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Home , Acicarpha, Anthotium, Arnaldoa, Barnadesia, Buphthalmum, Calycera

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Systematic studies of

DeVore, Melanie Lynn, Ph.D.

The Ohio State University, 1994

Copyright ©1994 by DeVore, Melanie Lynn. All rights reserved.

UMI 300 N. Zeeb Rd. Ann Arbor, MI 48106

SYSTEMATIC STUDIES OF CALYCERACEAE

DISSERTATION

Presented in Partial Fulfillment of the Requirements for

the Degree Doctor of Philosophy in the Graduate

School of the Ohio State University

By

Melanie Lynn DeVore, B.S.

*****

The Ohio State University

1994

Dissertation Committee: Approved by:

Daniel J. Crawford

Michael L. Evans

Tod F. Stuessy Adyisor-V Department of'Rlant Biology Copyright by

Melanie L. DeVore 1994 To Neil A. Harriman,

who along with serendipity, hooked me on vascular systematics.

11 ACKNOWLEDGMENTS

I gratefully acknowledge a number of individuals and institutions for their help throughout this work. Tod F.

Stuessy, my dissertation advisor provided help, as well as stimulating conversation and ideas concerning the early evolution of the Compositae and related families. I also extend appreciation to Daniel J. Crawford and Michael L. Evans for their advice and critical reading of the manuscript. I would also like to acknowledge Thomas and Edith Taylor who supported my first year of graduate studies at OSU and influenced some of my views regarding plant evolution. This dissertation would not have been possible without the support of faculty, graduate students, and staff, from the

Departamento de Botanica, Universidad de Concepcion,

Concepcion, Chile; Divisiones Entomologie, Plantas Vasculares, and Laboratorio de Sistematica y Biologie Evolutive (LASBE) of

Museo de La Plata; and Institute de Botanica Darwinion. All of these institutions contributed to this study by supplying field vehicles and support, access to herbarium collections and library research.

Ill Appreciation is expressed to; The American Society of Plant Taxonomists (Field Work Initiation Grant), The Ohio

State University (Beatley Herbarium Award from OS, Latin American Studies Travel Award, and Graduate Student Alumni Research Award), and Sigma Xi for financial support.

The following herbaria provided loans or access to herbarium collections: CONC, F, GH, LL, LP, MICH, MO, NY, OS, S, SI, TEX, US.

Conversations with the following individuals concerning , Calyceraceae, or were extremely helpful: K. Bremer, A.L. Cabrera, A. Cronquist, V. Funk, E.

Harris, T. hammers, C. Marticorena, R. Olmstead, and H. Robinson.

Appreciation is also expressed to Jorge Arriagada for photographing type material and Monica Taylor for editing segments of the manuscript.

Last but not least, I would like to thank my parents Don and Nancy DeVore for all their love, encouragement, and patience. Without their help, this work would not have been possible.

IV VITA

August 17, 1963...... Born - Warren, Ohio 1985...... B. S. Geology, The University of Wisconsin-Oshkosh

1985-1987...... Teaching Assistant, Depatment of Biology and Microbiology, The University of Wisconsin-Oshkosh

1987-1993...... Graduate Associate, Department of Plant Biology, The Ohio State University 199 0...... American Society of Plant Taxonomists, Field Work Initiation Grant

199 1...... Sigma Xi, Grant-in-Aid of Research

1991...... The Ohio State University Herbarium, Beatley Award

1991...... The Smithsonian Institution, 10-week Graduate Student Fellowship 1991 The Ohio State University, Latin American Studies Program Travel Award

1991...... The Ohio State University, Graduate Student Alumni Research Award 1993-1994 Lecturer, Department of Biological Sciences, Sam Houston State University V 1994 Botanist, Texas Regional Institute of Environmental Studies, Sam Houston State University

1994- Assistant Professor, Department of Biological Sciences, Sam Houston State University

PUBLICATIONS

DeVore, M. L. and N. A. Harriman. 1987. Juncus exil is Osterhout (Juncaceae), an overlooked synonym of J. confusus Coville. Sida 12: 243.

DeVore, M. L. and T. N. Taylor. 1988. Mesozoic seed : a organ from the Triassic of Antarctica [abstract]. Amer. J. Bot. 75: 105.

DeVore, M. L. 1990. The occurrence of tribuloides Juss. in Eastern North America [abstract]. Ohio J. Sci. 85: 5.

DeVore, M. L. 1990. The ballast distribution of Acicarpha tribuloides Juss. in eastern North America [abstract]. Amer. J. Bot. 77: 206.

DeVore, M. L. 1991. Acicarpha tribuloides (Calyceraceae) in eastern North America. Rhodora 93: 26-35. DeVore, M. L. and Tod F. Stuessy 1991. Time and place of origin of Asteraceae, Calyceraceae, and Goodeniaceae [abstract]. Amer. J. Bot. 78: 178.

DeVore, M. L. 1992. A new species of from northern Argentina [abstract]. Ohio J. Sci. 92: 17.

DeVore, M. L. 1993. The systematics and biogeography of Acicarpha (Calyceraceae) [abstract]. Ohio J. Sci. 93: 41.

DeVore, M. L. 1993. The systematics of Acicarpha (Calyceraceae) [abstract]. Amer. J. Bot. 80: 144

vi DeVore, M. L. 1994. Chromosome numbers in Calyceraceae [abstract]. Published abstracts. The Texas Academy of Science Meeting, Houston, TX, March 1994. DeVore, M. L. In press. Systematics of Argentinean Calycera (Calyceraceae) species [abstract]. Amer. J. Bot. 81: DeVore, M. L. In press. Calyceraceae and its relationships with Compositae [abstract]. Abstracts for The International Compositae Congress.

Stuessy, T. F., T. Sang, and M. L. DeVore. In press. The Phylogeny and biogeography of [abstract]. Abstracts for The International Compositae Congress. Pesecreta, T. C., V. I. Sullivan, and M. L. DeVore. In press. The connective base of Acicarpha tribuloides (Calyceraceae). Amer. J. Bot. 81:

DeVore, M. L. and T. F. Stuessy. In press. Time and place of origin of Compositae, Calyceraceae, and Goodeniaceae. In C. Jeffrey (ed.). Advances in Compositae. Academic Press, London.

FIELD OF STUDY

Major Field: Plant Biology

Studies in Plant Systematics with Tod F. Stuessy

V l l TABLE OF CONTENTS

ACKNOWLEDGMENTS...... iii

VITA...... V

LIST OF TABLES...... X

LIST OF FIGURES...... xi INTRODUCTION...... 1

CHAPTER PAGE

I. PLACE AND TIME OF ORIGIN OF ASTERACEAE, WITH ADDITIONAL COMMENTS ON CALYCERACEAE AND GOODENIACEAE ...... 5

Introduction ...... 5 Review of Morphology of Asteraceae, Calyceraceae, and Goodeniaceae...... 8 Present Distributions of Asteraceae and Goodeniaceae...... 20 Trends in Fossil Records of Asteraceae and Goodeniaceae...... 23 Tectonic Events in the Southern Hemisphere...... 27 Paleoclimatic D a t a ...... 30 References...... 36

II. CYTOLOGY OF CALYCERACEAE...... 55 Introduction ...... 55 Material and M e t h o d s ...... 56 Results...... 62 Cytology in relation to morphology and distribution of Acicarpha ...... 62 Cytology and Relationships Within Calycera...... 63

viii and its Relationship with Calycera...... 65 Estimate of Ancestral Base Number of Calyceraceae...... 68 Cytology and Biogeography...... 70 Literature cited...... 71

III. SYSTEMATICS OF ACICARPHA ...... 76 Introduction ...... 76 Taxonomic History ...... 77 Generic Relationships ...... 78 Phenetic studies of the Acicarpha tribuloides Complex ...... 79 Cladistic Analysis ...... 91 T a x o n o m y ...... 102 Refere n c e s ...... 132

IV. SYSTEMATICS OF CALYCERA...... 140 Introduction ...... 140 Taxonomic History...... 141 Generic Relationships ...... 142 Concepts of Species...... 143 Morphology and Taxonomic Criteria ...... 144 T a x o n o m y ...... 149 Refere n c e s...... 203

APPENDIX...... 208

The Distribution of Acicarpha tribuloides in Eastern North America ...... 209 LIST OF REFERENCES...... 219

IX LIST OF TABLES

TABLE PAGE

1. Previous chromosome reports and new chromosome counts (and voucher specimens) from taxa of Calyceraceae...... 57

2. List of 43 quantitative characters used in the phenetic studies of Acicarpha Section Acicarpha ...... 83

3. Eigenvectors for characters which load Factors (Axes) 1 - 3 ...... 89

4. Data matrix for cladistic analysis of Acicarpha...... 94 LIST OF FIGURES

FIGURE PAGE

1. Camera lucida drawings of meiotic chromosome counts from species of Calyceraceae...... 61

2. Principal Components Analysis (PCA) and Minimum Spanning based on 43 morphological characters...... 86

3. variation in Acicarpha tribuloides . . 88 4. Single most parsimonious tree of species of Acicarpha...... 99

5. Distribution in southern Brasil and adjacent northeastern Argentina of Acicarpha bupleuroides, A. itatiaiae, A. procumbens, and A. spathulata .... 110

6. Distribution of Acicarpha tribuloides . . . 128

7. Features of filaments and glands in Leucocera and Calycera...... 148

8. Calycera coronata...... 159

9. Distribution of Calycera coronata, C. leucanthema, and C. sympaganthera. . . . 161 10. Distribution of Calycera eryngioides, and C. sessiliflora...... 172

11. Distribution of Calycera crenata and C. pulvinata...... 180

12. Distribution of Calycera calcitrapa, C. herbacea, and C. involucrata...... 187

13. Distribution of Calycera spinulosa .... 196 xi INTRODUCTION

Calyceraceae is a southern South American of angiosperms (flowering plants) consisting of six genera.

Systematic studies of Calyceraceae conducted for dissertation research at The Ohio State University included estimating a place and time of origin of Asteraceae,

Calyceraceae, and Goodeniaceae (Chapter I). Other studies included cytological studies of Calyceraceae (Chapter II), and revisions of ?^cicarpha (Chapter III), and Calycera (Chapter IV).

Understanding the place and time of origin of

Asteraceae has been difficult because of a scanty fossil record and because of vagueness of relationships to other families within Asteridae. Recent chloroplast DNA (cpDNA) restriction site analysis, ribulose-1, 5-bisphospate carboxylase frbcL) data, and mapping studies of the inverted repeat region, suggest that Calyceraceae and Goodeniaceae are the families most closely related to Asteraceae. These macromolecular studies have also suggested (1) South

American Barnadesioideae to be the most primitive subfamily of Asteraceae; (2) Calyceraceae, also South American, to be more closely related to Asteraceae than to any other family; 2 and (3) Goodeniaceae, a family centered in Australia, to be more closely related to Asteraceae and Calyceraceae than to or Lobeliaceae. Based on relationships and distributions of Asteraceae, Calyceraceae, and Goodeniaceae, it is assumed that these three families derived from an ancestral lineage on the -Antarctica-Australia supercontinent. A late Eocene origin is estimated based on direct fossil evidence of Asteraceae and Goodeniaceae, morphological trends in fossil flowers, and time of appearance of pollination syndromes in the fossil record.

This age of origin is also supported by the early Eocene

(43-53 My a) separation of Australia and South America from

Antarctica. Paleoclimatic studies and measurements of evolutionary activity also are in agreement with a late

Eocene origin for Asteraceae, Calyceraceae and Goodeniaceae.

It is conceivable that more variable climatic regimes, as well as increased orogenic activity during late Eocene, may have enabled Asteraceae to expand into newly formed habitats.

Meiotic chromosome numbers were determined for one species of , five species of Calycera, and two species of Nastanthus. These numbers include counts for six species that have not previously been examined cytologically:

Calycera crassifolia (n = 21) C. eryngioides (n = 20-21), C. leucanthema {n = 17), C. spinulosa (n = 21), C. sympaganthera (n = 13), and Nastanthus andinus (n = 21). 3

Both new and previous counts for the family reveal that Acicarpha (n = 8) occurs at a diploid chromosome level, whereas species of Calycera (n = 13, 17, 20-21, 21), Boopis (n = 15 & 18), and Nastanthus are interpreted as being at the polyploid chromosome levels. Cytological data are not yet available for Gamocarpha or Moschopsis. An ancestral base number of r = 8 or 9 is suggested for the family based on correlations of morphological features, chromosome numbers, cytological trends in related families, and distributional history.

Taxonomic studies of Acicarpha (Chapter III) and

Calycera (Chapter IV) are the first complete treatments of any within Calyceraceae since Miers (1860). Acicarpha appears to be the most primitive genus within the family, as well as the most closely related genus to the primitive subfamily of Asteraceae (Barnadesioideae). Unlike other members of Calyceraceae, but similar to Asteraceae,

Acicarpha has capitula comprised of florets which mature towards the inside (centripetal sequence of development). Systematic studies of Acicarpha are therefore significant in providing clues regarding the early evolution of Asteraceae.

Calycera, containing 11 species, is one of the largest genera within Calyceraceae. The taxonomic treatment of

Calycera recognizes Argentinean and Chilean species groups as Subgenera Calycera and Anomocarpus. Completion of a taxonomic revision of Calycera provides the basis for 4 further evolutionary studies regarding spéciation within Calycera, as well as generic relationships within Calyceraceae. CHAPTER I

PLACE AND TIME OF ORIGIN OF ASTERACEAE, WITH ADDITIONAL

COMMENTS ON CALYCERACEAE AND GOODENIACEAE

Introduction

For more than a century, workers have speculated on the place and time of origin of Asteraceae (e.g., Bentham, 1873; Small, 1919; Cronquist, 1955, 1977; Raven and Axelrod, 1974;

Turner, 1977; Bremer, 1992, 1993a). Bentham (1873) examined areas of distribution of Asteraceae and concluded that Africa, Western America, and possibly Australia, were the homes of ancestral members of the family. He believed that

Africa contained several isolated remnants of extinct races, whereas the Andean Cordilleras and the Pacific Islands contained the most primitive extant members. Small (1919) estimated that Asteraceae originated during late Cretaceous or early Eocene in the Amazon and northern from a

Lobelioideae ancestor similar to Siphocampvlus Pohl or

Centropogon Presl. during major orogenesis and subsequent climatic evolution within the Andean Cordilleras. Cronquist

(1955, 1977) regarded as the most primitive tribe and indirectly suggested the highlands of central 6

Mexico as the area of origin by pointing out that this was the center of diversity for the tribe. Raven and Axelrod

(1974) offerred a mid-Oligocene and South American origin for the family. Turner (1977, p. 37) suggested that Asteraceae evolved from a lineage close to Calyceraceae during the middle Cretaceous in "westernmost Gondwanaland," and that tribal units appeared "along the leading edges" of drifting continents by the end of the Oligocene and diversified rapidly during Miocene climatic fluctuations.

Bremer (1992, 1993 a,b) estimated the ancestral area of Asteraceae by calculating the gain to loss ratio for each area represented on a composite cladogram of the family. A high gain/loss ratio and basal position on the cladogram suggested that a particular area may have been part of the ancestral region, and he estimated that this area included

South America and the Pacific Basin, but excluded Africa and most of Eurasia. Despite these numerous valuable perspectives, no definitive conclusions have been forthcoming on the place and time of origin of Asteraceae due to uncertainty on close familial relationships within Asteridae. Recent rbcL sequence studies (Olmstead et al., 1990, 1992) and restriction site mapping studies of the cpDNA inverted repeat (Downie and Palmer, 1992), however, have provided new insights to the familial relationships among Asteraceae and other families within Asteridae. To date, rbcL sequence 7 studies have shown that Calyceraceae and Goodeniaceae are sister groups of Asteraceae. Mapping studies of the inverted repeat region (Downie and Palmer, 1992) found extensive variation in sequence and length of cpDNAs from

Goodeniaceae. These preliminary data suggest that Calyceraceae may be the family most closely related to Asteraceae since they contain no major cpDNA rearrangements.

The results of cpDNA studies provide an opportunity to speculate on the place and time of origin of Asteraceae,

Calyceraceae, and Goodeniaceae. The Goodeniaceae are centered in Australia (Carolin et al., 1992), and Calyceraceae are limited to South America (DeVore, in prep).

Compositae are found on nearly every continent of the world, but recent cpDNA studies (Jansen and Palmer, 1987, 1988) have suggested South American Barnadesioideae (Bremer and

Jansen, 1992) as most primitive. This provides a South

American focus for the origin of this family. These data suggest that the three families may have originated from an ancestral lineage centered in the South America-Antarctica-

Australia supercontinent.

In view of new cpDNA data on relationships among

Asteraceae, Calyceraceae and Goodeniaceae, therefore, the purposes of the present paper are to: (1) review morphological relationships of the three families to test the suggestion from molecular data that Asteraceae and

Calyceraceae are more closely related to each other than 8 either is to Goodeniaceae; (2) discuss in more detail the present distributions of the three families; (3) comment on fossil records of Asteraceae and Goodeniaceae (fossils of Calyceraceae are unknown), with particular reference to morphological trends that bear on place and time of origin;

(4) present a summary of tectonic events in the southern hemisphere that may have influenced development of present distributional patterns; and (5) indicate relevance of paleoclimatic data to place and time of origin of the three families.

Review of Morphology of Asteraceae. Calyceraceae, and Goodeniaceae

If Barnadesioideae are primitive for Asteraceae, as suggested by Jansen and Palmer (1987), then it makes sense to base our discussion of the morphology of the family in relation to Calyceraceae and Goodeniaceae on this subfamily rather than on the entire diverse group. Asteraceae have been so successful evolutionarily in morphological adaptations, elaborations of secondary products, chromosomal evolution, gender strategies, dispersal abilities, etc., that relating variations in these factors for the entire family to the other two families would be difficult, if not misleading. Hence, our discussion will focus on

Barnadesioideae. This is facilitated by on-going studies in our laboratory on phylogeny and biogeography of the group 9 (e.g., Stuessy and Sang, 1993; Stuessy and Sagastegui, 1993.

Barnadesioideae consist of approximately 90 species in eight genera (Cabrera, 1959, 1977; Chung, 1965; Ezcurra, 1985: Arnaldoa Cabrera; Mutis ex L.f.; Chuouiraqa Juss.; Dasvphvllum Kunth; Doniophvton Wedd.; Fulcaldea

Poiret; Huarpea Cabrera; Schlechtendalia Less. The monotypic Duseniella Schumann has also been considered to be a part of this complex by some authors (e.g., Hansen, 1991a;

Bremer, 1992), but we exclude it here (it will be discussed fully in a forthcoming paper on the phylogeny and biogeography of the Barnadesioideae; Stuessy et al., in prep.). The following morphological summaries refer to these genera only.

Calyceraceae consist of eight genera and approximately

50 species. It was first mentioned as a family by Jussieu (1803), and later recognized by Cassini (1816) and Brown

(1817) as distinct from Asteraceae. Few workers have focused monographically on the family, but Pontiroli (1963) did produce a fairly detailed floristic treatment of the

Argentinean members. Although Calyceraceae do resemble

Asteraceae, some major differences also exist which will be detailed below.

Carolin has carried out extensive studies of Goodeniaceae including floral structure and anatomy

(Carolin, 1959), seed and fruit morphology (Carolin, 1966), trichome types (Carolin, 1970), cladistic analysis for all 10 11 genera within Goodeniaceae and (Carolin, 1978), and a treatment of the family for the

(Carolin et al., 1992). These studies, plus examination of available herbarium material, have been used to summarize the morphology (and distribution in the next section) of Goodeniaceae for this paper. For our purposes we will follow Carolin (1978) and include Brunonia.

A.

Barnadesioideae, as with the rest of Asteraceae, have tightly formed capitula with indeterminate centripetal development. The number of florets per head varies from nearly 100 in Arnaldoa to only one in Fulcaldea. The heads are mostly homogamous, with some differentiation into central and marginal florets in Barnadesia. This genus is suspected to be one of the most advanced within the complex (Stuessy and Sang, 1993), and the heterogamous tendency may reflect adaptations in part toward hummingbird pollination (Ortiz, in litt.: pers. observ.) with many long-exserted marginal florets and several reduced central ones (see

Chung, 1965). Phyllaries occur in several series, and the are closely imbricate with acute to acuminate or

cuspidate apices. The head is campanulate to cylindrical,

and receptacular bracts (pales) are absent. 11 All members of Calyceraceae are also characterized by capitula, but the majority are determinate, being composed of cymose subunits. Acicarpha. however, appears to be derived from a reduced spike (i.e., indeterminate), and this is one reason this genus is believed to have evolved in isolation from other members of the family (DeVore, 1993).

