SMITHSONIAN CONTRIBUTIONS TO BOTANY NUMBER 64 

A Palynological Study of the 

Liabeae (Asteraceae). 

Harold Robinson 

and Clodomiro Marticorena 

SMITHSONIAN INSTITUTION PRESS 

City of Washington 

1986 
ABSTRACT 

Robinson, Harold, and Clodomiro Marticorena. A Palynological Study of the 

Liabeae (Asteraceae). Smithsonian Contributions to Botany, number 64, 50 

pages, 208 figures, 1986.-Surface features and internal structure of the 

spines in the pollen of the Liabeae are illustrated by scanning electron 

microscopy and drawings made from oil immersion light microscopy. Observations are presented for all 16 genera of the tribe, Paranephelius and 

Pseudonoseris of the Paranephelinae, Astroliabum, Bishopanthus, Cacosmia, 

Chionopappus, Ferreyranthus, Liabellum, Liabum, Microliabum, Oligactis, and 

Sinclairia of the Liabinae, and Chrysactinium, Erato, Munnozia and Philoglossa 

of the Munnoziinae. No examples of echinolophate pollen are found in the 

tribe, but irregular groupings of spines are found to be characteristic of all 

but Munnozia and Chrysactinium in the Munnoziinae and Paranephelius and 

Pseudonoseris in the Paranepheliinae. The distribution of caveate forms in 

the Liabeae is clarified. The presence of a large cavus-like space interrupted 

by thin basal columuellae is confirmed in the genus Paranephelius, correcting 

early observations showing the genus as an exception to the basic uniformity 

of exine structure in the tribe. However, subcaveate forms have been discovered in the tribe in the genus Munnozia. SEM study of broken tecta in the 

genus Munnozia show previously unsuspected great potential for variation 

between closely related species and species groups. The structure of the exine 

supports views derived from floral structures regarding relationships with 

other tribes. No support is seen for a phyletic position near the Senecioneae, 

and closest similarity is seen to the Vernonieae. A basic difference is discovered in the arrangement of the basal columellae between the Liabeae and 

the Vernonieae, being clustered or forming a hollow cylinder under each 

spine in the former but forming a single solid basal columella under each 

spine in the most nearly comparable members of the latter. Genera of the 

Vernonieae shown for comparison include the non-lophate Blanchetia, Pseudostqftia, and Vernonia, and the lophate Bishopalea, Heterocypsela, and Phyllocephalum. 

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

recorded in the Institution's annual report, Smithsonian Year. SERIES COVER DESIGN: Leaf 

clearing from the katsura tree Cercidiphyllum japonicum Siebold and Zuccarini. 

Library of Congress Cataloging in Publication Data 

Robinson, Harold Ernest, 1932- 

A palynological study of the Liabeae (Asteraceae). 

(Smithsonian contributions to botany ; no. 64) 

Supt. of Docs. no. SI 1.29:64 

Bibliography: p. 

1 . Compositae-Pollen-Morphology-Atlases. 2. Compositae-Classification. 3. Palynotaxonomy. 1. Marticorena, Clodomiro. 11. Title. 111. Title: Liabeae (Asteraceae) IV. 

Series. 

QKlS2747 no. 64 [QK495.C74] 581 s [583'.55] 86-600032 
Contents 

Page 

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

Materials and Methods .................................... 2 

Pollen Structure of the Liabeae ............................. 3 

PARANEPHELIINAE ...................................... 4 

LIABINAE ............................................. 5 

MUNNOZIINAE ......................................... 6 

9 

Palynological Evidence on the Relationships of the Liabeae in the 

Asteraceae .......................................... 10 

Appendix .............................................. 14 

Literature Cited ......................................... 2 1 

Figures ................................................ 23 

Acknowledgments ...................................... 3 

Key to Types of Liabeae Pollen ............................. 

... 

111 

A Palynological Study of the 

Liabeae (Asteraceae) 

Harold Robinson 

and Clodomiro Marticorena 

Introduction 

A recent series of studies (Robinson and Brettell, 1973a, 1974; Robinson, 1978, 1983a) has 

explored the taxonomy and evolution of the 

geographically restricted neotropical tribe Liabeae, using mostly macroscopic and some microscopic structural features. Most recently the 

chromosome numbers of the tribe have been 

reviewed (Robinson et al., 1985). The present 

study of the pollen continues the effort to improve understanding of this previously most 

poorly understood tribe of the Asteraceae whose 

members were neither placed together nor recognized at the tribal level in the traditional treatments of the family. A summary of the pollen 

characters of the tribe is given based on examination of over half the species by at least oil 

immersion light microscopy. Generic patterns 

are noted, and detailed taxonomically significant 

variations in exine structure in closely related 

species groups are documented. The views presented in the recent studies by the senior author 

regarding relationships and distinctions between 

the Liabeae and other tribes such as the Senecioneae and Vernonieae are extended by knowledge of the pollen structure. 

Harold Robinson, Department of Botany, National Museum of 

Natural History, Smithsonian Institution, Washington, D. C. 

20560. Clodomiro Marticorena, Departamento de Botanica, Instituto de Biologia, Universidad de Concepcion, Chile. 

The Liabeae was not recognized at the tribal 

level or properly placed together in a related 

group until this century (Rydberg, 1927; Blake, 

1935; Sandwith, 1956; Cabrera, 1954; Robinson 

and Brettell, 1973a, 1974; Robinson, 1978, 

1983a). In the earliest classifications the members of the tribe with a capillary pappus were 

placed in the Vernonieae (Cassini, 1828). Later 

these were placed in the Senecioneae (Bentham 

and Hooker, 1873) while genera with reduced 

or plumose pappus forms were described in various other tribes such as the Heliantheae, Helenieae, and Mutisieae. The history of the tribe is 

dealt with more completely by Robinson (1 983a). 

The restricted neotropical distribution of the 

Liabeae and the failure to treat the group as 

taxonomically distinct resulted in their omission 

from the earliest palynological studies of the 

Asteraceae. It was in the study by Stix (1960) 

that the group was first included with observations on 14 species that are presently considered 

members of the genera Liabum, Austroliabum, 

Liabellum, Munnozia, Sinclairia, and Paranephelius. The study by Stix cited some variation in 

internal structure of the tectum in the pollen of 

the tribe. In 1966, the pollen of Cacosmia was 

described and compared with that of Liabum by 

Skvarla and Turner. Light microscope oil immersion observations of thick sections of Chionopappus pollen were used by Marticorena and 

Parra (1 974) to confirm the relationship of that 
2 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

genus to Liabum. Regularity of spine distribution 

and some variation in size of grains were mentioned by Robinson and Brettell (1974). The 

results of transmission electron microscope studies were included in the review of the family by 

Skvarla et al. (1 977) where apparent variations 

between non-caveate forms with basal columellae 

and forms without basal columellae were noted 

in the tribe. The apparently anomalous Paranephelius was shown to possess thin basal columellae 

by Feuer and Dillon in an oral presentation 

(1982). It remains for the present study to 

broaden the survey of the tribe to include all 16 

genera, introduce results of scanning electron 

microscope observations of whole and broken 

pollen grains, and to compare the results with 

conclusions from the simultaneous phyletic studies of the tribe (Robinson, 1983a). 

Materials and Methods 

A variety of techniques have been used for the 

study of pollen of the Liabeae. Many initial observations have been made of material mounted 

in Hoyer’s solution (50 cm3 distilled water; 30 

grams Gum Arabic, U.S.P. Flake; 200 grams 

Chloral hydrate; 20 cm3 Glycerin). The medium 

is regarded as temporary, eventually altering the 

index of refraction and making the exine essentially invisible. Whole grains have also been 

mounted in glycerin jelly. Paraffin sections have 

been prepared by Marticorena. All the above 

material was viewed with the light microscope 

with X40 or oil immersion objectives. Results 

were comparable in detail to those by Stix (1 960). 

The sections, which are often fragmentary and 

folded, have been used in the present study primarily for reference and only one photograph of 

a section of Chionopappus prepared during the 

study of Marticorena and Parra (1974), is presented herein. The slides of whole grains were 

used by Marticorena to produce ink drawings 

representing the internal spine structure in vertical view of over 100 species. These were 

grouped to show a range of columellar variation 

under different spines in grains from single collections. Representatives (Figures 1 -40), redrawn from the Marticorena camera lucida drawings, are included herein to provide the reader 

with a more convenient survey of the variation 

of such structure in the tribe than is available in 

the SEM micrographs. 

No observations have been made in this study 

with the transmission electron microscope, but 

data from previous TEM studies by Skvarla et al. 

(1 966, 1977) and Tomb (1975) are cited. In spite 

of some artifacts produced by breakage, the use 

of broken grains has proven much superior to 

TEM preparations for obtaining an understanding of the three-dimensional columellar structure. 

A primary effort in the present study has involved use of the scanning electron microscope 

(SEM). Pollen samples have been removed from 

herbarium material or in one case from a living 

plant. In a few cases the pollen was not acetolized, 

but most samples were acetolized according to 

the procedure outlined in Erdtman (1966). Acetolyzed grains were fractured by crushing with 

the edge of a glass slide, or more often while still 

in the centrifuge tube by using a needle. All 

samples were pipetted on to cover slips and allowed to dry. The samples on coverslips were 

thinly coated using an EFFA carbon coater, involving evaporation by the carbon-arc technique. 

They were then coated with gold-paladium using 

a DC sputtering unit. Specimens were examined 

and photographed with Cambridge Stereoscan 

MK IIA, Cambridge Stereoscan S4-10, or Coates 

and Welter field emission microscopes. 

The terminology for pollen structure used in 

the present paper follows mostly that of Skvarla 

et al. (1 977). Sizes of pollen grains are given over 

a broad range to account for variations due to 

method of preparation. Grains mounted in Hoyer’s solution tend to swell most, those in glycerine 

jelly are about the same size or slightly smaller 

than those in Hoyer’s, the grains in water are 

distinctly smaller, and air- or vacuum-dried material, such as that examined by SEM, is completely unexpanded and may be as small as Y3 the 

Hoyer’s Solution diameter. Actual overlap in 
NUMBER 64 3 

sizes in various genera should be less in any one 

medium than indicated by the general figures 

given. Measurements cited for the Liabeae pollen 

are of the equatorial diameters, but none of the 

grains of the tribe are sufficiently oblong or 

oblate in shape to produce a markedly different 

polar diameters. Since only the general sizes are 

mentioned for the Vernonieae the shorter polar 

diameters of the larger oblate grains of that tribe 

are not noted. Measurements in the study exclude the projecting parts of the spines. Specimens used in the study are listed in the appendix. 

graphs were prepared by the SEM Laboratory 

(supervised by Walter Brown) at the National 

Museum of Natural History (NMNH), Smithsonian Institution. The microscopes were operated 

by Mary Jacque Mann and Susann Braden. Dr. 

Joan Nowicke is to be thanked for use of the 

facilities of the Palynological Laboratory in the 

NMNH Department of Botany, and for suggestions and help in the traditional methods of 

breaking pollen grains. Janice Bittner of the Palynology Laboratory and Andrea Sessions helped 

by processing the pollen. Brian Kahn has helped 

in preparation of the plates. 

ACKNOWLEDGMENTS.-The SEM microPollen Structure of the Liabeae 

Stix (1 960) in the initial study of pollen of the 

Liabeae, recognized three variations in the tribe. 

