth. 

Nov. Gen. et Sp., ed. folio. 4:79. 1818. Il~~t~type: .Iiidrcimurhiu isolubilis Humboldt. Bonpland, and Kunth.] 
38 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

Scandent shrubs and vines, sparsely branching. 

Stems terete to strongly hexagonal, tomentose to 

glabrescent, apparently without milky sap. 

Leaves opposite, with or without nodal disks; 

petioles with or without wings, sometimes included in perfoliate leaf base; blade ovate to 

narrowly linear, base cuneate to rounded or 

slightly cordate, sometimes confluent into petiole 

wings, margins of blade never angulate, subentire 

to serrate, upper surface flat to slightly bullate, 

densely tomentose below, pinnately veined. Inflorescence terminal on branches or axillary, paniculate with spiciform to corymbose branches; 

pedicels less than 5 cm long, thinly to densely 

tomentose. Heads narrowly to broadly campanulate; 16-55 involucral bracts in 4-3 series, ovate 

to lanceolate, tips obtuse to narrowly acute, outer 

surface puberulous to hirsute; receptacle with 

laciniate chaff or prominent ridges. Flowers yellow. Rays 3-18 in l series; corolla with tube 

narrow, glabrous, or puberulous above, limb 

short, elliptical, glabrous; style branches not spiralled. Disk flowers 3-34, corollas narrowly funnelform, tube slender, sometimes slightly swollen 

and sparsely puberulous at base of throat, lobes 

without evident stomates, tips plain or with crests: 

filament of anther smooth; cells of anther collar 

with weak annulate thickenings on walls; anther 

thecae pale, digitate at base, median endothecial 

cells oblong with oval or strap-shaped sclerified 

bands, with a single thickening on the transverse 

walls; anther appendages oblong-ovate, smooth 

or papillose, with short-oblong to subquadrate 

cells: nectary short, not or scarcely lobed; style 

base with distinct node; style branches of disk 

flowers elongate, linear, 1.0-1.5 times as long as 

the hispidulous upper portion of the shaft (longer 

in subgenus Andromachiopszs) , tips narrowly obtuse, tapering in some species. Achenes prismatic 

with 5-8 ribs, bearing contorted setae and small 

glands, cells of achene wall with quadrate raphids; carpopodium short and distinct, usually 

annuliform with small subquadrate cells in 3-5 

series, walls of cells moderately thickened: pappus 

with 20-35 rather persistent inner setae often 

with broadened tips, with teeth of setae appearing 

mucronate-tipped in Hoyer's solution; with outer 

series of 7-20 short setae or squamellae. Pollen 

grains 25-35 pm in diameter, with spines rather 

unevenly dispersed, with distinct internal columellae grouped under spines. 

The relationship of Oligactis to Liabum is discussed under the latter genus. 

01igactis is divided into two subgenera, which 

have distinctive, scarcely overlapping distributions. The less-specialized subscandent subgenus 

Andromachiopsis, having heads in corymbose panicles, usually more than 18 flowers per head, and 

foliar disks on the nodes, is centered in Ecuador 

with extensions into southern Colombia. Three 

species have been found in northern Peru. Most 

of the species are restricted to elevations above 

2000 m, but 0. asplundii from the Pacific slope of 

the Ecuadorian Andes is found as low as 1200 m. 

One species, 0. cusalaguensi.r, has distinctive perfoliate leaves. 

The subgenus Oligactis is the most specialized 

element of the genus, with more scandent plants 

bearing heads in subglomerate, spiciform, or racemose branches in an axillary or terminal position. The anther appendages are unique in the 

tribe by the projecting cell-ends on the surfaces. 

The heads often have about 10 flowers and sometimes have as few as six flowers, three of each 

type. The subgenus occurs primarily in northern 

and central Colombia and extends into Venezuela, Panama, and Costa Rica. In most species, the 

leaves are ovate to narrowlv elliptical, but 0. 

garcia-barrigae has narrowly- linear leaves (Robinson, 1980a). 

Key to the Subgenera of Oligactis 

Inflorescence axillary or terminal, with a subglomerate, spiciform, or racemose 

arrangement of heads; heads usually with 6- 10 flowers; anther appendages 

with papillose surfaces subgenus Oligactis 
NUMBER 54 39 

Inflorescence terminal, with a corymbose arrangement of heads; heads with 

- 18-50 flowers; anther appendages smooth subgenus Andromachiopsis 

The genus contains approximately 12 species, 

which are disposed in the subgenera as follows. 

0ligacti.s subgenus Oligactis 

Oligacfi.\ garcia-barrigae H. Robinson, Phytologia, 46(2): 100, 

1980. Colombia. 

Oligacfis latzfolia (Hieronynius) H. Robinson and Brettell. 

Phytologia. 28(1):57. 1974. Colombia. 

Liaburn colubile var. latfolium Hieronymus. Engl. Bot. 

Jahrh., 28:622. 1901. 

Liaburn lati~olium (Hieronymusj Cuatrecosas. Fedde Rep. 

Spec. Nov.. 5.5: 129, 1953. 

Oligactis mikanioides (Blake) H. Robinson and Brettell, Phytologia. 28(1):57. 1974. Colombia. 

Liabum mikanioides Blake. Journ. Wash. Acad. Sci.. 1 7:294. 

1927. 

Oligactis .\emliflora (Humboldt. Bonpland, and Kunth) H. 

Robinson and Brettell, Phytologia, 28(1):58. 1974. Colombia. Venezuela. 

Andromachia sessilzflvra Humboldt, Bonpland, and Kunth, 

Nov. Gen. et Sp.. ed. folio. 4:79. 1818. 

Oligactis apodocephaln Cassini. Diet. Sci. Nat., 36: 17, 1825 

[noin. superfl.]. 

Liabum sessillflora (Humboldt, Bonpland, and Kunth) Lessing. Linnaea. 6:703. 1831. 

Liabum totarense Badillo. Bol. Soc. \'enez. Cienc. Nat., 

10:314. 1946. 

Uligaclis mleri (Standley) H. Robinson and Brettell, Phytologia. 28(1):58. 1974. Costa Rica, Panama. 

Liabum valeri Standley, Publ. Field Mus. Nat. Hist. Chicago. hot. ser.. 18: 1490, 1938. 

OIigac/is polubilis (Humboldt, Bonpland. and Kunth) Cassini, 

Dicl. Sci. Nat., 36: 17. 1825. Colombia, L'enezuela. 

Andromachia tolubili~ Humboldt. Bonpland, and Kunth 

Nov. Gen. et Sp., ed. folio. 4:80. 1818. 

Liaburn uduhile (Humboldt. Bonpland. and Kunth) Lessing, Linnaea. 6:704. 1831. 

Liabum hiattenuatimi Rushy. Descr. S. Amer. PI.. 159, 1920. 

Liabum bqpacense Cuatrecasas, Not. Fl. Colomb., 6:36, 1944: 

Liabum rneridense Badillo. Bol. Soc. Venez. Cienc. Nat., 

Oligactis biattenuala (Rusby) H. Robinson and Brettell. 

Oligactis hqgacensi.\ (Cuatrecasas) H. Robinson and Brettell, 

Rev. Acad. Colomb. 6:61. 1941. 

10:314. 1946. 

Phytologia, 28(1):57, 1974. 

Phytologia, 28(1):.57, 1974. 

OltgactiJ subgenus Andromachiopsis 

(H. Robinson and Brettell, Phytologia, 28( 10):58. 1974) 

Oligac/is asplundii H. Robinson. Phytologia, 35(3):200. 1977. 

Ecuador. 

Oligacfis cvriacea (Hieronynius) H Robinson and Brertell. 

Phytologia, 28( 1): 58, 1974. Colombia, Ecuador. Peru. 

Liabum corzaceum Hieronymus. Engl. Bot. .Jahrh. 29:58, 

1900. 

Liabum scandens Domke in Diels. Bibl. Bot. no.. 116: l(i7. 

1937. 

Liabum granatens~s Cuatrecasas. Fedde Rep. Sp Nov. 

Oligactia granalen3i.r (Cuatrecasas) €1. Robinson and HretOligactis scaridens (Domke) H. Robinson and Brettell. I'hyOligactis coriacea var. granatensi.r (Cuatrecasas) H. Robinson. 

Oligaclis cusalaguensis (Hieronynius) H, Robinson and BretLiabum cusalaguense Hieronymus. Engl. Bot. Jahrb., 29:.55. 

Oligactis ecuadoriensis (Hieronymus) H. Robinson and Brettell. 

Liabum ecuadoriense Hieronymus, Engl. Bot. Jahrh.. l9:hO. 

Oligactis ochracea (Cuatrecasasj H. Robinson and Brettell, 

Liabum ochraceum Cuatrecasas. Collect. Bot. Barcinone, 

Oligactis pastoensis (Cuatrecasas) H. Robinson and Brettell. 

Liabum pasloense Cuatrecasas. Not. Fl. Colomb., 6:N. 19.t-1; 

Oligacfis pichinchensis (Hieronymus) H. Robinsor) and BrerLiabum pichinchense Hieronymua, Engl. Bot. Jahrh., 29:.56, 

Liabicm ha1111 Hieronynius, op. cit.. 57. 

Olzgactis hallii (Hieronymus) H. Robinson and Brettell. 

128, 19 

tell, Phytologia, 28( 1):58, 1974. 

tologia, 28(1):59. 1974. 

Phytologia, 35(3):20 1, 1977. 

tell, Phytologia, 28( 1) :58. 1974. Ecuador. 

1900. 

Phytologia. 28( 1):58, 1974. Ecuador. 

1894. 

Phytologia, 28(1): 58. 1974. Peru. 

3:302. 19.53. 

Phytologia, 28( l):.58. 1974. Colombia. 

Rev. Acad. Colomb.. 6:62. 1944. 

tell, Phytologia. 28( 1):59, 1974. Ecuador. 

1900. 

Phytologia, 28( 1):.58. 1974. 

Incertae sedis 

Oligactis nubigena (Humboldt. Bonpland. and Kunth) Cassirii. 

Andromachia nubigena Humboldt. Bonpland. and Kunth. 

Liabum nubigenum (Humboldt. Bonpland. and Kunth) LessDiet. Sci. Nat., 36: 16. 1825. Ecuador.' 

Nov. Gen. et Sp.. ed. folio. 4:79. 1818. 

ing. Linnaea, 6:704, 183 I. 

There is no available treatment of the genus, 

but Robinson (1978a) has provided a key to the 

Ecuadorian species, and the two species from 

Venezuela have been treated by Aristeguieta 
40 SMITHSOSIAN CONTRIBUTIONS TO BOTANY 

FICIIRE 9.--012gac/i.s i~oI~bi/is (Humboldt. Bonpland. and Kunth) Cassini: ,A, habit. X ?$; B. head, 

X 7; c, ray corolla. X 13: D. disk corolla, X 13: E. disk style. X 13; P, anther, X 25: c;. achene, 

X 13. 
NUMBER 54 41 

(1964). Additional comments of value can be 

found in Cassini (1825: 16-18) and in Steyermark 

(1957:l 1.51). 

Sin cla iria 

S’rriciuirz/i Hooker and Arnott. Bot. Beech. Voy.. 433. 1811. 

[Type-species: Srrirfnirio dzscoior Hooker and Arnott: originally monotypic.] 

LSrrrc/uz/zufisr,r Rydberq. North Amer. Flora. 31(4):292. 1927. 

17’)ye-speciea: Lzcibutn X/n//ir B.L. Robinson and Greeniiiaii: ori~inall!. monotypic.] 

,Ll~,~u/iclhiitn Rydherg. North Amer. Flora. 34(4) :293, 1927. 

17’)-lx-species: T’moniu utzdrzeurir DeCandolle.] 

Erect to scandent subshrubs, shrubs, or vines, 

sparingly branched. Stems terete to slightly hexagonal, with sparing often evanescent arachnoid 

tomentum: with milky sap. Leaves opposite or 

ternate, often defoliated at anthesis, without nodal disks or sheaths, petioles distinct, narrow, not 

noticeably winged; blade ovate to suborbicular 

or triangular, base broadly cuneate to truncate, 

margin remotely mucro-denticulate to coarsely 

irregularly dentate, upper surface smooth to 

slightly roughened, with or without coarse hairs, 

thinly to densely whitish pubescent belo\v, 

strongly trinervate at or near base. Inflorescence 

terminal or partly axillary, corymbose to thyrsoid 

paniculate with corymbose branches, with few to 

many heads: pedicels 0.1-4.0 cm long, hirtellous 

to white-tomentose. Heads narrowly to broadly 

campanulate: 18-45 involucral bracts in 3-5 series, unequal, oblong to lanceolate, tips broadly 

rounded to narrowly acuminate, outer surface 

subglabrous to densely tomentose; receptacle 

plane, sometimes with minute spines, glabrous or 

with minute hairs. Flowers yellow. Rays when 

present 1-25 in 1 series; corolla with tube narrow, 

limb broadly to narrowly elliptical, sometimes 

obsolete, glabrous to minutely puberulous. minutely bilobed at tip: style branches elongate, not 

spiralled. Disk flowers 5-30; corollas narrowly 

funnelform, tube narrow, moderately expanding 

at base of throat, glabrous below, becoming 

densely puberulous above, lobes with stomata 

near margins, sometimes piliferous at tip: filament smooth. anther collar with few or no 

sub q u ad ra t e cells be 1 ow ? some t i nies wit h u nsc 1 erified lower cells, walls without obvious annular 

thickenings; thecae pale, minutely crenulate at 

base; median endothecial cells oblong, Lvith oval 

to strap-shaped sclerified shields, shields spirally 

ornamented with 1 to many nodes at lower end; 

appendages oblong, 1-2 times as long as \vide, 

smooth, cells elongate with mostly oblique end 

walls; nectary with irregular crenulations or lobules; style base usually without nodular expansion; style branches elongate, LO- 15 times as long 

as wide, often less than 1/2 as long as hispidulous 

upper part of shaft. Achenes prismatic, -8-10 

ribbed, up to 2.5-7.0 mm long, glabrous to 

densely setuliferous, cells of achene \Val1 m,ith 

elongate raphids; carpopodium short-cylindrical 

or annuliform, prominent, with - 15 rows of niinUte subquadrate cells, cell walls slightly thickened, densely beaded; pappus of 3-40 persistent 

setae, mostly 5-8 mni long, 1:3-13 nim long in 3 

species, tips not or slightly thickened. teeth simple. outer pappus with series of 15-20 short, 

narrow, usually squamiform setae. Pollen grains 

33-50 pm in diameter, with spines rather unevenly dispersed. with usually a few large internal 

columellae grouped under spines. 