One trend within the family is condensation of heads into large, compound units, such as in Nastanthus. reaching nearly 26 cm diam. (DeVore, pers. observ.). The involucral bracts are little more than slightly modified cauline . Receptacular bracts are present within all genera, and in Gamocarpha they are fused around cymose subunits.

Inflorescence types within Goodeniaceae include dichasia, thyrses, racemes, umbels, spikes, and heads.

According to Carolin (1967) the primitive inflorescence type in the family is the thryse, which through reductions of axillary cymes, gave rise to the common spikes and racemes of the family. Reduction of cymose and spike into heads occurs in R.Br. With regard to inflorescences, therefore, both

Asteraceae and Calyceraceae share indeterminate capitula, although most genera of the latter are determinate, except for Acicarpha. Both have involucral bracts which surround the heads, although they are much more reduced and modified in Asteraceae. Goodeniaceae have a diversity of inflorescence types, with flowers subtended by single 12 bracts/ and heads that occur are believed to be derived rather than primitive within the family (Carolin, 1967).

B . Flowers

Barnadesioideae vary in shape of corolla depending upon the genus. Common configurations are regular 5-lobed corollas, typical of disc corollas in the rest of the family

(e.g., Chucmiraaa. Doniophvton). Also common are four lobes fused on one side of the corolla, and one long and separate

lobe on the other side (typical in Barnadesia. Huarpea. and

Schlechtendalia). Dasvphvllum reveals more variation in

corolla features than in other genera, and nearly all types of conditions found in the subfamily occur in the genus

(Cabrera, 1959). The anthers are five, syngenesious, usually with apical appendages (except for most species of

Dasvphvllum), and usually short- or long-sagittate bases.

There are two fused carpels, a single locule with one basally attached , and short-bifid styles, either

smooth on the adaxial surface, or with very short papillae

(Hansen, 1991a). The pappus in Barnadesioideae, which we

assume to be a modified calyx, is uniformly plumose, persistent, in a single series, with antrorsely oriented

hairs.

The corolla in Calyceraceae is tubular and four- or

five-lobed. An unusual feature is a cuticular layer which

separates from the corolla, possesses stomata, and is 13 photosynthetic during bud development. In Acicarpha and some species of Calvcera. Gamocarpha and Nastanthus. this outer corolla layer becomes photosynthetic again during fruit development. There are four or five alternate with the , and the anthers are generally free, but sometimes slightly connate toward the base. Associated with filaments are nectaries, which in some cases occupy a position on the outside of the filamental column towards the base of the corolla. The is inferior with two fused carpels, a single locule, and a solitary, pendulous ovule.

The stigma is undivided, capitate, and papillate. In some members of Calyceraceae, the upper style and stigma extend nearly a centimeter beyond the tip of the corolla. The calyx is composed of four or five fused sepals which become arenchymous and sometimes "spine-like" and persistent on the fruit.

The basic corolla type in Goodeniaceae is tubular with an adaxial slit. Almost all genera within the family (with the exception of Cav.) are characterized by prominent outgrowths of the corolla lobe margins (called

"wings," Carolin, 1959). The following five corolla forms within the family can be recognized based on these two characters: (l) slit in the corolla tube, wings absent; (2) corolla tube spread open along adaxial slit, flowers fan­ shaped; (3) adaxial folded around indusium (to be described below); (4) 2-lipped corolla; and (5) 2-lipped 14 corolla with lower portion of adaxial corolla wings forming auricles surrounding the indusium. The androecium consists of five stamens which alternate with sepals. Filaments are typically free and anthers can be free or connate. In some species of . hairs are present on tips of anthers.

The ovary varies from superior, half-inferior, to completely inferior. Four carpels may be present (apparently the primitive condition; Carolin, 1959), but the two-locule (two-carpellate) condition prevalent in the family may have occurred through reduction (further reduction to a single locule has also occurred within Brunonia). The ovary has axile or parietal placentation in either basal or medial zones, or in both regions. The style may be simple, or have two or four stigmatic branches. An outgrowth of the style, the cup-shaped "indusium," is distinctive within the family and will be discussed in more detail with regard to pollination. Three or five sepals are present and may be free, connate, reduced to a rim, or absent; sometimes they are persistent. In comparing the three families in floral features,

Asteraceae and Calyceraceae are more similar to each other in their more regular corollas, fusion of at least part of the anthers, unilocular, uniovulate and uniformly inferior ovaries, and a tendency for a calyx modified into spines or hairs. 15

C. Pollen

Barnadesioideae pollen is unusual in Asteraceae in that the exine has a poorly developed foot layer and consists of fine internal rods, instead of larger columellae (Parra and

Marticorena, 1971; Skvarla et al., 1977; Hansen, 1991b).

The grains are round or ovoid, and sometimes with either small or large intercolpal concavities. The external sculpturing varies from smooth, spinulate, to lophate (the latter in Barnadesia and Huarpea; Gamerro, 1986).

Calyceraceae have tricolpate pollen grains with minute spines, or smooth surfaces, and with convex or concave intercolpal regions. Sometimes an exine layer is present along the colpi of the grain, and an exine pouch that extends toward the aperture also occurs in Acicarpha and Calycera.

Goodeniaceae have a wide range of surface variations of pollen grains from short spines to prominent striae (Skvarla et al., 1977). Pollen ultrastructural patterns within the family are rather consistent (with the exception of

Lechenaultia R. Br.) and are typically characterized by thick columellae which bifurcate distally.

The morphology of pollen of Calyceraceae has been used to argue a clear relationship with Asteraceae (Skvarla et al., 1977; Hansen, 1992). Pollen of Goodeniaceae, with its thick bifurcating columellae, are clearly divergent from the other two families. The intercolpar concavities in 16 Acicarpha and some species of Calvcera have been suggested

as a synapomorphy between Calyceraceae and Asteraceae

(Hansen, 1992). The pollen wall ultrastructure of the former has been described by Skvarla as nearly identical to the anthemoid-like pattern found in some members of

Barnadesioideae. It is possible, however, that these two characters are parallelisms and not synapomorphies, because other families approach the anthemoid pattern (e.g.,

Valerianaceae, Brunoniaceae, even Sapindaceae; Skvarla et al., 1977).

D. Pollen presentation mechanisms

In Barnadesioideae, as well as in the rest of

Asteraceae, pollen presentation is a pump-type mechanism

involving growth of the style up through syngenesious anthers (Leins and Ebar, 1990). With introrse dehiscence, the pollen is presented on the outer surface of the styles which, when the two stigma lobes open, leaves the pollen on the undersurface. In a few genera of the family (e.g., in some species of Gerbera L.), the pollen presentation is rapid and thigmotropic (Small, 1919; Pesacreta et al., 1993; pers. observ.). The filaments apparently are contractile and help "pull" the thecae down over the growing style.

The pollen presentation mechanism of Calyceraceae is

similar to the pump-type found in Asteraceae. One major difference is that anthers are typically not fused for their 17 entire length, and these bend outwards toward the corolla tube. The basal portions of the anthers are positioned near the style, and act as a cup to catch released pollen. In

Acicarpha. the connective base serves as a hinge, whereas other members of the family have hinge points located well below the anthers (Pesacreta, Sullivan and DeVore, in prep.).

Of all three families, the pollen presentation mechanism of Goodeniaceae is the most complex. While in bud, developing anthers are positioned beneath the indusium, which is a cup-like enlargement of the upper stylus area.

As the filaments grow upward, pollen is deposited into the indusium (or on the outside of the indusium in ^. The indusium closes, bends towards the center of the flower, and pollen is presented to the insect vector. Later the stigma grows upward, pushes the indusium open, and becomes receptive to pollen from other individuals. There are some variations of this basic pattern and additional structures (bristles) may be present to aid in pollen presentation.

Because of the complex pollen presentation mechanisms of Goodeniaceae (involving the indusium), Asteraceae and

Calyceraceae are much more closely related to each other. Both have at least or partially fused stamens and the mechanism of pollen presentation is pump-type in both cases. 18 E . Fruit

Achenes in Barnadesioideae at maturity usually have numerous antrorsely-oriented uniseriate hairs. Whether these are a unique type of trichome within the family, as suggested by Robinson (1987) and Hansen (1991a), remains to be seen with further surveying. Dispersal of of Barnadesioideae appears to be by wind.

The fruit type within Calyceraceae is an with persistent calyx lobes. Some members of the family (e.g.,

Nastanthus) have persistent sepals which expand and appear wing-like. The second fruit type has sepals which expand, enroll, and become spine-like (e.g., Calvcera). In both types, the sepals become arenchymous. Achenes appear to be dispersed by wind and water (DeVore, pers. observ.). In Acicarpha. achenes are fused to the and the entire capitulum disperses (DeVore, 1991). Calvcera has dimorphic achenes, those with small wing-like calyx lobes found toward the center of the head and dispersing individually, and those with elongated calyx lobes often fused to the involucral bracts, or receptacle, in which case the head disperses as a unit (DeVore, pers. observ.).

Fruit types within Goodeniaceae vary from drupes to capsules and nuts. Members of Sm., Selliera Cav., Sm., and Benth., produce seeds with wings which may aid in wind dispersal, serve as elaiosomes, or play a role in germination in dry habitats by helping to 19 absorb water. The genus with widest distribution, Selliera. is found in salt marshes in Australia, New Zealand, and

southern Chile. It has winged, mucilaginous seeds and

fleshy fruits; sea birds are believed to be the dispersal

agents (Carolin et al., 1992). Other members of the family are also dispersed by mammals and birds (Carolin et al., 1992) . Some species of Scaevola are dispersed by emus and several sections of the genus (sects. Scaevola.

Enantiophvllum. and two species placed in sect. Xerocarpa; Carolin, 1990) is characterized by fruits with fleshy mesocarps and presumptive animal dispersal.

There can be little doubt that the similar single­ seeded achenes of Calyceraceae and Asteraceae are more

indicative of a close relationship than the , drupes,

or nuts of Goodeniaceae. These differences in morphology

are accompanied by equally divergent mechanisms of dispersal.

F. Summary of morphological relationships

This brief morphological comparison of the three

families reveals clearly that Asteraceae are considerably closer to Calyceraceae than either is to Goodeniaceae. The

structure of corollas, anthers, ovaries (including placentation), pollen, pollen presentation, and fruit types and dispersal mechanisms are all extremely divergent in the

Goodeniaceae, whereas those of the other two families are 20 similar. Based on both careful considerations of trends within Barnadesioideae, Calyceraceae, and Goodeniaceae and current phylogenetic studies of Barnadesioideae (Stuessy et al., in prep.), the following features of Calyceraceae are potentially homologous with those of Barnadesioideae: (1) pollen ultrastructure (Skvarla et al., 1977); (2) anther microcharacters (connective bases; Pesacreta and DeVore, in prep.); (3) unilocular ovaries; (4) modified calyx (spine­ like, winged, or hairy); and (5) pump-type pollen presentation mechanism. A more detailed assessment of these, and additional homologies between Barnadesioideae and

Calyceraceae will be discussed in a future paper (Stuessy et al., in prep.). The utility of Goodeniaceae as an outgroup for morphological cladistic studies of Asteraceae is limited. The presence of such autapomorphic features in the family as winged corollas, indusia, diverse fruit types, and specialized trichomes makes recognizing homologies between

Goodeniaceae and Barnadesioideae difficult.

Present Distributions of Asteraceae and Goodeniaceae

Barnadesioideae are exclusively South American

(Cabrera, 1977). The largest genera are Barnadesia. Chucfuiraqa. and Dasvphvllum. and these have distinct ranges.

Barnadesia is usually found in moist environments at

elevations of 1500 to 3000 m along the northern Andes from

Colombia to Bolivia and northern Argentina (Chung, 1965; 21 pers. observ.). It is most speciose in Ecuador and Peru

(Chung, 1965). Chucmiraaa is abundant also along the Andes, but less so, with a stronger concentration in southern South America especially in Patagonia (Ezcurra, 1985). In fact, those species in the northern Andes are apparently hummingbird pollinated (Ortiz, in litt.) and appear derived from those insect-pollinated relatives to the south (Ezcurra and Crisci, 1987; Stuessy and Sang, 1993). Dasvphvllum is most common in southern South America, with a large concentration of species in southeastern Brazil (Cabrera,

1959). Cabrera (1977) believed that Archidasvphvllum. a Chilean section containing shrubs and large , was primitive within the genus. Doniophvton occurs on both sides of the southern Andes in Chile and Argentina over

3,000 m. The difficult-of-access Huarpea is found only in northern Argentina in and around the type locality in San

Juan Province. Schlechtendalia is known primarily from Uruguay, with a few scattered localities in adjacent southern Brazil (Stuessy, pers. observ.). Arnaldoa is restricted to northern Peru in and around the canyons of the

Rio Maranon. Taking all distributions into consideration, there is clearly a focus for Barnadesioideae in southern South

America. That Barnadesia is regarded as one of the most derived genera of this complex (Stuessy and Sang, 1993), especially in its heterogamous capitula, hummingbird 22 pollination syndrome (clearly derived within the family), lophate pollen, and hexaploid ploidy level, suggests that it came from somewhere toward the south rather than the reverse. If Schlechtendalia were regarded as the most primitive genus (Stuessy and Sang, 1993; it is the only diploid member, n = 8, known at this time; Cialdella and

Lopez, 1981), and if its present distribution suggested an origin for the subfamily, then Uruguay and southern Brazil would be indicated. All data at least suggest a concentration of present diversity within Chile, Argentina,

Uruguay, and Brazil. The concentrations of species in

Arnaldoa. Barnadesia. Chucmiraaa. and Dasvphvllum along the northern Andes are viewed as more recent northern migrations and morphological specializations.

Calyceraceae occur only in southern South America.

They inhabit coastal regions of southern Brazil and northern

Argentina, and the Andean range from the altiplano of southern Peru and Bolivia to Tierra del Fuego. One species of Nastanthus occurs in the Islas Malvinas (Falkland

Islands).

Of Goodeniaceae, approximately 377 of the 400 species are found in Australia. Scaevola has a distribution that extends into the Pacific islands and coastal regions of the

Atlantic and Indian Oceans. One species within the family, Goodenia pilosa. occurs in Indonesia, southern China, and the Philippines. 23

There is a definite focus, therefore, for Calyceraceae and Barnadesioideae in southern South America and Goodeniaceae largely in Australia. The data clearly support the close relationship of the former two families, which was demonstrated previously by morphological features.

Trends in Fossil Records of Asteraceae and Goodeniaceae

Although fragmentary, the fossil record for Asteraceae,

Calyceraceae and Goodeniaceae does offer some suggestions for their origins. Macrofossils, however, are disappointing. A number of isolated achenes (Knowlton, 1923; Berry, 1924; Segal, 1965; and Krassilov, 1973), leaves

(Cockerell, 1933; MacGinitie, 1933; Axelrod, 1937, 1950), and one compression fossil reported as a capitulum (Becker,

1969) have been attributed to Asteraceae but all inconclusively (Crepet and Stuessy, 1978). A literature search has failed to uncover macrofossil reports attributed to Calyceraceae or Goodeniaceae.

Microfossils provide better evidence for the origin of

Goodeniaceae and Asteraceae (no fossil pollen record for

Calyceraceae has yet been reported). Scaevola-tvoe pollen from the Oligocene similar to S. fructescens and described as Poluspissusites digitatus has been reported from Cameroon

(Salard-Cheboldaeff, 1978). The earliest report of

Asteraceae pollen has been from the Eocene (Romero, 1993). Five Asteraceae pollen types have been reported from the 24

Oligocene and Miocene. The earliest Asteraceae palynomorph of the Tubuliflorae-tvpe. occurs in the middle Oligocene of central Europe (Krutzsch, 1970), in Oligocene deposits of the United States (Leopold and MacGinitie, 1972) , and the upper Oligocene of Austria (Hochuli, 1978). Pollen grains assigned to the Liquliflorae-tvpe appear in the lower Miocene of Brazil (Pares Regali et al., 1974) and the lower

Miocene of Austria. Ambrosia-type pollen also first appears during the lower Miocene and has been reported from Brazil

(Pares Regali et al., 1974) and the United States (Leopold and MacGinitie, 1972). The first records of Artemisia-type pollen are from the middle Miocene of Tunis (Demarcq et al.,

1976), Hungary (Nagy, 1969), and the northwestern United

States (Gray, 1964). One uncertain record for Cirsium-tvpe pollen has been reported from the Paleocene of Germany

(Menke, 1976). These reports suggest at least an Oligocene origin for Asteraceae. The first appearance of a taxon in the fossil record, however, should not necessarily be considered equivalent to time of origin, unless fossil evidence shows transition from an ancestral type near the depositional environment (Muller,

1984). Axelrod (1970) and Stebbins (1974) argue that fossil pollen reflect when a group became established in lowland and coastal habitats since the preservation of pollen depends on: (1) quantity of pollen produced; (2) distance between source area and site of deposition; and (3) 25 pollination syndrome. Based on this argument, it is not surprising that an insect pollinated family like Goodeniaceae is represented by one record (Salard-

Cheboldaeff, 1978) of a palynomorph attributed to a coastal species, Scaevola fructescens. Lack of reports of

Calyceraceae is also supportive of Axelrod and Stebbin's view. Members of Calyceraceae inhabit microhabitats and form small, isolated, populations. Quantity of pollen produced by such populations would be reflected as a minor element of the pollen profile for a site. Furthermore, the majority of the family is pollinated by dipteran or lepidopteran visitors (i.e., not wind pollinated).

Unlike Calyceraceae and Goodeniaceae, Asteraceae pollen appeared contemporaneously on widely separated continents.

Turner (1977) believed this was sufficient evidence for a

Cretaceous origin for the family prior to major continental separations in the southern hemisphere. Muller (1984) indicates that most angiosperm pollen types appear suddenly in the fossil record and that transitional series in continuous stratigraphie sections are rare, perhaps due to mosaic character evolution. Since specialized pollen is easily recognized, unspecialized ancestral Asteraceae pollen may go unnoticed. Furthermore, pappus, capitulum, and other features of Asteraceae may have evolved prior to pollen specialization in the family, in which case more modern

Asteraceae pollen would seem to appear suddenly in the 26 fossil record. We believe, based on available direct fossil evidence, that Asteraceae probably appeared later than

Cretaceous and earlier than Oligocene. Even though the fossil record of Asteraceae and Goodeniaceae is incomplete, studies on general evolutionary trends of fossil angiosperm floral features (Friis & Crepet,

1987), fruits (Tiffney, 1984), and pollination syndromes (Crepet, 1984; Crepet and Friis, 1987; Crepet et al., 1991) can provide means to estimate time of origin for major groups with limited fossil records. By early Tertiary androecia had appeared with fused stamens, pseudanthia with sterile ray flowers (Euphorbiaceae), and flowers that were highly zygomorphic. Sympetalous flowers with long funnel- shaped corollas and narrow corolla tubes have been recovered from middle Eocene of Tennessee (Crepet, 1979). Crepet

(1984) also reported flowers with shorter funnel-shaped corollas and wider corolla tubes from the Eocene Green River

Formation. Crepet (1984) and Crepet and Friis (1987) have also assessed pollination syndromes in fossil flowers, but none as advanced as the stigmatic cup mechanism of

Goodeniaceae is known. These studies suggest that advanced pollen presentation mechanisms of Asteraceae, Calyceraceae, and Goodeniaceae probably evolved after middle Eocene.

Based on what is known about floral features of early

Tertiary angiosperms, therefore, a late Eocene time of origin for Asteraceae, Calyceraceae, and Goodeniaceae would 27 seem likely as there are no earlier records of tubular flowers aggregated into capitula, nor in any other feature diagnostic for any of the three families. One caution here is that no modern detailed studies have been completed for fossil floras in the southern hemisphere which obviously would be of value.