The first or andromachioides-type has large columellae restricted to the area under the spines, 

and most closely agrees with the type termed 

“Liaboid” by Skvarla et al. (1977). The species 

listed with this pollen type by Stix include five 

species now placed in Sinclairia, two species now 

placed in Liabellum, two species of Austroliabum, 

and Munnozia rusbyi. 

The second pollen type of Stix (1960), the 

umbellata-type, has smaller columellae under 

each spine and some basal columellae in the 

intervening parts of the exine. The three species 

cited by Stix include one now known as Munnozia 

hastfolia and the only two true Liabum species in 

her study. The pollen type is treated herein as a 

comparatively minor variation of the first type. 

The different examples seem to occur in species 

that are closely related to others of the andromachioides-type with larger columellae under 

the spines. 

The remaining pollen type of Stix (1 960), the 

ovatum-type, has the columellae partially to completely separate from the footlayer. The columellae have the form seen in the upper layer of 

the first and second types. This pollen type is 

cited by Stix from Liabum ovatum, which is a 

member of the genus Paranephelius. The type 

was examined by Skvarla et al. (1977) on the 

basis of a specimen presently determined as P. 

unijlora, and the suggestion of a caws and the 

more senecoid rather than anthemoid nature was 

emphasized. The apparent anomally of a senecoid pollen type in the Liabeae was partially 

resolved in the study by Feuer and Dillon (1 982), 

which discovered slender columellae under the 

spines that connect with the foot layer. 

The present understanding of the pollen structure in the Liabeae is reviewed herein according 

to the revised concept of the tribe with 16 genera 

distributed in three subtribes, Paranepheliinae, 

Liabinae, and Munnoziinae (Robinson, 1983a,b). 

The pollen of the Liabeae (Figures 1-180) is 

generally characterized as follows: 

Tricolporate; 25-50 Fm in diameter, spherical 

to slightly oblate-spheroid or in some cases where 

colpi are closed appearing oblong; ecto-apertures 

meridionally elongate (Figures 41, 50, 69, 74, 

104, 124) and endo-apertures transversely shortoval (Figure 121); echinate with spines often 

irregularly distributed and variously confluent 

with each other at the base (Figures 60, 62, 64, 

67, 94), sometimes regularly dispersed on the 

intercolpar area (Figures 49, 50, 140, 141, 147, 

148, 130). Internal structure basically with two 

levels (Figures 68, 88, 132, 179, 180) of columellae. Usually with few (Figures 1, 2, 46) to 

many (Figures 156, 160) large basal columellae 

clustered under spine bases, rarely with additional basal columellae dispersed in inter-spinal 

areas (Figure 1 18), columellae under spines 

sometimes coalesced into hollow cylinders (Fig- 
4 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

ures 175, 176); basal columellae sometimes reduced or separated from foot layer (Figures 162- 

167) with partial to complete development of 

cavus; upper layer of columellae usually regular, 

short, dense, with upper ends forming a perforate tectum, and covered below by unsupported 

or partially supported internal tecta forming a 

somewhat undulating or papillose surface in interspinal areas. 

Pollen size and spine distribution show variation in the Liabeae. The generic pair of Liabum 

and Oligactis has the smallest average pollen size 

in the tribe at 25-35 pm. A number of other 

members of the subtribe Liabinae, such as Cacosmia and Ferreyranthus, can have pollen as small, 

27-30 pm and 29-35 pm respectively, but one 

species of the latter ranges up to 43 pm. Still, in 

other Liabinae, such as Austroliabum, 33-40 pm, 

Chionopappus, 37-40 pm, and especially Sinclairia, 35-55 pm and Liabellum, 47-52 pm, the 

pollen is characteristically larger. The members 

of the other two subtribes are more consistent in 

size. Both genera of the Paranepheliinae have 

pollen mostly 35-45 pm in diameter. The four 

genera of the Munnoziinae, with rare exceptions, 

range from 30-40 pm in diameter. The primary 

exception is the typical element of Munnozia that 

ranges between 37-45 pm in diameter. For a 

sampling of size measurements see the appendix. 

Many genera of the Liabeae have a type of 

uneven distribution of the spines on the pollen 

that seems unique in the family. In these forms 

the spines seem to be in groups, sometimes in 

pairs or in short crests of three with confluent 

bases. This is especially evident in SEM views of 

some unacetolized grains (Figures 62, 63). Such 

grains seem to be best characterized by the irregular placement of the spines along the lateral 

margins of the colpi (Figures 72, 74). This pollen 

form is found throughout the subtribe Liabinae 

and in Erato and Philoglossa of the Munnoziinae, 

but it is lacking in Munnozia and Chrysactinium 

of the Munnoziinae and in the Paranepheliinae. 

The distribution in the tribe suggests that is the 

plesiornorphic condition in the tribe. It is correlated rather closely in its distribution in the tribe 

with the apparent plesiomorphic condition of the 

basal columellae under the spines. 

The spines are more evenly disposed on the 

surfaces of the pollen in two pairs of genera in 

two separate, comparatively derived groups, 

Munnozia and Chrysactinium in the Munnoziinae, 

and Paranephelius and Pseudonoseris in the Paranepheliinae. This arrangement is reflected in the 

regularly aligned spines along the lateral margins 

of the colpi (Figures 41, 50). The pattern is more 

evident in a preparation of unacetolized grains 

of Paranephelius (Figure 49), where the depressions between the bases of each of the spines are 

equal in spite of some displacement seen in the 

spines. The character can be observed under the 

light microscope. The separate and derived nature of the regular spine pattern in the two 

groups is indicated by the retention of the irregular pattern in two of the genera of the Munnoziinae and by the slight evidence of irregularity 

that survives along the margin of the colpus in 

Paranephelius. 

There is greater variation in the Liabeae in 

the internal structures of the pollen than in external features. The variations between many 

species of Munnozia are perhaps the greatest thus 

far known for internal exine structure of pollen 

in any such closely related group. Variations 

include tendencies toward the caveate condition 

seen in the Paranepheliinae, Philoglossa, and 

Munnozia, the fusion of columellae into hollow 

cylinders in Chrysactinium and Munnozia, and the 

production of an anomalous inner series of columellae seen in Munnozia subgenus Kastnera. 

These and other details of pollen structure in the 

Liabeae are best reviewed in each subtribe individually. The subtribes are treated herein in the 

following order: Paranepheliinae, Liabinae, and 

Munnoziinae. 

PARANEPHEL1INAE.-The subtribe Contains 

two genera that share long style branches, pale 

anthers, elongate raphids in the achene wall, 

narrow disk corollas, and leaves mostly in basal 

rosettes. The inflorescence has few heads and in 

the type-genus the heads are sessile in the rosette 

of leaves. The two genera, Paranephelius (Figures 
NUMBER 64 5 

1, 2, 41-49) and Pseudonoseris (Figures 50-53), 

have pollen structure that supports their status 

as a related pair. Both have grains that appear 

caveate, with spines rather evenly distributed 

over the intercolpar regions. The illustrations of 

Paranephelius (as Liabum ovatum) by both Stix 

(1960, Abb. 11) and Skvarla et al. (1977, pl. 15: 

fig. F) are equivocal regarding complete separation of the outer exine from the footlayer, and 

their preparations seem to have a columellate 

layer of the exine tightly contracted against the 

footlayer. A point in the TEM of Skvarla et al. 

(1977, pl. 15: fig. F) seems to show many columellae closely appressed against the foot layer 

and at least one clear example of a basal attachment. However, closer examination of the most 

obvious basally attached element shows that it is 

not directly attached to the overlying tectum. 

Preparations in Hoyer’s Solution, thick sections 

viewed with the light microscope, and SEMs of 

broken grains indicate that the columellate exine 

is normally less close to the foot-layer, and that a 

cavus-like area is present in the intercolpar parts 

of the grain. Studies of Paranephelius with oil 

immersion (Figures 1, 2) and SEM (Feuer and 

Dillon, 1982; Figures 45-48) show that the 

grains are not strictly caveate. They have a few 

narrow basal columellae under the spines that 

reach the foot layer. The SEM also shows occasional short pegs on the foot layer, but it is most 

likely that the basally attached structure seen in 

the TEM photo of Skvarla et al. (1 977) is part of 

one of the slender columellae that has been bent 

and cut. 

Broken pollen grains of Pseudonoseris have not 

been examined with the SEM, and oil immersion 

study does not show columellae of the type found 

in Paranephelius. If basal columellae are present 

in Pseudonoseris, they are more reduced than 

those of Paranephelius, which can by seen by light 

microscopy. The two genera, nevertheless, both 

show a tendency for great reduction in the basal 

columellae in pollen grains of comparatively 

large size. 

In its vegetative or floral characters, Pseudonoseris seems less specialized than Paranephelius, 

but the pollen seems more specialized. It is Paranephelius rather than Pseudonoseris that retains 

some irregularity of spine position and has more 

obvious remnants of basal columellae under the 

spines. 

The presence of slender basal columellae under the spines of Paranephelius pollen conforms 

to the anthemoid pattern of typical Liabeae and 

eliminates doubts about the senecioid pollen patterns or Senecionean relationships of the tribe. 

Nevertheless, the extremely slender form of the 

basal columellae represents a trend toward caws 

development that is interpreted herein as apomorphic within the tribe. The trend toward cavus development here and in some Munnoziinae 

indicates at least two separate instances of reduction in basal columellae in the Liabeae. 

Similarities between the pollen of the Paranepheliinae and Munnoziinae might indicate close 

relationship between the two, but this is not 

supported by other evidence. In fact, on the basis 

of such characters as style form, anther color, 

endothecial cells, and raphids of the achene wall, 

the two subtribes are at opposite extremes in the 

tribe. The surviving slight irregularity of spine 

distribution along the margin of the colpus that 

is evident in Paranephelius (Figure 4 1) supports 

the idea that the regularly disposed spine pattern 

of the Paranepheliinae arose from members of 

the tribe with more irregularly disposed spines 

on the pollen within the immediate relationship, 

and separately from that of the Munnoziinae. 

LIAB1NAE.-Members of the subtribe share 

pale anthers with fringed bases, narrow bases on 

the disk corolla throats, and leaves rarely or 

never in rosettes. The heads are often on short 

peduncles in crowded inflorescences. The style 

branches and the raphids in the achene wall may 

be long or short. The subtribe consists of ten 

genera, Austroliabum (Figures 3, 69-73), Bishopanthus (Figures 54-57), Cacosmia (Figures 4, 58- 

63), Chionopappus (Figures 5, 64-68), Ferreyranthus (Figures 6, 7, 78-82), Liabellum (Figures 

11, 94-98), Liabum (Figures 12-20, 108-123), 

Microliabum (Figures 74-77), Oligactis (Figures 

21-23, 99-107), and Sinclairia (Figures 8-10, 
6 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

83-93), ranging north to central Mexico and the 

Greater Antilles and south to northern Argentina. In all these genera the spines are unevenly 

distributed in small subgroups on the pollen surface (Figures 60, 62-65, 67, 69, 70). As far as 

known, this type of arrangement is unique to the 

Liabeae but apparently plesiomorphic in the 

tribe. The pattern is reflected in cross-sections 

(Figure 68; Skvarla et al., 1977) by the irregular 

thickness of the intercolpar exine and by the 

confluence between the bases of the adjacent 

spines. Size of the pollen grains, in contrast, is 

highly variable in the subtribe (Appendix). Size 

of pollen grains seems to be reflected in the 

subtribe in the size of the columellae, smallest in 

Liabum and Oligactis (Figures 12-23), and largest 

in Sinclairia and Liabellum (Figures 8, 10, 11). 