Sinclairia and Liabellum toget her form a distinctive element of the tribe that is nearly restricted 

to Mexico and Central America and sho\vs considerable diversity in that area. The diversity was 

the basis of four genera recognized by Rydberg 

(1927), two of which are not recognized here. 

Both of the Rydberg segregates synonymized here 

had heads with involucral bracts more obviously 

graduated and more acute than in typical Sinchiria. The genus .kfegaliabum had a normal short 

outer pappus but was notable for its coarse leaves 

and large heads. .kfegu/iubum included S. andrieiixii 

and S. pringlei, and the more recently described S. 

nzoorei would belong to the group. The genus 

Sinclairtopsis was distinguished by the lack of an 

outer pappus series and had heads of intermediate 

size with acute involucral bracts. The single species included in Sinclairiupsis, S. klattii, is notable 
42 SMITHSOXIAN CONTRIBCTIONS TO BOTANY 

FIGURE lO.-Sncluzrm discolor Hooker & Arnott: A, habit, X Yz; B, head, X 3%; c, ray corolla, 

X 8: D, disk corolla, X 8; E, disk style. X 10; F, bases of anther thecae, X 35; G, achene, X 8. 
NUMBER 54 4 :3 

for the lack of leaves at the time of anthesis, but 

this trait is developed to some extent in other 

members of Sinclairia. The seasonal foliage is reminiscent of the situation in Pachythamnus King and 

Robinson of the Eupatorieae and Pittocaulon Robinson and Brettell in the Senecioneae. both from 

the same geographic area. It is also a habit shared 

by many plants of other families such as Cnidoscolus in the Euphorbiaceae. 

Sinclairia has a few traits that are rather distinctive in the tribe. The presence of discoid heads 

in many species is a character found elsewhere in 

the Liabeae only in the closely related Liabellum. 

'4 character not found in any other genera of the 

tribe is the tendency for ternate leaves. Such 

leaves are not an absolute characteristic of any 

species, but they occur commonly in many members of the genus, including some material seen 

of S. (Megaliabum) pringlei. Unenlarged or slightly 

enlarged style bases are found in the subtribe only 

in Sinclairia and Liabellum, whereas elongate nectaries occur only in Sinclairia. The anther collar of 

Sinclairia is unique in the tribe by the nonquadrate or unsclerified basal cells. 

The distinctly petiolate and totally nonstipulate nature of the leaves in Sinclazrza prove to be 

one of the most useful characteristics of the genus. 

This feature distinguishes the genus completely 

from the closely related Liabellum in which the 

leaves are all sessile or lobed from the base or 

even perfoliate. Of the other genera of the Liabeae reaching Central America in at least Costa 

Rica, Liaburn usually has a nodal disk or a petiolar 

wing continuing across the node, and Munnoria 

often has either a winged petiole or a nodal lobe. 

The closest approach to the Sinclairza type of leaf 

insertion in a member of the Liabeae in Central 

America is in Oligactis, which is distinguished by 

its pinnately veined leaves. 

Sinclairia pobanthn has been known from Panama as far east as the Cerro Jeffe area, and there 

has been some suspicion that the genus occurred 

in the adjacent part of South America. Recently, 

an old collection of the species has been seen from 

the Departmento de Caldas in Colombia, Sneidem 

524C, La Selva, 1600-1900 m "I 8 1946.'' 

The genus contains -25 species. which are 

almost entirely restricted to Mexico and Central 

America. The species are as follows. 

Siniluiriu udenoitirha (Greenman) R\.dherq. North Amer. 

Liabuin udetiu/richzirn Greenman. Field h111s. hb1. Hot.. 

Sinciartiu andriei/.rii (DeCandolle) H. liohinsoti aiid Bretrell. 

Flora. 34(4):300. 1927. llexico. 

2:" 1912. 

Phytologia. 28(1):60. 1974. hlexico. Guatemala. 

Yernoritu atidrim\.// DeCandolle. Prodr.. i: l(i. lii.i(j. 

Liuburn atidrieicuzi (DeChidolle) Benthani atici 1 iooker 1. ex 

Hemsley. Biol. Centr. Aiiier.. Bot.. 2:231. 1881. 

.I.ff,gn //a bum atidrimt i I ( DeC a ti doll c I< y cl tier q. Nc )r t h :\ nler. 

Flora. 34(4):293. 1927. 

Smciairiu utidriirnaihiuidr.J (Lessing) Schulrz-Bil,oiititi~i~ ex 

Rydberq. Sorth Amer. Flora. .34(.11:2W~ 1927. Mexico. 

li.rtioiiia undruinnrhioIdea Lessing. Liiinaea. (i::\97. 183 1. 

Liuhurn undrornuchioid~~ (Lessing) Benr ham and 1 looker f. ex 

Hemsley. Biol. Centr. Arner.. Bot.. 2:2:31. 1881. 

Sinciairiu biakei H. Robinson and Brertell. Phyroli)~ja. 

28(1) :60. 1974 lnon Sinc/uiriu /ij/io/tit~~ [C;reenniaii) IZydberg]. hlexico. 

C7tr~iunru hjpoieuru DeCandolle. Procir.. 5:27. l8:3(i. 

Liuburn hJpoieuiurn (DeCandolle) Blake. f'roc. Biol. Soc. 

\I'ash., 39: 144? 1926. 

Siniiairiu brui/?/iu.\ Rydherg. Sorth Amer. Flora. 34(4):2W. 

1927. Guatemala. 

Srtiriuiticc hl-uorneur H. Robinson [as h)o(,ti/ti 1. Ptiytoloqia. 

.33(4):287. 1976: correction. Ph~~roloqia, 41(1 I: 17. 1078. 

Mexico. 

S/nc/uiriu cuduc!fi)/ia (B.L. Robinson and Banletr) I<) dlxrq. 

North Amer. Flora. 24(4):?99, 1927. blexico. 

Liaburn raduiiJoiiiim B.L. Robinson and Kartle~t. Proc. 

Anier. Acad. 4.3: 39. 1907. 

SIriciuiriu deunziz (B.L. Robinson and Barrlett ) IZycIl~er~, Sort11 

Amer. Flora, 34(1):299. 1927, Guatemala. Belize. tlonduraa. El Sal\.ador. 

Liuburn dearnii B.L. Robinson and Bartlett. I'roc. hnier. 

Acad.. 43:60. 1907. 

Liuburn suh~clatiduiurr Blake. Contr. L-.S. Sat. Iierh.. 24:.3l, 

1922. 

Siiiikuiriu suh~giunduiuri, (Blake) R\ dberg. Nort h Amer 

Flora. 34(4):298. 1927. 

Sinilairiu deppeatia (Lessing) Rydberg. North rimer. Flora. 

34(4):300. 1927. Mexico. 

.Iridrornuchiu deppeatia LesinS. Linnaea. (i:401. 183 I. 

Liuburn depp~ariuvi (Lessing) Henislev. Biol. Chitr. Ainer.. 

S'/riiiuzriu dirnidia (Blake) H. Robinson and Brettell. PhytoloLiahutri dirtiidiutn Blake. Journ IVasti. .kid. Sci.. 22:W). 

Sini/uiriu disculor Hooker arid .\rnoit. Bot. Beech. \'(I\. .. -1:i.l. 

Bot.. 2:232. 1881. 

gia, 28(1):6l, 1927. Guatemala. 

1932. 
44 SMITHSONIAS CONTRIBUTIONS TO BOTANY 

184 I. Xlexico. Guatemala. BeliLe. Honduras. El Salvador. Nicaragua. Costa Rica. 

Liahurn dzscolor fHooker and Arnott) Bentham and Hooker 

f. ex Henisley. Biol. Centr. Amer.. Bot.. 2:232. 1881. 

.Yinclainci glublu (Hemsley) Rydberg, North .4mer. Flora. 

34(-1.):297. 1927. Mexico. 

Liuhurn glabrum Henisley. Biol. Centr. Amer.. Bot.. 2:232. 

1881. 

Liuhurn ,qlubrurn var. hlpoleucurn Greenman. Proc. Amer. 

Acad.. 32:294. 1897. 

Si/ic/airiu /gpoleuca (Greenman) Rydberg. North Amer. 

Flora. 34(4):297. 1927. 

S'iricluiriu /{jpoch/vra (Blake) Rydberg, North .Anier. Flora. 

34(4):301. 1927. Mexico. Guatemala. 

Liahlrin /ypochlol-irm Blake, Contr. Gray Herb.. n.s.. 53:27. 

1918. 

Sinciuzriu Xluttzi (B.L. Robinson and Greenman) H. Robinson 

Liahuni kla//ii B.L. Robinson and Greenman. Amer. Joum. 

LSinc/airiup,w klallii (B.L. Robinson and Greenman) RydSinciairiu liebinannil (Klatt) Schultz-Bipontinus ex Rydberg. 

arid Brettell. Ph>-tologia. 28(1):61, 1974. Mexico. 

Sci.. ser. 3. 1:156. 1893. 

berg. North .Anier. Flora. 34(4):293. 1927. 

North Xnier. Flora. 34(4):300, 1927. Mexico. 

Liabum liehmannii Klatt, Leopoldina, 23: 146. 1887. 

Brertell, PhytoloSia. 28(1):61. 1974. Mexico. 

27(4):252. 1973. 

1927. Costa Rica. 

North .r\nier. Flora. 34(4):296. 1927. Mexico. 

Siticluiriu rnoor~z (H. Robinson and Brettell) H. Robinson and 

Liuhum nioorei H. Robinson and Brettell. Phytologia. 

Sincluiria pi//itii Rydberg. North Anier. Flora. 34(4) :300. 

.Sinc/uiim /i/a/ylepi,\ (Schultz-Bipontinus ex Klatt) Rydberg, 

Liuhuni plo!l /epic Schultz-Bipontinus ex Klatt. Leopoldina. 

Sinciairia pojyanthu (Klatt) Rydberg. North Xnier. Flora. 

:34(4):299. 1927. Guatemala, Belize. Costa Rica. Panama. Colombia. 

Liaburnpolyati//ium Klatt. Bull. Soc. Roy. Bor. Belg., 31:209, 

1892. 

Liahum /onduzii B.L. Robinson, Proc. Bost. Soc. Nat. Hist., 

31:270. 1904. 

Sinciairza londuzzi (B.L. Robinson) Rydberg, Norrh Amer. 

Flora. 34(4):?98. 1927. 

Sinclairiu priti,<lpi (B.L. Robinson and Greenman) H. 

Rohinson and Brettell. Phytologia, 28(1):61, 1974. MexLiaburn pritiglri B.L. Robinson and Greenman. Proc. Amer. 

.i4r,qaliaburn pringlei (B.L. Robinson and Greenman) Ryd- 

.Sinclairia ,srrirolr/iU (Hemsley) Rydberg. North Amer. Flora, 

23: 146, 1887. 

1co. 

Acad., 32:49, 1896. 

berg-. North Anier. Flora, 34(43:294, 1927. 

3 &(4):30 1, 1927. Mexico. 

Liabum seriroiepis Hemsley. Biol. Centr. Amer., Bot. 2: 

188 1. 

S'iricluiria rzmi1i.s (McL'augh) H. Rohinson and Brettell. Phytologia. 28(1):62. 1974. Mexico. 

Liabum simiit Mc\.augh. Contr. L'niv. Mich. Herb., 9468, 

1972. 

Sinchiria sublvbata (B.L. Robinson) Rydberg. North Amer. 

Flora, 34(4):297. 1927. Guatemala. El Salvador. Honduras, Nicaragua. 

Liabiirn sublobalunz B.L. Robinson, Proc. .-\nier. .Acad., 

~51:539. 1916. 

Sinchiria tajurnulcensis (Standley and Steyermark) H. 

Robinson and Brettell. Phytologia. 28( 1):62. 1974. Guatemala. 

Liubum lajumulcense Standley and Steyermark. Publ. Field 

Mus. Nat. Hist.. Chicago. hot. ser., 'L3:27. 1943. 

Siizclairiu raguns (Blake) H. Robinson and Brettell. Phytologia, ?8( 1):62. 1974. Guatemala. 

Liaburn vagaris Blake, Brittonia. 2:354. 1937. 