Tectonic events in the Southern Hemisphere

Tectonic history of continents in the southern hemisphere may have influenced evolution of Asteraceae, Calyceraceae, and Goodeniaceae in two ways. First, distribution of these families (Barnadesioideae and

Calyceraceae in South America, and Goodeniaceae in Australia) strongly suggests that separation of Antarctica from Australia and South America may explain present distributional patterns and provide estimates of earliest ages of origin for the families. Second, tectonic events are also strongly interactive with atmospheric and hydrospheric systems (Condie, 1989) which can lead to changing climates and also plant distributions. It is not our purpose to give a detailed description of Tertiary tectonic history of the southern hemisphere, but to discuss only those tectonic events which may have influenced origin and distribution of Asteraceae, Calyceraceae, and

Goodeniaceae. 28

The presence of a South America-Antarctica-Australia continental connection has been reflected in a wide range of terrestrial animal distributions including ratite birds (Keast, 1981), scorpionflies (Williams, 1981), annelids

(Jamieson, 1981), mole crickets (Tindale, 1981), scorpions

(Koch, 1981), scarib beetles (Howden, 1981), midges

(Brundin, 1966), and spiders and pseudoscorpions (Main,

1981). Thorne (1972, 1978, 1986), listed 48 genera and seven families of gymnosperms and angiosperms that have distributions restricted to temperate Australia and South America. These examples illustrate how strongly separation of Australia and South America from Antarctica influenced evolution and distribution of taxa in the southern hemisphere. In the case of Asteraceae, Calyceraceae, and

Goodeniaceae, it seems likely they, too, originated after continental separation.

The separation of Australia from Antarctica is dated from 53 million years ago in late Paleocene or early Eocene

(Cande and Mutter, 1982; Jones and Fitzgerald, 1984) to 43 million years ago during late Eocene (Weissal and Hayes, 1972; Weissel et al., 1977; Ha11am, 1980). Based on fossil marsupial distributional data, Marshall (1980) suggested that faunal interchange existed between Antarctica and Australia until early Eocene. 29 The connection and reseparation of South America to and from the Australia-Antarctica supercontinent are difficult to date. Western Antarctica consists of a number of smaller crustal blocks (e.g., Antarctic Peninsula and Ellsworth- Whitmore mountains) which during early to middle Jurassic were located 60° to 50° N along the western edge of eastern Antarctica (Grunow et al., 1987). Connection of the

Antarctic Peninsula to South America to form a continuous

Andean Cordillera occurred after this time. Exact timing is problematic due to problems in reconstruction of areas which are composites of crustal blocks: (1) it is difficult to correlate paleomagnetic data from such blocks with normal­ sized plate tectonic scenarios due to rotational movement

(Dalziel, 1983; Cox and Hart, 1986); (2) suture zones can often be difficult to identify; and (3) geometry of smaller crustal blocks may have been altered as a result of deformation. In regard to Antarctica specifically, there is the additional logistic problem in an area covered by a continental glacier of obtaining oriented core samples for paleomagnetic studies and collecting field data needed for correlations with geophysical data.

Breakup of the cordillera system between Antarctica and South America occurred when a spreading rift system collided with the western edge of the continents. This event, which resulted in the formation of the Scotia Arc, occurred late

Paleocene-early Eocene (Dalziel and Elliot, 1973; Dalziel, 30

1983) to Oligocene (Hallam, 1980; Craddock, 1982).

Findings of tectonic studies cited above fit with late

Eocene origin for Asteraceae, Calyceraceae, and

Goodeniaceae. Furthermore, since Australia was the first continent to rift from the supercontinent, we would expect more disparity in morphology and molecular characters of lineages isolated there. The unique floral features of Goodeniaceae (Carolin et al., 1992) and rearrangements of the chloroplast genome in Goodeniaceae (Downie and Palmer,

1992) suggest that the family is not as closely related to Barnadesioideae or Calyceraceae as these two South American groups are to each other.

Paleoclimatic Data

Recent paleoclimatic studies (McGowran, 1990; Parrish,

1990) have looked at paleoclimates as parts of an evolving exogenic system, which is defined as the interactions of the atmosphere, hydrosphere, or biosphere with the crust and mantle (Condie, 1989). The atmospheric part of the exogenic system appears to be changed by both major tectonic events and astronomical factors (Condie, 1989). Our interest focuses on what major changes occurred in the atmospheric exogenic system during Eocene and how these events may have influenced evolution of Asteraceae, Calyceraceae, and

Goodeniaceae. 31 Frakes (1979) used paleoclimatic indicators (abundances of coals and evaporates) and theoretical models of climatic evolution to estimate mean global temperature and precipitation regimes throughout geologic time. Based on

Frakes' curves, Paleocene and Eocene were the warmest periods of the Cenozoic. Frakes' curve for worldwide precipitation suggests a marked increase in humidity towards the Paleocene-Eocene boundary and throughout Eocene before diminishing in Oligocene. Both mean global temperature and precipitation curves illustrate variable climatic regimes after Eocene.

Parrish, Ziegler, and Scotese (1982) estimated global rainfall patterns using circulation maps constructed by

Parrish and Curtis (1982) for late Cretaceous and middle Eocene. Parrish (1987) incorporated rainfall patterns with paleogeographic reconstructions by Ziegler, Scotese, and

Barrett (1983) to illustrate climatic zonation for the Lutetian (middle Eocene). According to Parrish's estimations, central Australia possessed a to

Mediterranean climatic belt which was flanked by a humid subtropical to marine zone to the east. A tropical rainforest or savanna was believed to be present on the western edge of the continent which extended into the northern coast of eastern Antarctica. Eastern Antarctica was largely covered by a steppe to Mediterranean climatic belt. Parrish estimated that a humid subtropical or marine 32 climate existed over much of western Antarctica. Finally, most of southern South America had a climate similar to eastern Antarctica, with a northern arid zone isolating a second Mediterranean and rainforest belt at the southernmost tip of the continent.

The previously mentioned studies are based on qualitative paleoclimatic indicators. Paleotemperatures can also be estimated using such quantitative indicators as and isotype ratios present in calcareous, marine microfossils (McGowran, 1990). High carbon and low oxygen isotope ratios indicate warming events, while low carbon and high oxygen isotope ratios reflect cooling events. Measurements of both isotope ratios can be obtained for surface and bottom waters by segregating planktonic forms from benthic forms before analysis. Another advantage to these quantitative indicators is that the microfossils used to obtain the isotope ratios are powerful index fossils and provide a ready means of correlating the ratios with a chronologic scale.

Shackleton (1986) examined curves of carbon and oxygen isotope ratios for the Paleogene. Based on his data,

McGowran (1990) suggests that increased bottom water temperatures in early Eocene sharply dropped at the beginning of middle Eocene (49.5 mya) and sharply decreased a second time at the end of late Eocene (39.5 mya). Based on these quantitative indicators, global warming of early 33

Eocene was followed by two episodes of global cooling and an overall decrease in global temperature during Oligocene. McGowran (1990) looked at correlations of paleobiological signals (changes in extinction and rate of appearances curves or percentage of faunal change over geologic time) with tectonic and paleoclimatic data.

Included were data from benthic to planktonic marine organisms and terrestrial floras and European mammals. All these curves show evolutionary activity high during late Paleocene and early Eocene, relatively low in middle Eocene, and high again during late Eocene.

Muller (1981) constructed a curve showing rate of appearance of angiosperm families by estimating the number of new families recognizable in the fossil pollen record within a unit of geological time and dividing this value by the duration of each unit. Like curves presented in McGowran (1990), Muller's curve of rate of appearance of angiosperm families shows high rates in late Paleocene and early Eocene, followed by low rates in middle Eocene and a second increase in evolutionary activity in late Eocene. This repeating pattern indicates that some major earth historical events were influencing evolution of many types of organisms during Eocene. No doubt the ancestral lineage which gave rise to Asteraceae, Calyceraceae, and

Goodeniaceae was influenced by these events as well. 34

Another factor which could possibly have influenced major evolutionary activity, and particularly during Eocene, might be extraterrestrial impact events. The best physical evidences for an impact event are craters bounded by reversed stratigraphie sequences, spherulite particles produced at impact, high polymorphs of quartz, and actual

interplanetary debris. Impacts have generated considerable

interest since Alvarez et al. (1982) proposed a model for Cretaceous-Tertiary boundary extinctions. Hut et al. (1987)

found that estimated ages of impact structures suggest major impacts at 35, 65, and 99 million years ago, which correlate reasonably well with approximate 26 million-year intervals

for mass extinctions. These data suggest mass extinctions at the Cretaceous-Tertiary boundary, and as well as the latter portion of the late Eocene. Each of these events is

estimated to have caused 50-95% extinction of lower taxa (progressing from least to most ecologically tolerant) during a 1-3.5 million-year time span (Hut et al., 1987).

It appears, therefore, that late Eocene impact events,

along with tectonic activity and major paleoclimatic

changes, could have stimulated the rapid evolution of

Calyceraceae, Goodeniaceae, and Asteraceae. These factors may have particularly stimulated Asteraceae which have

subsequently colonized every continent on earth (except

Antarctica) and radiated into numerous tribes with more than

20,000 species— one of the most successful families of 35 angiosperms on earth.

ACKNOWLEDGEMENTS

We wish to thank: The American Society of Plant

Taxonomists for an herbarium travel grant (MLD), an Ohio State University Beatley Herbarium Award and Alumni Graduate

Student Research Award (both to MLD); a Grant-in-Aid from Sigma Xi (MLD); and The National Geographic Society (grant

No. 4459-91; TFS). Conversations with the following individuals concerning Asteraceae, Calyceraceae, or Goodeniaceae were extremely helpful: K. Bremer, A.L.

Cabrera, A. Cronquist, V. Funk, E. Harris, T. Lammers, C.

Marticorena, R. Olmstead, and H. Robinson. 36

REFERENCES

Alvarez, L. W., Alvarez, W., Asaro, F., and Michel, H. V.,

et al. (1982). Current status of the impact theory for the terminal Cretaceous extinction. Geol. Soc. Amer. Spec. Paper 190;305-315.

Axelrod, D. I. (1937). A Pliocene flora.from the Mount

Eden beds, California. Publ. Carnegie Inst. Wash. 476.

. (1950). Further studies of the Mount Eden flora.

southern California. Publ. Carnegie Inst. Wash. 476.

(1970). Mesozoic paleogeography and early angiosperm history. Bot Rev. 36: 277-319.

Becker, H. F. (1969). Fossil plants of the Tertiary

Beaverhead Basins in southwestern Montana.

Palaeontographica 127: 1-142.

Bentham, G. (1873). Notes on the classification, history, and geographic distribution of Compositae. J. Linn. Soc. Bot. 13: 335-577. 37 Berry, E. W. (1924). The middle and upper Eocene floras of southeastern North America. U.S. Geol. Surv. Prof. Paper 92.

Bremer, K. (1992). Ancestral areas: a cladistic

reinterpretation of the center of origin concept. Syst. Biol. 41: 436-445.

. (1993a). Intercontinental relationships of African

and South American Asteraceae: A cladistic

biogeographic analysis. In P. Goldblatt (editor). Biological Relationships between Africa and South

America, pp. 105-138. Yale University Press, New Haven.

_. (1993b). Asteraceae: Cladistics and Classification. Timber Press, Portland.

and Jansen, R. K. (1992). A new subfamily of the

Asteraceae. Ann. Missouri Bot. Gard. 79: 414-415.

Brown, R. (1817). Observations on the natural family of plants called Compositae. Trans. Linn. Soc. 12: 76- 142. 38 Brundin, L. Z. (1966). Transantarctic relationships and

their significance as evidenced by chironomid midges.

K. Sven. Vetenskapsakad. Handl. 11; 1-472.

Cabrera, A. L. (1959). Revision del género Dasvphvllum

(Compositae). Rev. Mus. La Plata (new ser.) 9: 21-

100.

_____ . (1977). Mutisieae— systematic review. In V. H.

Heywood, J. B. Harborne & B. L. Turner (editors). The

Biology and Chemistry of the Compositae, pp. 1039-1066. Academic Press, London.

Cande, S. C. and Mutter, J. C. (1982). A revised

identification of the oldest seafloor spreading

anomalies between Australia and Antarctica. Earth

Planet Sci. Lett. 58: 151-160.

Carlquist, S. (1969). Wood anatomy of Goodeniaceae and the

problem of insular woodiness. Ann. Missouri Bot. Gard. 56: 358-390.

Carolin, R. C. (1959). Floral structure and anatomy in the

family Goodeniaceae. Proc. Linn. Soc. N.S.W. 84: 242-

255. 39

(1966). Seeds and fruit of the Goodeniaceae. Proc. Linn. Soc. N.S.W. 91:58-83.

(1967). The concept of the inflorescence in the

order . Proc. Linn. Soc. N.S.W. 92: 7-26.

(1970). The trichomes of Goodeniaceae. Proc. Linn. Soc. N.S.W. 96: 8-22.

(1978). The systematic relationships of Brunonia.

Brunonia 1: 9-29.

(1990). Nomenclatural notes, new taxa and the systematic arrangement in the genus Scaevola

(Goodeniaceae) including synonyms. Telopea 3: 477-515.

Morrison, D. A. and Rajput, T. (1992). Flora of

Australia 35. Australian Government Publishing

Service, Canberra.

Cassini, H. (1816). Acicaroha (Bot.). Diet. Sci. Nat. 1, Suppl: 32-33.

Chung, I. (1965). Revision of Barnadesia (Compositae—

Mutisieae). Chicago: published by author. 40

Cialdella , A. M. and Lopez de Riesling, A. G. (1981). Cariologla de Schlechtendalia luzulaefolia

(Compositae). Darwiniana 23:357-360.

Cockerell, T. D. A. (1933). A fossil goldenrod. Torreya

33: 72.

Condie, K. C. (1989). Plate Tectonics and Crustal Evolution. Pergamon, New York.

Cox, A. and Hart, R. B. (1986). Plate Tectonics: How it

works. Blackwell Scientific, Oxford.

Craddock, C. (1982). Antarctica and Gondwanaland. In C. Craddock (editor). Antarctic Geoscience, pp. 3-13.

Univ. Wisconsin Press, Madison.

Crepet, W. L. (1979). Some aspects of the pollination

biology of Middle Eocene angiosperms. Rev. Palaeobot. Palynol. 25: 241-252.

. (1984) . Advanced (constant) insect pollination

mechanisms: pattern of evolution and implications vis-

a-vis angiosperm diversity. Ann. Missouri Bot. Gard.

71: 607-630. 41 and Stuessy, T. F. (1978). A reinvestigation of the

fossil Vicmiera cronouistii (Compositae). Brittonia 29: 137-153.

and Friis, E. M. (1987). The evolution of insect

pollination in angiosperms. In E. M. Friis, W. G.

Chaloner & P. R. Crane (editors). The Origins of

Angiosperms and their Biological Consequences, pp. 181- 202. Cambridge Univ. Press, Cambridge.

_, _____ and Nixon, K. (1991). Fossil evidence for the evolution of biotic pollination. Phil. Trans. Roy. Soc. bond. B33: 187-195.

Cronquist, A. (1955). Phylogeny and of the Compositae. Amer. Midi. Naturalist 53: 478-511.

_____ . (1977). The Compositae revisited. Brittonia 29: 137-153.

(1981). An Integrated System of Classification of

Flowering Plants. Columbia Univ. Press, New York. 42 Dalziel. I. W. (1983). The evolution of the Scotia Arc; A

review. In R. L. Olivier, P. R. Jones & J. B. Jago

(editors). Antarctic Earth Science, pp. 238-288. Cambridge Univ. Press, Cambridge.

_____ and Elliot, D. H. (1973). The Scotia Arc and

Antarctic margin. In A. E. M. Mairn & R. G. Stehl

(editors). The Ocean Basins and Margins, Vol. l. The

South Atlantic, pp. 171-246. Plenum Press, New York.

Demarcq, G., Méon-Vilain, H., Miquet, R. and Kujawski, H.

(1976). Un bassin paralique Néogène: celui de Skanes-

Monastir (Tunisie orientale). Notes Serv. Géol.

Tunisie 42: 97-147.

DeVore, M. L. (1991). Acicarpha tribuloides in eastern

North America. Rhodora 93: 26-35.

(1993). The systematics of Acicarpha

(Calyceraceae). Ohio J. Sci. 92:41. Abstr.

Downie, S. and Palmer, J. (1992). Restriction site mapping

of the chloroplast DNA inverted repeat: a molecular

phylogeny of the Asteridae. Ann. Missouri Bot. Gard.

79: 266-283. 43

Ezcurra, C. (1985). Revision del genéro Chucmiraaa

(Mutisieae, Compositae). Darwiniana 26: 219-284.

and Crisci, J. V. (1987). Relaciones fenéticas entre las especies del género Chucruiraaa ( Compos itae- Mutisieae): un analisis numérico. Darwiniana 28: 219- 229.

Frakes, L. A. (1979). Climates Throughout Geologic Time.

Elsevier, Amsterdam.

Friis, E. K. and Crepet, W. L. (1987) . Time of appearance

of floral features. In E. M. Friis, W. G. Chaloner &

P. C. Crane (editors). The Origins of Angiosperms and

their Biological Consequences, pp. 145-179. Cambridge

Univ. Press, Cambridge.

Gamerro, J. C. (1986). Morfologia del polen de Huarpea y

su relaciôn con Barnadesia (Mutisieae, Compositae). Darwiniana 26: 219-284.

Gray, J. (1964). Northwest American Tertiary palynology:

the emerging picture. In L. M. Cranwell (editor). Ancient Pacific floras, pp. 21-30. Univ. Hawaii Press,

Honolulu. 44 Grunow, A. M., Dalziel, I. W. D. and Kent, D. V. (1987). Ellsworth-Whitmore Mountains crustal block, western

Antarctica: New paleomagnetic results and their tectonic significance. In Gondwana Six: Structure,

Tectonics, and Geophysics, pp. 161-171. Geophysical

Monograph Series Vol. 40. Washington DC: American Geophysical Union.

Hallam, A. (1980). A reassessment of the fit of Pangaea

components and the time of their breakup. In D. W.

Strangway (editor). The Continental Crust and its

Mineral Deposits, pp. 375-387. Geol. Assoc. Canada Spec. Paper 20.

Hansen, H. V. (1991a). Phylogenetic studies in Compositae

tribe Mutisieae. Opera Bot. 109:1-50.

_____ . (1991b). SEM-studies and general comments on pollen in tribe Mutisieae (Compositae) sensu Cabrera. Nord.

J. Bot. 10:607-623.

_. (1992). Studies in the Calyceraceae with a

discussion of its relationship to Compositae. Nord. J.

Bot. 12: 63-75. 45

Hochuli, P. A. (1978). Palynologische Untersuchungen im Oligozan und Untenniozan der Zentralen und Westlichen

Paratethys. Beitr. Palaont. Ost. 4: 1-132.

Howden, P.A. (1981). Zoogeography of some Australian

Coleoptera as exemplified by the Scarabaeoidea. In A.

Keast (editor). Ecological Biogeography of Australia, vol. 2, pp. 1007-1035. W. Junk, The Hague.

Hut, P. et al. (1987). Comet showers as a cause of mass

extinctions. Nature 329; 118-127.

Jamieson, B. G. M. (1981). Historical biogeography of Australian . In A. Keast (editor).

Ecological Biogeography of Australia, vol. 2, pp. 885-

921. W. Junk: The Hague.

Jansen, R. K. and Palmer, J. D. (1987). Chloroplast DNA

from lettuce and Barnadesia (Asteraceae): structure, gene localization, and characterization of a large

inversion. Curr. Genet. 11: 553-564.

and _____. (1988). Phylogenetic implications of

chloroplast DNA restriction site variation in the

Mutisieae (Asteraceae). Amer. J. Bot. 75: 751-764. 46 Jones, J. B. and Fitzgerald, M. J. (1984). Extensive

volcanism associated with the separation of Australia

and Antarctica. Science 226: 346-348.

Jussieu, A. L. (1803) . Mémoire sur I'Acicaroha et le

Boopis. deux genres nouveaux de plantes de la famille

des Cinarocephales. Ann. Mus. Nat. Hist. 2: 345-350.

Keast, A. (1981) . The evolutionary biogeography of Australian birds. In A. Keast (editor). Ecological

Biogeography of Australia, vol. 2, pp. 1587-1635. W. Junk, The Hague.

Knowlton, F. H. (1923). Revision of the flora of the Green

River Formation, with descriptions of new species. U.

S. Geol. Surv. Prof. Paper 131: 133-182.

Koch, L. E. (1981). The scorpions of Australia: aspects of

their ecology and zoogeography. In A. Keast (editor).

Ecological Biogeography of Australia, vol. 2, pp. 873-

884. W. Junk, The Hague.

Krassilov, V. A. (1973). Mesozoic plants and the problem

of angiosperm ancestry. Lethaia 6: 163-178. 47 Krutzsch, W. (1970). Die stratigraphisch verwertbaren Sporen- und Pollenformen des mitteleuropaischen

Alttertiars. Jahrb. Geol. Bot. 3: 309-379.