The two closely related genera with the smallest 

pollen grains, Liabum and Oligactis (25-35 pm), 

are also distinctive in the shorter stouter tips of 

the spines. This is evident in both the Andean 

(Figures 99-107) and West Indian (Figures 108- 

11 1) material. 

The SEMs of broken pollen in Austroliabum, 

Ferreyranthus, Liabum, and Sinclairia, and sectioned grains of Chionopappus (Marticorena and 

Parra, 1974), all indicate a distinct group of large 

basal columellae under the spines. Nevertheless, 

oil immersion studies by Marticorena (Figures 3, 

7, 11) show variations toward a fused hollow 

cylinder in each of these genera. The fusion 

might be the result of poorer resolution or of the 

vertical direction of the observation in the oil 

immersion studies. Still, the pollen in one SEM 

preparation of Microliabum (Figure 77), which is 

closely related to Austroliabum, may be interpreted as an example of a fused hollow cylinder. 

There is no other SEM evidence of the trait in 

the subtribe. 

The pollen form called the umbellatum-type 

by Stix (1 960) has been examined in Liabum 

(Figures 12, 13, 18, 20, 118, 123) using SEM 

preparations. The pollen type differs from the 

more common form in the tribe by the basal 

columellae being somewhat smaller and more 

numerous, and extending into the areas between 

the spines. Although named after Liabum umbellatum (L.) Schultz-Bipontinus of Jamaica, the 

character does not occur in that species. The 

character does occur in species in Hispaniola that 

have sometimes been included in a broader concept of L. umbellatum. Two species from Hispaniola have been examined. In L. subacaule Rydberg the columellae under the spines become 

narrower distally while those between the spines 

are often constricted at the base (Figure 118). 

There are also a few projections from the under 

surface of the internal tectum among the interspinal columellae. In L. barahonense Urban the 

supposed columellae between the spines are seen 

to be mostly projections from the under surface 

of the internal tectum and they are not connected 

to the foot-layer (Figures 122, 123). From the 

evidence, the umbellatum-type pollen in West 

Indian Liabum has columellae of basically different structure in the interspinal areas from those 

under the spines. This particular design of columellae is probably restricted to the genus Liabum. 

The umbellatum-type pollen was also credited to 

Munnozia hastfolia by Stix (1 960, as Liabum), but 

the latter seems to be different in the details of 

its structure. 

Most of the remaining genera of the subtribe 

have massive columellae clustered under the 

spines, but the survey of material under oil immersion by Marticorena has revealed a variant in 

Sinclairia deamii (B.L. Robinson and Bartlett) 

Rydberg having only small columellae (Figure 9). 

Examination with the SEM shows these columellae to be further distinguished by their common 

failure to reach the foot-layer (Figures 91, 92). 

The variant seems to represent an extreme reduction of a type that is rare in the subtribe 

Liabinae, though it does occur commonly in the 

Munnoziinae. As interpreted herein on the basis 

of the exposed unbroken basal columellar surface, the SEM photograph of Microliabum (Figure 

77) represents another example in the subtribe 

of basal columellae that are incompletely fused 

to the foot-layer. 

MUNNOZI1NAE.-The subtribe contains four 

genera, Erato, Philoglossa, Chrysactinium, and 
NUMBER 64 7 

Munnozia that share short style branches, black 

anthers, quadrate raphids in the achene wall, 

somewhat expanded bases in the throats of the 

disk corollas, and a tendency for long-pedunculate heads. Nevertheless, for purposes of discussion of the pollen, the subtribe falls into two 

distinct subgroups consisting of two genera 

apiece. On the basis of pollen alone, the two 

subgroups would not be considered closely related. 

Erato (Figures 24, 124-1 33) and Philoglossa 

(Figures 25, 26, 134-139) both have the irregularly clustered pattern of spines on the pollen 

surface (Figures 124-1 27, 134-1 36) that is characteristic of the larger subtribe Liabinae. Thick 

sections and whole grains examined with oil immersion, and broken grains viewed with the 

SEM, show that Erato further resembles the Liabinae by the large clustered basal columellae 

under each spine (Figures 24, 129, 131, 132, 

133), all firmly attached to the foot layer. The 

presence of such pollen in Erato in the subtribe 

Munnoziinae strongly indicates that the form is 

plesiomorphic in the tribe. 

In Philoglossa the pollen is externally the same 

as Erato; however, oil immersion and SEM studies show that the basal columellae under the 

spines are reduced in size and number. Such 

reduction is independent of the size of the grain 

that is the same size as in Erato. The oil immersion and SEM studies show that basal columellae 

are present (Figures 25, 26, 138, 139), a fact 

demonstrated also by Feuer and Dillon (1982). 

Still, the SEM preparations indicate that the columellae are weakly attached to the foot-layer and 

that a near-caveate condition exists (Figures 138, 

139). The broken grains show a large expanse of 

papillose foot-layer with little evidence of columellar scars. Philoglossa is the only member of 

the tribe characteristically combining such reduction of the basal columellae with an irregular 

disposition of spines on the surface. 

The remaining two genera of the subtribe, 

Chrysactinium (Figures 27, 28, 140-146) and 

Munnozia (Figures 29-40, 147-1 80) have spines 

regularly distributed on the surface of the pollen 

with no merging of spine bases. Internally the 

columellae show a strong tendency for reduction, 

although a number of species (Figures 15 1 - 160) 

including the type of the genus, M. lanceolata, 

show the massive type seen in Erato and the 

Liabinae. Nevertheless, partial or complete columellae are clearly present in all members of the 

two genera that have been examined by light 

microscope or SEM. In this respect the pollen 

differs from that found in the Paranepheliinae. 

As seen in SEMs, however, there is a capacity in 

Munnozia for the most complete separation of 

the columellae from the foot-layer and thus the 

most technically complete caws development in 

the tribe. 

In the Chrysactinium-Munnoziu complex, the 

detailed structure of the columellae and tecta 

shows great variation, and the variations seem to 

correlate well with taxonomic groups within a 

genus. As such, the variations are of more taxonomic value than variations in other parts of 

the tribe. In Chrysactinium, according to oil immersion studies (Figures 27, 28), the basal columellae vary from separate strands to a single 

hollow cylinder, but they are rather small and 

regularly arranged. An SEM view shows that the 

columellae connect with the foot-layer (Figures 

144, 145). The examples seen in the SEM preparation are all fused, with the hollow center 

evident in one view (Figure 146). The form of 

the columellae in the genus is not precisely 

matched in size, symmetry, attachment to the 

foot-layer, or variable dissection in any member 

of Munnozia. 

Munnozia itself has many different subgroups 

that can be distinguished by the internal structure of the pollen walls. The type-species, Munnozia lanceolata Ruiz and Pavon (Robinson and 

Brettell, 1974), and a series of close relatives, M. 

cardenasii, M. foliosa, M. olearioides, M. peruensis, 

and M. rusbyi, all having rather coriaceous leaves, 

appear to have a characteristic cluster of large, 

separate, basal columellae in a ring under each 

spine (Figures 29, 33, 151-155). No members of 

the group have the columellae completely fused 

into a hollow cylinder. In this typical group, the 
SMITHSONIAN CONTRIBUTIONS TO BOTANY 

columellae, as observed in thick section and SEM, 

are attached to the foot-layer. A singularly distinctive species, M. perfoliata, also has columellae 

grouped and unfused under the spines (Figure 

156), but these columellae are less massive and 

are sometimes mixed with smaller columellae 

inside the circle. The even smaller circle of columellae in M. lyrata, as seen with oil immersion 

and SEM, seems to be unique to that distinctive 

species, with the enlarged proximal ends that do 

not reach the foot-layer (Figures 34, 161-165). 

The foot-layer characteristically bears a small peg 

under each spine below the hollow center. 

The dissected pattern of columellae in Munnozia lyrata is in sharp contrast to the fused 

cylinder, which is mostly separated from the footlayer, and seems to be characteristic of the 

subgenus Kastnera (Figures 39,40, 172-180), M. 

senecionidis (Figure 35), and the latter’s relatives, 

M. affinis, M. angusta, M. convencionensis, M. corymbosa, M. glandulosa, M. longifolia, M. oxyphylla, 

M. pinnulosa, M. silphioides (Figure 36), and M. 

venosissima (Figure 38). These fused columellae 

seem usually eccentric, and as seen by SEM, the 

eccentric form in M. senecionidis may be associated with partial eccentric attachment to the footlayer (Figure 166, 167). It is evident, on the basis 

of pollen, that the M. senecionidis group is more 

closely related to the subgenus Kastnera (Figures 

39,40) than to typical Munnozia (Figures 29,33). 

According to Stix (1960), and as seen in this 

study in thick sections examined under the light 

microscope and in some broken grains under 

SEM, Munnozia hastifolia has columellae in the 

interspinal areas of the pollen wall. For this reason, the species was included in the umbellatumtype pollen by Stix (1 960). Such a design seems 

anomalous in the group, because the two most 

closely related species, M. maronii and M. subviridis, have columellae of the Kastnera-M. senecionidis-type according to oil immersion studies (Figure 37). 

One additional distinctive pollen type is found 

in Munnozia jussieui. As observed under oil immersion by Marticorena (Figures 30-32) the columellae under the spines are of two forms, with 

numerous smaller ones surrounding a large central mass. The central mass seems to be solid, but 

on the basis of the outline of many samples, the 

central mass seems to consist of several large 

basal columellae that may be slightly separated. 

It seems unlikely that the species would be the 

only member of the tribe lacking a hollow or 

open space under the center of the spine. SEMs 

of broken grains (Figures 157-160) show that all 

the basal columellae are completely fused at the 

bases to the foot-layer. 

In Munnozia tenera of the subgenus Kastnera, 

there is, as usual, an outer layer of short uniform 

columellae, but there also is a second inner layer 

of similar short uniform columellae between the 

spines and adjacent to the foot-layer (Figures 

172-1 80). Close examination shows that the 

proximal ends of the short columellae in both 

series are narrow and not, or only weakly, attached to the structure below them. Thus, the 

inner layer is not, or only weakly, attached to the 

foot-layer, and more surprisingly the outer layer 

is not, or only weakly, attached to the “internal 

tectum” that caps the inner layer of short columellae. This differs from the structure of the 

“internal tectum” seen in other members of the 

tribe and described by Skvarla et al. (1977:156), 

which is attached to, or gives rise to, the outer 

layer of columellae. The inner layer of columellae is at a level equivalent to that occuppied under 

the spines in most Liabeae by the larger basal 

columellae attached to the foot-layer. In M. tenera, it is notable that the cylindrical structure 

formed by the basal columellae under the spines 

is like the smaller columellae in not being attached to the foot-layer. The character may be 

more widely distributed in the subgenus Kastnera, but it is not found in M. senecionidis, which 

is the closest relative outside of the subgenus that 

has been examined under SEM in this study 

(Figures 166, 167). 
NUMBER 64 

Key to Types of Liabeae Pollen 

9 

1. Pollen with spines arranged in irregular groups on the surface, not evenly aligned along margin 

2. Exine nearly or truly caveate, with only slender basal columellae present under the spines 

2. 

of colpus, with bases of spines variably confluent .................................. .2 

............................................................. Philoglossa 

Exine not caveate, with stout or numerous basal columellae present under the spines ... .3 

Pollen 25-35 pm in diameter; spines short with stout tips. .................... .4 

the spines (umbellata-type of Stix, 1960) ... Liabum barahonense and some other 

With large columellae restricted to area under the spines ................... 