Most of the species of Sznclairia can be identified 

using the key by Rydberg (1927), and the Guatemalan species have recently been treated by 

Nash (1976). Greenman (1897) provided an older 

key of some interest to the Mexican species of 

Liabum that included 10 species of Sinchiria. 

PARANEPHELIINAE, newsubtribe 

Paranepheliinae, subtribus nov. Plantae rosuliformes vel breviter caulescentes perennes; folia 

pinnatinervata vel leniter trinervata. Capitula 

sessilia vel in pedunculo elongato saepe solitaria; 

corollae disci in tubis valde elongatae; antherae 

pallidae; thecae base non denticulatae, cellulis 

endothecialibus in partibus scleroideis non contiguis ovalis vel taeniatis in parietibus transversalibus pauce noduliferis; rami stylorum elongati 

interdum spiraliter contorti; raphides achaeniorum elongatae. Grana pollinis regulariter spinulifera, spinis intus fere vel omnino caveata. 

Type genus: Paranephelius Poeppig and Endlicher. 

The subtribe is restricted to the central Andes 

in Peru, Bolivia, and northernmost Argentina. 

The subtribe contains the following two genera. 
NUMBER 54 45 

Paranephelius 

FIGURE 11 

Paranephelius Poeppig and Endlicher, Nov. Gen. et Sp., 3:42, 

pl. 248, 1843. [Type-species: Paranephelius unifZ0rus Poeppig 

and Endlicher; originally monotypic.] 

Liabum sect. Paranephelius (Poeppig and Endlicher) Bentham and Hooker f., Gen. PI., 2(1):436, 1873. 

Perennial acaulescent or shortly caulescent 

herbs, single or in clusters, with distinct tap-root 

and often with prominent fusiform lateral roots; 

sometimes with milky sap. Leaves in rosette or 

rather congested, petiolate or sessile with petioliform bases, usually narrowly winged, not 

sheathing; blade ovate to oblanceolate, crenate to 

pinnately lobed, trinervate to pinnately veined, 

smooth to prominently bullate, glabrous to pilose 

above, whitish tomentose below. Inflorescence 

with 1 or a few clustered sessile heads. Heads 

broadly campanulate; involucral bracts 40-50 in 

about 4 series, slightly to strongly unequal, oblong 

to lanceolate, outer bracts digitately lobed to 

blunt or acute, inner bracts acute to acuminate, 

outer surface white-tomentose to pilose; receptacle with high ridges enclosing bases of achenes, 

without chaff. Flowers yellow. Ray flowers 

-20-35, in 1 series, corolla with tube hirsute 

above, limbs linear with sparse to dense tomentum on abaxial surface, style branches spiralled. 

Disk flowers -20-35, corollas mostly 15-23 mm 

long, narrowly funnelform, not abruptly expanded at base of throat, tube very long, upper 

tube and most of throat puberulous; lobes with 

stomates along distal margin, glabrous or with a 

few stipitate glands near tip; walls of cells of 

anther collars not strongly annulated; anther thecae pale, not digitate at base, median endothecial 

cells elongate, oblong with strap-shaped to oval 

sclerified shields, attenuate and often uncinate at 

upper and lower ends; appendages smooth, cells 

mostly short-oblong with mostly truncate end 

walls; nectary short, cylindrical, unlobed; style 

base scarcely noduliferous, not or scarcely narrowed above into shaft of style; style branches 

slender, elongate, up to 5 mm long, abaxial hairs 

not or scarcely extending below bases of branches 

onto shaft of style. Achenes when mature 4-5 mm 

long, prismatic with -10 ribs, glabrous or with 

sparse setae or tomentum, cells of achene walls 

containing elongate raphids; carpopodium a narrow rim of -3-5 rows of rather small, subquadrate cells, cell walls slightly thickened, porose; 

pappus with 45-80 capillary setae, 1-2 cm long 

in 2-3 series, with a few shorter, often finertipped, outer setae, without distinct outer series, 

teeth of setae simple. Pollen grains 35-45 pm in 

diameter, spines mostly evenly dispersed, without 

columellae or with 1-3 slender strands. 

The sessile rosette of Paranephelius is a habit 

shared by a number of other genera of Asteraceae 

in the Andes, most notably Wernerza and a few 

species of Senecio such as S. macrorrhizus Weddell 

and S. wernerioides Weddell in the Senecioneae 

and a number of species of Hypochaeris in the 

Lactuceae. The members of the Senecioneae can 

be distinguished from Paranephelius most easily by 

the single series of equal involucral bracts, which 

are often partially fused in Werneria. Closer examination will also show truncate style branches 

in the disk flowers, short-lobed, glabrous, disk 

corollas, and narrowed appendages on the anthers. Hypochaeris is readily distinguished by the 

ligulate corollas and by the plumose setae of the 

pappus. 

The fusiform lateral roots seen on many species 

of Paranephelius are unique in the tribe but are 

approached in form by roots of many genera 

specialized for similar habitats. Such roots have 

not been seen in some species of the genus, such 

as P. ferreyrii, and are comparatively sparse or 

lacking in P. ouatus, but the apparent differences 

may be more the result of faulty collecting of 

underground parts of the plants. 

Paranephelius is the only genus in the tribe with 

tomentum on the outer surfaces of the ray corollas. Such tomentum was lacking in P. ovatus according to Weddell (1855-1857). The latter species does show only slight tomentum on the rays 

of most specimens, but tomentum is apparently 

present in all the species of the genus. On younger 

rays the tomentum is usually dense, but it may 
SMITHSOYIAh COhTRIBCTIOhS TO BOTASY 

FICI:RE 1 l.--i’~~rirctnrl/hei2u tinyfirus Poeppig & Endlicher: A. habit, X I/?: B. head. X 3%: c. ray 

corolla. X 10: D. disk corolla. X 10; E, disk sryle. X 10: F. bases of anther thecae. X 50: C, 

achenc, X 2’12: H. achene. X .i. 
NUMBER 54 47 

not be obvious on the fully expanded mature 

corollas. 

The achenes of Paranephelius are larger than 

those of most other members of the tribe and are 

present in mature form in a number ofcollections. 

The latter feature is not seen in specimens of most 

other Liabeae. Because of the habitat of the genus 

and the size of the achenes, it is suspected that 

achenes of Paranephelius have a longer viability 

than those of other members of the tribe. 

The following seven species are recognized 

from Peru, Bolivia, and Argentina. 

Piir(ii/r/ihe/iu,\ qtw~/i~/m (Muschler) H. Robinson and Brettell. 

Phytoloqia. 28( lj:59. 1974. Bolivia. Argentina. 

Licthrrni a.\;t,rrtjii/2urn Muschler. Engl. Bot. Jahrh. 50. Beibl.. 

3:78, 191.1. 

/'arai~e/i/re/rii.i /iu//a/u.\ .A. Gray ex Weddell. Chlor. And.. 1 :2 14, 

185.5. Peru. 

I'artriir/ilw//u.s Jerrrprii H. Robinson, Phytologia. 35(3) :235. 

1977. Peru. 

/'ur(i/itphr/iii.\ jelskii (Hieronynius) H. Robinson and Brettell. 

I'hytoloSia. 28( 1):59. 1974. Peru. 

/~r/r~~~/r/r,~r/.\kii Hieronymus. Engl. Bot. Jahrb.. 36:499. 1905. 

/'crrnric;t,/ip/iic oralus U'eddell. Chlor. And., 1:214. pl. 37B. 

/,rcihron or~a/im (LVeddell) J. Ball. Journ. Linn. SOC.. 22:46, 

/'ureriiyi/re/iii\ untj7oru.r Poeppig and Endlicher. No\-. Gen. et 

Li(rhrrrn rirr!jlorum (Poeppig and Endlicher) Schultz-Bipon- 

/'arcrnr/i/rp/iu\ ir~itrd~rki~ H. Robinson. Phytologia. 35(3):236, 

18,55. Peru. Bolivia. 

1883. 

Sp.. .1:42. pl. 248, 1843. Peru. Bolivia. 

tinus. Flora, 36:34, 1853. 

1977. Peru. 

Distinctions of many of the species are discussed by Robinson (1977b), but no key has been 

prepared. 

Pseudonoseris 

FIC~JRL 12 

/?\pudorio.ctri.i H. Robinson and Brettell, Phytologia. 28( 1) :59% 

1974. [Type-species: Liahiirn ~/rzaturn Ferre)~a.] 

Perennial acaulescent or short-stemmed herbs 

usually in clusters, with distinct tap root, with 

milky sap. Leaves opposite, sometimes in rosettes, 

sessile, oblanceolate, base slightly auriculate, margin crenate to deeply lobed, pinnately veined, 

upper surface rugose to nearly smooth, whitetomentose below. Inflorescence scapose or subscapose, with sparse alternate branches above. Heads 

broadly campanulate; involucral bracts -40 in 

-4 series, unequal, lanceolate, outer bracts narrowly acute, inner bracts long-attenuate. pilose to 

stipitate-glandular on outer surface; receptacle 

weakly alveolate, subglabrous. Flowers yellow or 

reddish. Ray flowers 15-20 in 1-2 series, corolla 

with tube puberulous, limbs linear. usually yellow 

(reddish in P. srysryloieicrii) : style branches elongate, not spiralled. Disk flowers 25-55, narrowly 

funnelform, not abruptly expanded at base of 

throat, tube long, upper tube and lower throat 

hirsute: lobes with stomates along margins, glabrous or with single glands distally on outer 

surface, walls of cells of anther collars not strongly 

annulated; anther thecae pale, not digitate at 

base, shields of median endothecial cells oval with 

nodular thickenings at upper and lower ends: 

appendages shortly ovate to oblong, 1- 1 '/2 times 

as long as wide, smooth, cells rather elongate with 

truncate or oblique end walls: nectary short, not 

lobed: style base with a large node tapering into 

the broad shaft; style branches slender. -3-4 nim 

long, filiform, tapering distally, hispidulous surface not or scarcely extending onto upper part of 

shaft. Achenes prismatic, obovate, slightly narrowed below, with -10 strong ribs, sparsely setuliferous, cells of achene wall containing elongate 

raphids: carpopodium short-cylindrical, with 

rather large cells in 5-10 series, walls of cells 

noduliferous; pappus with 25-:30 somewhat persistent inner setae in a single series, with outer 

series of short, narrow. squamiform setae, teeth of 

setae simple. Pollen grains 35-45 pni in diameter. 

spines evenly dispersed, without columellae or 

with minute tenuous columellae under spines. 

The usually subrosulate habit, the long tubes 

of the disk corollas, the long slender branches of 

the disk styles, and the elongate raphids of the 

achene wall are all consistent with the assumed 

relationship of Pseudonoseris with Paranephelius. 

The occurrence at generally lower elevations, the 

presence of latex. the erect branching inflorescences, the simple low-alveolate surface of the 
48 SMITHSONIAX CORTRIBLTIONS TO BOTANY 

E'IGIIRE l".-l'.reud~r2osr~~~ disctiitir (Muschler) Robinson B Breltell: A. habit. X I%: B. head. X 3]%: 

c. ray corolla, X 8: D, disk corolla. X 8: E. disk style, X 8; F, bases of anther thecae, X 35: G. 

median endothecial cells. X 400: H. achene. X 8. 
NI'MBER 54 49 

receptacle, and the more differentiated outer series of the pappus bvould all indicate a less specialized condition of Pseudotioseris in the subtribe. 

Nevertheless, similarities between all three of the 

widely separated species of Pseudonoseris indicate 

that they are more closely related to one another 

than any are to Paranephelius and that they are 

not simply relicts of an older stock of the subtribe. 

In Pseudonoseris. the leaves all have blades continuing to the base and forming auricles at the 

node, and the venation is strictly pinnate. The 

inflorescence always has a lax alternate branching 

ivith long-stalked glands on the pedicels and in- 

\,olucre, and the involucre is multiseriate with 

narro\v, often attenuate bracts. The two northern 

species have lobed leaves and achenes with heavily corticated ribs, in contrast to the more southern P. discolor. One species, P. szyszy lowicrii, has 

reddish or reddish-orange flowers. 

,411 three species of Pseudono.reris are found in 

Peru, but they are from widely separated localities; P, discolor is from Pun0 near the Bolivian 

border, P, sqszylou~iczii is from the eastern slope 

of the Andes in the Dept. of Amazonas, and P. 

strictn is from the coastal range in Lambayeque. 

The three species are as follows. 

/',\mdoi/o\?~/5 dr\cir/iir (Muschler) H. Robinson and Brettell. 

Phytologia. 28( l):60, 1974. Peru. 

Oti(l.\?ir\ dixudor Xluschler. Engl. Bot. Jahrh. 50. Beibl.. 

:$:!I I, 19 13 111011 Licihrtm dr\co/or Huniholdt. Bonpland. 

aiid Kilnth]. 

Lrrthrori /crrrcilrtm Ferre!.ra. Bol. soc. Peru. Bot.. 1: 17. 1948. 

P\tudhrwri.\ .s/riu/u (Cuatrecasas) H. Robinson and Brettell. 

Licihrirtr ilriiiiron C:i~atrecasas. Collect. Bot.. Barcinone. 

/',\mdorio \t>r it \;y rqioii ,iczr i (Hieron)mus) €3. Ro hinson and 

Lrcchrtrrr cj~~~j~/oi~~~c~ii Hieronynius. Engl. Bot. Jahrb.. 

Phytologia. 28( l):GO. 1974. Peru. 

.3:.iO6. l95:3. 