Leins, P. and Erbar, C. (1990). On the mechanisms of

secondary pollen presentation in the Campanulales-

Asterales-complex. Botan. Acta 103: 87-92.

Leopold, E. B. and MacGinitie, H. D. (1972). Development

and affinities of Tertiary floras in the Rocky Mountains. In A. Graham (editor). Floristics and

Paleofloristics of and Eastern North America, vol.

2, pp. 147-200. Elsevier, Amsterdam.

MacGinitie, H. D. (1933). The Trout Creek flora of

southeastern Oregon. Publ. Carnegie Inst. Wash. 416.

Main, B. Y. (1981). A comparative account of the

biogeography of terrestrial invertebrates in Australia: some generalizations. In A. Keast (editor).

Ecological Biogeography of Australia, vol. 2, pp. 1055

1077. W. Junk, The Hague. 48

Marshall, L. G. (1980). Marsupial paleobiogeography. In

L. L. Jacobs (editor). Aspects of Vertebrate History:

Essays in Honor of Edwin Harris Colbert, pp. 345-386.

Museum of Northern Arizona Press, Flagstaff.

McGowran, B. (1990). Fifty million years ago. Amer. Scientist 78: 30-39.

Menke, B. (1976). Pliozane und altestguartare Sporen- und

Pollenflora von Schleswig-Hollstein. Geol. Jahrb. A32: 3-197.

Muller, J. (1981). Fossil pollen records of extant

angiosperms. Bot. Rev. 47: 1-42.

. (1984). Significance of fossil pollen for

angiosperm history. Ann. Missouri Bot. Gard. 71: 419- 443.

Nagy, E. (1969). Palynological elaborations of the Miocene

layers of the Balcony Mountains (Valpalota) of Hungary.

Acta Bot. 8: 153-163.

Nowicke, J. W. and Skvarla, J. J. (1974). A palynological

investigation of the genus Tournefortia

(Boraginaceae). Amer. J. Bot. 61: 1021-1036. 49 Olmstead, R. G., Jansen, R. K., Michaels, H. J., Downie, S.

R. and Palmer, J. D. (1990). Chloroplast DNA and phylogenetic studies in the Asteridae. In S. Kawano

(editor). Biological Approaches and Evolutionary

Trends in Plants, pp. 119-134. Academic Press, London.

_, Michaels, H. J., Scott, K. M. and Palmer, J. D.

(1992). Monophyly of the Asteridae and identification of their major lineages inferred from DNA sequences of

rbcL. Ann. Missouri Bot. Gard. 79: 249-265.

Pares Regali, M. de S., Uesugui N. and Santos, A. de S. (1974). Palinologia dos sedimentos meso-cenozoicas do

Brasil (I, II). Bol. Teen. Petrobras, Rio de Janeiro. 17: 177-191, 263-301.

Parra, 0. and Marticorena, C. (1971). Granos de polen de

plantas chilenas. II. Compositae-Mutisieae. Gayana 21: 3-107.

Parrish, J. T. (1987). Global palaeogeography and

palaeoclimate of the Late Cretaceous and Early

Tertiary. In E. M. Friis, W. G. Chaloner & P. R. Crane (editors). The Origins of Angiosperms and their

Biological Consequences, pp. 51-74. Cambridge

University Press, Cambridge. 50 (1990). Gondwanaland paleogeography and

paleoclimatology. In T. N. & E. Taylor (editors).

Antarctic Paleobiology, pp. 15-26. Springer-Verlag, New York.

_ and Curtis, R. L. (1982). Atmospheric circulation. upwelling, and organic-rich rocks in the Mesozoic and

Cenozoic. Palaeogeogr. Palaeoclim. Palaeoecol. 40: 31-

66.

_, Ziegler, A. M. and Scotese, C. R. (1982). Rainfall

patterns and the distribution of coals and evaporates in the Mesozoic and Cenozoic. Palaeogr. Palaeoclimat.

Palaeoecol. 40: 67-101.

Pesacreta, T. C., Sullivan, V. I. and Hasenstein, K. H.

(1993). The connective base of Cirsium horridulum

(Asteraceae): description and comparison with the visoelastic filament. Amer. J. Bot. 80:411-418.

Pontiroli, A. (1963). Flora Argentina, Calyceraceae. Rev. Mus. La Plata, sec. Bot. 9: 175-241.

Raven, P. H. and Axelrod, D. I. (1974). Angiosperm

biogeography and past continental movements. Ann. Missouri Bot. Gard. 61:539-673. 51

Robinson, H. (1987). Some suggestions regarding the significance of chloroplast DNA variation in the

Asteraceae. Phytologia 65:316-324.

Romero, E. G. (1993). South American paleofloras. In P.

Goldblatt (editor). Biological Relationships between

Africa and South America, pp. 62-85. Yale University Press, New Haven.

Salard-Cheboldaeff, M. (1978). Sur la palynoflore

Maestrichtienne et Tertiaire du bassin sédimentaire

littoral du Cameroun. Pollen et Spores 20: 215-260.

Segal, R. (1965). New fossil fruit (Compositae) from the

Pliocene of western Kansas. Amer. Midi. Nat. 73: 430-

432.

Shackleton, N. J. (1986). Paleogene stable isotope events.

Palaeogeog. Palaeoclimat. Palaeoecol. 57: 91-102.

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

(1977). Pollen morphology in the Compositae and in

morphologically related families. In V. H. Heywood, J.

B. Harborne & B. L. Turner (editors). The Biology and

Chemistry of the Compositae, pp. 141-248. Academic

Press, London. 52 Small, J. (1919). The origin and development of the Compositae. Reprint of New Phytologist No. 11.

Stebbins, G. L. (1974). Flowering Plants: Evolution Above the Species Level. Harvard University Press,

Cambridge.

Stuessy, T. F. and Sagâstegui A., A. (1993). Revision de

Arnaldoa (Compositae, Barnadesioideae), género endémico

del norte del Peru. Arnaldoa 1: 9-21.

and Sang, T. (1993). Phylogeny of Barnadesioideae

and early evolution of Compositae. Ohio J. Sci. 92:41. Abstr.

Thorne, R. F. (1972). Major disjunctions in the geographic

ranges of seed plants. Quart. Rev. Biol. 47: 365-411.

. (1978). Plate tectonics and angiosperm

distribution. Notes Royal Bot. Gard. Edinburgh 36: 297-315.

_. (1986). Antarctic elements in Australasian

rainforests. Telopea 2:611-617, 53

Tiffney, B. (1984). Seed size, dispersal syndromes, and

the rise of the angiosperms: evidence and hypothesis. Ann. Missouri Bot. Gard. 71: 551-576.

Tindale, N. B. (1981). The origin of Lepidoptera relative

to Australia. In A. Keast (editor). Ecological

Biogeography of Australia, pp. 955-976. W. Junk, The

Hague.

Turner, B. L. (1977). Fossil history and geography. In V. H. Heywood, J. B. Harborne & B. L. Turner (editors).

The Biology and Chemistry of the Compositae, pp. 21-39.

Academic Press, London.

Weissal, J. K. and Hayes, D. E. (1972). Magnetic anomalies

in the Southeastern Indian Ocean. In D. E. Hayes (editor). Antarctic Oceanography II: Australia-New

Zealand Sector, pp. 234-249. Antarctic Res. Ser., Vol.

19. American Geophysical Union, Washington, D.C.

_____ , and Herron, E. M. (1977). Plate tectonic

synthesis; the displacements between Australia, New

Zealand, and Antarctica since Late Cretaceous. Marine Geol. 25: 231-277. 54 Williams, W. D. (1981). Aquatic insects: an overview. In

A. Keast (editor). Ecological Biogeography of

Australia, vol. 2, pp. 1213-1229. W. Junk, The Hague.

Ziegler, A. M., Scotese, C. R. and Barrett, S. F. (1983). Mesozoic and Cenozoic paleogeographic maps. In P.

Brosche & J. Sundermann (editors). Tidal Friction and

the Earth's Rotation, vol. 2, pp. 240-252. Springer- Verlag, Berlin. CHAPTER II CYTOLOGY OF CALYCERACEAE Introduction

Calyceraceae are a family of six genera and approximately 50 species of temperate and subtropical

southern South America. Although in relative obscurity for more than a century, Calyceraceae have attracted renewed

interest due to suggestions that it may be closely related to the Compositae (Olmstead et al., 1992; DeVore & Stuessy, in press).

Cytological data have been helpful in resolving taxonomic and evolutionary problems in Compositae (Solbrig,

1977; Turner et al., 1979), Goodeniaceae (Peacock, 1963),

and Lobeliaceae (Lammers, 1988), but few studies have been done on Calyceraceae. The sparsity of cytological information from the family may be attributable to three

reasons: (1) difficulty of obtaining meiotic counts from

collections of species inhabiting high elevations within the

Andean chain; (2) difficulty of germinating achenes of

Andean species and growing many members of the family in

greenhouses; and (3) lack of comprehensive systematic

studies. Previous chromosome numbers reported in the family

include: 2n = 16, 30, 36, and 42; n = 8, 15, 18, and 21

55 56 (Sugiiira, 1936, 1937; Moore, 1983; Rahn, 1960; Rodrigues et al., 1977; and Turner, 1977). Based on these reports,

Stebbins (1977) suggested three possible base numbers of x = 7, 8, 9.

This report utilizes previous and new cytological data to assess generic delimitations of Calycera and Nastanthus, to compare present cytological data from Calyceraceae with counts obtained from related families, and to comment on the presumptive base number of the family.

Materials and Methods

Collections studied of species of Boopis, Calycera, and

Nastanthus are listed in Table 1. Meiotic chromosome counts were obtained from florets dissected from heads collected in the field, fixed in 3:1 (ethanol:acetic acid) for 16-18 hrs,

stored in 70% ethanol at -5°C, and stained with acetocarmine. Young anthers were squashed and made semi­ permanent using Hoyer's mounting medium. Camera lucida drawings were made of preparations.

Heads were obtained and fixed from several individuals

from each population studied. Voucher specimens were collected from each population (Table 1) and deposited in

CONC, LP, OS, and OSH. 57

Table 1. Previous chromosome reports and new chromosome counts (and voucher specimens) from taxa of Calyceraceae. Numbers in parentheses indicate number of cells studied.

Taxon n 2n Reference or voucher

ASlgarghft tribuloides 16 Sugiura, 1936; 1937 Juss.

a. spathulata R. Br. 8 Rodrigues, 1977

Boopis anthemoides 15 Rahn, 1950 Juss.

australis Decne. 36 Moore, 1983a

S> gracilis Phil. 18 36 Rahn, 1960; Argentina, Mendoza: 20 km NW of El Sosneado on Route 220, DeVore 1747 (3); 5 km. S of Hotel Termas, DeVore 17SS (2).

Calvcera calcitrapa 21 Turner, 1971 Griseb.

crassifolia (Hiers) 21 Argentina, Buenos Aires: Cabo Hicken San Antonio, DeVore & Delucci (13). 58

Tzüsle 1 (continued) *£. ervnqioides Kemy 22 Chile, Santiago: Lagunillas, DeVore 1513 (7).

£. herbacea Cav. 21 42 Covas and Schanck, 1947; Rahn, 1960; Argentina, Hendosa: 2 km. E of Puente del Inca, DeVore 1721 (4); DeVore 1722 (6).

*£. leucanthema (Poepp. 17 Chile, Talca: 10 km. S of ex Hiers) Ktintze Vilches Alto, DeVore 1486 (14). Bio-Bio: Village of Recinto, DeVore 1431 (14).

*£. spinulosa C. Gillies 21 Argentina, Mendoza: 10 km. NE ex Hiers of Los Arboles, DeVore 1729 (6); 3 km. NE of Los Arboles, DeVore 1765 (5); 3 km. NE of Rincon del Atuel, DeVore 1765 (4).

*S. svnpaqanthera (Ruiz 13 Chile, Xalleco: Piedra del & Pav.) O. Kuntze Aguila, DeVore & Baeza 1611

(9). 59

Table 1 (continued) N. falklandicus D. M. 20-21 Moore, 1967, 1968 Moore

IL. spathulatus 21 42 Rahn, i960; Argentina, (Phil.) Hiers Xendosa: Las Cuevas, DeVore 1723 (4)

First report for teuton. FIGURE 1. Camera lucida drawings of meiotic chromosomes from species of Calyceraceae. 1-1, Boopis gracilis, n = 18.

1-2, Calycera crassifolia, n = 21. 1-3, C. eryngioides, n = 21-22. 1-4, C. herbacea, n = 21. 1-5, C. leucanthema, n =

17. 1-6, C. spinulosa, n = 21. 1-7, C. sympaganthera, n

= 13. 1-8, Nastanthus andina, n = 21. 1-9, JT. spathulatus, n = 21. All same scale. All diakinesis except telophase I in Fig. 1-1 and 1-6.

60 61

• »o

. t . r *

lOuni

. ' . 4 «r V " ^ I % oi'-O «• *.'* 4 ■■■ ^ ""a

a ;.* iKv L ®

Figure i 62 Results

Meiotic chromosome numbers of 15 species belonging to four genera are given in Table 1. Numbers for six species represent first reported counts, while the remaining three new meiotic counts confirm previously reported numbers. Regular meiosis with normal bivalent formation was observed in all cases (Fig. 1). Bivalents in diakinesis ranged from

1.8 jum long in Nastanthus andinus to 3.0 /xm in Calycera sympaganthera (Fig. 1). Fragments and bridges were clearly visible during Anaphase II in Calycera eryngioides , (DeVore

1513), C. herbacea {DeVore 1722), C. leucanthema (DeVore 1431) , and C. sympaganthera (DeVore & Baeza 1611) .

Cvtolooy in relation to morphology and distribution of Acicarpha.

The present survey found no new diploid species within

Calyceraceae. Diploid members of Calyceraceae have only been reported as n = 8 in Acicarpha spathulata (Rodrigues et al., 1977) and 2n = 16 in Acicarpha tribuloides (Sugiura,

1936, 1937). Both of these counts indicate that Acicarpha, as classically delimited (Jussieu, 1803), has a base number of X = 8.

The cytological distinctness of Acicarpha correlates with both morphological and distributional differences found within the genus. The presence of such autapomorphic features as (1) centripetally developing heads; (2) connate 63 upper portions of filaments; (3) thickened filaments adjacent to the connective; and (4) expanded connective bases, indicate that Acicarpha is morphologically divergent from other genera within Calyceraceae. Acicarpha also has a distribution centered in southern Brasil, Uruguay, and northern Argentina, whereas other genera within the family (Boopis, Calycera, Gamocarpha, Moschopsis, and Nastanthus) have distributions centered along the Central and Southern

Andean Chains.

Cvtolocrv and relationships within Calycera.

New counts for Calycera, along with previous reports

(Covas and Schnack, 1947; Rahn, 1960; Turner, 1973),

indicate that Argentinean species (C. calcitrapa, C.

crassifolia, C. herbacea, and C. spinulosa) are all

polyploid with n = 21. These cytological data are congruent

with uniform morphological features found in Argentinean

species of Calycera. Members of the Argentinean species group are characterized by the following morphological

characters (1) large (> 4 cm in diam), globular, receptacles; (2) long (6-9 mm), constricted corolla tubes;

(3) filamental glands positioned near the base of the

anthers; and (4) anthers lacking connective bases, tails, or

apical appendages. 64

In contrast to Argentinean species of Calycera, chromosome numbers are more variable in Chilean species counted with n = 21-22 (C. eryngioides), n = 17 (C. leucanthema), and n = 13 (C. sympaganthera). The Chilean species are readily distinguished from one another by autapomorphic features, such as tetramerous flowers, filamental gland position, anthers with tails, and apical anther appendages. Although some of these characters are shared by some of the species, not one of these features is found in all Chilean taxa of Calycera species. Chromosome counts from these species were not useful in providing any additional insights on their relationships.

Other Andean genera, such as Pappoholus Blake (Panero, 1992), are not defined by clear synapomorphies, but are clearly monophyletic. Likewise, Calycera illustrates the potential danger of defining holophyletic taxa (sensu

Ashlock, 1971) by a set of synapomorphic features shared by all members of a group. In Calycera, it is best to delimit species on the bases of characters found in the majority of the species. There is no need to break up taxa into two or more genera simply to meet the requirements of holophyly

(Charig, 1982; Mayr and Ashlock, 1991).

In light of known chromosome numbers within Calycera, and morphological differences between Argentinean and

Chilean groups of species, one could make a case for segregating Calycera into two genera. In fact, Miers (1860) 65

erected the genus Anomocarpus to include almost all the

Chilean species of Calycera, except C. sympaganthera, based on the presence of the following characters; (1) single, axillary heads on short peduncles; (2) thin, membranaceous, cup-shaped, five-lobed involucres; (3) short receptacles;

and (4) shorter calycine lobes. Argentinean species and C.

sympaganthera were maintained within Calycera.

Current taxonomic work with Calycera (Chapter 4) recognizes Argentinean and Chilean species groups as

distinct subgenera, rather than genera. Although there are

clear and consistent differences between them, both

Argentinean and Chilean species of Calycera are still united

in having: (1) heads composed of cymose subunits; (2)

dimorphic achenes; 3) achenes lignified; and 4) pollen with intercolpar concavities. The combination of these four

character states serves as evidence that Calycera is a monophyletic genus and should be maintained taxonomically.

Nastanthus and its relationship with Calycera

Counts obtained from Nastanthus, as well as the Argentinean Calycera species, are all n = 21. Besides

having the same chromosome numbers, Nastanthus and the

Argentinean Calycera species share the following

morphological characters: (l) glands positioned distally

inside the filamental column; (2) corolla tube usually over

6 mm long and constricted; (3) globular receptacles; and (4) 66 anthers without collars or tails. The presence of these

four shared character states, as well as identical chromosome numbers in Nastanthus and Argentinean Calycera species, suggest a close evolutionary relationship between these two genera.

Some evolutionary insights can be gained by pointing out the differences and similarities in reproductive and vegetative features present in Argentinean Calycera species and Nastanthus at different stages of their life histories. Unlike mature Argentinean Calycera species, Nastanthus maintains a rosette growth at maturity, and produces soft, not highly lignified, achenes which lack elongate calycine

lobes. Argentinean Calycera species form rosettes in the

juvenile stage of their lifespan which develop branching stems in the flowering stage. In the field, the juvenile

growth forms of Calycera and Nastanthus are nearly

indistinguishable (DeVore, pers. observ.). The vegetative morphological similarities between the juvenile Argentinean

species of Calycera and Nastanthus, as well as the presence

of similar floral morphologies and identical chromosome

numbers suggest a close relationship. Since Nastanthus maintains the juvenile features of a closely related taxon,

one would suspect that heterochrony may have been a factor

in the evolution of Nastanthus. 67 Heterochrony has been defined as evolutionary change in the time of development of an organism (Gould, 1977).

One morphological expression of heterochrony is paedomorphosis. An adult organism which expresses paedomorphosis possesses characteristics typical of its ancestor's juvenile form. Paedomorphosis can arise via neoteny or progenesis (Alberch et al., 1979), retardation of vegetative or acceleration of reproductive development, respectively. An ancestor to Nastanthus, similar to the

Argentinean Calycera species, may have been capable of accelerating timing of flowering and fruiting, while still in the juvenile, vegetative growth form. This would, therefore, have been progenesis.

Paedomorphosis has been suggested as enabling a group to change from one specialization and habitat to another

(Takhtajan, 1959; 1991). Progenesis may have been correlated with several adaptive, morphological changes.

The production of smaller diaspores, without lignified, elongated, calycine lobes could have resulted in a considerable reduction of photosynthate invested in fruit production. Such changes in fruit size and investment of photosynthate would decrease the amount of time needed for the completion of reproductive cycles at higher elevations

(3000-4000 m) characterized by shorter growing seasons.

Another feature which may have enabled Nastanthus to speciate at high elevations is the development of extensive, 68 water-rich parenchyma within the stem and fused capitula.

One would expect that Nastanthus, like Espeletia, may be able to utilize this stored water during morning hours at high elevations (3000-4000 m.) when water is frozen and unavailable for plant use (Monasterio, 1986). Character changes (reduced diaspore size, shorter flowering and fruiting times, water-rich parenchyma cells) through progenesis could possibly explain the development of an adaptive strategy which permitted Nastanthus to radiate within the highest vegetation zones of the southern Andes.