.......................................... Oligactis and most Liabum; 

3. 

4. With small numerous basal columellae under both the spines and areas between 

Liabum species from Hispaniola 

4. 

Pollen mostly 30-50 pm in diameter; spines with narrow tips. ................. .5 

5. Pollen less than 40 pm in diameter. .... Austroliabum, Cacosmia, Chionopappus, 

Erato, Feweyranthus, and Microliabum 

5. Pollen mostly 40-50 pm in diameter .............. Liabellum, Sinclairia, and 

one Ferreyranthus species 

Pollen with spines regularly arranged on surface of intercolpar area and mostly aligned along 

sides of colpus, each spine separated to equal depth on all sides ....................... .6 

6. Pollen 35-45 pm in diameter, with no or only very slender basal columellae under the 

spines ................................................................. 7 

7. Spines regularly aligned along sides of colpi; basal columellae not visible under the 

spines using light microscopy ................................. Pseudonoseris 

7. Spines slightly unaligned along sides of the colpi; a few slender basal columellae present 

under the spines .......................................... Paranephelius 

Pollen 27-45 pm in diameter, with basal columellae forming clusters, rings, or fused 

cylinders under the spines ................................................ .8 

8. Basal columellae under the spines completely or almost completely separated from footlayer at the base .................................................... .9 

9. Pollen 40-42 pm in diameter; basal columellae in a circular unfused cluster under 

each spine. .......................................... Munnoria lyrata 

9. Pollen 27-37 pm in diameter; basal columellae under each spine usually fused into 

asingleunit .................................................... 10 

10. With two layers of short, dense columellae in interspinal areas of tectum 

10. With only one layer of short dense columellae in interspinal area of tectum 

.......................................... Munnoria senecionidis 

Basal columellae under the spines attached at bases to foot-layer ............... 11 

11. Basal columellae under each spine in small single symmetrical series about 

2 pm in diameter ...................................... Chrysactinium 

11. With at least some massive basal columellae under each spine, the clusters mostly 

3-4pmindiameter .............................................. 12 

3. 

1. 

6. 

..................................... Munnoria subgenus Kastnera 

8. 
10 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

12. Basal columellae under the spines of two types, many small peripheral columellae surrounding a central, massive, fused column ................... .............................................. Munnoria jussieui 

12. A simple cluster of mostly large basal columellae under each spine ...... 13 

13. Pollen 28-30 pm in diameter; with basal columellae in both interspinal 

and spinal areas of tectum .................... Munnozia hastifolia 

13. Pollen 37-45 pm in diameter; with basal columellae only under the 

spines .................................................. 14 

14. Cluster under each spine with individual basal columellae often 

2-3 pm in diameter. ..................................... ............... typical Munnozia, M. lanceolata and its relatives 

14. Cluster under each spine with individual columellae not more than 

1.5 pm in diameter ...................... Munnoria pe?foliata 

Palynological Evidence on the Relationships 

of the Liabeae in the Asteraceae 

The pollen structure in the Liabeae reinforces 

recent conclusions regarding the two conflicting 

placements of the group in the family by various 

authors. Cassini (1 828), Robinson and Brettell 

(1 973b), Nordenstam (1977), Jansen and Stuessy 

(1980), and Robinson (1983a) have placed the 

genera of the Liabeae in or near the Vernonieae. 

In contrast, Bentham and Hooker (1873) followed by Hoffmann (1 894) and Cronquist (1 955) 

have placed the broadly interpreted genus Liabum in the Senecioneae. The two tribes involved 

have the distinction of belonging to two different 

subfamilies of the Asteraceae as interpreted by 

Carlquist (1 976) and Robinson (1 977). The Vernonieae share most characters with the Arctoteae, Eremothamneae, Echinopsideae, Cynareae, 

Mutisieae, and Lactuceae in a group best recognized as the subfamily Cichorioideae, while the 

Senecioneae belong with the Eupatorieae, Heliantheae, Inuleae, Calenduleae, Astereae, and 

Anthemideae in a group best recognized as the 

Asteroideae. For comparisons of characters see 

Robinson and Brettell(1973b), Wagenitz (1976), 

Carlquist (1976), and Robinson (1977). As such 

the two tribes with which the Liabeae have been 

placed are not closely related, and there are many 

structural features by which the closest relationships of the tribe can be tested. Results of evidence from floral anatomy have most recently 

indicated that the Liabeae are close to the Vernonieae and other members of the Cichorioideae 

(Robinson and Brettell, 1973b; Nordenstam, 

1977; Robinson, 1983a). Pollen evidence should 

reflect a similar relationship. 

A general trend has been noticed by Robinson 

and Brettell (1 97313) where all pollen surfaces 

that depart from an evenly distributed echinate 

pattern seem to belong to the subfamily Cichorioideae. These variations include some distinctive lophate and oblong psilate types, but also 

include most members of the Vernonieae having 

less obvious sublophate forms in which the spines 

are only arranged around depressed areoles 

rather than restricted to well-defined ridges. 

The pollen in the Liabeae is echinate without 

obvious lophate or psilate variations, but the 

spines of most Liabeae are not evenly distributed 

on the surface of the grains. In all genera of the 

Liabeae except Chrysactinium, Munnozia, Paranephelius, and Pseudonoseris, the spines tend to 

be grouped in small series or crests with somewhat confluent bases. The gaps within the series 

are slightly but distinctly smaller than those between the series. The grouping of spines in these 

Liabeae is unlike the variations seen in any other 

tribe of the Asteraceae, but by its irregularity it 

would seem to be excluded from the subfamily 

Asterioideae, and would fall into the Cichorioideae. 

Internal structure of the pollen wall shows 

more characters that can indicate relationship in 

the Asteraceae. Skvarla et al. (1966, 1977) have 

described features they termed a “cavus,” ‘” internal foramina,” and “internal tecta,” and on the 

basis of these structures they have recognized 

four basic pollen types in the family: helianthoid, 

senecioid, arctotoid, and anthemoid. Each of 
NUMBER 64 11 

these types shows some broad systematic correlations in the family and furnish a useful basis for 

comparison with the Liabeae. 

The Liabeae do not have “internal foramina” 

in the pollen wall and therefore do not fall into 

the helianthoid classifiaction of Skvarla et al. 

(1 977) that is defined by the presence of internal 

foramina with a cavus. The character is almost 

restricted to the subfamily Asteroideae, occurring especially in such tribes as the Eupatorieae 

and Heliantheae, which have never been considered close relatives of the Liabeae. Internal foramina are notably undeveloped in a few of the 

most highly evolved members of the Asteroideae, 

such as most true Senecioneae, the Anthemideae, 

and most Ambrosiinae of the Heliantheae. The 

fact that there are no internal foramina in the 

Liabeae conforms with placement in the Cichorioideae but does not in itself preclude relation 

to some Senecioneae. 

The Liabeae seem to lack a “cavus” in the 

pollen wall in their plesiomorphic form. The lack 

of the cavus provides a further basis for excluding the Liabeae from the helianthoid grouping 

of Skvarla et al. (1977). The cavus is also characteristic of two of the other major pollen wall 

patterns described by Skvarla et al. (1 966, 1977), 

the senecoid, and arctotoid. A caws extends 

through most members of the subfamily Asteroideae including the Senecioneae and is the best 

character after that of spine distribution for excluding the Liabeae from the Senecioneae and 

the Asteroideae. The “lack” of a caws is, in fact, 

the same as the “presence” of basal columellae 

connecting the outer exine to the foot layer in 

the intercolpar area, and this is the principal 

characteristic of the fourth or anthemoid pollen 

wall pattern of Skvarla et al. (1977). 

The distribution of the non-caveate or anthemoid pollen of the type found in the Liabeae is 

of considerable interest, especially since it is the 

type found in most members of the subfamily 

Cichorioideae excluding the Arctoteae. Of the 

various pollen types, it is the one that most nearly 

aligns with the presently accepted subfamily classification of the Asteraceae. There are important 

exceptions in the distribution of the anthemoid 

pollen type, however, including its presence in 

the Anthemideae of the Asteroideae and its absence in various Cichorioideae such as the Arcto teae. 

In one way, it is unfortunate that the noncaveate pollen type has been called anthemoid 

by Skvarla et al. (1977), because the type is much 

more characteristic of members of the subfamily 

Cichorioideae. The Anthemideae is the one major group in the Asteroideae in which the anthemoid pattern occurs. It should be emphasized 

further that the Anthemideae is not primitive 

within the subfamily Asteroideae and is not close 

to the Cichorioideae (Robinson, 198 1). Caveate 

pollen, the type predominant in the Asteroideae, 

is found even in what seems to be a primitive 

element of the Anthemideae, in Ursinia (Robinson and Brettell, 1973b; Skvarla et al., 1977). 

One could conclude from foregoing evidence 

that the non-caveate pollen of the Anthemideae 

represents a development of basal columellae in 

the intercolpar area from ancestral caveate types 

that did not have such columellae. The anthemoid pollen pattern thus would have at least two 

separate origins in the Asteraceae. 

In the subfamily Cichorioideae, where the anthemoid pollen pattern predominates, there are 

some caveate forms such as those within the 

arctotoid pattern, some Lactuceae having an interrupted cavus, and a few Mutisieae such as 

Dasyphyllum (which is caveate along the colpar 

margins). Bolick (1978) cites caveate pollen in 

Ethulia, Hoplophyllum, and Corymbium of the Vernonieae and Centaurea in the Cynareae, and indicates that the Arctoteae is approximately 

equally divided between caveate and non-caveate 

types. To these examples may be added those 

now known from the Liabeae. In these cases, the 

caveate pollen seems to be derived from noncaveate ancestors, and it has evidently been derived more than once. Apparently, the two 

subfamilies of the Asteraceae have the two different types of pollen as basic, but each type, 

caveate and non-caveate, seems able to give rise 

to the other. The fact that the anthemoid pollen 

wall type occurs in the one specialized tribe of 

the Asteroideae, in the absence of any other 

supporting characters, does not particularly favor the idea of asterioid relationships for the 

Liabeae. 

In addition to the cavus, the presence of an 
12 

“internal tectum” is used as a basis for differentiating the arctotoid pollen pattern within the 

subfamily Cichorioideae (Skvarla et al., 1977). 

This involves the most characteristic form of 

internal tecta that almost equally divides the region of solid columellae. The arctotoid pollen is 

also claimed in Eremothamnus (Liens, 1970) of 

the tribe Eremothamneae (Robinson and Brettell, 1973c), but this should be checked. Illustrations of Skvarla et al. (1977, pl. 24: fig. F, H) 

suggest an arctotoid pattern in some Lactuceae 

such as Rafinesquia and Lapsana. Internal tecta 

of this type occur in pollen of other Cichorioideae, seeming to be most highly developed in the 

tribes Cynareae and Mutiseae. The latter two 

tribes show relationship to the Arctoteae in the 

form of the style, but they differ and are excluded from the Brctotoid type by Skvarla et al. 