Breitell. Ph)toloqia. 28( 1):GO. 197-1.. Peru 

36 : 50 3. 1 903. 

MUNNOZIINAE, new subtribe 

Munnoziinae, subtribus nov. Plantae herbaceae perennes vel raro annuae interdum subscandentes vel procumbentes; folia saepe in marginis 

angulatae plus minusve trinervatae vel palmatinervatae. Capitula solitaria aut in paniculo disposita: corollae disci in faucibus saepe abrupte 

expansae: antherae plerumque nigrescentes, thecis base non denticulatis, cellulis endothecialibus 

in partibus scleroideis plerumque contiguis; rami 

stylorum breves: raphides achaeniorum quadratae. Grana pollinis regulariter vel irregulariter 

spinosa, spinis intus columellatis vel variabiliter 

caveatis. 

Type genus: 1Wunnozia Ruiz and Pavon. 

The subtribe can be divided into two groups, 

each containing two genera. Cfirysactinium and 

Munnozia have tomentum on their stems and 

leaves as in most Liabeae but have pollen grains 

with regularly dispersed spines. Erato and Philoglossa have large stiff hairs on their stems and 

leaves and have pollen with irregularly dispersed 

spines. The subtribe is mostly Youth American, 

but three species reach Costa Rica. The four 

genera are as follows. 

Chrysactinium 

Flf;liRE 1:I 

Chy.\ac/iri2urn (Huniboldr. Bonpland. and Kunth) \Veddell, 

Chlor. .And.. l:?I?. 18.56. 

'4 rdrortdi ia sect . Ch ryJ ur / 1 rr 1 urn H II IN bu 1 d I . B o 11 p 1 and , and 

Kunth. Nov. Gen. et Sp.. ed. folio. k:77. 1818. [Lectotype: 

:1 iidrorrinch ia acau/i.t Hum bol d t . Bon 11 I and. and Ku 11 t h . ] 

Small perennial herbs; without reports of milky 

sap. Leaves opposite, in basal rosettes, or crowded 

on short stems, sessile with cuneate or petioliform 

bases; blades obovate, trinervate from near basal 

fourth, entire to dentate, green above with or 

without pilosity or arachnoid pubescence, tonientose below. Inflorescence with long unbranched 

erect peduncles bearing solitary heads, peduncles 

covered with arachnoid pubescence, with scattered purplish hairs. Heads broadly campanulate. 

Involucral bracts 40-60 in -4 series. strongly 

unequal. narrowly ovate to lanceolate, tips 

sharply acute to acuminate, outer surface glabrous to glabrescent; receptacle with chaff prominent, narrowly lanceolate. Flowers yellow. Rays 

30-60, tubes of corollas puberulous, limbs narrowly elliptical to linear, glabrous, slightly trilobed at tips: style branches elongate, not spi- 
50 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

F~c:ri~t I~~.~~,’/~~~.\or/~~~u7~ orriulr (Ilurnholdr. Bonpland. and Kurith) Weddell: A, habir, X ‘2: I$, 

head. X ?I.: c. ray corolla. X 8: L). disk corolla. X 13; E. anther collar and I~aaea of thecae. X 4,j: 

F. disk stvle. X 13; c;, achene. X 13. 
NUMBER 54 5 1 

ralled. Disk flowers 50-100?: upper tube and 

lower throat of corolla hirsute on outer surface; 

throat rather abruptly expanded at base; lobes 

with stomates near margin, with only subsessile. 

short-stalked glands on outer surface: cells of 

anther collar densely annulated, cross walls obscure: thecae all or partly black, not digitate at 

base: median endothecial cells oblong with simple 

broad oval or strap-shaped sclerified band, with 

single thickening on transverse wall; appendages 

smooth, cells rather elongate with truncate or 

oblique end bvalls: nectar) not or slightly lobed; 

style base with distinct node: branches short, less 

than 1/2 as long as the hirtellous upper part of 

the shaft, with rounded tips. Achenes prismatic, 

puberulous. cells of achene wall containing small 

quadrate raphids: carpopodium distinct, annuliform. with subquadrate to suboblong cells in 3-5 

series. cell walls slightly thickened: pappus white, 

with 30-60 hispid slender-tipped capillary setae. 

fragile at base. teeth of setae simple, without 

shorter setae in outer series. Pollen grains 30-40 

pni in diameter, with spines regularly dispersed, 

with distinct columellae under spines. 

The nanie "Chr1,sactznium" is unfortunately close 

to that of another asteraceous genus, Chyxzctinza 

A. Gray of the Heliantheae (Pectidinae). The 

Iirtmzationa/ Code ?/'Botanical ,Vomenclature does not 

prohibit such names when the endings differ in 

gender. but such similar names would not have 

been chosen intentionally. especially for members 

of the same family. 

The genus Chr),sactiniurn is closely related to 

h'iintioriu. from which it is superficially most distinct in the reduced acaulescent habit with longscapose heads. .-I more complete study shows that 

the habit does not accurately distinguish between 

the genera. some species of Ch~~~xactiniurn having 

more elonqate \.egetative stems similar to those in 

sonic of the smaller species of iWunnoria such as 

.W. jkrrtqrii, ,LI. (Kastnera) niiiea, and .W. ,be foliata. 

The appearance of intergradation in habits. itself, 

is misleading. however, since the genera are completely distinct in a number of details that seem 

to indicate a significant phyletic gap. In Chrq'sac- 

~inzwri. the cells of the anther collar always have 

dense annular thickenings, whereas none are seen 

in .Wunnoria, and the corolla lobes of Chcpsactiniurn 

have only small subsessile glands, never the longer 

hairs or glands usually present near the lobe tips 

in .bfunnozia. It is convenient that the pappus of 

Chysactinium is pure white, byhereas that in all 

species seen in Munnozia is darkened or discolored 

in some way, thus providing the most easily observed reliable distinction between the genera. 

In Chvsactinium, as in :Llunnotia, the stout hairs 

or pilosity on the upper surface of the leaves has 

been used as a species character, and here. as in 

Munnotia, the character has been found unreliable. An apparently reliable species character is 

the greater length of the projections on the receptacle in C. hieracioides reported by Robinson 

(1978a), but the feature is of limited use. sirice it 

is usually hidden within the flowering heads, and 

specimens rarely have more than one or two 

heads. Hieronymus (1895) described the achenes 

of C. longiradiatziin as glabrous, but this is presumed to be an error based on immature material. 

No glabrous achenes have been seen in the genus. 

The following six species are knmvn. ranging 

from central Ecuador south to central Peru. 

C%r J \uc/iriiitrri acciiiii~ [€ Ium bolclt . Boil pl anti. anel Kii iit ti ) Li'etl- 

.4ndrorriuchiu uruuiis Huiiiboldr. Koiiplaricl. ancl Kiiiith. 

Liuhirrri uruitic (Humholdt. Bonplantl. iiiitl Kiiiir ti) Ix\\iiiq. 

Liuhitri or//iro//irit Blake. Joiirn. LVasti. .\cacl. Sci.. 17:21111. 

L%risnr/irriurit cil-/hrc~//iri.x (Blake) €1. liohinson and Krc~rell. 

k I, li o ti i nc 111 a I i cl 13 rc t t c I I . 

dell. Chlor. .hd.. l:212. 1857. LciratioI. 

So\.. Gen. et Sp.. ed. folio. +:i7. 1818. 

Linnaea. 6:696. 183 1 

1927. 

Phpologia, 28(1 j:49. 197 &. 

(.% (1 \ uc/i,i i iirri mnp/io//i ri x ( B I a ke 

Phytoloyia. 28fH:M. 197-k. Peru. 

Liaburii cim/iho//zri.t Blake. Joiirn. \\ash. ,icatl. Sci.. 1i:YO. 

C'h rJ,\cir/irii itm cait/r.\cm ( Hi eron )- 111 11s ) 1 I . Ii ob i n wii ii I i ti K rct - 

Liubrcrrz tnit/r.\crri.\ Hieron! iiius. Eiiql. €hit. ~Jahrli.. ,36:,NO. 

Liaburri hicoior Blake. Journ LVash. :\cad. Sci.. I7:ZK). 

C/iqsar/iniiirn hiroior (Blake) €1. Kohin\oii and Hretrrll. 

(;/I!) \ur/iriiitrn /i/rrucioirir\ (Iluiiiholcit, 13ciiipI~iiitl. iiiid Kiiiir ti) 

1927. 

tell. Phytologia. 28( l):jO, 197-k. Lciiatioi-. Pcrii. 

1905. 

192i. 

Phytoloqia. 18( l):49. 1971. 
52 SMITHSOXIAN CONTRIBUTIONS TO BOTANY 

H. Robinson and Brrttell. Phytologia, 28( 1):50, 1974. 

Ecuador. Peru. 

ilndromachia liieraciuzdts Huniboldt. Bonpland. and Kunth. 

Nov. Gen. et Sp.. ed. folio. 4:77. 1818. 

Liubuin hieru~~oidt~ (Huniboldt. Bonpland. and Kunth) 

Lessing, Linnaea. 6:699, 1831. 

C%2~yac/iniutn longiradiaium (Hieronymus) H. Robinson and 

Brertell, Ph>-tologia. 28(11:30. 1971. Ecuador. 

Liabuni longirudiu/urn Hieronynius. Engl. Bor. Jahrb.. 

21:3.52? 1895. 

Liuhuvi /unuiuc Blake lab tenuior], Journ. \Vasti. Acad. Sci., 

17:289, 1927. 

Chrysaclinium lenuius (Blake) H. Robinson and Brettell, 

Phytologia. 28(1):30, 1974. 

Ch~y.~u~/i~7iu7n rosu/a/urn ( H ieron y ni Lib) H . Robinson and Brer - 

tell. Phytologia. 28(1):50. 1971. Peru. 

Liuburn rosulatuin Hieronynius. Engl. Bot. Jahrb., 36:50 1, 

1903. 

The species occurring in Ecuador have been 

treated by Robinson (1978a). 

Erato 

Ercr/o DeCandolle, Prodr., 5:3 18. 1836. IT>.pe-species: Erato 

.tfuunnoaa subg. Erato (DeCandollcj H. Robinson and Brettell. 

pojyn~nioides DeCandolle: originally monotypic.] 

Phytologia. 28( 1):.56, 1974. 

Coarse perennial herbs or subshrubs. Stems, 

stipule-like sheaths, both surfaces of leaves, and 

peduncles pilose or strigose with stiff bulbousbased hairs. Stems with milky sap, lengths of 

internodes variable without pattern. Leaves opposite; stipule-like bases broadly oblong, usually 

deeply emarginate: petioles unhinged: blades 

somewhat succulent with 5-9 radiating veins 

from base, margins of larger leaves coarsely 

toothed or serrate, upper surface green, lower 

surface pale green. without tomentum. Inflorescence cymose to subumbellate: peduncles elongate. 

Heads broadly campanulate. Involucral bracts 

40- 100 in -4 series, unequal. broadly ovate to 

lanceolate, outer bracts fleshy at base, with obtuse 

herbaceous tips often tomentose at tip, margins 

densely ciliate with stiff hairs: receptacle foveolate 

with unlobed or slightly lobed puberulous ridges. 

FlOWers yellow. Ray flowers 75-230, in 2-3 series, 

tubes usually puberulous above, limbs linear and 

glabrous, deeply notched at tip, often 3-lobed: 

style branches elongate, not spiralled. Disk flou.- 

ers 20-150; tube and lower throat of corolla 

hirsute on outer surface, throat abruptly expanded at base, lobes with large stomates near 

margin, strongly spiculiferous at tip; cells of anther collars not or weakly annulated on walls: 

thecae black, not digitate at base; median endothecial cells oblong with sparse minute thickenings mostly on vertical walls; appendages oblong (sometimes tapering in E. pob:mnioides), 

smooth, with basal cells rather elongate, apical 

cells shorter to subquadrate; nectary elongate, 

slightly lobed: style base with distinct node: style 

branches short, usually less than l/2 as long as 

the hirtellous portion of the upper shaft. with 

obtuse to rounded apices. Achenes mostly 4-sided. 

glabrous or hispidulous, cells of achene wall with 

small quadrate raphids: carpopodium distinct 

with large subquadrate cells in -5 series, cell 

walls thin: pappus setae 25-50, capillary or short 

and awn-like, sordid white to broLvnish-red, 

strongly spreading and persistent or easily deciduous, scabrid with simple teeth. Pollen grains 30- 

40 pm in diameter, Lvith spines somewhat unevenly dispersed, with distinct stout internal columellae grouped under spines. 

The genus Erato was originally placed in a 

position remote from other Liabeae by DeCandolle (1836) and close to elements now included in Baccharis of the Astereae. The genus 

was first placed in Liabum by Bentham in Bentham and Hooker (1873), along with all other 

elements now placed in the Liabeae that have a 

basically capillary pappus. Erato was placed in a 

broad concept of Munnotia in the generic revision 

of the tribe Liabeae by Robinson and Brettell 

(1974b) but was restored to separate generic status during a study of the members of the tribe in 

Ecuador (Robinson, 1976d, 1978a). The position 

of the genus is obviously closest to Philoglossa, 

with which it shares the same type of stiff hairs 

on the stems and leaves. The achenes of both 

genera also share a reduced number of ribs or 

angles, four in Erato and two in Philoglossa. 