Estimate of Ancestral Base Number of Calvceraceae

New chromosome counts, in combination with data from current systematic studies (DeVore, in prep.) and cytological trends within Campanulales (hammers, 1992), can be used to estimate an ancestral base number for

Calyceraceae. Counts reported here support Stebbins' (1977) general viewpoint that the potential base numbers within the family are % = 7, 8, or 9. Cytological trends present in related families may provide clues as to which of the possible base numbers of Calyceraceae might be ancestral.

Sequence studies using rJbcL have suggested that Calyceraceae are most closely related to Goodeniaceae and Asteraceae

(Olmstead et al., 1992). Both Raven (1975) and Solbrig

(1977) estimated the base number of Asteraceae as x = 9. hammers (1992) suggested that since Barnadesioideae are 69 considered to be primitive members of Asteraceae (Jansen &

Palmer, 1988), the ancestral base number of that family might be % = 8 or 9.

Peacock (1963) conducted a thorough survey of

chromosome numbers of Goodeniaceae. He estimated that potential ancestral base numbers for the family were x = S or 9 and believed n = 7 was derived through descending aneuploidy. If one carefully correlates Peacock's cytological data (1963) with the results of Carolin's cladistic analysis (1978), it appears that those genera estimated to be primitive within Goodeniaceae ( Smith, Brunonia Smith, Dampiera R. Brown, and Lechenaultxa

R. Brown) all are n = 9. The correlation of n = 9 with these primitive genera suggests that this could be the ancestral base number for Goodeniaceae.

Since Goodeniaceae is the outgroup for Calyceraceae, then one could argue that the ancestral number of

Calyceraceae may be x = 8 or 9. If Acicarpha (n = 8) is ancestral (DeVore, in prep.) to either Calycera or

Nastanthus (n = 7), then it appears that descending aneuploidy may have occurred. The remaining genera within the family (Boopis, Gamocarpha, and Moschopsis), have not been thoroughly surveyed. Some counts for Boopis (Table 1) could be interpreted as being based on n = 9. More counts are needed from Boopis, as well as unsurveyed genera within the family (Gamocarpha and Moschopsis), to clarify 70 cytological patterns within the entire family.

Cytology and bioaeocrraphv

Cytological data can also be correlated with some aspects of distributional history of Calyceraceae. The polyploid species Boopis australis (2n = 36) occurs in Tierra del Fuego, in the southernmost distribution of the family. Likewise, B. gracilis (2n = 36) is found in glaciated regions at high elevations (1800-3000) centered toward the Argentinean border. Argentinean Calycera species and Nastanthus (both polyploids) all inhabit recent environments (moraines, alluvial deposits, and coastal dunes) which were made available directly, or indirectly, by

Quaternary glaciations (DeVore, per. observ.). One would suspect that the highest ploidy levels within Calyceraceae would be found in the most recent habitats resulting from

Quaternary glaciations (Stebbins, 1971; Lokki and Saura,

1980). Moore (1983a,b) speculated that few genera underwent spéciation in the extreme southern Andes since the flora was eradicated during Quaternary glaciations. Additional chromosome counts from other Calyceraceae species endemic to Tierra del Fuego would be invaluable in testing this hypothesis. 71

Literature Cited

Alberch, P., S. J. Gould, G. F. Oster, & D. B. Wake, 1979.

Size and shape in ontogeny and phylogeny. Paleobiology

5: 297-317.

Ashlock, P. D. 1971. Monophyly and associated terms.

Syst. Zool. 20: 430-438.

Carolin, R. C. 1978. The systematic relationships of

Brunonia. Brunonia 1: 9-29.

Charig, A. J. 1982. Systematics in biology: A fundamental

comparison of some major schools of thought. Pp. 363-

440 in K. A. Joysey and A. E. Friday (editors),

Problems of Phylogenetic Reconstruction. Academic

Press, London.

Covas, G & B. Schanck, 1947. Nûmero de cromosomas en

antofitas de la region de Cuzco (Repûblica Argentina). Rev. Argent. Agron. 13: 153-166.

DeVore, M. L. 1993. The Systematics of Acicarpha

(Calyceraceae). Amer. J. Bot. 80. (Abstract). 72 , and T. F. Stuessy. In press. Time and place of

origin of Compositae, Calyceraceae, and Goodeniaceae. In C. Jeffrey (ed.). Advances in Compositae. Academic

Press, London.

Gould, S. J. 1977. Ontogeny and Phylogeny. Harvard

University Press, Cambridge, Mass.

Jansen, R. K., & J. D. Palmer. 1988. Phylogenetic implications of chloroplast DNA restriction site

variation in the Mutisieae (Asteraceae). Amer. J. Bot.

75: 751-764.

Lammers, T. G. 1988. Chromosome numbers and their

systematic implications in the Hawaiian Lobelioideae

(Campanulaceae). Amer. J. Bot. 75: 1130-1134.

. 1992. Circumscription and phylogeny of the

Campanulales. Ann. Missouri Bot. Gard. 79: 388-413.

Lokki, J. & A. Saura. 1980. Polyploidy in Insect

Evolution. Pp. 277-312 in W. Lewis (editor),

Polyploidy. Plenum Press, New York

Miers, J. 1860. On Calyceraceae. Contr. Bot. 2: 1-42. 73 Monasterio, M. 1986. Adaptive strategies of Espeletia in

the Andean desert paramo. Pp. 49-80 in F. Vuilleumier

& M. Monasterio (editors), High Altitude Tropical Biogeography. Oxford University Press, Oxford.

Moore, D. M. 1967. Further records for the vascular flora of the Falkland Islands. Bot. Notiser 120: 2-24.

1968. The vascular flora of the Falkland Islands. Brit. Antarc. Surv. Sci. Rep. 60: 1-202.

1983a. The flora of the Fuego-Patagonia Cordillera:

its origins and affinities. Rev. Chil. Hist. Nat. 56:

123-136.

. 1983b. Flora of Tierra del Fuego. Missouri

Botanical Garden, St. Louis, Missouri.

Olmstead, R. G., H. J. Michaels, K. M. Scott, and J. D.

Palmer. 1992. Monophyly of the Asteridae and

identification of their major lineages inferred from

DNA sequences of rJbcL. Ann. Missouri Bot. Gard. 79:

249-265.

Panero, J. L. 1992. Systematics of Pappobolus (Asteraceae-

Heliantheae). Syst. Bot. Monographs 36: 1-195. 74 Peacock, W. J. 1963. Chromosome numbers and cytoevolution

in the Goodeniaceae. Proc. Linn. Soc. New South Wales

88: 8-27.

Rahn, K. 1960. Chromosome numbers in some South American

angiosperms. Bot. Tidsskr. 56: 117-127.

Raven, P. H. 1975. The bases of angiosperm phylogeny:

cytology. Ann. Missouri Bot. Gard. 62: 724-764.

Rodrigues, C., W. T. Ormond, and M. C. Bezerra Pinheiro.

1977. Contribucao à citologia de Acicarpha spathulata

R. Brown (Calyceraceae). Bol. Mus. Nac. Nova Serie 45:

1-6.

Solbrig, O. T. 1977. Chromosomal cytology and evolution in the family Compositae. Pp. 264-281 in V. H. Heywood,

J. B. Harborne & B. L. Turner (editors), The Biology and Chemistry of the Compositae. Academic Press,

London.

Stebbins, G. L. 1977. Development and comparative anatomy of the Compositae. Pp. 91-109 in V. H. Heywood, J. B.

Harborne & B. L. Turner (editors), The Biology and

Chemistry of the Compositae. Academic Press, London. 75 Sugiura, T. 1936. A list of chromosome numbers in

angiospermous plants. II. Proc. Imp. Acad. Tokyo 12: 144-146.

. 1937. Studies on the chromosome numbers in higher plants, with special reference to cytokinesis. Fujii Jub. Vol., pt. 2: 845-849.

Takhtajan, A. 1959. Die Evolution der Angiospermen.

Gustav Fischer, Jena.

_____ . 1991. Evolutionary Trends in Flowering Plants.

Columbia University Press, New York.

Turner, B. L. 1973. lOPB chromosome number reports LXII.

Taxon 27: 519-535.

, B. L., J. Bacon, L. Urbatsch, and B. Simpson. 1979. Chromosome numbers in South American Compositae. Amer.

J. Bot. 66: 173-178. CHAPTER III THE SYSTEMATICS OF ACICARPHA

Introduction

Acicarpha is a genus of annuals and perennials distributed from southern Brasil and northern Argentina to the altiplano of Bolivia and Peru. One species, Acicarpha tribuloides, is widely distributed, while remaining species within the genus (A. bupleuroides, A. itatiaiae, A. procumbens, and A. spathulata) have restricted distributions. The species share heads with centripetal development of florets, white corollas with constricted tubes, upper parts of filaments fused into a cylinder, anther connectives with flared bases, filaments thickened at junction with connective bases, and pollen with smooth intercolpar concavities. The last taxonomic study of

Acicarpha was by Miers (1860), who considered the genus closely related to Calycera.

After examining different genera within Calyceraceae (i.e., Acicarpha Juss., Boopis Juss., Calycera Cav.,

Gamocarpha DC., Moschopsis Phil., and Nastanthus Miers), it became apparent that Acicarpha might be a useful starting point for monographic studies on the family. It is the only

76 77 genus within Calyceraceae with centripetally developing heads, and based on this feature, plus its diploid chromosome number (the only diploid reported so far in the family; DeVore, Chapter II), it may be the most primitive genus within the family. This would obviously be a good

starting point for phylogentic studies in the family.

Furthermore, Acicarpha has a distribution centered outside the Andean ranges (DeVore, 1993). It seems logical, therefore, to look at Acicarpha first, and then attempt to relate it to other genera within Calyceraceae which have

speciated along the Andes.

This treatment of Acicarpha expands Miers' generic

concept, employs numerical taxonomic methods to delimit

species within the Acicarpha tribuloides complex (A.

procumbensf A. spathulata, and A. tribuloides), and utilizes cladistic analysis to clarify relationships among species

and to reconstruct phylogeny of the genus.

Taxonomic History

A. L. Jussieu examined collections made by Commerson in

1766 in the region of Buenos Aires, Argentina and

Montevideo, Uruguay. One particular specimen, which

possessed fruits resembling Tribulus, with undivided

stigmas, and sterile, central flowers, represented an

undescribed genus. Jussieu (1803) named the new genus

Acicarpha faci = spines, carpha = straw-like) and described 78 a single species, A. tribuloides. Although Jussieu (1803) suggested that Acicarpha may represent a new family, he placed it within Compositae. A second species, Acicarpha spathulata, was described by Robert Brown (1817), and he placed it within a new family, Calyceraceae.

Richard (1820), who produced the first monographic work on Calyceraceae, accepted Acicarpha as a genus.

Acicarpha was also accepted as a genus in later treatments of Calyceraceae including Candolle (1836), Miers (1860),

Bentham (1873), Hock (1894), Hicken (1919), and Pontiroli (1963). Five additional species were described by Lessing

(1831).

Generic Relationships

Both morphological features, as well as cytological data, show Acicarpha to be a distinct genus within Calyceraceae. The capitulum that develops centripetally is the most distinctive character of Acicarpha. Both Brown

(1817) and Richard (1820) recognized that the capitular morphology of Acicarpha segregated the genus from both

Boopis and Calycera. Other diagnostic characters of the genus are lateral fusion of filaments above nectaries, flared anther connective bases, and prominent thickening of cells at the filament-connective base junction. Other characters not shared by all members of Acicarpha, but unique to the Acicarpha tribuloides complex, include female- 79 sterile central florets within capitula, as well as presence of a capitular diaspore composed of the fused, outer achenes and the receptacle. Chromosome numbers reported from Acicarpha spathulata of n = 8 (Rodrigues et al., 1977) and 2n = 16 (Sugiura, 1936, 1937) for Acicarpha tribuloides, indicate that Acicarpha, in contrast to Boopis, Calycera, and Nastanthus, is the only diploid genus within the family (Devore, 1993).

Acicarpha, on rare occasion, has been confused with

Calycera. Miers (1860) believed, based on achenial morphology, that a close relationship existed between

Acicarpha and Argentinean species of Calycera. He cited the greater length of calycine lobes and the presence of freely dispersed achenes as the only characters distinguishing

Calycera from Acicarpha. Recently, Hansen (1992) regarded Calycera, which has achenes with elongated calycine lobes throughout the capitulum, as being part of a morphocline leading to the evolution of the fused achene-capitular diaspore condition found within Acicarpha.

Phenetic studies of the Acicarpha tribuloides complex

Numerical, or phenetic methods, provide a means of recognizing taxa based on their affinities (Sneath and

Sokal, 1973). Numerical taxonomic studies are still an excellent means for treating taxa with close morphological similarities or which show continuous patterns of variation 80

(e.g., McClintock and Waterway, 1994). In the case of Acicarpha, species within the Acicarpha tribuloides complex

{A. procumbens, A. spathulata, and A. tribuloides) exhibit a wide range of variation in leaf shape and habit. Previous workers, who only had a limited number of specimens available, recognized more than three species within the

Acicarpha tribuloides complex (e.g., Miers, 1860, with five). Because a considerable variation exists within this complex, numerical analyses were utilized to help determine species boundaries. Sixty-seven individuals from throughout the ranges of Acicarpha procumbens, A. spathulata, and A. tribuloides were selected from over 200 available specimens for evaluation of

43 quantitative characters (Table 2). All specimens used in the analysis are indicated by an asterisk (*) in the taxonomic treatment. Floral morphology is conservative within Acicarpha, whereas vegetative and achenial morphology are variable. Characters utilized for the analyses were therefore selected from leaves (basal, medial, and upper cauline) and achenes. Ratios are included to translate leaf shapes (linear and spathulate) and calycine lobe shape

(acicular and cuspidate) into quantitative characters. Ratios are not used here to scale characters, as might be done to remove the size the component of variation. This approach has been shown to be statistically unreliable

(Atchley et al. 1976). 81 Principal components analyses and minimum-spanning tree branch calculations (Figure 2) were accomplished using NTSYS

(Rohlf, 1988). A minimum-spanning tree based on Manhattan

Distance was superimposed on a projection of Factor 1 and

Factor 2 (Figure 2-2). This style of presentation is helpful by utilizing the minimum-spanning tree to show similarity between nearest neighbors, while illustrating the similarity among clusters of OTU's in the principal components analysis (Crovello, 1974).

The results of the analyses (Figure 2) indicate that 24.4% of the variance is contained in the first three principal components axes. Characters that load Factor 1 (Axis 1) are primarily characters of the basal and cauline leaves (Tables 2 and 3), whereas achenial characters load

Factor 2. Factor 3 is loaded by the total length and length from widest region to base of basal and cauline leaves.

These results demonstrate the following: 1) Acicarpha spathulata is the most phonetically discrete species within the section; 2) Acicarpha procumbens is most similar to individuals of Acicarpha tribuloides collected in Peru; and

3) OTU's from Peru (located between OTU's representing

Acicarpha tribuloides and A. procumbens) are the only group of specimens within the highly variable Acicarpha tribuloides complex which group geographically. However, it is important to note that Peruvian specimens of Acicarpha tribuloides used in the study were all collected from two 82 localities in the Province of Cuzco. In my opinion, in light of the tremendous variation present within Acicarpha tribuloides, this is not an adequate sample size to convincingly allow varietal recognation.

Based on the combination of these analyses, three species of Acicarpha {A. procumbens, A. spathulata, and A. tribuloides) can be recognized. Average Manhattan Distances given for OTU's (Figure 2-2) indicate that the most variable species within Acicarpha is A. tribuloides (average

Manhattan Distance between OTU's 0.620), but this variation is not correlated with geographic distribution (Figure 3), and therefore no infraspecific taxa are recognized. 83 TABLE 2. List of 43 quantitative characters used in the phenetic studies of the Acicarpha tribuloides complex . All

Measurements are in mm.

Basal Leaves. 1. Length. 2. Length from widest portion to

base. 3. Width at widest portion. 4. Width at middle. 5. Width between middle and apex. 6. Ratio of length from widest portion to base and total length. 7. Ratio

of length to width at widest portion. 8. Number of

mucronate apices and lobes.

Cauline Leaves. 9. Length. 10. Length from widest portion

to base. 11. Width at widest portion. 12. Width at midsection. 13. Width between midsection and apex.

14. Ratio of length from widest portion to base and

total length. 15. Ratio of length to width at widest

portion. 16. Number of mucronate apices and lobes.

Leaves at Base of Heads. 17. Length. 18. Length from

widest portion to base. 19. Width at widest portion. 20. Width at midsection. 21. Width between midsection

and apex. 22. Ratio of length from widest portion to

base and total length. 23. Ratio of length to width at

widest portion. 24. Number of mucronate apices and

lobes.

Heads. 25. Length. 26. Diameter at base. 27. Diameter at tip. 28. Ratio of diameter at base to length. 84 Table 2 continued. Involucral Bracts. 29. Length. 30. Length from widest portion to base. 31. Width at widest portion. 32.

Width at midsection. 33. Width between midsection and

apex. 34. Ratio of length from widest portion to base

and total length. 35. Ratio of length to width at

widest portion. 36. Number of mucronate apices and lobes.

Calycine Lobes of Achenes. 37. Length. 38. Length from

widest portion to base. 39. Width at widest portion.

40. Width at midsection. 41. Width between midsection

and apex. 42. Ratio of length from widest portion to

base and total length. 43. Ratio of length to width at widest portion. FIGURE 2. 2-1, Principal Components Analysis (PGA) and

Minimum-Spanning Tree based on 43 morphological characters.

Cluster P = Acicarpha procumbens, S = Acicarpha spathulata, and T = Acicarpha tribuloides. 2-2 Minimum-Spanning Tree based on Manhattan Distance superimposed on Principal

Components Analysis. Numerical values flanked by lines represent average Manhattan Distances between OTUs representing specimens of p = Acicarpha procumbens, s =

Acicarpha spathulata, and t = Acicarpha tribuloides.

Distances labeled on tree represent the Manhattan Distance values between clusters of all three species.

85 86

SPATHULATA TRIBULOIDES I 1 J

PROCUMBENS

•394 a7 o•« to

/— 64S

.478 .482 .620

Karl IT I

Figure 2 FIGURE 3. Leaf variation in Acicarpha tribuloides. Representative basal (left) and cauline (right) leaf morphology from specimens of Acicarpha tribuloides collected in 14 localities distributed throughout its range from southern Brasil and northern Argentina to Peru (see Figure

6). Vouchers are indicated by double asterisks (**) in the specimens cited in the taxonomic treatment.

87 88

I

f i: T.

40« MO KWCTCW

5JNU50C*:, »RCJCZZCN

Figure 3 89

TABLE 3. Eigenvectors for characters which load Factors

(Axes) 1-3. The number beneath each set of eigenvectors

is the eigenvalue equal to the variance along the principal axis.

FACTOR 1

CHARACTER EIGENVECTOR

3 0.847

4 0.709

5 0.786

8 0.766

11 0.700 12 0.765

13 0.726

16 0.763

24 0.742

27 -0.816 10.8

FACTOR 2

CHARACTER EIGENVECTOR

2 0.732

25 0.714

38 0.809

40 0.744

42 0.721 90 Table 3 continued.

7.98

FACTOR 3

CHARACTER EIGENVECTOR 1 -0.792 2 -0.786

9 -0.860

10 -0.796

5.64 91

Cladistic Analysis

Phylogenetic systematics is a methodology used to construct explicit classifications of taxa and estimate branching patterns of evolution (Hennig, 1966). Hennig's method, now renamed cladistics (Ashlock and Mayr, 1991), has been viewed in two ways. The traditional viewpoint, based on Hennig (1966), is that the cladogram reflects the phylogeny of the study group. The second viewpoint, held by

Patterson (1980) and Platnick (1985), is that branching patterns of evolution are unknowable and the cladogram represents only how characters may have changed within the taxon. The author takes the viewpoint that the topology of the cladogram represents the best estimate of the branching component of evolution.

Phylogeny, in the eyes of the strict cladist, is confined to the branching component of phylogeny (Hennig,

1966; Mayr 1969; Funk and Brooks, 1981; Nelson and Platnick, 1981). In this work, the term phylogeny is used to define a pattern of evolution produced by both cladogenesis and anagenesis (Mayr and Ashlock, 1991, pps. 229-230). Both autapomorphic and synapomorphic characters are used in cladistic analysis in estimating phylogeny for Acicarpha.

Deleting autapomorphic characters from the analysis could have limited the usefulness of the cladogram in generating a

classification of Acicarpha and masked evolutionary gaps 92 present among species.