(1977) because of the lack of a “cavus.” The 

Liabeae and Vernonieae lack the elaborate internal tecta of the arctotoid type or the Cynareae 

and Mutisieae type, but enlargements at the 

proximal ends of the short outer columellae form 

a layer at the lower limit of the outer columellae 

that is also referred to as an internal tectum by 

Skvarla et al. (1 977: 156). The latter layer is often 

prominent in the areas between the spines where 

there may be no basal columellae, and the structure thus appears very different from the structure referred to by the same name in the Arctoteae and is structurally equivalent to the layer 

below the internal tectum of the Arctoteae. 

Nevertheless, the two structures are formed in 

the same manner and no other better term seems 

to have been provided as yet. 

Within the anthemoid pollen pattern Skvarla 

et al. (1977) have described a modified pattern 

they termed “Liaboid.” It is distinguished by the 

basal columellae being much broader (Figures 

68, 89, 90, 113, 114, 120) than those noted for 

the typical anthemoid pollen pattern. The pattern is cited from ”Liabum,” Cacosmia, and the 

Vernonieae and is cited elsewhere by Skvarla et 

al. only in Scolymus of the Cichorioid Lactuceae 

on the basis of TEM studies by Tomb (1975). 

Observation during the present study show that 

Gundelia, of the prospective Cichorioid tribe 

SMITHSONIAN CONTRIBUTIONS TO BOTANY 

Gundelieae, has similar massive columellar structure under the spines, but these do not reach the 

foot-layer. The limitation of the liaboid pollen 

pattern to members of the subfamily Cichorioideae seems to provide confirmation of the placement of the Liabeae in that subfamily. The distribution of the liaboid pattern within the 

subfamily seems to parallel evidence of stylar 

form in suggesting closest relationship of the 

Liabeae to the Vernonieae and Lactuceae, especially the former. 

Of the tribes that appear to be most closely 

related to the Liabeae, the pollen of the Lactuceae can be distinguished by various characters 

such as lophate surfaces or more complex internal tecta. In the cases where Tomb (1975) and 

Skvarla et al. (1977) show internal structure 

more like that of the Vernonieae and Liabeae 

the structure is under the ridges of specialized 

echinolophate grains. Among the non-lophate 

members of the Lactuceae, there seem to be 

cavus-like areas periodically interrupted by columellae reaching the foot layer, a form seen in 

comparatively few derived members of the Liabeae. 

The Vernonieae is regarded here as the probable closest relatives of the Liabeae, and the two 

tribes have been combined by some recent authors (Nash, 1976; Turner and Powell, 1977; 

Jansen and Stuessy, 1980). The two groups were 

placed adjacent to each other in the pollen study 

by Stix (1 960) because of their similar wall structure. It is of interest, therefore, that the pollen 

of the Vernonieae and Liabeae is in fact distinguishable, and almost without exception the 

tribes show totally different trends in the details 

of their anthemoid structure. 

Surface features alone could be used to distinguish most Vernonieae from the Liabeae. Only 

the recently described Pseudostqftia (Robinson, 

1979) in the Vernonieae appears to have no trace 

of a lophate surface pattern (Figures 181, 182). 

The remainder of the Vernonieae have various 

degrees of areole formation (Keeley and Jones, 

1979). Most members of the tribe have a form 

termed the Lychnophora-type by Stix (1 960), 

where the tectum is continuous between the 
NUMBER 64 13 

colpi, but where slight areolae are present (Figures 190-192). The polygonal areolae are more 

obvious under the light microscope where the 

columellae are visible to accentuate the pattern. 

The ridges in some genera, such as Bishopalea, 

are more obvious with the tectum still continuous 

across the areolae (Figures 200-202). More 

strongly lophate pollen forms in the tribe, such 

as Heterocypsela, have regularly arranged ridges 

with a restricted tectum (Figures 203, 204). The 

extreme form of lophate grain in the tribe has 

little or no tectum and has the columellae visible 

from a lateral view of the ridges. This is seen in 

Elephantopus and its relatives, as well as in some 

Paleo-tropical elements of the subtribe Vernoniinae, such as Cyanthillium and Phyllocephalum 

(Figures 205-208; Skvarla et al., 1977; Keeley 

and Jones, 1979; Jones, 1981; Kirkman, 1981). 

The trend toward a lophate pollen surface seems 

basic to the Vernonieae, but is absent from the 

Liabeae. 

Size of pollen grains alone offers some distinction between the tribes. Pollen of the Liabeae is 

mostly 25-40 pm in diameter with some up to 

50 pm. The pollen of the Vernonieae is usually 

40 Fm in diameter or more, and it is often over 

50 pm. 

Based on a limited sample, Skvarla et al. (1977) 

indicated some differences in the detailed internal structure of the liaboid pattern in the Vernonieae and Liabeae. On page 156 they stated 

the following. 

In Liabum the lateral branches of the distal ends of the 

columellae (sic basal columellae) form a uniform internal 

tectum. From this internal tectum a short, uniform set 

of columellae arise which in turn are capped by a perforate tectum. Commonly, the tectum areas between the 

large columellae show gentle undulations. In the Vernonieae the major distinction from Liabum is that the 

lateral branches are frequently (but not always) a thick 

network of solid rods or tubules with complex anastomosing patterns. 

The distinction is evident to some degree in the 

examples of broken pollen grains seen in the 

present study. Many Vernonieae, including Blanchetia (Figures 197-199) and Pseudostfltia (Figures 184-189), show a branching pattern that 

completely fills the inter-spinal areas, while Vernonia fuertesii shows that even an unsupported 

internal tectum in the tribe does not bear a layer 

of regular short columellae as in the Liabeae 

(Figures 193-195). 

The most important difference in the internal 

structure of the pollen wall was first noted by 

Stix (1 960) and has been confirmed in the present study by the light microscope and SEM views 

of broken grains. In the Vernonieae with welldeveloped spines, each spine seems to have a 

single solid basal columella centered under the 

spine, which is continuous with the apex of the 

spine (Figures 193-195, 197-199). The only 

exception seems to be Pseudostgtia with basal 

columellae distributed more randomly throughout the spinal and interspinal areas with branches 

of more than one basal columella contributing to 

each spine (Figures 184-1 89). Though unlike 

any other Vernonieae, Pseudostqfiia is not like 

any Liabeae. In the Liabeae, the basal columellae 

are either grouped under the spines (Figures 12- 

16, 29, 46, 89, 163-165) or are coalesced into a 

single hollow cylinder (Figures 40, 145, 146, 

166, 167, 174-1 76). The coalesced form, which 

seems continuous with the apex of the spine, 

resembles the single columella of the Vernonieae 

except for the hollow center. Single solid columellae centered under the spines do not seem to 

occur in the Liabeae except possibly in a specialized form seen in Munnozia jussieui. In the latter 

case the oil immersion studies (Figures 30-32) 

suggest a characteristically higher degree of coalescence than any other member of the tribe, 

but the SEM seems to show narrow separation 

(Figures 157-160) that was overlooked at the 

lower magnification. The difference between the 

basal columellae in the two tribes is regarded 

herein as fundamental, and observation of the 

range of variation in the Liabeae indicates that 

the cluster of separate columellae is plesiomorphic in the tribe. The form most resembling 

the Vernonieae is apparently an isolated derivative. 
Appendix 

List of specimens examined palynologically in the study with figures in 

which they appear and with measurements (in pm) of pollen in glycerine 

mounts and Hoyer’s (H) mounts. 