Erato almost totally lacks the arachnoid pubes- 
FI~:~.RE 14.--Erulo pojyninioide~ Decandolle: ,A. habit. X I,$: B, foliar hairs. X 50: c. head. pat 

anrhesis. X 3?$; u, ray corolla. X 10: t. disk corolla, X 10: F. disk style, X 10: c;, achene, X 10. 

5 :3 
5 4 SMITHSONIAN CONTRIBUTIOXS TO BOTANY 

cence that is characteristic of most of the tribe, 

but all the species except Erato pobmnioides retain 

slight tomentosity on the tips of the involucral 

bracts. The species E. sodiroz is singularly distinct 

in the short, multiseriate, deciduous pappus awns. 

The genus ranges rather widely, with Erato 

mlcanica reaching Costa Rica in the north and E. 

po/ymniozdes reaching Bolivia in the south. In Peru, 

E. pobmnioides occurs widely in the inner mountain ranges, but the endemic E. stenolepis is restricted to the coastal area near Lima. The center 

of diversity is in Ecuador, where three of the four 

species occur. 

The availability of greenhouse plants of Erato 

pohmnioides has allowed observation of the latex 

production over a period of time. The amount of 

latex \,aria, especially with the amount of water 

furnished to the plant. On a few occasions the 

plants might have been judged as lacking latex. 

The observed variation in Erato is cause for caution in judging absence of latex in other genera. 

especially the closely related Philoglossa. 

The four species of the genus are as follows. 

Eru/o po/>.tnniotde.s DeCandolle. Prodr., 5:3 18. 1836. Ecuador. 

Perl1. Boli\ia. 

LictOrctn pa//a/anprrrte Hieronymus, Eql. Bot. Jahrb.. 29:iiO. 

1 !)OO. 

.\friuri~iu ,/io/prnitrorde.\ (LleCandolle) H. Robimon and Brettell. Phytologia. 28(1):56. 1974. 

/<?a/() todiror (Hieronynius) €3. Robinson. Phytolog-ia. 34 

(k):.<79. 1976. Ecuador. 

Erci/o \/triolf/i/\ (Blake) ti. Robiiison. Phytologia. 34(-k:,?Ii9. 

19i6 Peru. 

/,ro/irit/i .s//jrio/t/iis Blake. Joiirn. \\:ash. Acad Sci.. 17::IO'L. 

l5127, 

.1fiiniioziu .\/tno/rpi.s (Blake) H. Robinson and Brettell, Phyroloqia. 28( l):57. 1974. 

Eici/o mitotircx (Klatt) €3. Robinson, Phytologia. 34(4) :379. 

I<)T(j, Costa Rica. Colombia. \'enezuela. Ecuador. 

Licihruti irclc-anirron Klatt. EngI. Bot. Jahrh.. 8:47, 1887. 

/,citburrt cinri/rnci Benoist, B~ill. Soc. Bot. France. 84:633. 

/,icihion ~ii.\r,gnc Badillo. Bol. Soc. \.enez. Cienc. Nat.. 

.\luniiozr~i anci/irrn (Benoist) €1. Robinson and Bretrell. Ph!.- 

.L'Irirttio:~~ i~itlcuntcu (Klart) €1. Robinson and Brettell, Phy19:VI. 

10:.31:3. 1946. 

toloqia. 28( 1):56. 1974. 

toloqia. 28(1):.57. 1974. 

A key and descriptions have been provided by 

Robinson (1978a) for the three species that occur 

in Ecuador. 

Munnozia 

FIGI'RE 15 

.Vfunno;ia Ruiz and Pavon. PI. Peruv. Chil. Prodr.. 108. 1794. 

[ Lectot ype: ,bluiiiiozza lariceolata Ruiz and Pavon.] 

illtbum Lessing, Syn. Comp.. 132. 1832. [Type-species: ;I/zhum 

1iaboide.r Leasing.] 

/'rionole/ii.i Poeppig and Endlicher. Nov. Gen. et Sp.. 3:55. pl. 

261. 1845. [Type-species: /'riono/epi.s .si/,bhiciides Poeppiq and 

Endlicher.] 

Ltaburn sub,q. Ch~y.\ai/rur~i M'illdenow ex Schultz-Bipontinus. 

Flora. 36:37. 1833. Ilectotype: Lrabum hagit/a/utn SchultLBipontinus (= .Wunnozia ,\entcton/dzs Bentham).] 

k'as/riera Schultz-Bipontinus. Flora. 36:37. 18 

cies : Ka.i liierri ten em Sc hu It L- B i pon t i nus. ] 

O'hrysaJ/rum \Villdenow ex LVeddell. Chlor. ;\nd.. 1:21 I. 1837 

[in nota: iioni. nud. in s!-n.]. [T!,pe-species: C:h~jJcc~truni 

su,gi/ta/iini 1villdenow ex \Veddell.j 

Munnozia subg. kh~inrra (Schultz-Bipontintis) H. Robinson 

and Brettell. Phytologia. 'L8( 1):37. 19i4. 

.4nnual or perennial herbs or subshrubs, low or 

creeping to subscandent. Stems with distinct internodes, with milky sap. Leaves opposite. Petiole 

bases often auriculate, sometimes forming nodal 

disk, not sheathing; petioles distinct, sometimes 

winged; blade with base rounded to hastate, trinervate or pinnately veined, margin serrulate to 

deeply lobed or pinnatisect, upper surface tomentose to pilose or glabrous, lower surface tomentose 

or rarely nearly glabrous. Inflorescence terminal. 

sparingly to densely corymbose to subcyniose, 

often with ultimate dichasia: peduncles short to 

very long, tonientose to stipitately glanduliferous. 

Heads broadly campanulate; involucral bracts 

17-70 in 2-4 series. subequal to unequal. outer 

bracts ovate to oblong with acute herbaceous tips, 

margins not densely pubescent, glabrescent to 

tomentose or stipitately glanduliferous on outer 

surface: receptacle with prominent chaff of lacerate squamae (Lvithout chaff in subgenus Kastnera). Flowers yellow (in 2 species w-ith ray and 

sometimes disk corollas Tvhitish, becoming lavender). Rays 6-70; corolla with upper tube often 

hirsute, limbs linear, usually with some hairs or 

glands. with short lobes at tip; style branches not 
NUMBER 54 
36 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

spiralled. Disk flowers 9-83: upper tubes and 

bases of throats often densely pubescent. throat 

rather abruptly to bulbously expanded at base: 

lobes with large stomates along lateral veins, 

sometimes bearing stout hairs at tip: walls of cells 

of anther collars not strongly annulated: thecae 

usually black, pale in 3 species, not digitate at 

base: median endothecial cells with polar thickenings on transverse walls (radial thickenings in 

subgenus Kastnera) ; appendages smooth, cells 

rather elongate with truncate or oblique end 

walls: nectary short, not or slightly lobed: mature 

style base with distinct node; style branches of 

disk flowers short, usually less than 1/2 as long as 

the hirtellous portion of the upper shaft, with 

obtuse to rounded tips. Achenes prismatic with 

6-10 ribs, setuliferous, cells of achene wall containing small quadrate raphids: carpopodium annuliform with 3-12 rows of small to mediumsized rather thick-walled cells; pappus with 3-53 

sordid or reddish capillary inner setae, scabrid 

with simple teeth, with fe\v to many outer setae 

or with a series of distinct outer squamae. Pollen 

grains 27-40 pm in diameter, with spines regularly dispersed, with internal columellae under 

spines variously fused or separated from foot 

layer. 

The identity of the genus iVunnozia apparently 

has never been in doubt, even when the genus 

was included in the synonymy of Liabum: however, the identity of the four species described by 

Ruiz and Pavon from Peru has been in considerable doubt. The types, which still exist in Madrid, were comparatively unavailable for many 

years. The problems have been resolved recently 

because of the visit of Dr. Jose Cuatrecasas to 

Madrid, where he photographed and studied the 

types. All four species now have been matched 

with more recent collections, and it is notable 

that none of the four represent any of the most 

common species of the genus. 

The species of Ruiz and Pavon most readily 

identified was iM. corymbosa, which is represented 

by a duplicate at Paris and a small fragment 

obtained by S.F. Blake in the U.S. National 

Herbarium. The species is closely related to the 

common .M. senecionidis, but it has more densely 

lanate stem and leaf surfaces, cordate rather than 

hastate bases on the leaf blades, and disk corolla 

lobes lacking apical hairs. It appears to be the 

same as Liabum eriocalyx that Blake described from 

the same area of Peru on the basis of a MacBride 

collection. 

Munnotia venosissima Ruiz and Pavon is also in 

the relationship of Ad. senecionidis, but it has abundant purple hairs and petioles toothed distally, 

and it sometimes has prominent stipuliform lobes. 

The species seems identical to Liaburn pulchrzirn 

Blake, based on a MacBride collection from the 

same region in Peru. 

Munnozia trinervis Ruiz and Pavon is also a 

species with triangular leaf blades but with the 

blade scarcely longer than wide and with the 

petiole short and narrowly winged. More recent 

collections include Vargas 4/09 from Cuzco (F), 

and Killip and Smith 22247 from Ayacucho (US). 

Munnozia lanceolata Ruiz and Pavon, with its 

solitary heads, has very short petioles and firm, 

narrowly elliptical blades. A specimen collected 

by Hutchison and Wright 5928 from Huanuco, Peru 

(US) has been determined as this species by Dr. 

R. Ferreyra and appears to be the species, although it lacks the minute basal auricles seen on 

the leaf blades of the type. 

Ruiz and Pavon (1794) described the genus 

Munnozia with mention of only two species, which 

were not named. The same authors later (1 798) 

named four species. As noted by Cabrera (1960) 

in his study of the types, only two of the four 

types were annotated by Ruiz and Pavon as 

representing a new genus. Of these two, Robinson 

and Brettell (1974b) selected iM. lanceolata as the 

lectotype of the genus. 

Within the typical subgenus there are a number of distinctive elements. The type-species is 

one of a group restricted to Peru and Bolivia, 

having coriaceous leaves and pinnate venation. 

Most members of the genus have herbaceous or 

even membranaceous trinervate leaves. Two species. M jussieui and .M. campii, have flowers that 

are usually whitish, sometimes turning lavender. 

Apparently, rare yellow-flowered forms of '1.I. jus- 
SL'MBER 54 5 7 

sieui are knowm. Three species, M. hastifilia, .%if. 

maronii. and iz/I. subciridis, characteristically have 

pale anther thecae, lacking the blackish color that 

is otherwise characteristic of the subtribe. 

The subgenus Kastnera is a completely natural 

group originally distinguished because of the lack 

of chaff or projections on the receptacle. Subsequent investigation shows differences from other 

.Wunnoziae in the consistent lack of connation betlveen the stipule-like lobes of paired leaves, in 

the strongly differentiated, often enlarged outer 

pappus elements. in the reduced number of inner 

pappus setae, and in the radial pattern of nodular 

thickenings on both transverse and vertical walls 

of the endothecial cells. In spite of these differences, the close relation and close siniilarity to 

other parts of Munnozia are evident, and separate 

generic status seems unnecessary and counterproductive. For others of a different opinion, it 

should be noted that the name "Alibum Lessing" 

would take precedence at the generic level. 

In the revision of the tribe by Robinson and 

Brettell (1974b). the type-species of Alibum, A. 

liaboides Lessing of Ecuador, was mistakenly regarded as the same as M. bratn of Peru. On 

further examination, the two species prove thoroughly distinct, with M. lyrata not being a member of the subgenus Kastnera. 

The subgenus Kastnera contains a number of 

species that are notable for their pinnately lobed 

or pinnatifid leaves. One of these. M. annua, has 

uniquely tnembraneous and scarcely pubescent 

leaves, and it and M. (Kastnera) niilea are two of 

the few annuals in the tribe. 

.I.lunnozia is remarkable for the many taxonomically significant structural variations in the interior of the pollen wall (Robinson and Marticorena, in prep.). The pollen characters indicate that 

the iW. senecionidis-;W. coymbosa group is more 

closely related to the subgenus Kastnera than it is 

to the typical element of the genus. Other distinctive pollen types are seen in ;%if. lyrata and M. 

jussieui. 

The genus ranges from Costa Rica and Panama into Andean South America from L'enezuela 

to Bolivia. 

Key to the Subgenera of Munnozia 

Receptacle with crests and projections: outer series of pappus with narrow 

segments; inner pappus bristles numerous: endothecial cells with nodular 

thickenings usually few or polarized: nodes of stem with stipuliform lobes 

often connecting between paired leaf bases subgenus Munnozia 

Receptacle without crests or projections; outer series of pappus often with 

rather broad segments: inner bristles 5-20, endothecial cells with radial 

thickenings: wings or lobes at bases of leaves never connecting across 

nodes 

The genus contains 40 known species, which 

are disposed in the subgenera as follows. 

.Mr~ntioau subgenus Munnozzu 

.Llunnci--iu </]inid (Blake) H. Robinson and Brettell. PhytoloL,inhiirn </fine Blake. Journ. IVash. Acad. Sci.. 17:301, 1927. 

.Miinnoziu crngus/a (Blake) H. Robinson and Brettell, PhytoLruDrriii angus/urn Blake. Journ. Wash. Acad. Sci.. 17:295. 

qia. 28:(1):54. 1974. Peru. 

loqia. ?8(l):X. 1974. Peru. 

1927. 

.. subgenus Kastnera 

.Munnoriu cumpii H. Robinson, Phytoloqia. 3,5( 1):138. 1976. 

.Munnorra canarensis (Cuatrecasas) H. Robinson and Brettell. 