Cladistic analysis can only be utilized when one is certain that the study group is monophyletic. Acicarpha is a monophyletic genus based on the following synapomorphies:

1) heads with centripetal development of florets; 2) well- developed filamental thickenings at the base of the anthers;

3) anthers with flared connective bases; 4) pollen with broad, smooth, intercolpar concavities; and 5) the presence of distinct pouches of exine over the aperture of the pollen grains. Of all of the currently recognized genera within

Calyceraceae (Boopis, Calycera, Gamocarpha, Moschopsis, and

Nastanthus), Acicarpha seems to be the most natural.

Outgroup analysis is a useful method for determining primitive character states within the study group (Crisci and Stuessy 1980; Stevens, 1981; and Wheeler, 1981), even though difficulties are known to occur (Stuessy and Crisci,

1984; Disney, 1993). Selection of an outgroup for Acicarpha was especially difficult since the genus appears to be primitive within the family (DeVore, 1993). By definition, the outgroup would have to be found within Goodeniaceae since that family is the sister group to Calyceraceae

(Olmstead et al., 1992). Using the Goodeniaceae as an outgroup for cladistic analyses of Calyceraceae is not practicable due to the large number of autapomorphic characters which define both families (Chapter I). In light of these difficulties, group trends (Crisci and Stuessy, 93

1980) within Boopis were used to polarize character state transformation series within Acicarpha. Since parallelisms exist within Calyceraceae, as well as in Asteraceae and other angiosperm families (Leppik, 1977), these parallel evolutionary trends can suggest evolutionary directionality.

Boopis was selected, because some members of the genus have filamental glands of a similar size and in the same size position within the corolla tube as Acicarpha. Secondly, the majority of the species have the same corolla form as Acicarpha. Since no other genus within Calyceraceae exhibits heads with centripetal floret development,

Dampiera (Goodeniaceae) was utilized as an outgroup to polarize the character transformation series for receptacle shape (character 6), since the genus shows a transition from spikes to heads with centripetal heads (Carolin et al., 1992).

After examining the entire range of variation within and among all five species of Acicarpha, eleven character state trees were selected for cladistic analysis. Only characters which could be confidently polarized using the group trends or outgroup methods were used in the analysis. The polarized series are listed below with some comments regarding the evolutionary significance of each character and its distribution within other genera of Calyceraceae.

Table 4 shows the final data matrix block entered into PAUP

3.0 (Swofford, 1991), a microcomputer program used to find 94

TABLE 4. Data matrix for cladistic analysis of Acicarpha

Character

Taxon 1 2 3 4 5 6 7 8 9 10 11

Outgroup* 0 0 0 0 0 0 0 0 0 0 0

A. bupleuroides 0 0 1 0 0 1 0 1 0 0 0

A. itatiaiae 0 0 1 0 0 0 0 1 0 0 0

A. procumbens 2 0 0 1 1 0 0 0 1 1 1 A. spathulata 1 1 0 1 0 0 1 0 0 1 1

A. tribuloides 0 2 0 2 0 0 1 0 0 1 1

- = Boopis (Characters 1-5 and 7-11), Dampiera (Character

6) . 95

the shortest, most parsimonious cladogram.

1. Habit— Annual (0) ; woody-based perennial (1); or stoloniferous perennial (3) . The woody-based perennial habit has evolved only one other time within Calyceraceae (in Calycera calcitrapa Grisebach). Stoloniferous species are more abundant and can be found within all genera currently recognized within the family. Acicarpha procumbens is the only species within Acicarpha that predominately reproduces vegetatively via stolons.

2. Leaf Margins— Entire (0); mucronate tips occasionally present on upper one-third of margin (1); present throughout margin (2). This character has evolved within the Acicarpha tribuloides complex.

3. Leaves— Monomorphic (0); dimorphic (1). Acicarpha bupleuroides and A. itatiaiae possess cauline leaves with leaf shapes different from those of the rosette. This character is only present within Acicarpha.

4. Rosette leaf bases— Long, narrow, petiolate (0); sessile with a membranous wing (1); broad, clasping, bases

(2). The Acicarpha tribuloides complex possesses leaves with membranous wings. Only Acicarpha tribuloides has rosette leaves with clasping bases. 5. Number of heads— 20-40 (0) ; 5-11 (1). Only Acicarpha procumbens has less than 11 separate heads. 96 6. Receptacle— Elongate (0); or convex (1). This character was polarized using Dampiera (Goodeniaceae), since this genus has some members with centripetal heads which appear to be derived from spikes (e.g., R.

Brown; Carolin et al., 1992). Acicarpha has heads with centripetal development of florets which may have been derived from a reduced spike. Therefore, one might asume that an elongated receptacle may be a primitive character state within the genus.

7. Pales (receptacular bracts)— Unlignified, clearly visible in fruiting heads (0); lignified, not visible in fruiting heads (1) . Two members of the Acicarpha tribuloides complex (A. spathulata and A. tribuloides) have receptacular bracts which are lignified and fused between the calycine lobes of adjacent achenes. This character is absent in all other species within the family.

8. Number of Floral parts— Pentamerous (0); tetramerous (1). Calycera leucanthema is the only other species within the family which has tetramerous flowers.

Two species of Calycera (C. pulvinata and C. sessilifora) possess occasional tetramerous florets interspersed among pentamerous florets within the heads, but this intermediate condition is not found in any taxa of Acicarpha. 9. Corolla limb length (in relation to total corolla length)— Limbs long, greater than 1/5 total corolla length

(0); limbs short (2). Short corolla limbs are also found 97 within Moschopsis and Nastanthus.

10. Calycine lobes— Absent (0) ; elongate (1). Achenes crowned with elongate calycine lobes is one of the characteristics of all species of Calycera. This character state has developed within Acicarpha and is a major synapomorphy which unites the Acicarpha tribuloides complex. 11. Achenes— Free and dispersed at maturity (0) ; fused with one another and the receptacle (1) . This is a second synapomorphy which defines the Acicarpha tribuloides complex. A similar condition is approached within Calycera

(C. eryngioides Remy), but unlike Acicarpha, C. eryngioides possesses groups of two achenes fused to the peripheral portion of the receptacular bracts.

The data matrix was formatted and treated with both the ordered and unordered (Fitch) character type options of PAUP

3.0 (Swofford, 1991). The latter option treats any multistate character as being unordered and disregards the polarity implied by the symbols in the data matrix. The exhaustive search method was employed to evaluate every possible cladogram and select those which are optimal (most parsimonious, or the cladogram with the shortest tree length). All the tree(s) found in the exhaustive search were then rooted using the designated outgroup.

The exhaustive searches using ordered and unordered character state options resulted in a single, most parsimonious cladogram of 14 steps with no reversals (Figure FIGURE. 4. Single most parsimonious tree of species of Acicarpha. Numbers on branches represent number of character state changes.

98 99

Boopis

bupleuroides

itatiaiae

procumbens

spathulata

tribuloides

Figure 4 100

4). The consistency index (Kluge and Farris, 1969), retention index, and rescaled consistency index (Farris, 1989) are three indices commonly used for measuring the

"fit" of characters to cladograms. Since all three indices equal one, the cladogram found in the exhaustive search is, in theory, the cladogram that best describes the data.

The cladistic analysis of Acicarpha implies that two sections can be recognized within the genus based on the presence of two major clades, one supported by two synapomorphies, and the second supported by three synapomorphies. One clade contains A. bupleuroides and A. itatiaiae, two species placed within Boopis. The second clade consists of A. procumbens, A. spathulata, and A. tribuloides, species which embody the previous generic concept of Acicarpha (Jussieu, 1803).

Based on the cladistic analysis, evolution within

Acicarpha has proceeded along two lines. One lineage (A. bupleuroides and A. itatiaiae) is characterized by tetramerous florets, dimorphic leaves, freely dispersed achenes, and annual habit. The second lineage (A. procumbens, A. spathulata, and A. tribuloides) is united by achenes with persistent, elongated, calycine lobes, which are fused with one another and the receptacle. This lineage, which has more derived fruits, has diversified more than the lineage with tetramerous florets. The cladogram suggests that two evolutionary units (A. procumbens and A. 101 spathulata-tribuloides) can be recognized within the Acicarpha tribuloides lineage. Acicarpha procumbens is a species endemic to the coastal region and Uruguay River of

Argentina and Uruguay and is characterized by a reduced number of heads and vegetative reproduction. The results of the cladistic analysis suggest the following: 1) Acicarpha procumbens split from the common ancestor of A. spathulata- tribuloides j 2) Acicarpha procumbens has undergone significant anagenesis as indicated by three autapomorphic characters present in the analysis; and 3) the close phenetic relationship of Acicarpha procumbens to A. tribuloides represents parallel development of leaf characteristics within herbaceous species in the genus. Within Acicarpha the evolution of a capitular dispersal unit (characters 10 and 11) may be viewed as a significant evolutionary development. Since each head contains 10-25 viable achenes, several individuals may develop from a single dispersal unit. In general, the time of flowering of individuals is staggered within populations of Acicarpha and many other members of Calyceraceae (DeVore, pers. observ.).

Since members of the family are protandrous, this dispersal syndrome could potentially insure the establishment of a cross-pollinating population via a single, dispersed head.

In most cases, seeds produced by plants are dispersed close to the maternal individual (Raven, 1976). Since the capitular dispersal unit can be transported by both flowing 102 water, wind, and possibly animals (DeVore, 1991), it is likely the species within the Acicarpha tribuloides complex are dispersed further from the maternal plant. The establishment of small, isolated, interbreeding populations may account for some of the variation within the complex.

This hypothesis could be tested in the future by examining genetic variation within and among populations using isozymes or genetic markers.

Taxonomv

Acicarpha Juss., Ann. Mus. His. Nat. (Paris) 2: 347-348.

1803. Acicarpa Miers, Ann. Mag. Nat. Hist. Ser. 3, 5:

399. 1850. Orthogr. var. TYPE SPECIES: Acicarpha tribuloides Juss.

Cryptocarpha Cass., Aperçu gen. Synanth. 5: 85. 1817.--

LECTOTYPE SPECIES (HERE CHOSEN): Cryptocarpha

spathulata Cass. = Acicarpha spathulata R.Br.

Acanthosperma Veil., El. Flum. 8. t. 152. 1820. TYPE

SPECIES: Acanthosperma littorale Veil. = Acicarpha

spathulata R.Br.

Sommea Bory, Ann. Gén. Sci. Phys. 6: 92. 1820.---TYPE

SPECIES: Sommea calcitrapa Bory. = Acicarpha spathulata R.Br. 103

Echinolema J.Jacq. ex DC., Prod. 3: 3. 1836.---TYPE SPECIES: Echinolema arenarium J.Jacq. ex DC. =

Acicarpha spathulata R.Br.

Annual to perennial herbs. Stems erect or procumbent, terete, with few to many branches, finely ribbed, glabrous.

Leaves simple, alternate, exstipulate; blades spathulate to linear; upper surfaces glabrous, dull; lower surfaces glabrous, dull, paler; margins entire, dentate, or lobed; leaf bases attenuate to auriculate; apices obtuse, pungent, retuse, or subulate, entire or mucronate. Heads solitary, terminal or axillary; florets not part of a cymose subunit; central florets with aborted ovaries in A. procumbens, A. spathulata, and A. tribuloides. Involucre composed of 4-5 bracts, similar in appearance to uppermost cauline leaves.

Receptacle conical or low-convex; pales lanceolate to linear, light-dark-green, lignified at maturity in A. procumbens, A. spathulata, and A. tribuloides. Florets 15-

65, perfect, epigynous, persistent on achenes; corolla funnelform, base usually tightly constricted (more open in

A. procumbens), white, with lobes 4-5. Stamens 4-5, alternate with corolla lobes, fused to lower 2/3 of corolla; filaments connate nearly to anther bases; filamental nectaries 4-5, ovate, yellow-brown, located at point where filaments are free from corolla tube; anthers with flared. 104 connective bases. Ovary inferior, 2-carpellate with a single locule and one basal ovule; style slender, terete, glabrous; stigma capitate. Fruit accessory, consisting of achene and persistent calyx; calycine lobes 4 or 5, enrolled and spine-like with tissue lignified outside, parenchymous inside; pericarp thin; white.

Key to the species of Acicarpha

A. Flowers 4-merous; persistent calyx lobes <0.5 mm long on mature achenes; achenes dispersed separately at maturity. Section Itatiaiae.

B. Leaves on branched portion of stem linear or

narrowly oblanceolate, at base attenuate; receptacle

elongate, conical.

...... 1. A. itatiaiae

B. Leaves on branched portion of stem pandurate, with auriculate, clasping bases; receptacle convex.

2. A. bupleuroides

A. Flowers 5-merous; persistent calyx lobes 1-6 mm long on mature achenes; achenes fused and dispersed with receptacle. 105

Section Acicarpha.

C. Leaves linear to narrowly oblanceolate, with margins entire; corolla limbs approximately 1/5 the length of corolla...... 3. A. procumbens

C. Leaves spathulate, with lower 2/3 of dentate

entire, or leaves broadly to narrowly oblanceolate with

dentate or lobed margins; corolla limbs at least 1/4 the length of corolla.

D. Decumbant to suberect perennials with woody-

fibrous stem bases; leaf bases scarious,

persistent on stem and bases of major branches;

leaves spathulate with margin usually entire, occasionally dentate toward the tip

...... 4. A. spathulata

D. Erect annuals, sometimes persisting from a

thickened base; leaf bases herbaceous, deciduous;

leaves oblanceolate with margin dentate or lobed...... 5. A. tribuloides

I Section Itatiaiae DeVore, sect. nov. TYPE SPECIES:

Acicarpha itatiaiae Dusén. 106

Annuae, folia dimorphica; petala 4; calix 4-dentatus; achenia libra.

Annuals; leaves dimorphic; florets tetramerous; achenes with four short (1-3 mm) calycine lobes, freely dispersed.

Species 1-2.

II Section Acicarpha. DeVore, sect. nov. TYPE SPECIES:

Acicarpha tribuloides Juss.

Annuae-perennes, folia monomorphica; petala 5; calix 5- dentatus; achenia connata.

Annuals or perennial herbs; leaves monomorphic; florets pentamerous; achenes with five, long (5-10 mm) calycine lobes, fused. Species 3-5.

1. Acicarpha itatiaiae (Dusén) DeVore, comb, nov.— Boopis

itatiaiae Dusén, Arch. Mus. Nac. Rio de Janiero 13:24.

1903.-- TYPE: BRASIL, Rio de Janeiro, planalto of

Itatiaia [Martius, 1906; Lanjouw, 1945], Feb-Mar, 1894 Ule s.n. (holotype, probably at S, not located;

possibly destroyed at B or deposited at IH). 107

Annuals. Stems erect, 15-50 cm long, 0.1-2 cm diam, green-pink in color. Rosette leaves 6-14 cm long, 0.5-4.5 cm wide, petiolate; blade lanceolate, spathulate on late appearing leaves, 2.5-5.5 cm long, 0.5-4.0 cm wide; apex obtuse-acute; margin serrate-dentate; midvein prominent, with 5-8 distinct secondary veins. Cauline leaves lanceolate-linear, sessile, 1-6 cm long, 0.3-2.5 cm wide; base attenuate; apex acute; margins broadly dentate-serrate to shallowly lobed. Heads 5-12 in number, 0.5-2.5 cm tall,

0.5-1.2 cm diam. Involucral bracts 4-5 in number, linear- lanceolate; 0.3-4 cm long, 0.1-0.5 cm wide; base attenuate; apex acute; margins broadly dentate. Receptacle conical, 6-

10 mm tall, 5-8 mm diam. Pales lanceolate, 0.5-3.0 mm long,

0.2-0.5 mm wide; bases attenuate; apices mucronate; margins entire. Florets 15-40, tetramerous; corolla funnelform, 4- lobed, 2-6 mm long, 0.5-2 mm diam, white; limb 0.6-1.3 mm long, 0.2-0.6 mm wide. Achenes 4-sided, 2-5 mm long, 0.5-3 mm diam; calycine lobes 3-4 in number. Chromosome number unknown.

Phenology. Flowering from October to December.

Distribution. Endemic to the planalto of Itatiaia,

Brasil; 1900-2200 m. Figure 5.

Specimens examined. Brasil. RIO DE JANEIRO: Itatiaia, Parque Nacional de Itatiaia, Barth s.n. (US); Itatiaia, planalto. Brade 15126 (G), 15585 (G, MO, NY); Ayuruoca, 108

Serra Frio, Simart 16211 (US); Itatiaia, Abrigo Reboucas, Strang 745 & Castellanos 25739 (F). FIGURE 5. Distribution in southern Brasil and adjacent northeastern Argentina of Acicarpha bupleuroides (triangles), A. itatiaiae (hexagons), A. procumbens

(circles), and A. spathulata (squares).

109 20

30

Figure 5 M H O Ill

Acicarpha itatiaiae can easily be recognized from other members of the genus by its lanceolate-linear cauline leaves with broadly dentate-serrate margins, and elongated receptacles bearing tetramerous flowers. Acicarpha itatiaiae can be distinguished from A. tribuloides collected

in early flowering stages by the absence of mucronate points

along the leaf margins in A. itatiaiae.

A personal search for type material at S, and a written request for material to IH, failed to locate any types of Acicarpha itatiaiae. Stafleu and Cowan (1976) list Dusén's

collections as also being deposited at B, but if so, were

likely destroyed in 1943 (Lack, 1987). Dusen's description

provides enough details to insure correct application of the

name (e.g., similarity to A. bupleuroides, linear-lanceolate

cauline leaves with dentate margins, and cylindrical receptacles).

2. Acicarpha bupleuroides Less., Linnaea 6: 527. 1817.

Boopis bupleuroides (Less.) C.A.Mull., Flora Bras. 6,

4: 355. 1885. Boopis bupleuroides (Less.) C.A.Mull, var. genuina C.A.Mull., Flora Bras. 6, 4: 355. 1885.—

- TYPE: BRASIL, Parana, specific locality unknown, but

probably collected in either Castro or Fortaleza during

Sellow's inland expedition [Herter, 1945, Papavero,

1971], "Brasilia meridionali locis ulginosis", Oct-Dec 112 1828, Sellow 4949 (holotype, probably at CW, not

located, letters sent to B, FI, KIEL, L, LE, LY, and MW

[Stafleu and Cowan, 1976]).

Boopis bupleuroides (Less.) C.A.Mull. var. microphylla.

C.A.Mull., Flora Bras. 6, 4: 356. 1885.--- TYPE: BRASIL, Santa Catarina, specific locality unknown,

probably collected in either Desterro or Laguna, Dec

1827, [Herter, 1945], Sellow 4434 (holotype probably at

CW, not located, letters sent to B, FI, KIEL, L, LE, LY, and MW).

Annuals. Stems erect to decumbent, 25-70 cm long, 0.1-

2 cm wide, green-reddish tan in color. Rosette leaves 6-17 cm long, 0.5-6 cm wide, petiolate; blade spathulate- lanceolate, 3-12 cm long, 2-5 cm wide; apex obtuse with a point; margins serrate-dentate; midvein distinct; secondary veination highly reticulate. Cauline leaves pandurate, 2-8 cm long, 1-3.5 cm wide, sessile; base auriculate; apex obtuse; margins dentate. Heads 7-20 in number, 0.6 to 1.5 cm tall, 0.5-1.3 cm wide. Involucral bracts 3-5 in number, pandurate, 0.5-3.5 cm long, 0.3-1.8 cm diam; bases fused.

Receptacle low-convex, 4-6 mm tall, 3-11 mm in diam. Pales lanceolate, 0.7-3.0 mm long, 0.3-0.5 mm wide; bases attenuate; apices mucronate; margins entire. Florets tetramerous; 20-40 in number; corolla funnelform, 2.5-6.5 mm 113 long, white; limb 0.6-1.5 mm long (to apex), 0.2-0.5 mm wide. Achenes 3-4 sided, 0.5-1.2 cm long, 0.4-0.8 cm diam; calycine lobes 4 in number, 2-5 mm long, 0.5-1 mm diam.

Chromosome number unknown. Phenology. Flowering from October to January.

Distribution. Grasslands and fields in the Province of

Parana, Brasil; 300-1000 m. Figure 5.

Specimens examined. Brasil. PARANA: Terrinha, Dusén 8559

(F, GH [2], MICH, NY [2]), Dusén 17653 (GH, MO); San Mateus do Sul, Tesoura, Hatschbach 3808 (US); Jardin Natalia,

Curitiba, Hatschbach 33621 (F); Guarituba, Piraquara,

Hatschbach 41110 (F); Aqua Doce, 22 km S of Horizonte, Smith

& Klein 15595 (F, NY).