LIABEAE 

Austroliabum 

A. candidum (Grisebach) H. Robinson & Brettell 

A. eremophyllum (Cabrera) H. Robinson & Brettell 

A. polymnioides (R.E. Fries) H. Robinson & Brettell 

0. Kuntze s/n, Argentina, Figures 69-72 

Venturi 6580, Argentina 

Pflanz 4 102, Bolivia, Figure 3 

Venturi 5280, Argentina 

Wall & Sparre 622, Argentina, Figure 73 

Bishopanthus 

B. soliceps H. Robinson 

King 9280, Peru, Figures 54-57 

Cacosmia 

C. rugosa Humboldt, Bonpland, & Kunth 

Barclay 8e Juajibioy 8305, Ecuador 

Barclay 8e Juajibioy 8339, Ecuador 

Wurdack 1016, Peru, Figures 4, 58-61 

Chionopappus 

C. benthamii (Bentham) Blake 

Asplund 10996, Peru, Figures 5, 64-67 

Ferreyra 7663, Peru 

Ferreyra 1 1078, Peru 

Macbride 8e Featherstone 162, Peru 

Pennell 14480, Peru 

Chrysactinium 

C. acaule (Humboldt, Bonpland, & Kunth) Weddell 

Camp E-2231, Ecuador, Figures 27, 144-1 46 

Haught 3261, Ecuador, Figures 140-143 

C. amphothrix (Blake) H. Robinson 8c Brettell 

Ferreyra 948 1, Peru 

C. caulescens (Hieronymus) H. Robinson & Brettell 

Jameson s/n, Ecuador, type of Liabum bicolor Blake 

Wurdack 764, Peru 

Wurdack 11 13, Peru, Figure 28 

Fagerlind 8e Wibon 1563, Ecuador 

Haught 326 1, Ecuador 

Holmgren 560, Ecuador 

E. polymnioides DeCandolle 

C. hieracioides (Humboldt, Bonpland, & Kunth) H. Robinson & Brettell 

C. longzradiatum (Hieronymus) H. Robinson & Brettell 

Erato 

Ferreyra 8 168, Peru 

H 38-40 

H 33-37 

35-37 

33-37 

30(-35), H 35 

H 27-30 

H 27 

H 40 

H 35-37 

30 

H 40 

30-32 

30-34 

H 35-37 

H 35-37 

14 
NUMBER 64 15 

King 655 1, Ecuador, Figures 124- 130 

Lug0 24 1, Ecuador, Figure 24 

Barclay & Juajibioy 83 14, Ecuador, Figures 13 1-1 34 

Rose & Rose 22413, Ecuador 

E. stenolepis (Blake) H. Robinson 

Hutchison 11 90, Peru 

E. vulcanica (Klatt) H. Robinson 

Cuatrecasas 256, Colombia 

Steyermark 5598 1, Venezuela, type Liabum insigne Badillo 

v. Sneidern A.504, Colombia 

E. Sodiroi (Hieronymus) H. Robinson 

Ferrepranthus 

F. excelsus (Poeppig & Endlicher) H. Robinson & Brettell 

Hutchison & Wright 3854, Peru, Figure 6 

F. rugosus (Ferreyra) H. Robinson & Brettell 

Wurdack 469, Peru, type, Figures 82, 83 

F. vaginans (Muschler) H. Robinson & Brettell 

Vargas 11050, Peru 

F. verbascfolius (Humboldt, Bonpland, & Kunth) H. Robinson & Brettell 

Camp E-3906, Ecuador 

Hutchison & Wright 5133, Peru, Figures 78-81 

Lehmann 7958, Ecuador, type of Liabum salvifolium Hieronymus 

F. vernonioides (Muschler) H. Robinson & Brettell 

Hutchison &c Wright 5176, Peru, Figure 7 

Kunkel 975, Peru 

Liabellum 

L. angustissimum (A. Gray) Rydberg 

Pringle 2501, Mexico 

Rose 3406, Mexico 

Pringle 4398, Mexico 

Barnes 8c Land s/n, Mexico, Figures 94-98 

Pringle 2328, Mexico 

L. cervinum (B.L. Robinson) Rydberg 

L. palmeri (A. Gray) Rydberg 

Liabum 

L. acuminatum Rusby 

Williams 1605, Bolivia 

Woytkowski 34392, Peru 

Klug 3 183, Peru 

Howard 12087, Dominican Republic, Figures 12, 121-123 

Jimenez 3284, Dominican Republic, Figures 13, 108-1 11 

Barclay & Juajibioy 8316, Ecuador, type 

Skutch 4052, Costa Rica, Figure 14 

Skutch 4730, Costa Rica 

Standley & Valerio 5 186 1, Costa Rica 

L. crispum Schultz-Bipontinus 

Ekman 15643, Cuba 

L. cubense Schultz-Bipontinus 

Ekman 1542, Cuba 

L. domingense Rydberg 

L. amplexicaule Poeppig & Endlicher 

L. barahonense Urban 

L. barclapae H. Robinson 

L. bourgeaui Hieronymus 

35, 36 

35-40, H 35-37 

36-38 

32-34 

29-35, H 35-37 

H 35 

30-34 

39-41 

H 43 

H 50 

50-52 

47-50 

28-31 

30-34 

34,35 

H 34-36 

34,35 

27-32 
SMITHSONIAN CONTRIBUTIONS TO BOTANY 

Valeur 994, Dominican Republic 

Camp E-3 196, Ecuador 

Fagerlind & Wibon 290, Ecuador, Figure 15 

Haught 3446, Peru 

Smith 2012, Colombia, type, Figure 16 

Hutchison, Wright & Straw 5948, Peru 

Camp E-3 196, Ecuador 

Hutchison & v. Bismark 6337, Peru 

Camp E-3123, Ecuador 

Rose, Pachano & Rose 23232, Ecuador, type of L. amplexans Blake 

Heilborn 534, Ecuador, Figures 11 2-1 17 

Lehmann 4896, Ecuador, type of L. lehmannii Hieronymus 

Sydow 484, Ecuador 

Dryander 391 72, Colombia 

Humbert et al. 25795, Colombia 

L. nigro-pilosum Hieronymus 

Mexia 6686, Ecuador 

L. nudicaule H. Robinson 

Hutchison 1191, Peru 

L. oblanceolatum Urban & Ekman 

Loigier 15402, Dominican Republic 

L. ovatfolium Urban 

Ekman H-12578, Dominican Republic, type 

L. polycephalum Urban 

Ekman H-5346, Haiti 

L. poiteaui (Cassini) Urban 

Ekman H-3713, Haiti, Figure 17 

L. sandemanii H. Robinson 

Sandeman 4420, Peru, type 

L. selleanum Urban 

Ekman H-1548, Haiti, type, Figure 18 

L. solidagineum (Humboldt, Bonpland, & Kunth) Lessing 

Humbert 31002, Peru 

Hutchison 1129, Peru 

Macbride & Featherstone 1519, Peru 

Ekman H-1870, Haiti, Figure 19 

Valeur 56, Dominican Republic, Figures 20, 118-120 

Maxon 277, Jamaica 

Maxon 8575, Jamaica 

L. vargasii H. Robinson 

Vargas 10182, Peru, type 

L. wrightii Grisebach 

Ekman 506 1, Cuba 

Ekman 14745, Cuba 

L. wurdackii Ferreyra 

L. eggersii Hieronymus 

L. falcatum Rusby 

L. ferreyrii H. Robinson 

L. jloribundum Lessing 

L. grandzjlorum (Humbolt, Bonpland, & Kunth) Lessing 

L. igniarium (Humboldt, Bonpland, & Kunth) Lessing 

L. kingii H. Robinson 

L. melastomoides (Humboldt, Bonpland, & Kunth) Lessing 

L. subacaule Rydberg 

L. umbellatum (L.) Schultz-Bipontinus 

27-30 

H 30-32 

H 27 

24-46 

32-35 

27-30 

H 30-32 

25-27 

33-35 

33-35 

30-33 

H 35 

32-35 

H 25-27 

32, 33 

32-35 

30-32 

30-32 

32-35 

3 1-34 
NUMBER 64 

Hutchison & Wright 5850, Peru 

Microliabum 

M. humile (Cabrera) Cabrera 

Okada 5953, Argentina, Figures 74-77 

Munnozia subgenus Munnozia 

M. affinis (Blake) H. Robinson & Brettell 

Macbride 4337, Peru, type 

M. angusta (Blake) H. Robinson & Brettell 

Mexia 8152, Peru 

M. canarensis (Cuatrecasas) H. Robinson & Brettell 

Camp E-2836, Ecuador, type 

iM. cardenasii (Cabrera) H. Robinson & Brettell 

Herzog 2 149, Bolivia 

M. conuencionensis (Cuatrecasas) H. Robinson & Brettell 

Vargas 4446, Peru, type 

h4. corymbosa Ruiz & Pavon 

ex herb Pavon, possible type 

M. foliosa Rusby 

Bang 1195, Bolivia, type 

Bang 158 1, Bolivia, type of M. chrysanthemoides Rusby, Figure 29 

Vargas 2761, Peru, type of Liabum herrerae Cabrera 

Metcalf 30754, Peru 

M. gigantea (Rusby) Rusby 

Bang 2379, Bolivia, type 

M. glandulosa (0. Kuntze) Rusby 

Cardenas 683, Bolivia 

M. hastifolia (Poeppig & Endlicher) H. Robinson & Brettell 

Uribe 3149, Colombia 

Hutchison 1191, Peru 

Lehmann 8450, Colombia, type of Liabum nonoense var. microcephalum 

Triana 1145, Colombia, Figures 30, 157-160 

v. Sneidern 1974, Colombia, Figure 31 

Humbert 30952, Peru, Figure 33 

Hutchison, Wright & Straw 5928, Peru, Figures 147-155 

M. longziolia Rusby 

Buchtien 3079, Bolivia, type 

M. lyrata (A. Gray) H. Robinson & Brettell 

Ferreyra 6968, Peru, Figures 34, 161-165 

iM. maronii (Andre) H. Robinson 

Mandon 240, Bolivia 

M. olearioides (Muschler) H. Robinson & Brettell 

Weberbauer 4417, Peru, type frag. 