Ecuador. 

Phytologia. 28( 1):54, 1971. Ecuador. 

Lrnbum cunarensp Cuatrecasas, Brittonia, 8:46. 195-1. 

,Munnorfa cardenasii (Cabrera) H. Robinson and Brettell. PhpLiahuin curdenu~ii Cabrera. Sot. hlua. La Plata. Bot.. 

.Mmmiu r/iirnboruc~n.si.s H. Robinson. Phytoloqia. 35(3): 109. 

Munnoau conuencionensis (Cuatrecasas) H. Robinson and Brettologia. 28( l):.%, 1974. Bolivia. 

11:19l, 1949. 

1977. Ecuador. 

tell. Phytologia. 28(1):51. 1974. Peru. 
SMITHSONIAN CONTRIBUTIOXS TO BOTANY 

Liabum convencionense Cuatrecasas, Collect. Bot., Barcinone, 

3:300, 1953. 

Munnoria corymbosa Ruiz and Pavon, Syst. Veg. Peruv. Chil., 

195, 1798. Peru. 

Liabum coymbosum (Ruiz and Pavon) Schultz-Bipontinus, 

Flora, 36:34, 1853. 

Liabum eriocalyx Blake, Journ. Wash. Acad. Sci., 17:297, 

1927. 

Munnozia eriocalyx (Blake) H. Robinson and Brettell, Phytologia, 28(1):54, 1974. 

Munnozia ferreyrii H. Robinson, Phytologia, 34(4):381, 1976. 

Peru. 

Munnozia foliosn Rusby, Bull. Torrey Bot. CI., 54:312, 1927. 

Peru, Bolivia. 

Munnotia chrysanthemoides Rusby, Bull. Torrey Bot. Cl., 

54:313, 1927. 

Liabum herrerae Cabrera, Rev. Univ. Cuzco, 33(87): 119, 

1945. 

Liabum foliosum (Rusby) Ferreyra, Bol. SOC. Peru. Bot., 

1: 18, 1948.-Cabrera, Not. Mus. La Plata. Bot., 14: 193, 

1949. 

Munnozia herrerae (Cabrera) H. Robinson and Brettell, 

Phytologia, 28(1):55, 1974. 

Munnoria fosbergii H. Robinson, Phytologia, 34(4):382, 1976. 

Colombia. 

Munnozia gigantea (Rusby) Rusby, Bull. Torrey Bot. CI., 

54:312, 1927. Peru, Bolivia. 

Liabum giganteum Rusby, Bull. N.Y. Bot. Gard., 4:391, 

1907. 

Munnozia glandulosa- (0. Kuntze) Rusby, Bull. Torrey Bot. 

CI., 54:314, 1927. Bolivia. 

Liabum glandulosum 0. Kuntze, Rev. Gen., 3(2): 163, 1898. 

Munnozia hastifolia (Poeppig and Endlicher) H. Robinson and 

Brettell, Phytologia, 28(1):55, 1974. Colombia, Venezuela, Ecuador, Peru, Bolivia, Argentina. 

Liabum hastifolium Poeppig and Endlicher, Nov. Gen. et 

Sp., 3:43, 1843. 

Heterotheca deltoidea Klatt, Ann. Naturh. Hofmus. Wien: 

9:358, 1894. 

Liabum laticifrum Badillo, Bol. SOC. Venez. Cienc. Nat., 

10:312, 1946. 

Munnotia hirta (0. Kuntze) Rusby, Bull. Torrey Bot. Cl., 

54:314, 1927. Bolivia. 

Liabum hirtum 0. Kuntze, Rev. Gen., 3(2): 163, 1898. 

Munnozia jussieui (Cassini) H. Robinson and Brettell, Phytologia, 28( 1):55, 1974. Colombia, Ecuador [original citation by Cassini from Peru evidently in error]. 

Andromachia jussieui Cassini, Bull. SOC. Philom., 1817: 184, 

1817. 

Conyza stipulata Vahl ex Cassini, loc. cir. [nom. nud. in 

syn.]. 

Liabum jussieui (Cassini) Cassini, Dict. Sci. Nat., 26:205, 

1823. 

Liabum nonoense Hieronymus, Engl. Bot. Jahrb., 29:59, 

Liabum nonoense var. microcephalum Hieronymus, Engl. Bot. 

Liabum tabanense Cuatrecasas, Caldasia, 3:425, 1945. 

Munnozia nonoensis (Hieronymus) H. Robinson and Brettell, 

Munnozia tabanensis (Cuatrecasas) H. Robinson and BretMunnozia karstenii H. Robinson, Phytologia, 35(6):490, 1977. 

Munnozia lanceolata Ruiz and Pavon, Syst. Veg. Peruv. Chil., 

Liabum lanceolatum (Ruiz and Pavon) Schultz-Bipontinus, 

Munnozia longifolia Rusby, Bull. Torrey Bot. CI., 54:313, 1927. 

Liabum hexagonum Blake, Journ. Wash. Acad. Sci., 17:300, 

Munnozia lyrata (A. Gray) H. Robinson and Brettell: PhytoLiabum lyratum A. Gray, Proc. Amer. Acad., 5:ll5, 1861. 

Munnozia maronii (Andre) H. Robinson, Phytologia, 35 

(3):200, 1977. Bolivia. 

Andromachia maroni Andre, Revue Horticole, 1887(2 1):496, 

1887. 

Liabum corymbosum Schultz-Bipontinus ex Klatt, Annal. 

Naturh. Hofmus. Wien, 9:363, 1894. [Not L. rovmbosum 

(Ruiz and Pavon) Schultz-Bipontinus.] 

Munnozia klattii H. Robinson and Brettell, Phytologia, 

28(1):55, 1974. 

Munnozza olearioides (Muschler) H. Robinson and Brettell. 

Phytologia, 28(1):55, 1974. Peru. 

Liabum olearioides Muschler, Engl. Bot. Jahrb. 50, Beibl., 

3:82, 1913. 

Munnozia oxyphylla (Cuatrecasas) H. Robinson and Brettell, 

Phytologia, 28(1):55: 1974. Peru. 

Liabum oxyphyllum Cuatrecasas, Collect. Bot., Barcinone, 

3:303, 1953. 

Munnozia pefoliata (Blake) H. Robinson and Brettell, Phytologia, 28(1):55, 1974. Colombia. 

Liabum pefoliatum Blake, Journ. Wash. Acad. Sci., 17:291, 

1927. 

Munnotza peruensis (Cuatrecasas) H. Robinson and Brettell. 

Phytologia, 28(1):56, 1974. Peru. 

Liabum peruense Cuatrecasas, Collect. Bot., Barcinone, 

3:304. 1953. 

Munnozia pinnulosa (0. Kuntze) H. Robinson and Brettell. 

Phytologia, 28(1):56, 1974. Bolivia. 

Liabum pinnulosum 0. Kuntze, Rev. Gen., 3(2): 163, 1898. 

Munnotia deltoidea Rusby, Bull. Torrey Bot. CI., 54:315, 

Munnozia muricata Rusby, Bull. Torrey Bot. CI., 54:315, 

1900. 

Jahrb., 28:625, 1901. 

Phytologia, 28(1):55; 1974. 

tell, Phytologia, 28(1):55, 1974. 

Colombia. 

196, 1798. Peru. 

Flora, 36:34, 1853. 

Bolivia. 

1927. 

logia, 28(1):55, 1974. Peru. 

1927. 

1927. 
NUMBER 54 

AMunnu:ia rusbli (Britton) Rusby. Bull. Torrey Bot. CI., 

Liaburn rushy Britton. Bull. Torrey Bot. CI., 19:263, 1892. 

.Munnu:ia .\enecionidu Bentham. PI. Hartweg., 134. 1844. Costa 

Rica. Panama. Colombia. \"enezuela, Ecuador. Peru, 

Bolivia. 

,54:312. 1927. Bolivia. 

Lrabutn su~gittaturn Schultz-Bipontinus. Flora. 36:37, 1853. 

Liaburn megucephalurn Schultz-Bipontinus, Flora, 36:38. 

.Munnuzia Jagilfata Weddell, Chlor. And., 1:21 I, 1857. 

C,'hrt..~astrum ,iu,gi/kz/urn \.\.'illdenow ex b'eddell. Ioc. cit, [nom. 

Lzaburn ha.\la/urn Britton, Bull. Torrey Bot. CI.. 19:263. 

Munnu:ia ha.itata Weddell ex Britton, op. cit., 263 [nom. 

.2.lrtnno:za ari.\f~~jo.rephz Rusby. Bull. Torrey Bot. Cl.. 54:314. 

.Wurinuzza aftmuafa Rusby. Bull. Torrey Bot. CI., ~54:315. 

,Munnuzia strzgulosa Rusby. Bull. Torrey Bot. Cl.. 54:317, 

.MunnoriaJi/z/~es Rusby. Bull. Torrey Bot. CI.. 54:318, 1927. 

,Wunnozia laxqlura Rusby. Bull. Torrey Bot. CI., 54:319. 

Liabum zsodon/utn Blake, Journ. \.\.'ash. Acad. Sci.. 17:298. 

Liubutn tarniutrichurn Blake. Journ. Wash. Acad. Sci.. 

.Munnuzia z.\odun/a (Blake) H. Robinson and Brettell. PhyIM~nnu~~a nzegacephaia (Schultz-Bipontinus) H. Robinson 

,blunriuziu Jugitfuta (Schultz-Bipontinus) H. Robinson and 

.bfunnuziu tarriiotrirhu (Blake) H. Robinson and Brertell, 

ALlunnozru .rzlphzozdes (Poeppig and Endlicher) H. Robinson 

Przuuole/ii i Liiphiuides Poeppig and Endlicher. Nov. Gen. et 

Lzcihurn ~zlphiuid~s(Poeppig and Endlicher) Blake, Journ. 

,Munrro:zu subcrridis (Blake) H. Robinson and Brettell, PhytoLiubuin subr,iiide Blake. Journ. LVash. Acad. Sci.. 17:294. 

.Llunizo,-zu tiinenzi Ruiz and Pavon. Syst. Veg. Peruv. Chil.. 

Liahurn /rznerce (Ruiz and Pavon) Schultz-Bipontinus, 

Munnuzia uenurissimu Ruiz and Pavon, Syst. Veg. Peruv. Chil., 

1853. 

nud. in syn,]. 

1892. 

nud. in syn.: error for .W. sagittafa Weddell]. 

1927. 

1927. 

1927. 

19'7. 

1927. 

17:298. 1927. 

tologia. 28(1):55. 1974. 

and Brettell. Phytologia. 28(1):55. 1974. 

Brettell. Ph!-rologia. 28(1):56. 1974 Inom. illeg.]. 

Ph>-tologia, '.'8(1):56. 1974. 

and Brertell, Phytologia. 28(1):56. 1974. Peru. 

Sp.. :3:3.5. pl. 261, 1843. 

\.\.'ash. hcad. Sci.. 17:3U1. 1927. 

logia, 28(1):56. 1974. Peru. Bolivia. 

1927. 

19.i. 1798. Peru. 

Flora. 36:34. 18.53. 

19.5. 1798. Peru. 

Liabum venosissimutn (Ruiz and Pavon) Schultz-Bipontinus. 

Liabum pulchrum Blake. Journ. Wash. Acad. Sci.. 17:299, 

Munnozia pulchra (Blake) H. Robinson and Brettell, Phy- 

.Wunnozia ~ilburiz H. Robinson. Phytologia. 39(5):33 I, 1978. 

Flora, 36:34, 1853. 

1927. 

tologia. 28(1):56, 1974. 

Costa Rica. 

.Munnoria subgenus Kustriera 

Munnozia acostae (Chung) H. Robinson and Brettell. Phytologia, 28(1):54. 1974. Ecuador. 

Liabum acostae Chung, Phytologia, l4((j): 

'Munnozia annua (Muschler) H. Robinson and Brettell, PhyLiabum annuum Muschler, Engl. Bot. Jahrb. 50, Beibl.. 3:84, 

;Munnoria asplundii H. Robinson, Phytologia, 34(4j :380. 1976. 

Munnoria liaboides (Lessing) H. Robinson. Phytologia. 35 

tologia. 28(1):57. 1974. Ecuador. Peru. 

1913. 

Ecuador. 

(1):38. 1976. Ecuador. 

Alibum liaboides Lessing, Syn. Comp., 1.52. 1832. 

Lzabum /iaboides (Lessing) Hieronymus. Eiigl. Bot. Jahrb.. 

19:63, 1894. 

Liabum alibum FIieronymus. Engl. Bot. Jahrb.. 28527, 

1901. 

Munnoria nivea (Hieronymus) H. Robinson and Brettell, Phytologia. 28(1):57. 1974. Colombia. Ecuador, Peru. 

Liabum niueum Hieronymus, Engl. Hot. Jahrb.. 19:62, 1894. 

Liabum stuebclii Hieronymus. EnSI. Bot. Jahrh., 21:3 

1895. 

tell, Phyrologia. 28( 1):57, 1974. Ecuador. 

29:62. 1900. 

tell, Phytologia. 28( 1): 57. 1974. Colombia. 

'Wunnozia pinnatipartita (Hieronymus) H. Robinson and BretLiabum pinnatipar/ifum Hieronyniiis. Engl. Bot. Jahrb.. 

AMunnozia tenera (Schultz-Bipontinus) H. Robinson and BretKastnera tenera Schultz-Bipontinus. Flora, 36:38. 18.53. 