Acicarpha bupleuroides is characterized by pandurate leaves with clasping auriculate bases and convex receptacles. The species is superficially similar to A. tribuloides, but the former can readily be distinguished by its dentate leaf margins with blunt teeth, tetramerous florets, and capitula with fertile central florets. In contrast, Acicarpha tribuloides has leaf margins with mucronate teeth, pentamerous flowers, and capitula with female-sterile central florets. 114 Letters requesting loans for type material of Acicarpha bupleuroides were sent to CH, FI, KIEL, L, LE, LY, and MW. To date, no Sellow specimen has been located. Lessing's observations of clasping leaves and tetramerous florets on the type material strongly suggests that the name is applied correctly.

3. Acicarpha procwnbens Less., Linnaea 6: 527-528. 1831.--

TYPE: BRASIL, specific locality not indicated on type

or in protologue, but collection possibly made from 2 Jan-Jul 1822 when Sellow sailed from Montevideo to Buenos Aires along the Rio Uruguay [Papavero, 1971, p.

71], Sellow s.n. (L!).

Acicarpha procumbens Less. var. viridiflora C.A.Mull, in

Mart. FI. Bras. 6, 4: 357. 1885.-- TYPE: Brasil, Concepcion del Uruguay, Oct 1877, Lorentz 1178 (holotype GOET, isotype GHl).

Annuals, 4-30 cm tall, stoloniferous. Stems erect to decumbent, 3-28 cm long, 0.1-1.0 cm diam, usually 1-3 branched, occasionally 5-9 branched, green in color. Leaves 1.0-11.5 cm long, 0.2-1.5 cm wide, sessile to briefly petiolate; blades linear to lanceolate 0.9-10.0 cm long; base truncate; apex subulate, mucronate; margins entire; midvein prominent, with two lateral veins extending up into 115 the apex. Heads usually one to five, occasionally up to nine (many branched individuals), spheroid, 0.6-1.2 diam.

Involucral bracts triangular, 2.5-6.5 mm long, 1-7 mm diam; base truncate; apex pungent to acuminate; margins entire.

Pales oblanceolate to linear-lanceolate, 1-4.3 mm long, 0.6- 2.3 mm diam; base attenuate; apices mucronate; margins entire. Florets 30-60, pentamerous; corolla funnelform- campaulate, 5-7 mm long, 0.8-2 mm diam, white; limb 0.8-1.4 mm long (to apex), 0.45-0.60 mm wide. Achenes five sided,

3-9 mm long, 2-4 mm diam; persistent calycine lobes 1.5-3.0 mm long, base expanded 0.5-1.0 diam, terminating abruptly into a fine point, tan-brown. Chromosome number unknown.

Phenology. Flowering from December-January.

Distribution. Riverbanks in the vicinity of Rio

Uruguay and Rio La Plata, Argentina and Uruguay, 0-600 m.

Figure 5.

Specimens examined. Argentina. BUENOS AIRES: Bancalari, Burkhart 4320' (GH); between La Plata and Ensenada, Cabrera

1777 (GH, LP); Monte Veloz, Rio de la Plata, Cabrera 1979

(LP); La Plata, Praderas Salada de Tolosa, Cabrera 3409

(NY*, LP) , Cabrera 9820 (F*, GH, LP) ; San Vincente, 39 km SE of Buenos Aires, Eyerdam & Beetle 23196' (GH); CORRIENTES: Rio Mirihay and Ruta 23, Schinini & Quarin 14452' (UC) ;

Cuatia, Ruta 12, 17 km E of Paso Lopez, Tressens et al. 544'

(UC) ; ENTRE RIOS: Gualeguay, Burkhart 18055' (UC). Uruguay. 116 SALTO: Picada de Peryia, Roseugurt B1013* (GH).

Acicarpha procumbens is distinguished by its stoloniferous habit, linear grass-like leaves, reduced number of capitula and unlignified receptacular bracts. The capitula of A. procumbens also tend to be more globular than those of other species of Acicarpha.

Specimens of Acicarpha tribuloides collected in Bolivia and Peru [e.g., Asplund 3503 (LL); Brunei 139 (GH); King et al. 274 (F)] have grass-like leaves similar to those of

Acicarpha procumbens. These specimens of A. tribuloides can readily be distinguished from A. procumbens by the presence of large basal leaves and elongate receptacles.

4. Acicarpha spathulata R.Br., Trans. Linn. Soc. London,

12: 129. 1817. Cryptocarpa spathulata (R.Br.) H.

Cass., Aperçu gen. Synanth. 5: 85. 1817.-- TYPE:

BRASIL, locality unknown, probably in vicinity of Rio

de Janeiro, 1814-1815 [Papavero, 1971], Sellow s.n.

(holotype BM, photo OS!, isotype S!). 117

Soimea calcitrapa Bory, Ann. Gén. Sci. Phys. 6: 92. 1820--- TYPE: Brasil, locality, date, collector unknown; description was written and published while Bory was

exiled from France [De Virville, 1954] (holotype perhaps at P, not located).

Acanthosperma littorale Veil., FI. Flum. 8. t. 152. 1820.- — TYPE: BRASIL, Sao Paulo, date unknown, Vellozo s.n.

(holotype presumed lost; material used for Flora

fluminensis has not been traced to LISC, P, or R

[Stafleu and Cowan, 1986]).

Echinolema arenarium J.Jacg. ex DC., Prod. 5: 3. 1836.-- TYPE: BRASIL, locality, date, and collector unknown (holotype W, not located), J.Jacq. may have described

material collected in Brasil by J. Milkan, J. Pohl, J.

Natterer, H. Schott, 1817-1821 or obtained via one of

his numerous exchanges with European herbaria (D'Arcy,

1970).

Acicarpha spathulata R.Br. var. glauca DC., Prod. 5: 3.

1836.-- TYPE: SWITZERLAND, Botanical Garden of Geneva,

9 Oct 1832, (holotype G, microfiche OS!). 118

Perennial herbs with taproot 2-15 cm long, 2-15 mm diam. Stems decumbent, 10-55 cm long, 2.5-16 mm diam, usually many branched (12-24, older individuals) or few branched (1-6, young individuals), woody or fibrous at base, green-pink in color. Leaves 1.2-8 cm long, 0.7-2.8 cm wide, sessile; blade spatulate 1-7 cm long, 0.8-2.9 cm wide; apex obtuse to retuse, usually prominently mucronate; margins commonly entire, occasionally with up to ten, shallow, mucronate teeth along the upper one-third, and with a thin band of scarious tissue along upper margin, expanding towards base to form a slight wing; midvein prominent, usually with two other major parallel veins from the leaf base to 1/3-1/2 length of leaf. Heads 3 (young plants) to fifteen (old), 8-13 mm tall, 4-10 mm wide, with tip of head with undeveloped achenes usually deciduous. Involucral bracts spatulate, 4-10 mm long, 1.5-6 mm wide, spatulate; base attenuate; apex obtuse. Pales oblanceolote, 1-2 mm long, 0.2-0.7 mm wide, often fused with receptacle at maturity. Florets 46-60 in number, pentamerous; corolla funnelform, 3.8-6.5 mm long, green in bud, white in anthesis, changing to green during fruit development; limb

1-1.4 mm long (to apex), 0.25-0.38 mm wide. Achenes 4-5 sided, 3-6 mm long, 0.5-3 mm wide; calycine lobes 4-7 mm long, base expanded 0.5-1.3 mm, commonly brown-red.

Chromosome number n = 8 (Rodrigues et al., 1977).

Phenology. Flowering throughout the year. 119 Distribution. Foreshore dunes of coasts and outlets of

Atlantic Ocean from Santa Catarina north to Bahia, Brasil; 0-200 m. Figure 5.

Specimens examined. Brasil. BAHIA: Nova Viçosa, Davis et al, 47081 (F), Hatschbach & Guimarâres 47081" (NY); 5 km S of Santa Cruz Cabrâlia, Resinga by the sea, Harley 17126 (MO); Salvador, Itapoa, Santos 1906 & Sacco 2167 (NY, A) ;

ESPIRITO SANTO: 5 km S of Santa Cruz Cabrâlia, Harley et al.

17126" (NY); PARANA: Guaratuba, Dombrowski 1127, Salto 9444

& Pereira 385 (AH), Dusén 13673 (F, GH, MICH, NY);

Paranagua, Pontal do Sul, Hatschbach 14384 (F, NY*) , Hatschbach & Fontella 20861" (F, NY, UC) , Hatschbach & Guimarâres 40311 (F); Morro do Cristo, Krapovlckas &

Cristobal 40311 (AH, F*, UC) ; Praia dos Ferroviarios,

Matinhos, Kummrow 296 (UC) ; Matinhos, Llndeman & de Hass 102

(NY) ; RIO DE JANEIRO: Recreio dos Bandeirantes, Black &

Alder 55-18106 (UC); Praia da Gavea, Brade 15275" (NY) ; Praia Morta, Lagoa da Conceicâo, Duarte & Duarte 3342 (NY); Marambaia, Estado da Guanabara, Jclyaso 44 (NY); Picao da

Praia, Leltz 3529 (F); Restinga de Itapeba, Recreio do

Banderirantes, Segadas-Vlaima 3633 (US); Casimiro de Abreu, near Barra de Sao Joao, Segadas-Vlanna et al, 361 (US) ;

Recreio dos Banderiantes, Silva & Alves 61 (NY); Praia da 120

Gavea, Smith 1305 (G); SANTA CATARINA: Praia Morta, Duarte

3342 (GH, MO*, NY) ; Ihla de Santa Catarina, Eunt 6369 (F,

MO, NY*); Praia Braba, Itajal, Klein 381 (F, UC, US), Klein & Smith 381 (F, NY) ; Barra Vehla, Itajuba Beach, Landirum 2583 (MICH, NY*) ; Guanabara-Barra da Tijuca, Lems s.n. (NY) ;

Restinga de Tijuca, Machado s.n. (NY); Gunabara, Reitz & Klein 1060 (NY); Itajal, Campo do Massiambû, Palhoça, Reitz

& Klein 701* (US) , Reitz & Klein 1223 (NY, US) ; Barra do

Sul, Araquarl, Reitz & Klein 1476 (UC, US) ; Itajal, Praia de

Itajal, Smith & Reitz 6075 (US); SAO PAULO: Municlpio de Peruibe, Casellani 90* (F) ; Beach a few km S of Ubatuba,

Davis et al. 59858 (F); Guarijâ, Dusén 18119 (GH); Municlpio de Cananéia S tip of the island of Ilha Comprida along shore of the bay outlet Barra Cananéia, Eiten & Clayton 6119 (US) ;

Praia Grande, Hoehne s.n. (MO, NY); Santos, Hoehne s.n.'

(NY); Itanhaem, Kuhlman s.n. (NY); Conceiçâo de Itanhaen, Smith 2065 (F, GH).

The only species of Acicarpha that overlaps in distribution with A. spathulata is A. tribuloides.

Acicarpha spathulata is easily distinguished from A.

tribuloides by the presence of woody stem bases with prominent leaf scars, leaves with entire margins (no more than 12 teeth), and persistent calycine lobes with flared bases. 121

5. Acicarpha. tribuloides Juss., Ann. Mus. His. Nat. (Paris)

2; 347-348. 1803. Cryptocarpha tribuloides (Juss.)

Cass., Aperçu gen. Synanth. 5; 85. 1817.--- TYPE:

ARGENTINA, Buenos Aires, 1766 [De Virville, 1954], Coimerson s.n. (holotype P, isotype F!).

Acicarpa pinnatifida Miers, Ann. Mag. Nat. Hist. ser. 3, 5:

403. 1850. Acicarpha tribuloides Juss. var.

pinnatifida Kuntze, Rev. Gen. 3, 2: 126. 1898.--

TYPE: ARGENTINA, Buenos Aires, 1819-25, Miers s.n. (holotype BM, photo OS!).

Acicarpa runcinata Miers, Ann. Mag. Nat. Hist. ser. 3, 5:

403. 1850.--- TYPE: URUGUAY, Banda Oriental, 1832,

Tweedie s.n. (holotype K, not located).

Acicarpha laxa R.E.Fr., Arkiv for Bot. 6: 2. 1907.-- TYPE:

BOLIVIA, Tarija, 1904, Fries 1233 (lectotype, here

designated Si). Fries 1223 was chosen as lectotype

because of more ample material. The other syntype.

Fries 271, "Sierra Santa Barbara in regione Podocarpi",

1904, is here relegated to paralectotype status

(following terminology of Hansen and Seberg, 1984). 122 Acicarpha tribuloides Juss. var. dentata Kuntze, Rev. Gen.

3, 2: 126. 1898. TYPE: ARGENTINA, Dec 1891-Jan 1892

or Oct-Nov 1892 [Lanjouw, 1945], O. Kuntze s.n. (G,

holotype not located).

Annuals, occasionally persisting from a woody base. Stems erect to decumbent, 5-50 cm long, 1-20 mm diam, 12-30 branched, green-tan. Leaves 2-17 cm long, 0.5-4 cm wide, sessile or shortly petiolate; blades spathulate, 1.5-16 cm long, 1.1 to 4 mm wide; bases auriculate; apex obtuse, dentate, with a mucronate tip; margins commonly dentate- pinnatifid, with mucronate tips; midvein prominent with numerous secondary and tertiary veins extending up into lobes. Heads 10-35 in number, 7-15 mm tall, 3-14 mm wide, with tip of head with undeveloped achenes usually deciduous.

Involucral bracts 3-20 mm long, 1.5-4 mm diam, linear- lanceolate, margin entire, occasionally minutely dentate, fused with the receptacle at maturity. Pales oblanceolate to linear, 0.5-1.2 mm wide, lignified on mature heads.

Florets 46-70 in number; pentamerous; corolla funnelform,

3.5-7 mm long, green in bud, white in anthesis, changing to green during fruit development; limb 0.8-1.5 mm long (to apex), 0.3-0.5 mm diam. Achenes 5 sided, 2-7 mm long, 2-4 mm wide; calycine lobes 2-15 mm long, base expanded, 0.4 to

1.3 mm diam, commonly yellow-tan to brown-red. Chromosome number 2n = 16 (Sugiura, 1936, 1937). 123

Phenology. Flowering October through February.

Distribution. Southern Brazil, Uruguay, Paraguay, Bolivia, Peru, and northeastern-central Argentina; 0-4000 m.

Figure 6.

Specimens examined. Argentina. BUENOS AIRES: San Antonio de Arcio, estancia "El Ombû", Boelcke 4566"'** (US); San Isidro, coastal marsh, Bartlett 19256 *•** (US) ; Xsla

Santiago, Cabrera 3373 (NY); Between Rio Santiago and Palo

Blanco, Rodrigues 572* (A) ; CHACO: Margarita Belen, Aguilar

675 (S) ; Resistencia, Jorgensen 2351*’** (GH) ; CORDOBA: Pampa de Achala, Burkhart 10450 (MO); Cerro el Chorito, Foil 430*

(F); La Calera, Legname & Montenegro 169 (TEX); Sierra Chica on bank of Rio Primero, 5 km W of La Calera, Soloman &

Soloman 4065*'** (MO)} Cruz del Eje, Villafane 225 (TEX);

CORRIENTES: Rio Guaquiraro, 18 km S of Sauce, Cristobal et al. 1557 (MO, UC) ; Loreto, Mroginski 43*'**, (LL) ; Estancia

Gauruchos, Pederson 9215*'** (US) ; Estancia La Yela, Pederson 12482*'** (NY) ; Paraisal, Pierotti 6673 (TEX) ; Esquina,

Rodrigo 954 (NY); Establecimiento "La Yerba", Schwarz 167

(NY*, TEX); ENTRE RIOS: Estancia La Invernada, Burkhart &

Bacigalupa 21435* (MO); Crespo, Burkhart & Troncoso 28056

(TEX); JUJUY: Camino to Valle, Abra de Cahas, Cabrera &

Fabris 16099 (TEX*, LP) ; Ledesma, Parque Nacional Calilegua, Cabrera et al. 32118 (MO); Lagunas de Yala, Krapovlckas &

Cristôbal 17471 (F), Schinini et al. 10150 (UC); Perico, 124

Parodi 9036* (GH) ; SALTA: Paso de les Gauchos, Venturi 9970 (GH) ; SANTA FE: San José del Rencon, Alvarez 866* (NY) ; La Guardia, Huedobro 3088 (TEX*, NY); Road from Santa Fé to

Laguna Strobel, Job 682* (NY) ; TÜCÜMAN: Roadside 1-5 Jem S of

Anta Muerta, on the road to Villa Nougués, Conrad 2595*

(MO) ; Rio Monteros, S of Tucuman, Killip 3950** (US) ;

Infiernillo, Krapovlckas & Cristôbal 20530 (F*, GH) ; Monteros, Jem 39-41 on road from Acheral to Tafi del Valle,

Landmn et al. 5781 (MO*, TEX); Tafi, Sala Chaquivil, Olea

179*’** (NY) ; Equina, Terribile 318 (GH) ; Famailla, Rio

Colorado, Venturi 425 (GH); Cumbre de Tafieillo, Venturi

5888 (GH); Alto Terobe, Ville 671 (F). Bolivia.

COCHABAMBA: Independencia, Beck 7456 (UC) ; JLA PAZ: Copacabana, l^plund 3503 (LL); TITICACA: Island of Titicaca,

Bandelier 26 (GH) ; Tacoma, Sarecapa, La Paz, Cardenas 5342**

(GH). Brasil. PARANA: Punta Grassia, Dusén 7952* (LL),

Dusén 10300 (F*'**, GH, MICH) (duplicate also OTU) ; Castro

Rio lapo, Hatschbach 11758* (F) ; Fazenda Campo Real,

Guarapuava, Reitz & Klein 17787 (NY, US) ; RIO GRANDE DO SUL: Arroyo near Pelotas, Archer 4279 (GH*, NY) ; N. Humburgo,

Bornmiiller 342* (GH) ; Viamao, Morro da Grota, Bueno 1958

(F); Santa Rita, Rambo 45668 (US); Osorio, Rambo 48907

(TEX); near Xanxerê, pasture near Rio Xanxerê, Smith & Klein

13271* (NY); Viamao, Itapua, Lagao dos Patos, Sobral 2461*

(F); André da Rocha, Sobral 3517 (F); Pantano Grande, Rio

Pardo, Sobral & Folz 3060 (F) ; SANTA CATARINA: Praia da 125

Teresa, near city of Laguna, Hatschbach 52347 & Kummrow

(MO) ; Bom Jesus, Rambo 34944 (MO*, NY) ; Bom Retiro, Fazenda Campo dos Padres, Smith et al. 7681 (US); Xanxere, along Rio Xanxere, Smith & Klein 13271 (MICH*, US) ; Agua Doce, Campos de Palmas, 28.5 km SE of Horizonte, Smith & Klein 13452

(GH*, NY, UC, US); SAO PAULO: Campos do Jordao, Parque

Estadual, Mattos 15948 (NY). Paraguay. ALTO PARANA:

Between Cacarapegua and Acahay, Casa 3520 & Molero (NY) ; CENTRAL: Ypacaray, Hassler 12313 (MO); Nemby, road to Pozo de Senasa, Vavrek & Vavrek 298 (US); LA ROSADA: near entrance to Ybycui National Park, Gentry et al. 51913 (MO);

Cordillera de Altos, Fiebrig 235 (F*, GH) ; MISIONES:

Estancia La Soledad, Santiago, Pedersen 5241 (AH). Peru.