M. oxyphylla (Cuatrecasas) H. Robinson & Brettell 

Woytkowski 34165, Peru, type 

iI4. perfoliata (Blake) H. Robinson & Brettell 

King & Guevara 6 184, Colombia, Figure 156 

M. peruensis (Cuatrecasas) H. Robinson & Brettell 

Woytkowski 34275, Peru, type 

M. pinnulosa (0. Kuntze) H. Robinson & Brettell 

Holway 6 16, Bolivia 

M. rusbyi (Britton) Rusby 

Cardenas 6278, Bolivia 

M. jussieui (Cassini) H. Robinson & Brettell 

Hieronymus, Figure 32 

M. lanceolata Ruiz & Pavon 

17 

31,32 

H 35 

H 37-40 

33-35, H 35 

42-45 

H 37 

H 42-45 

H 43-45 

28-30 

35 

37 

H 40-42 

30-35 

H 40-44 

H 38-40 

H 45 
18 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

M. senecionidis Benth 

Garcia-Barrigd 125 18, Colombia 

Cardenas 3277, Bolivia 

Haught 3295, Ecuador, Figure 35 

King & Almeda 7842, Ecuador, Figures 166, 167 

Pennell 769, Colombia, type of M. attenuata Rusby 

Schneider 7, Colombia 

v. Sneidern 2778, Colombia 

Swingle 183, Peru, Figure 36 

Cook & Gilbert 1365, Peru, type, Figure 37 

Macbride 43 12, Peru, type of Liabum pulchrum Blake, Figure 38 

M. silphioides (Poeppig & Endlicher) H. Robinson & Brettell 

M. subuiride (Blake) H. Robinson & Brettell 

M. venosissima Ruiz & Pavon 

Munnozia subgenus Kastnera 

Asplund 9586, Ecuador 

Hutchison & Wright 5084, Peru, Figures 39, 168-171 

Ferreyra 6968, Peru 

Haught 3294, Ecuador 

M. niuea (Hieronymus) H. Robinson & Brettell 

Killip & Varela 34590, Colombia, Figure 40 

Weberbauer 6022 

Barclay 8c Juajibioy 5760 

Killip 9775, Colombia, Figures 172-180 

M. acostae Chung 

M. annua (Muschler) H. Robinson & Brettell 

M. liaboides (Lessing) H. Robinson 

M. tenera (Schultz-Bipontinus) H. Robinson & Brettell 

Oligactis subgenus Oligactis 

0. lat$olia (Hieronymus) H. Robinson & Brettell 

0. sessilz/7ora (Humboldt, Bonpland, & Kunth) H. Robinson & Brettell 

v. Sneidern 354, Colombia 

Allart 370, Venezuela, type of Liabum touarense Badillo 

Ariste-Joseph A-237, Colombia 

Cuatrecasas 12855, Colombia, Figures 99-1 03 

Gehringer 427, Venezuela, type of Liabum meridense Badillo 

Perez-Arbelaez & Cuatrecasas 8 164, Colombia, type of Liabum boyacense 

Smith 20 13, Colombia, type of Liabum biattenuatum Rusby, Figure 2 1 

0. uolubilis (Huniboldt, Bonpland, & Kunth) Cassini 

Cuatrecasas 

Oligactis subgenus Andromachiopsis 

0. coriacea (Hieronymus) H. Robinson & Brettell 

Camp E-4190, Ecuador, Figure 22 

Mexia 7476, Ecuador 

Lehmann 4897, Ecuador, type 

Townsend A-193, Peru 

Cuatrecasas 11949, Colombia, type 

Mexia 7697, Ecuador, Figures 104-107 

Sydow 605, Ecuador, Figure 23 

0. ecuadoriensis (Hieronymus) H. Robinson & Brettell 

0. ochracea (Cuatrecasas) H. Robinson & Brettell 

0. pastoensis (Cuatrecasas) H. Robinson & Brettell 

0. pichinchensis (Hieronymus) H. Robinson & Brettell 

27-30 

30-32 

H 34-37 

H 35-37 

H 33-35 

H 35-37 

27-30 

25-27 

33-37 

H 33-35 

H 33-35 

30-32, H 35 

27-30 
NUMBER 64 19 

Paranephelius 

P. asperifolius (Muschler) H. Robinson & Brettell 

Fiebrig 3538, Bolivia, type 

P. bullatus A. Gray ex Weddell 

Macbride & Featherstone 1656, Peru 

Macbride & Featherstone 2 13 1, Peru 

Wurdack 760, Peru, Figure 1 

Wurdack 1240, Peru, Figure 48 

Pennell 13363, Peru 

Cerrate 1367, Peru, Figures 45-47 

Ferreyra 5580, Peru, Figures 41-44 

Herzog 1840, Bolivia, Figure 2 

P. jelskii (Hieronymus) H. Robinson & Brettell 

P. ovatus Weddell 

P. unzj7orus Poeppig & Endlicher 

Philoglossa 

P. mimuloides (Hieronymus) H. Robinson & Cuatrecasas 

Dodson & Thien 1467, Ecuador 

Firmin 52, Ecuador 

King 6633, Ecuador 

Bristol 348, Colombia, type, Figure 25 

Asplund 13735, Peru, Figures 138, 139 

Ferreyra 4022, Peru, Figures 26, 134-137 

P. pterocarpa Sandwith 

Soukup 4219, Peru 

P. mimuloides forma sapida (Bristol) H. Robinson & Cuatrecasas 

P. peruviana DeCandolle 

Pseudonoseris 

P. discolor (Muschler) H. Robinson & Brettell 

P. striata (Cuatrecasas) H. Robinson & Brettell 

Metcalf 3053 1, Peru, Figures 50-53 

Weberbauer 7107, Peru, type 

Sinclairia 

S. andrieuxii (DeCandolle) H. Robinson & Brettell 

Cronquist & Sousa 10457, Mexico 

S. andromachioides (Lessing) Rydberg 

Pringle 5905, Mexico 

Purpus 2942, Mexico 

Pittier 1886, Guatemala, type 

King 8c Soderstrom 4988, Mexico 

Palmer 245, Mexico 

S. deamii (B.L. Robinson & Bartlett) Rydberg 

Edwards 568, Honduras, Figures 9, 91-93 

Standley 19695, El Salvador, Figures 10, 88-90 

Botteri 1177, Mexico 

Bartlett 12602, Guatemala, type 

Molina et al. 17514, Costa Rica 

Ton 780, Mexico, Figures 84-87 

S. brachypus Rydberg 

S. caducifolia (B.L. Robinson & Bartlett) Rydberg 

S. deppeana (Lessing) Rydberg 

S. dimidia (Blake) H. Robinson & Brettell 

S. discolor Hooker & Arnott 

38-40 

35-37 

47-50 

36-40 

42-45 

H 35 

H 35-37 

H 30 

H 37-40 

H 38-40 

H 50-52 

H 37-42 

35 

H 37-42 

37. H 42-45 

37 
20 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

Williams & Molina 13728, Honduras 

S. glabra (Hemsley) Rydberg 

Hartman 120, El Salvador 

Mexia 8797, Mexico 

S. hypochlora (Blake) Rydberg 

Skutch 203 1, Guatemala 

S. klattii (B.L. Robinson & Greenmarl) H. Robinson eC Brettell 

Conzatti 1751, Mexico, Figure 8 

Pringle 6059, Mexico 

S. moorei (H. Robinson & Brettell) H. Robinson & Brettell 

Moore 5518, Mexico, type 

S. pittieri Rydberg 

Pittier 9093, Costa Rica, type 

S. platylepis (Schultz-Bipontinus ex Klatt) Rydberg 

Schipp S-738, Belize 

S. polyantha (Klatt) Rydberg 

Allen 2886, Panama 

Skutch 3488, Costa Rica 

Standley 42513, Costa Rica 

S. pringlei (B.L. Robinson & Bartlett) H. Robinson & Brettell 

Pringle 62 14, Mexico 

S. sericolepis (Hemsley) Rydberg 

Bourgeau 21 77, Mexico, type 

S. vagans (Blake) H. Robinson & Brettell 

Skutch 191 3, Guatemala, type 

VERNONIEAE 

Bishopalea 

B. erecta H. Robinson 

King & Bishop 8729, Brazil, Figures 200-202 

Blanchetia 

B. heterotricha DeCandolle 

Schery 736a, Brazil, Figures 196-199 

Heterocypsela 

H. andersonii H. Robinson 

Anderson et al. 9223, Brazil, type, Figures 203, 204 

Phyllocephalum 

P. scabridum (DeCandolle) Kirkman 

Stocks 127, India, Figures 205-208 

Pseudostifftia 

P. kingii H. Robinson 

King et al. 8145, Brazil, Figures 181-189 

Vernonia 

V. fuertesii (Urban) H. Robinson 

Ekman H-735 1, Haiti, type of Vernonia barkeri Ekman ex Urban, Figures 

190- 195 

35-37 

42-45 

48-50 

H 53-55 

33-37 

38-40 

H 37-40 

42, H 50-52 
Literature Cited 

Bentham, G., and J.D. Hooker 

1873. Ordo 88: Compositae. In Genera Plantarum, 

2( 1): 163-533, 536, 537. 

The Genus Chionopappus of Bentham (Asteraceae). Journal of the Washington Academy of Sciences, 

Blake, S.F. 

193.5. 

25( 1 1):488-493. 

Bolick, M.R. 

1978. Taxonomic, Evolutionary, and Functional Considerations of Compositae Pollen Ultrastructure and 

Sculpture. Plant Systematics and Evolution, 

1301209-2 18. 

Cabrera, A.L. 

1954. Compuestas sudamericanas nuevas o criticas, 11. 

Notas del Museo, llniversidad Nacional de Eva PeTon, 17(84):71-80. 

Tribal Interrelationships and Phylogeny of the 

Asteraceae. Aliso, 8(4):465-492. 

Vernoniees. In G. Cuvier, editor, Dictionnaire des 

Sciences Naturelles, 57:338-347. Paris [Reprinted 

1975 in R.M. King and H.W. Dawson, editors, 

Cassini on Compositae, 3: 180 1-1 8 1 1. New York: 

Oriole Editions.] 

Phylogeny and Taxonomy of the Compositae. The 

American Midland Naturalist, 53(2):478-5 1 1. 

Pollen Morphology and Plant Taxonomy: Angiosperms. xii + 553 pages. Stockholm: Almquist and 

Wiksell. 

Feuer, S.M., and M.O. Dillon 

Pollen Morphology and Ultrastructure of the Liabeae (Asteraceae). Botanical Society of America Miscellaneous Publication, 162:93. 

Compositae. In A. Engler and K. Prantl, editors, 

Die naturlichen P’anzenfamilien, 4(5):87-387. 

Leipzig. 

Chromosome Counts of Compositae from Latin 

America. American Journal of Botany, 67(4):589- 

594. 

Synoptic Classification and Pollen Morphology of 

Vernonia (Compositae; Vernonieae) in the Old 

World. Rhodora, 8 3( 8 3 3): 59-7 5. 

Carlquist, S. 

1976. 

Cassini, H. 

1828. 

Croiiquist, A. 

1955. 

Erdtinan, G. 

1966. 

1982. 

Hoffinann, 0. 

1894. 

Jansen, R.K., and T.F. Stuessy 

1980. 

Jones, S.B. 

198 1. 

Keeley, S.C., and S.B. Jones 

1979. Distribution of Pollen Types in Vernonia (Vernonieae: Compositae). Systematic Botany, 4(3): 195- 

202. 

Kirknian, L.K. 

Taxonomic Revision of Centratherum and Phyllocephalum (Compositae: Vernonieae). Rhodora, 

198 1, 

83(833): 1-24. 

Liens, P. 

1970. Die Pollenkorner und Verwandtschaftsbeziehungen der Gattung Eremothamnus (Asteraceae). Mitteilungen der Botanischen Staatssammlung, Munchen, 7:369-376. 

Morfologia de 10s granos de polen y posicion 

sisteniatica de Anisochaeta DC., Chionopappus 

Benth., Fedea Urb. y Gochnatia glomerijlora Gray 

(Compositae). Boletin Sociedad Biologia de Concepcion, 47:187-197. 

Tribe I: Vernonieae. In D.L. Nash and L.O. Williams, Compositae. Flora of Guatemala, 24( 12):4- 

32. 

Senecioneae and Liabeae: Systematic Review. In 

Heywood, Harborne, and Turner, editors, The 

Biology and Chemistry of the Compositae, pages 799- 

830. London and New York: Academic Press. 

An Analysis of the Characters and Relationships 

of the Tribes Eupatorieae and Vernonieae (Asteraceae). Systematic Botany, 2(3): 199-208. 

1978. 190 (2), Compositae-Liabeae, Flora of Ecuador, 

1979. Two New Genera of Vernonieae (Asteraceae) 

from Brasil, Heterocypsela and Pseudost$tza. Phytologia, 44(7):442-450. 

A Revision of the Tribal and Subtribal Limits of 

the Heliantheae (Asteraceae). Smithsonian Contributions to Botany, 5 1 : 102 pages. 

1983a. A Generic Review of the Tribe Liabeae (Asteraceae). Smithsonian Contributions to Botany, 54: 69 

pages. 

1983b. Studies in the Liabeae (Asteraceae), XVI: New 

Taxa from Peru. Phytologia, 54(1):62-65. 

1973a. Tribal Revisions in the Asteraceae, 111: A New 

Tribe, Liabeae. Phytologia, 25(6):404-407. 

Marticorena, C., and 0. Parra 

1974. 

Nash, D.L. 

1976. 

Nordenstam, B. 

1977. 

Robinson, H. 

1977. 

8: 1-62. 

1981. 

Robinson, H., and R.D. Brettell 

21 
22 

1973b. Tribal Revisions in the Asteraceae, VIII: A New 

Tribe, Ursinieae. Phytologia, 26(2):76-85. 

1973c. Tribal Revisions in the Asteraceae, XI: A New 

Tribe, Eremothamneae. Phytologia, 26(3): 163- 

166. 

Studies in the Liabeae (Asteraceae), 11: Preliminary Survey of the Genera. Phytologia, 28( 1):43- 

63. 

Chromosome Numbers in Compositae, XV: Liabeae. Annals of the Missouri Botanical Garden, 

72:469-479. 

1974. 

Robinson, H., A.M. Powell, R.M. King, and J.F. Weedin 

1985. 

Rydberg, P.A. 

1927. (Carduales) Carduaceae, Liabeae, Neurolaeneae, 

Senecioneae (pars). North American Flora, 

34(4):289-360. 

Sandwith, N.Y. 

1956. Contributions to the Flora of Tropical America, 

LXI: Notes on Philoglossa. Kew Bulletin, 

1956:289-293. 

Skvarla, J.J., and B.L. Turner 

1966. Systematic Implications from Electron Microscopic Studies of Compositae Pollen: A Review. 

SMITHSONIAN CONTRIBUTIONS TO BOTANY 

Annals of the Missouri Botanical Garden, 53:200- 

256. 

Pollen Morphology in the Compositaeand in Morphologically Related Families. In Heywood, Harborne, and Turner, editors, The Biology and Chemistry of the Compositae, pages 141-248. London 

and New York: Academic Press. 

Skvarla, J.J., B.L. Turner, V.C. Patel, and A.S. Tomb 

1977. 

Stix, E. 

1960. Pollenmorphologische untersuchungen an Compositen. Grana Palynologica, 2(2):4 1-1 04, plates 

1-21. 

Pollen Morphology in Tribe Lactuceae (Compositae). Grana, 15( 1/3):79-89. 

Helenieae: Systematic Review. In Heywood, Harborne, and Turner, editors, The Biology and Chemistry of the Compositae, pages 699-737. London 

and New York: Academic Press. 

Systematics and Phylogeny of the Compositae. 

Plant Systematics and Evolution, 125:29-46. 

Tomb, A.S. 

1975. 