Liabum tenerum (Schultz-Bipontinusj Blake, Journ. Wash. 

Acad. Sci.. 17:303. 1927. 

There is no general treatment of the genus. 

The Ecuadorian species, including all but one of 

the members of subgenus Kastnera, have been 

treated by Robinson (1978a). 

Philoglossa 

FIGURE 16 

Philoglossa DeCandolle, Prodr.. 5:567, 1836. [Type-species: 

Philoglossa peruviana DeCandolle; originally monotypic.] 

Jaurneopsis Hieronymus. Engl. Bot. Jahrb., 29:53. 1900 Inom. 
60 SMITHSOKIAN CONTR1BI;TIONS TO BOTANY 

nud.]: 28:619. 1901 [nom. nud.]. [Type-species: Juumea 

mirnuloides Hieronyrnus; originally monotypic.] 

Small erect, decumbent, or creeping herbs. 

Stems, stipule-like leaf bases, both surfaces of 

leaves. and peduncles pilose or strigose with stiff 

bulbus-based hairs. Stems with little or no milky 

sap, basal branch internodes of P. peruriana up to 

18 cm long, followed by short internodes and 

leaves in false-whorls. Leaves opposite: stipulelike bases broadly oblong, usually deeply emarginate: petioles unwinged; blades herbaceous, 

ovate to elliptical, trinervate, margins serrulate to 

serrate, upper and lower surfaces green, without 

tomentum. Inflorescence from axils of normal 

leaves or from axils of leaves in false-whorls of 4 

leaves. Heads solitary on slender and sometimes 

clustered peduncles, broadly campanulate; involucral bracts 20-30, unequal to subequal, oblong 

to oblong-lanceolate; outer bracts fleshy at base 

with obtuse herbaceous tips, margins densely ciliate with stiff hairs: receptacle without chaff, 

crests unlobed or slightly lobed. Flowers usually 

yellow, disk flowers red in 1 species. Ray flowers 

2 1-70, tubes hirsute, limbs linear and glabrous, 

slightly notched at tip, bilobed; style branches 

elongate, not spiralled. Disk flowers 30-60; corolla tube and lower throat hirsute on outer surface; throat abruptly expanded at base, lobes 

with large stomates near margins, with spicules 

or stiff hairs at tip; walls of cells of anther collars 

without annulations; thecae black, not digitate at 

base; median endothecial cells subquadrate in 

series with numerous thickenings on lateral walls, 

transverse walls very faint: appendages smooth, 

with entire to denticulate margins, cells rather 

elongate with truncate or oblique end walls; nectary not or slightly lobed; style base not or 

scarcely enlarged; style branches short, less than 

1/2 as long as the hirtellous portion of the upper 

shaft, with obtuse to rounded tips. Achenes compressed, with 2 ribs, mostly glabrous, sometimes 

hirtellous or winged on ribs, cells of achene wall 

containing small quadrate raphids, carpopodium 

poorly differentiated with many series of quadrate to short oblong cells, walls of cells scarcely 

thickened; pappus of small, usually deciduous 

squamellae or awns, or lacking. Pollen grains 35- 

37 pm in diameter, with spines rather unevenly 

disposed, with small or indistinct internal columellae. 

Philo<glossa was described originally by DeCandolle (1836) on the basis of material from 

near Lima, Peru, and was placed in the proximity 

of Encelia and Kudbeckia. members of the Heliantheae. The genus was later named but not validated by Hieronymus (1901) under the name 

’.Jaumeopsis” based on material from Ecuador and 

Bolivia. Hieronymus also associated the genus 

with the Heliantheae-Helenieae series. Bentham 

and Hooker (1873) had placed Philoglossa in the 

Heliantheae in the subtribe Melampodiinae. 

Such a relationship was understandable, given 

the black anthers and the compressed achenes 

with reduced pappus. The position in the Melampodiinae seems to derive from the frequently 

incomplete fertilization or incomplete development of the central flowers, often given the appearance of a sterile disk. It was Sandwith (1956) 

who first cited the unpublished conclusions of 

S.F. Blake regarding the relationship of Philoglossa 

to Liabum, a position confirmed by Robinson and 

Cuatrecasas (1973) and Robinson and Brettell 

(1974b). The close relationship to Erato, with 

which it shares the distinctive pubescence and the 

reduced number of ribs on the achene, kvas not 

fully recognized until the study of the tribe Liabeae in Ecuador by Robinson (1978a). 

The genus \vas originally credited with a single 

species (DeCandolle. 1836) and continued to be 

regarded as monospecific even after the description of a second species by Hieronymus (1900, 

190 l), which he failed to recognize as a Philoglossa. 

Sandwith (1956) described a new species and 

suggested that there were three species in the 

genus. Robinson and Cuatrecasas (1973) provided a summary of the genus with four species, 

but shortly afterward a fifth species was described 

(Robinson, 1976b). 

The individual species show some distinctive 

characters, including the elongate internodes followed by false-whorls of four leaves and the spinose crested tips of the disk corolla lobes in P. 
NUMBER 54 

FIGIIRE i(j.-f'hi/oghu ,/irrii~~iam~ DeCandolle: A. habit. X '5: B, foliar hairs. X 50: i:. head. X :I: 

D. ray corolla. X 16: E, disk corolla. X 16: F. disk style. X 16: (;. anther. X 20: 1-1. mediaii 

endorhecial cells, X 400; I. achenes. side and top views. X 16. 

fi 1 
62 SMlTHSONIAN CONTRIBUTIOKS TO BOTATiY 

peruczana. the winged outgrowths of the achene 

margin in P. pterocarpa. the procumbent habit in 

P. rnirnuloides, and the numerous, short. barbellate 

or laciniate pappus squamellae in P. blukei. The 

most distinctive species. P. purpureodisca. has purple disk flowers: such as are found in the genus 

Chionopappus but which are otherwise not known 

in the tribe. The last species also has a greater 

number of flowers in the head, up to 70 rays and 

60 disk flavers. a well-developed pappus of two 

awms and intervening squamellae, and large setae 

on the tips of the disk corolla lobes similar to the 

hairs found on the vegetative parts of the plant. 

The floiver color in P. purpureodisca is approached 

in some specimens of P. peruuiana that have purple 

tips on the disk corollas. It seems significant that 

the only examples of such a color pattern in the 

tribe occur in the same area of coastal Peru, a 

comparatively small part of the range of the tribe. 

The genus is mostly limited in distribution, 

with four of the five species concentrated in the 

coastal region of Peru. The single species P. miinuloides occurs outside of that area. extending 

from southernmost Colombia south through Ecuador into the interior mountain ranges of Peru. 

The latter species has one outlying population in 

Bolivia (Hieronymus, 1901). Some of the extreme 

distribution of the latter species may derive from 

its use by natiL’e populations. Bristol (1964) indicates that the leaves of the plants are apparently 

eaten for flavoring in various boiled foods, and 

the noncultivar is frequently boiled and fed to 

pigs. 

The genus contains the following five species. 

fYrilo</o\ \(i blcihri €1, Robinson and Ciiatrecasas. PhytoloSia. 

/’////f~~//J\\// rnirnu/ordri (IIieron? nius) H. Robinson and Cua21ii,-)i:385. l97,3. Peru 

trecases. Phytologia. 26(5):384. 1973. Colombia. Ecuador. Peru. Bolivia. 

,/aiivira mrmuloidei Hieronynius. Engl. Bot. Jahrb.. 29:.52. 

1900. 

JaurnrupJri mzrnu1oide.r Hieronymus. Engl. Bot. Jahrb.. 

28:619. 1901 [noni. nud.]. 

,/(iiiinrol).si.s mimuloides fornia .subirile,.yrjiliro Hieron).nius. 

hgl. Bot. Jahrb.. 28:619. 1901 [noni. nud.]. 

I’hiloglossn /j~rriiiann var. snpidu Bristol. Bot. hlus. Lfts. 

Harvard Lniv,. 20:326. 1961. 

l%r/o,glossu ~nrniuloides var. .supidu (Bristol) H. Robinson and 

Cuatrecasas. Phytologia, 26(5):38.i. 1973. 

1’/ii/u~~/il,\sa rniinulordes fornia ~ihl~~/~,{i~/~~/iO Hieronymits ex 

H Robinson and Cuatrecasas. Phytoloqia. 26(j):.38.5. 

1973. 

I’/rilug/os,\u pcriii~icoin DeCandolle. Prodr.. 3:567. 1836. Peru. 

f’hrio,gloss(i ptrrocurpha SandLvith. Kew Bull.. 1936:292, 1956. 

I%i/o,oli~,\su /iur,/iurroditca H. Robinson. Phytoloqia. 311 1 i:54. 

Four of the species are treated with a key and 

descriptions by Robinson and Cuatrecasas (1973). 

Peru, 

1976. Peru. 

Species Excluded from the Tribe 
Literature Cited 

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Peruciano et in terra Amatonica annis mdcccxxvii ad 

mdcccxxxiz legit Eduardus Poeppig et cum Stephano Endlirher descripsit, iconibusque illustracit. Volume 3. 

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1980. 

Nash. D.L. 

1976. 

Nordenstam. B. 

Norlindh, T. 

1977. 

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1976a. Studies in the Liabeae (Asteraceae), 111: Notes on 

the Genus Cacosmia. Phytologia, 34( 1) :46-52. 

1976b. Studies in the Liabeae (Asteraceae), IV: Additions 

to the Genus Philoglossa. Phitologia, 34( 1):53-56. 

1976c. Studies in the Liabeae (Asteraceae), V: New Andean Species of Liabum. Phytologia, 34(3):285-300. 

1976d. Studies in the Liabeae (Asteraceae), VI: Notes on 

the Genus Erato. Phytologia, 34(4):378-379. 

1976e. Studies in the Liabeae (Asteraceae), 1’11: Additions to the Genus Munnotia. Phytologza, 34(4):380- 

1976f. Studies in the Liabeae (Asteraceae), L’III: Additions to Liabum and ’Wunnozia. Phytologia, 35(1):37- 

42. 

1977a. Studies in the Liabeae (Asteraceae), IX: Additions 

to Munnoria and Oligactis. Phytologia, 35(1): 199- 

204. 

1977b. Studies in the Liabeae (Asteraceae), X: Additions 

to the Genus Paranephelius. Phytologia. 35(3):233- 

240. 

1977c. Studies in the Liabeae (Asteraceae), XI: New Spe387. 

cies in the Kew Herbarium. Phytologia, 35(6):488- 

496. 

1978a. 190 (2), Compositae-Liabeae. Flora of Ecuador, 

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1978b. Studies in the Liabeae (Asteraceae), XII: A New 

Species of Munnotia from Costa Rica. PhFtologia, 

39 (5) : 3 3 1 - 33 3. 

1978~. Studies in the Liabeae (Asteraceae), XIII: A New 

Species of Liabellum from Nayarit, Mexico. Plytologia. 41(1):45-49. 

1980a. Studies in the Liabeae (Asteraceae), XIV: New 

Species of Liabum and Oligactis. Phytologia, 46(2): 

99-103. 

1980b. New Species of L’ernonieae (Asteraceae). \-: Additions to Lkrnonia from Brazil. Phytologia. 

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A Revision of the Tribal and Subtribal Limits of 

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Robinson, H., and J. Cuatrecasas 

1973. 

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1798. 
NUMBER 54 65 

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Index 

(Synonyms and page numbers of main entries of taxa of Liabeae in italics) 

Adenoiin. 7 

Aiibum. I, 54. 57 

Allendea. 2. 31. 32 

.4mellus. 1 

Andromarhia, 1. 2, 31. 32 

liaboides, 54, 57, 59 

ianceolata, 3 1. 34 

umbeilatus, 3 1, 35 

acauits. 49. 51 

deppeana. 43 

excelsa , 2 9 

grandifliora, 35 

hierartoides, 52 

igniaria. 31, 35 

melastomoides. 35 

nubigena, 39 

poiteaui. 35 

stssiiiflora. 39 

solidaginea: 35 

uerbasifolia, 27, 29 

colubilis, 37: 39 

JuSsieU2, 58 

Andromachiopsis (subgenus of OligacAngelianthus, 35 

humilis. 37 

Austroliabum, 6, 9. 11, 13, 15. 16, 17. 

tis), 7, 11, 38, 39 

19, 20, fig. 1 

candidum. 20, 22, fig. 2 

eremophilum. 22 

mulgediifoltum, 22 

polymnioides, 22 

Baccharis. 1, 52 

Blepharispermum, 4 

Cacalioid. 7 

Cacosmia. 1. 2. 6, 7, 11. 13, 14. 15. 16, 

19, 20, 22, 24. 27, 29. 32, fig. 1 

harlingii, 24 

hieronymi, 24 

quinqueneruia, 25 

rugosa. 22, 24, 25. fig. 3 

rugosa var. arachnoidea, 24 

rugo5a var. kingii, 25 

rugosa var. nivea, 25 

Centratherum. 8 

Chaenactidinae, 7 

Chionopappus, 2, 4, 6. 9, 11, 13, 14. 15. 

16, 19, 20, 24, 25, 29, 62. fig. 1 

benthamii. 15, 25, 27, fig. 4 

Chresta. 9 

Chrysactinia. 51 

Chrysactinium, 2, 5, 6, 11. 12, 14, 15, 

16. 17. 18. 19. 49, 51 

acaule. .?I, fig. 13 

amphothrix, 51 

arthrothrix, :TI 

bicolor. 51 

caulescens. 51 

trigeroides. 62 

hieracioides, 51 

longiradiatum. 52 

rosulatum. 52 

/enuius, 52 

Chypactinium (section of Andromachia). 