Turner, B.L., and A.M. Powell 

1977. 

Wagnitz, G. 

1976. 
NUMBER 64 23 

'd :- 0 

:a. 

14 

FIGURES I-l7.-Basal columellar patterns of the tribe Liabeae in vertical view (for citation of 

specimens see appendix): 1, Paranephelius jelskii; 2, P. uniflorus; 3, Austroliabum polymnioides; 

4, Cacosmia rugosa; 5, Chionopappus benthamii; 6, Ferreyranthus excelsus; 7, F. vernonioides; 8, 

Sinclairia klattii; 9, 10, S. deamii; 11, Liabellum angustissimum; 12, 13, Liabum barahonense; 14, 

L. bourgeaui; 15, L. eggersii; 16, L. falcatum; 17, L. poiteaui. 
24 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

...... , .*-,:.- .......... ... .:.. .... :, *. '--a. 2Q 1-1 0: 0%: 

ec :- 18 

FIGURES 18-40.-Basal columellar patterns of the tribe Liabeae in vertical view: 18, Liabum 

selleanum; 19, 20, L. subacaule; 21, Oligactis uolubilis; 22, 0. (Andromachiopsis) coriacea; 23, 0. 

(A,) pichinchensis; 24, Erato polymnioides; 25, Philoglossa mimuloides f. sapida; 26, P. peruviana 

DC; 27, Chrysactinium acaule; 28, C. hieracioides; 29, Munnozia foliosa; 30-32, M. jussieui; 33, 

M. lanceolata; 34, M. lyrata; 35, M. senecionidis; 36, M. silphioides; 37, M. subuiridis; 38, M. 

uenosissima; 39, M. (Kastnera) annua; 40, M. (K.) niuea. 
NUMBER 64 

FIGURES 4 1-46.-Scanning Electron Micrographs of tribe Liabeae, Paranephelius unzjlorus 

pollen: 4 1, colpar view; 42, polar view; 43, colpus; 44, intercolpar area; 45, broken tectum; 

46, broken tectum. (Scale lines 41 = 10 pm; 42-46 = 5 pm.) 

25 
26 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

FIGURES 47-53.--SEM of tribe Liabeae, Paranephelius and Pseudonoseris pollen: 47, Paranephelius unijlorus; 48, P. jelskii; 49, P. unijlorus, unacetolized grain. Pseudonoseris discolor: 50, 

colpar view; 51, polar view; 52, colpus; 53, spines. (Scale lines 47, 48 = 2 pm; 49-51 = 10 pm; 

52 = 5 pm; 53 = 4 pm.) 
NUMBER 64 

FIGURES 54-59.--SEM of tribe Liabeae, Bishopanthus and Cacosmia pollen. Bishopanthus 

soliceps: 54, equatorial view with colpus; 55, polar view; 56, intercolpar region; 57, broken 

tectum. Cacosmia rugosa: 58, colpar view; 59, polar view. (Scale lines 54-56, 58, 59 = 10 pm; 

57 = 2 pm.) 

27 
28 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

FIGURES 60-65.--SEM of tribe Liabeae, Cacosmia and Chionopappus pollen. Cacosmia rugosa: 

60, colpus; 61, spines; 62, 63, unacetolized grains. Chionopappus benthamii: 64, colpar view; 65, 

polar view. (Scale lines 60, 61 = 5 pn; 62-65 = 10 pm.) 
NUMBER 64 29 

FIGURES 66-7 1 .-SEM of tribe Liabeae, Chionopappus and Austroliabum pollen. Chionopappus 

benthamii: 66, spines; 67, colpus; 68, microphotograph of microtome section. Austroliabum 

candidum: 69, colpar view; 70, polar view; 71, spines. (Scale lines 66 = 2 pm; 67 = 5 pm; 68- 

70 = 10 pm; 71 = 4 pm.) 
30 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

FIGURES 72-77.--SEM of tribe Liabeae, Austroliabum and Microliabum pollen. Austroliabum 

candidum: 72, colpus; A. polymnioides: 73, broken grain. Microliabum humile: 74, colpar view; 

75, polar view; 76, spines; 77, broken grain. (Scale lines 72, 76 = 5 pm; 73 = 2 pm; 74, 75 = 

10 pm; 77 = 1 pni.) 
NUMBER 64 31 

FIGURES 78-83.-SEM of tribe Liabeae, Ferreyranthus and Sinclairia pollen. Ferreyranthus 

uerbasczjcolius: 78, colpar view; 79, polar view; 80, colpus; 81, spines. F. rugosus: 82, broken 

grain. Sinclairia discolor: 83, colpar view. (Scale lines 78, 79, 83 = 10 pm; 80, 81 = 5 pm; 

82 = 2 pm.) 
32 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

FIGURES 84-89.--SEM of tribe Liabeae, Sinclairia pollen. Sinclairia discolor: 84, polar view; 

85, colpus; 86, spines; 87, unacetolized grain. S. deamii: 88, 89, Standley 19695, broken grains. 

(Scale lines 84, 87 = 10 Fm; 85, 86 = 5 Fm; 88, 89 = 2 wm.) 
NUMBER 64 

FIGURES 90-95.--SEM of tribe Liabeae, Sinclairia and Liabellum pollen. Sinclairia deamii: 90, 

Standley 19695; 91-93, Edwards 568. Liabellum palmeri: 94, colpar view; 95, polar view. (Scale 

lines 90 = 2 gm; 91-93 = 1 gm; 94, 95 = 10 pm.) 

33 
34 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

FIGURES 96- 10 1 .-SEM of tribe Liabeae, Liabellum and Oligactis pollen. Liabellum palmeri: 96, 

polar view; 97, colpus; 98, spines. Oligactis volubilis: 99, colpar view; 100, 101, polar views. 

(Scale lines 96, 97, 99-101 = 10 pn; 98 = 5 prn.) 
NUMBER 64 

FIGURES 102-107.--SEM of tribe Liabeae, Oligactis pollen. 0. volubilis: 102, colpus; 103, 

spines. Oligactis (Andromachiopsis) pichinchensis: 104, colpar view; 105, polar view; 106, colpus; 

107, spines. (Scale lines 102, 106 = 5 pm; 103, 107 = 2 pm; 104, 105 = 10 pm.) 

35 
36 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

FIGURES 108-1 13.--SEM of tribe Liabeae, Liabum pollen. L. umballatum: 108, colpar view; 

109, polar view; 110, colpus; 11 1, spines. L. igniarium: 112, 11 3, broken grains. (Scale lines 

108,111, 112, 113=5pm; 109,110=10pm.) 
NUMBER 64 

FIGURES 1 14-1 19.--SEM of tribe Liabeae, Liabum pollen, broken grains. 1 14-1 17, L. igniarium. 11 8, 119, L. subacaule. (Scale lines 114 = 5 Fm; 115-1 19 = 2 gm.) 

37 
38 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

FIGURES 120-1 25.--SEM of tribe Liabeae, Liabum and Erato pollen. L. subacaule: 120, broken 

grains. L. barahonense: 12 1 - 123, broken grains. Erato polymnioides: 124, colpar view; 125, polar 

view. (Scale lines 120 = 2 pm; 121-123 = 1 pm; 124, 125 =10 pm.) 
NUMBER 64 

FIGURES 126-1 33.-SEM of tribe Liabeae, Erato pollen. Erato polymnioides: 126 polar view; 

127, colpus; 128, spines; 129, 130, broken grains. E. sodiroi: 131-133, broken grains. (Scale 

lines 126 = 10 pm; 127 = 5 pm; 128-133 = 2 pm.) 

39 
40 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

FIGURES 134-1 39.-SEM of tribe Liabeae, Philoglossa peruuiana pollen: 134, near polar view; 

135, colpar view; 136, colpus; 137, spines; 138, 139, broken grains. (Scale lines 134, 135 = 

10 pm; 136 = 5 pm; 137, 138 = 2 pm; 139 = 1 pm.) 
NUMBER 64 41 

FIGURES 140- 146.-SEM of tribe Liabeae, Chrysactinium acaule pollen: 140, colpar view; 14 1, 

polar view; 142, colpus; 143, spines; 144-146, broken grains. (Scale lines 140, 141 = 10 pm; 

142 = 5 pm; 143-146 = 2 Fm.) 
42 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

FIGURES 147-1 53.--SEM of tribe Liabeae, Munnoria lanceolata pollen: 147, colpar view; 148, 

polar view; 149, colpus; 150, spines; 151-153, broken grains. (Scale lines 147, 148 = 10 pm; 

149, 150 = 5 pm; 151-153 = 2 pm.) 
NUMBER 64 

FIGURES 134- 160.-SEM of tribe Liabeae, jZlunnozia pollen, broken grains: 154, 155, Munnozia 

lanceolata. 156, M. perfoliata. 137-160, iM. jussieui. (Scale lines 154-156, 158 = 2 pm; 157, 

159, 160 = 1 pm.) 

43 
44 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

FIGURES 161-167.-SEM of tribe Liabeae, Munnozia pollen, broken grains: 161-165, M. 

lyrata. 166, 167, M. seneczonidis. (Scale lines 161, 164, 165 = 1 pm; 162, 163, 166, 167 = 

2 pm.1 
NUMBER 64 

FIGURES 168-1 73.-SEM of tribe Liabeae, Munnozia pollen (subgenus Kastnera). M. (Kastnera) 

annua: 168, colpar view; 169, polar view; 170, colpus; 17 1, spines. M. (Kastnera) tenera: 172, 

173, broken grains. (Scale lines 168, 169 = 10 pm; 170, 171 = 2 Wm; 172, 173 = 1 pm.) 

45 
46 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

FIGURES 174- 180.-SEM of tribe Liabeae, Munnoria (Kastnera) tenera, pollen, broken grains 

(1 79, arrows showing two layers of small columellae). (Scale lines = 1 pn.) 
NUMBER 64 

FIGURES 18 1-1 89.-SEM of tribe Vernonieae, Pseudost@tza kingzz, pollen: 18 1, colpar view; 

182, polar view; 183, spines; 184-189, broken grains. (Scale lines 181, 182, 184 = 10 pm; 

183, 185, 187 = 5 pm; 186, 188, 189 = 2 pm.) 

47 
48 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

FIGURES 190-1 95.--SEM of tribe Vernonieae, Vernoniafuerteszz pollen: 190, colpar view; 19 1, 

polar view; 192, spines; 193-195, broken grains. (Scale lines 190, 191 = 10 gm; 192, 194, 

195 = 2 pni; 193 = 0.5 pm.) 
NUMBER 64 

FIGURES 196-202.-SEM of tribe Vernonieae, Blanchetia and Bishopalea pollen. Blanchetia 

heterotricha: 196, surface with spines; 197-1 99, broken grains. Bishopalea erecta: 200, whole 

grains; 201, polar view; 202, surface. (Scale lines 196, 197, 202 = 5 pm; 198, 199 = 2 pm; 

200, 201 = 10 pm.) 

49 
50 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

FIGURES 203-208.--SEM of tribe Vernonieae, Heterocypsela and Phyllocephalum pollen. Heterocypsela andersonii: 203, equatorial view,; 204, broken tectum. Phyllocephalum scabridum: 205- 

207, grains; 208, reticulum, showing two strata. (Scale lines 203, 205-208 = 10 pm; 204 = 

2 pm.)