Ch ysastrurn, 54 

Chysastrum (hubgenus of Liabum). 54 

Ciairuiliea, 22 

Conyra stipulata. .78 

49 

sa,git/aturn. 54. 59 

quinquenervia, 22, 24 

Dahlia, 7 

Encelia! 1, 60 

Erato. 1. 2. 4% 5. 6. 7. 10, 11. 12, 14, 16. 

18, 19, 49, 52. 54. 60, fig. 1 

polymnioides. 16, 52, 54, fig. 14 

sodiroi. 15. 54 

stenolepis. 54 

vulcanica, 54 

Erato (subgenus of Munnozia). 52 

Eremanthus. 9 

Eremothamnus. 8, 9 

Eupatorium hitchcockii, 62 

Eutetras, 9 

Ferreyranthus, 4, 6, 11, 13, 14, 15, 17, 

19, 20, 24, 27, 29, fig. 1 

excelsus. 29 

fruticosus, 29 

pseudosaluitfoiius, 29 

66 

ramonii, 29 

rugosus, 29 

touarii, 29 

vaginans, 29 

verbascifolius, 4, 15. 29, fig. 5 

vernonioides. 29 

Gaillardiinae, 7 

Gongrothamnus. 2 

Gynoxys boliviana. 62 

columbiana, 62 

Heterotheca deltoidea, 58 

Hieraceum-type, 15 

Hypochaeris, 45 

Jaumea mimuiotdes. 60, 62 

Jaumeopsis, 59, 60 

mimuioides, 62 

mtmuloides forma subintegrtfolza. 62 

Kastnera, 54 

Kastnera (subgenus of Munnozia), 5. 

6. 7. 14. 15. 16. 54, 56. 57. 59 

Krigia. 9 

(enera, 59 

Liabelium Cabrera, 2; 35 

Liabellum Rydberg, 2,6, 10, 11, 12, 13, 

14, 16, 17, 19, 29, 31, 41, 43, fig. 1 

humile. 35, 37 

angustissimum, 31 

cervinum, 31 

gentryi, 31 

palmeri, 31, fig. 6 

Liabinae, 4, 6, 7, 10, 11. 13, 14, 15, 16, 

19, 24 

Liabum. 1, 2, 6, 9. 10. 11. 12, 13, 14, 

15, 16. 18, 19. 20, 24, 25, 27, 29, 

31, 32, 38, 43, 44, 52, 56, 60, fig. 1 

acaule, 51 

acostae, 59 

acuminatum, 34 

acutifolium, 35 

adenotrichum. 43 

a&, 57 

alibum. 59 
NUMBER 54 67 

arnphothrix, 51 

amplexans. 35 

amplexicaule, .?4 

anatina. 54 

andrieuxii. 43 

andromachioidrs, 4.3 

anguslissimum. .?I 

angustum. 57 

annuurn, 59 

arthrothrix, 51 

asclepiadeum. 34 

asperifolium, 47 

aiiric-rilritum. 22 

t~arationene. ..If 

barclayae. 34 

biatlmualum. 39 

bicolor. 51 

boltL~ianum. 62 

bonplandii, 35 

bourgeaui, 11. 34 

boyacense. .?9 

brownei, 3 1. 35 

caduciyolium: 43 

caliense, ,711 

canarense. j7 

candidum. 22 

candidum var. subcirrhosum. 22 

cardenasii, 57 

caulescens. 51 

ceruinum. 31 

columbianum, 62 

conuencionense. 58 

coriaceum, 39 

cocymbosurn (R. & P.) Sch. Bip.. 58 

coymbosuin Sch. Bip., 58 

crispum. .34 

cubense. 34 

cusalaguensis. 39 

deamii, 43 

deppeanum. 43 

diehlii. 32, .?4 

dimidium. 43 

discolor, 44. 49 

domingense, 35 

ecuadoriense, 3.9 

eggersii, 34 

eremophiium. 22 

erigeroides. 62 

eriocabx, 56. 58 

eriocaulon, 34 

eupatorioides. 62 

excelsum. 29 

falcatum. 34 

ferrevrii, 34. .35 

floribundum. 35 

frutirosum, 29 

fuluotomenlosum, 35 

gigan teum, 58 

glabrum, 44 

glabrum var. hypoleucum, 44 

glandulosum. 58 

granatensis, 39 

grandiflorum, 35 

hallii, 39 

hastatum. 59 

hastlfolium, 58 

hexagonum, 58 

hieracioides, 52 

hirtum, 58 

homogamum, 62 

hypoleucum, 43 

igniarium. 32, 34. 35 

insigne, 54 

isodontum, 59 

jelskii, 47 

jussieui. 58 

kingii, 35 

klattii, 41, 44 

lanatum, 49 

lanceolatum, 34, 58 

lattciferum, 58 

latifilium, 39 

lehmannii, 35 

liebmannii, 44 

longipes. 35 

longiradiatum, 52 

bratum, 58 

macbridei, 35 

megacephalum, 59 

melastomoides, 32, 35 

meridense, 39 

mikanioides, 39 

moorei, 44 

mulgedii folium, 22 

nigro-pilosum, 35 

niueum, 59 

nonoense, 58 

nonoense var. microcephalum, 58 

nubigenum, 39 

nudicaule, 35 

oblanceolatum, 35 

ochraceum, 39 

oleariotdes, 58 

ovatifolium, 35 

ouatum, 47 

oxyphyllum, 58 

pa lla t angense , 54 

palmeri, 29, 31 

pastoense, 39 

perfoliatum, 58 

peruense: 58 

pinnat iparti tum. 5.9 

pinnulosum. 58 

plaQlepis, 44 

poiteaui. .?5 

pobanthum. 44 

polycephalum, 35 

polymnioides. 22 

pringlei, 44 

pseudosalaiifolium. 29 

pulchrum, 56, .i9 

rosulatum, 52 

rugosum, 29 

rusbiii. 59 

sagtltatum, 54. 39 

saloyense, 35 

salci~filium, 29 

sandemanii, 11, 32, 34. 35 

scandens. 39 

selleanum. 3'5 

sericolepis, 44 

sessiliylora, 39 

si lp hioides. 59 

simile. 44 

solidagineum, 32, 34. 33' 

steinbachii, 11, 3j 

stenolepis, 54 

stipulatum, 3.j 

J/rin(uiri. 47. 119 

stuebeli I, 59 

suhacaule. .I.! 

.subrirrhosum. 22 

subglandulare. .4.? 

sublobatum. 44 

subumbellatum, 34 

suboiride, 59 

srpyloruiczii. 49 

tabanense, 58 

taeniotrichum, 59 

tajumulcense. 44 

tenerum, 59 

tenuius (tenuior). 52 

tondurii, 44 

toisarense. 39 

touarit. 29 

trianae, 35 

trinerur, 59 

ulei. 34 

umbellarum, 31, 35. fig. 7 

uniylorum. 47 

uagans, 44 

uaginans, 29 

caleri. 39 

vargasii. 35 

uenosisst mum, 59 
68 SMITHSONlAN CONTRIBUTIONS TO BOTANY 

terbasctfolium. 1'9 

wlubilr, 39 

uulcanicum, 54 

icddmwi, 35 

wrightii. 35 

wurdackii. 35 

.We,falia bum 4 1 

andrieuxii. 43 

pringlei. 44 

Melampodiinae, 2, 60 

Microliaburn. 2, 11, 13, 14, 15, 16, 17, 

19, 20, 35, fig. 1 

humile. 37. fig. 8 

Munnozia, 1. 2, 5: 10. 11, 12. 15, 16, 

17, 18. 19. 'LO, 43, 49. 51, 52, 54, 

36. -37. fig. 1 

acostae, 59 

affinis. 57 

anatina. 54 

angusta. 57 

annua. 57, 59 

aristejosephi, 59 

asplundii, 59 

attenuala, 59 

campii. 15. 56, 57 

canarensis, 57 

cardenasii. 57 

chimboracensis, 57 

chysunthemoides. 58 

convencionensis. 57 

corynibosa. 56, 57. 58 

delloidea, 58 

ferreyrii, 51, 58 

Ji'iiprs, 59 

foliosa. 58 

fosbergii. 58 

gigantea. 58 

glandulosa. 58 

hnstata. 59 

hastifolia, 4. 15. 17. 57, 58 

herrerue. 58 

hirta, 58 

isodonta, 59 

jussieui, 15, 56, 57, 58 

karstenii, 58 

kiat ti i, 58 

lanceolata. 54, 56, 58 

laxiylora. 59 

liaboides, 59 

longifolia. 58 

lyrata, 57. 58 

rnaronii, 4. 10, 57, 58 

megacepha la, 59 

mu ri cata . 58 

nivea. 51. 57. S9 

nonoensis, 58 

olearioides, 58 

oxy-phylla, 58 

perfoliata, 51, 58 

peruensis, 58 

pinnatipartita. 59 

pinnulosa, 58 

pohmnioides, 54 

pulchra. 59 

rusbyi. 59 

qittala (Sch. Bip.) H. Robins. & 

.qttla/u \Veddell. 59 

senecionidis, 54, 56. 57, 59, fig. 15 

silphioides, 59 

stenolepis, 54 

slriguiosa, 59 

subviridis, 4, 57, 59 

tabanensis, 58 

taeniotricha, 59 

tenera, 59 

trinervis. 56, 59 

venosissirna, 56, 59 

culcanica, 54 

wilburii. 59 

Brettell, JY 

Munnoziinae, 4, 5, 6, 10, 11, 12, 13. 14, 

1.5. 16. 19, 24, 49 

h'assauviinae, 8 

.Veomirandea hitchcockit, 62 

Neurolaena. 2 

hornogama, 62 

Oligactis. 1, 6, 7, 9. 10, 11, 12, 13, 14, 

15. 16, 17, 18, 19, 24, 25, 27, 29, 

32, 34. 37, 38. 39, 43, fig. 1 

upodocepha la, 39 

asplundii. 38, 39 

biattenuata, 39 

bymnsis, 39 

coriacea, 39 

coriacea var. granatensis, 39 

cusalaguensis, 38, 39 

ecuadorensis, 39 

fruticosa, 29 

garcia-barrigae. 39 

granatensis, 39 

hallii. 39 

latifolia. 39 

rnikanioides. 39 

nubigena, 39 

ochracea, 39 

pastoensis, 39 

pichinchensis, 39 

scandens, 39 

sessilifolia, 39 

valeri, 39 

volubilis. 39, fig. 9 

Oltqncirr (section of Andromachia), 37, 

Oiioseris discolor, 49 

Orithrophium peruvianum, 62 

.YCr 

Paranepheliinae. 1. 6. 10. 11. 13. 11. 

Paranephelius, 2, 6, 9, 10. 11. 12, 13, 

16. 19, 24, 44 

11. 15. 16. 19. 'LO. 44. 4.5, 47. 49. 

fig. 1 

asperifolius, 4i 

bullatus. 47 

ferreyrii, 43% 47 

jelskii, 47 

ovatus. 4.5. 47 

uniflorus, 13. 47. fig. 11 

wurdackii. 47 

Paranephelius (section of Liabum). 4.5 

Pectis, 9 

Philoglossa. 1. 2, 4, 5, 6, 7. 11, 12, 13, 

14. 16, 18, 19, 25. 49, 52, 54, 59, 

60, fig. 1 

blakei, 62 

rnimuloides. 15. 62 

mimuloides forma subintegrifolia, 62 

peruviana, 15, 59. 60. 62. fig. 16 

pterocarpha, 62 

purpureodisca, 15, 62 

siiphioides, 54, 59 

Prionolepis, 54 

Pseudonoseris, 6, 10, 13, 14. 15. 16: 17. 

19. 20, 47. 49. fig. 1 

discolor. 49 

striata. 49 

szyszylowiczii, 15. 47, 49 

Pseudostifftia, 8 

Psiadia, 1 

Rudbeckia, 1, 7, 60 

Sartwellia, 9 

Schistocarpha. 2 

eupatorioides. 62 

sinforosi, 62 

macrorrhizus, 45 

wernerioides, 45 

Senecio, 45 

Sinclairia, 2. 6, 9, 10, 11, 12, 13, 16. 18, 

19. 23, 31, 41, 43, 44, fig. 1 

adenotricha, 43 
NUMBER 34 69 

andrieuxii. 31. 4.3 

andromachioides. 4.3 

blakei. 4.3 

brachypus, 43 

broomae (broomei), 4.3 

caducifolia. 4,? 

deamii, 43 

deppeana, 4.3 

dimidia. 4.7 

discolor. 41, 4.3, fig. 10 

glabra, 44 

hypoleuca, 43. 44 

klattii, 44 

liebmannii, 44 

moorei, 41. 44 

pittieri. 44 

platylepis, 44 

polyantha. 43- 44 

pringlei, 41. 44 

sericolepis, 44 

similis. 44 

subglandularis. 4.7 

sublobata, 44 

tajumulcensis, 44 

tonduzii. 44 

vagans. 44 

Sinclairiopsis. 41 

klattii, 4 1, 44 

Solidago, 1 

Starkea, 1. 31 

umbellata. 3.5 

Vernonia. 9 

andrieuxii, 4 1. 43 

andromachioider, 4 7 

ViLiania. 32 

bicolar. 32 

Werneria, 45