CUZCO: Chaccan, Brunei 39' (MO) , Brunei 139 (GH*, MO) , Bxrunel 355 (GH, MO*) , Brunei 59l''" (MO) ; Ruins of Inca fortress, Sacsahuaman, above Cuzco, Ferreyra 2646' (US) ;

Chincheros, along side trail. King et al. 274 (F, NY, TEX),

Pennell 13584 (F*, NY) , Soukup 46' (F) , Soukup 6327 (US) ,

Ugent 3786' (UC); Calea, Marin 596' (F) ; PUNO: Muelapata near Moho, Shepard 112 (NY, US). United States. ALABAMA:

Mobile, Burk s.n. (F [2], NY); FLORIDA; Pensacola, Curtiss s.n. (GH); Walton County, Curtiss s.n. (GH); PENNSYLVANIA:

Philadelphia Burk s.n. (F [2], GH); NEW JERSEY: Camden,

Parker s.n. (NY); NORTH CAROLINA: no locality indicated,

McCarthy s.n. (US [2]); SOUTH CAROLINA; no locality indicated, McCarthy s.n. (GH). Uruguay. CALONIA: Arroyo de 126

Pintos, Artilleros, near Puerto Platero, Bartlett 21226

(NY); CANELONES: Rio Sta. Lucia, Paso Cuello, Gallinal et

al, 2404 (NY); ENTRE RIOS: Concordia, Salto Grande, Casa de Piedra, Renvoize 2867' (NY) ; Ceonpo dos Padres, Bom Retiro,

Reitz 2702 (NY); MONTEVIDEO: Santiago Vasquez, Herter 86a

(F*'**, GH, MO, NY, UC) ; PAYSANDU: Arroyo Quebracho, S of

Quebracho, Bartlett 21178 (NY); SALTO: Daymen, Osten 5249''"

(US) ; SAN JOSE: Barra Santa Lucia, Osten 22161' (GH); Barra, Herter 86b (MO, UC) . FIGURE 6. Distribution of Acicarpha tribuloides in south- central South America. Ephemeral introductions to the

United States and New Zealand are not shown (see DeVore 1991 for details of these collections).

127 ;

N > Figure 6 03 129

Acicarpha tribuloides is a distinctive member of the genus because of its dentate-pinnatifid leaf margins and teeth with mucronate tips. Acicarpha tribuloides differs from other pentamerous species (A. procumbens and A. spathulata) by its herbaceous, many-branched growth form and clearly distinguishable rosette and cauline leaves.

Of all the species within Calyceraceae, Acicarpha tribuloides has the widest range of morphological variation.

Numerical taxonomic studies (see previous section) suggest that no infraspecific taxonomic units should be recognized within A. tribuloides. The interpretation of these data is based on the belief that subspecies and varieties are distinct sets of populations with geographic integrity (e.g,

Lewis, 1955; Stuessy, 1990). This is clearly not the case with A. tribuloides.

Acicarpha tribuloides is the only species within Calyceraceae known to have medicinal use. One herbarium label {King et al. 274) documents that Padre Lobon, of

Chinchero, Peru, collected "estrella khishka" to make mate for altitude sickness.

Specimens of Acicarpha tribuloides have been collected from New Zealand and the United States from outside its native range (DeVore 1991). These specimens represent waifs. No naturalized populations of A. tribuloides have been documented. 130 Doubtful and excluded species

Buphthalmum bonariense Pers., Syn. pi. 2: 474. 1807.

Acicarpha bonariense (Pers.) Herter, Rev. Sudamer. Bot. 7: 233.-- TYPE: Brasil, date, collector, and locality unknown, (holotype P, or possibly L). Based on the description, it would be difficult to refer the plant to Acicarpha.

Acicarpha crassifolia Miers, Ann. Mag. Nat. Hist. ser. 3,

5: 402. 1850.-- TYPE: ARGENTINA: Buenos Aires

province, 1832 [Lanjouw, 1945], Tweedie s.n. (K, not located). = Calycera spinulosa Gilles ex Miers, Ann. Mag. Nat. Hist., Ser. 3, 6: 400. 1850.

Acicarpha lanata Lag. ex Pers., Syn. pi. 2: 488. 1816.--

TYPE: MEXICO: "Nova Hispaniâ" (P, not located). DC.

(1836) examined the type and found insufficient

material to refer the plant to Acicarpha. Based on the

description alone, I can find no features which would suggest that the specimen is, indeed, Acicarpha.

Furthermore, Mexico is out of the natural range of the

genus. 131

Acicarpha rosulata N.E.Br., Hook. le. Pl. 27; pl. 2636 B. 1900. = Moschopsis rosulata (N.E.Br.) Dusén,

Gefassplf. Ost. U. Südpat. 42. 1907.--- TYPE: unknown (Stafleu and Cowan, 1976). Based on both Brown and

Dusén's description, which both include references to poorly a developed involucre, the species clearly belongs within Moschopsis 132 References

Atchley, W., C. Gaskins, and D. Anderson. 1976. Systematic

Zoology 25:137-148.

Bentham, G. 1873. Calyceraceae. Pp. 161-163 in Genera plantarum, vol. 2, eds. A. Engler and K. Prantl. Leipzig: W. Engelmann.

Brown, R. 1817. Some observations on the natural family of plants called Compositae. Transactions of the Linnean

Society of London 12:76-142.

Candolle, A. L. P. P. de. 1836. Calyceraceae. Pp. 1-3 in

Prodomus systematis naturalis regni vegetabxlis, vol. 5. Paris: Treuttel et Würtz.

Carolin, R., D. A. Morrison, and T. Rajput. 1992. Flora of

Australia 35. Canberra: Australian Government Publishing Service.

Cassini, H. 1816. Acicarpha. Pp. 32-33 in Dictionnaire des

sciences naturelles, ed. F. Cuvier. Paris: Le Normant. 133

Crisci, J. V. and T. F. Stuessy. 1980. Determining primitive character states for phylogenetic reconstruction. Systematic Botany 5:112-135.

Crovello, T. J. 1974. Analysis of character variation in

systematics. Pp. 451-484 in Systematics,

eds. A. E. Radford, W. C. Dickison, J. R. Massey, and C. R. Bell. New York: Harper & Row.

D^Arcy, W. G. 1970. Jacquin names, some notes on their typification. Taxon 19: 554-560.

DeVore, M. L. 1991. The occurrence of Acicarpha

tribuloides (Calyceraceae) in eastern North America. Rhodora 93:26-35.

. 1994. The systematics of Acicarpha (Calyceraceae)

American Journal of Botany 80:144. [abstract].

, and T. F. Stuessy. In press. Time and place of origin of Compositae, Calyceraceae, and Goodeniaceae. In C. Jeffrey (ed.). Advances in Compositae. Academic

Press, London.

De Virville, D. 1954. Histoire de la Botanique en France.

Paris: Société D'Édition D 'Enseignement Supérieur. 134 Disney, R. H. 1993. Mosaic evolution and outgroup

comparisons. Journal of Natural History 27:1219-21.

Farris, J. S. 1989. The retention index and rescaled

consistency index. Cladistics 5:417-419.

Funk, V. and D. Brooks. 1981. Advances in Cladistics. New

York: New York Botanical Garden.

Hansen, H. 1992. Studies in the Calyceraceae with a

discussion of its relationship to Compositae. Nordic

Journal of Botany 12: 63-75.

. 1984. Paralectotype: a new type term in botany. Taxon: 33: 707-711.

Hennig, W. 1966. Phylogenetic Systematics. Urbana:

University of Illinois Press.

Herter, W. 1945. Auf den Spuren der Naturforscher Sellow

und Saint-Hilaire. Botanische Jahrbiicher 74:119-149.

Hicken, C. 1919. Catalogo de las Caliceraceas Argentinas.

Primera Reunion Nacional de Ciencas Naturales, 1918- 1919: 238-253. 135 Hock, F. 1894. Calyceraceae. Pp. 84-86 in Die natiirlichen

Pflanzenfamilien, tell 4, abt. 5, eds. A. Engler and K. Prantl. Leipzig: W. Engelmann.

Jussieu, A. L. 1803. Sur 1' Acicarpha et le Boopis, deux

generes noveaux de plantes de la famille de Cinarocephales. Annales du Muséum d' Histoire

Naturelle 2:345-350.

Kluge, A. J. and J. S. Farris. 1969. Quantitative phyletics and the evolution of anurans. Systematic

Zoology 18:1-32.

Lack, H. W. 1987. Opera magna in Berlin plant taxonomy.

Englera 7:253-282.

Lessing, C. F. 1831. Calyceraceae Rich. Linnaea 6:258-

260.

Leppik, E. E. 1977. The evolution of capitulum types of

the Compositae in light of insect-flower interaction. Pp. 61-90 in The Biology and Chemistzy of the Compositae, eds. V. H. Heywood, J. B. Harborne, and B.

L. Turner. London: Academic Press. 136 Lewis, H. 1955. Specific and infraspecific categories in

plants. Pp. 13-20 in Biological Systematics. Corvalis: Oregon State University.

Martius, C. F. P. 1906. Flora Brasiliensis 1 pt. 1. Munich: Fleischer.

Mayr, E. and P. Ashlock. 1991. Principles of Systematic Zoology. New York: McGraw-Hill.

McClintock, K. A. and M. J. Waterway. 1994. Genetic differentiation between Carex lasiocarpa and C. pellita

(Cyperaceae) in North America. American Journal of Botany 81: 224-231.

Miers, J. 1860. On the Calyceraceae. Contributions to Botany 2:1-42.

Nelson, G. J. and N. I. Platnick. 1981. Systematics and

Biogeography: Cladistics and Vicariance. New York: Columbia University Press. 137 Olmstead, R. G., R. K. Jansen, H. J. Michaels, S. R. Downie,

and J. D. Palmer. 1990. Chloroplast DNA and

phylogenetic studies in the Asteridae. Pp. 119-134 in Biological Approaches and Evolutionary Trends in

Plants, ed. S. Kawano. London: Academic Press.

Papavero, N. 1971. Essays on the histoxry of Neotropical Dipterology. Sao Paulo: Emprêsa Grafica da Revista dos Tribunais.

Patterson, C. 1980. Cladistics. Biologist 27:234-240.

Pesecreta, T. C., V. I. Sullivan, and M. L. DeVore. In

press. The connective base of Acicarpha tribuloides. American Journal of Botany.

Platnick, N. I. 1985. Philosophy and the transformation of cladistics. Cladistics 1:87-94.

Pontiroli, A. 1963. Flora Argentina, Calyceraceae. Revista

Museo de La Plata 9:175-241.

Raven, P. H. 1976. Systematics and plant population biology. Systematic Botany 1:284-316. 138 Richard, M. 1820. Mémoire sur une famille de plantes dites les Calycérées. Mémoires d/Histoire Naturelle Botanique 6:28-82.

Rodrigues Correia, M. C., W. T. Ormond, and M. C. Bezerra

Pinheiro. 1977. Contribucao à citologia de Acicarpha spathulata R. Brown, Calyceraceae. Boletim do Museu Nacional, Nova Serie 45:1-6.

Rohlf, F. J. 1988. NTSYS-pc. Numerical taxonomy and

multivariate analysis system. Version 1.50. Setauket.

New York: Exeter Publishing, Ltd.

Sneath, P. H. A. and R. R. Sokal. 1973. Numerical

Taxonomy: The Principles and Practice of Numerical

Classification. San Francisco: Freeman.

Stafleu, F., and R. S. Cowan. 1976-1988. Taxonomic Literature, 2nd ed. 1-7: A-Z. Utrecht: Bohn, Scheltema & Holkema.

Stevens, P. F. 1981. On ends and means, or how polarity

criteria can be assessed. Systematic Botany 6: 186-188.

Stuessy, T. F. 1990. Plant Taxonomy. New York: Columbia University Press. 139 .. and J. Crisci. 1984. Problems in determination of

evolutionary directionality of character-state change for phylogenetic reconstruction. Pp. 71-87. in

Cladistics: Perspectives on the Reconstruction of

Evolutionary History, eds. T. Duncan and T. F. Stuessy. New York: Colimbia.

Sugiura, T. 1936. À list of chromosome numbers in angiospermous plants. Proceedings of the Imperial

Academy Tokyo 12: 144-146.

. 1937. Studies on the chromosome numbers in higher

plants, with special reference to cytokinesis. Fujii Jubilee Volume, pt. 2: 845-849.

Swofford, D. L. 1991. PAUP: Phylogenetic analysis using

parsimony, version 3.0, Champaign: Illinois Natural

History Survey.

Wheeler, Q. D. 1981. The ins and outs of character

analysis: A response to Crisci and Stuessy. Systematic Botany 6:297-306. CHAPTER IV SYSTEMATICS OF CALYCERA Introduction

Calycera is a genus of annuals and perennial herbs with a principal distribution along the Andean cordiliera of

Argentina, Chile, and Peru. One species, Calycera spinulosa, is widely distributed in central Argentina. A second species, C. sympaganthera, is only found on the

Coastal Range of Chile. The genus is distinguished from other genera of

Calyceraceae by a combination of cymose capitular subunits, free pales, and dimorphic achenes. Other diagnostic characters such as pentamerous or tetramerous florets, presence or absence of tails and apical anther appendages, are helpful in distinguishing members of the genus. The

last taxonomic study of both Argentinean and Chilean species of Calycera was by Miers (1860).

The lack of a comprehensive taxonomic treatment of Calycera has impeded formation of evolutionary hypotheses regarding the genus. Results from the present study suggest, first, that there is a split within the genus between Argentinean and Chilean species (recognized as

subgenera Calycera and Leucocera respectively). This

140 141 suggests that Argentinean and Chilean species have evolved in isolation from one another. Secondly, Chilean species show more variation in floral morphology than do Argentinean species. It appears that spéciation may have proceeded at a faster rate within the Chilean species group, or perhaps these species are older. More data, particularly macromolecular studies, are needed to test these hypotheses.

The present study incorporates both field and herbarium studies to produce a new classification of Calycera.

Taxonomic History

The taxonomic history of Calycera begins with the publication of Calicera and Calicera herbacea by Cavanilles

(1791), described from material collected by A. Née near Portillo, Chile. Cavanilles' name for the genus, Calicera, was never adopted since calyx is usually spelled with a "y" and not an "i". Jussieu (1803), Brown (1817), and Richard

(1820) all used the name Calycera in their treatments of the family. Proposal 1010 (Reveal and Hoagland 1991) to conserve Calycera was accepted during the Tokyo Congress

(Brummitt, 1994).

Subsequent workers recognized Calycera as a genus.

These include Bentham (1873), Hock (1894), Hicken (1919), and Pontiroli (1963). Additional species of Calycera were described by Remy (1846, 1847), Philippi (1870), Grisebach

(1874), and Fries (1905). 142

Miers (1860) placed most of the Argentinean species and

C. sympaganthera (from Chile) into Calycera. Miers erected a second genus, Anomocarpus, for mainly Chilean species characterized by single axillary heads, cup-shaped involucres, smaller receptacles, and shorter calycine lobes.

Since most subsequent workers (e.g., Reiche, 1900;

Pontiroli, 1963) only treated members of the genus from Chile or Argentina, Mier's keen observations regarding the differences between the two groups of species were never noted. The present treatment recognizes both species groups as subgenera Leucocera and Calycera.

Generic Relationships

Calycera is easily recognized by its heads composed of cymose subunits and the presence of achenes with both long- and short calycine lobes within capitula. Within Calycera, there are two recognizable groups of species. One group, subg. Calycera, characterized by large (3.5 cm in diam. or

less), globular receptacles, corollas with tightly constricted tubes, and anthers without tails or apical appendages, contains Argentinean members of the genus. The

second group, subg. Leucocera, which includes Chilean species, is distinguishable by planar or convex receptacles

(semi-spherical in C. sympaganthera), small (less than 3.5 cm diam) capitula, glands located towards the midsection of the filaments, apical appendages, and tails. 143

Recently, Hansen (1992) hypothesized that Calycera was

closely related to Acicarpha since both genera possess elongated calycine lobes. However, I believe that elongate calycine lobes probably evolved in parallel within Acicarpha

(see Chapter 4). The presence of tetramerous florets and pollen grains with smooth intercolpar concavities are potential synapomorphies shared by Acicarpha and Chilean

species of Calycera.

Concepts of Species

Calycera presents problems in circumscription of

species because of extensive variability found in quantitative vegetative characters. Field observations

reveal considerable quantitative morphological differences

among individuals in single populations. Species within

Calycera have been recognized on the bases of qualitative vegetative or reproductive characters. Differences in morphological characters, it is assumed, reflect genetic and

reproductive gaps (Stuessy, 1990).

Varieties and forms are not recognized here. In many cases, populations of Andean species of Calycera occur

isolated from one another at high elevations. In the case

of C. crenata, which displays continuous variation in both vegetative form and degree of capitular fusion, recognition

of forms would create a cumbersome infraspecific

classification. The approach taken in this treatment is to 144 informally describe such patterns of variation. Recognizing all minor morphological variations within a species is not always useful, particularly when the variation has no geographic component (Davis and Heywood, 1963; Stuessy, 1990).

Morphology and Taxonomic Criteria

Calycera species display a wide range of variation within and among populations. Only characters which were constant were utilized to delimit species within the genus.

Duration and Habit. Calycera comprises annuals, biennials and perennials. Biennials usually exist as a rosette during the first growing season, and produce a branching stem and capitula the second season. One perennial species, Calycera calcitrapa, is woody at the base, whereas the remaining species are suffrutescent.

Calycera species have erect, ascending, or decumbent branches. The range of growth forms present within Calycera makes this character valuable for distinguishing species.

Roots. Most species of Calycera have thick, somewhat fleshy roots. Calycera eryngioides, C. leucanthema, and C. sessiliflora have rather shallow, narrower taproots. Root morphology is one feature that distinguishes subg. Calycera

(thick-rooted) from Leucocera (shallow, narrower taproots). 145 Leaves. Species of Calycera have three to five major veins arising from the base of the (if present).

The blade outline is spathulate, elliptical, or pinnatifid.

Leaf bases are usually attenuate, but may be nearly auriculate (C. herbacea) or cuneate (C. eryngioides and C. sessiliflora). Apices are acuminate, acute, or obtuse. The leaf margin present within Calycera may be entire, lobed, dentate, crenulate, or incised.

Leaf morphology can vary within and among individuals of a population, as well as among populations. This is especially true for species distributed along the Andean chains {Calycera crenata, C. eryngioides, C. herbacea, C. involucrata, C. pulvinata, and C. sessiliflora).

Vesture. As a family, Calyceraceae are nearly devoid of vesture. Within Calycera, velutinous to villous hairs can be found on the involucral bracts and pales of C. sympaganthera, and sparsely on upper nodal regions of C. sessiliflora. Dense villous hairs are present on the upper stem region of C. leucanthema. Within Calyceraceae, only these three Chilean species possess vesture.

Inflorescence. All species of Calycera have capitula arising from the axils of leaves. Fusion of capitula is common in Calycera crenata and C. pulvinata.

Heads and Involucral Bracts. Calycera, like most members of Calyceraceae (with the exception of Acicarpha), has heads composed of cymose subunits consisting only of 146 florets with tubular corollas. Each head is subtended by 4- 7 involucral bracts which are usually partially fused at their bases. In Calyceraceae, involucral bracts are not highly modified, as is the case with Asteraceae, and they are nearly identical to those of the upper cauline leaves.

Head size is taxonomically important in Calycera.

Members of subgenus Leucocera tend to have small heads (0.5- 2.5 cm diam). In contrast, most of the species within subgenus Calycera have large, globular heads at least 3.5 cm diam. All diameters of heads in the descriptions do not include the extending, spine-like, calycine lobes.

Involucral bracts are of taxonomic utility in the

Chilean species group. Cup-shaped involucres is one feature that segregates Calycera pulvinata and C. sessiliflora. Presence of vesture on involucral bracts in Calycera sympaganthera helps distinguish the species from C. coronata.

Pales. Most species possess persistent pales of medium texture. The pales tend to be lanceolate, with a mucronate apices. One species, Calycera sympaganthera, has villous hairs on the margins of the pales. Color is green.

Florets. Taxonomically useful characters are found in tetramerous vs. pentamerous florets within heads, degree of constriction of the corolla tube, and filamental adnation and connation (Figure 7). Species of Calycera possess a filaments which can be defined in three sections. The upper FIG. 7. Features of filaments and glands in Calycera. A, nomenclature for filament structure (FS = segment of filament separate; FC = segment of filament connate; FCA =

segment of filament connate and adnate to corolla). B-C, Filamental differences between Subgenera Leucocera (B) and Calycera (C).

147 B

I

FS

• '• a

FC

"FCA

Figure 7

00 149 portion is distally separate (FS). The second section, in which the filaments are connate (FC), is located just above the filamental glands. The lower portion of the fillaments is connate and adnate to the corolla (FCA). The lengths of the FCA, FC, and FS sections are important taxonomic characters in subgenus Leucocera.

Achenes. The achenes of all species possess spine-like projections which are interpreted to be persistent calycine lobes. The developmental sequences from typical greenish- calycine segments to modified calycine lobes are clearly visible in young capitula (DeVore, pers. observ.). Two types of achenes are produced in each head. At an early developmental stage (visible in bud), the calycine segments of some achenes expand, enroll, and become lignified. In other achenes, the calycine lobes are persistent, but do not expand into spine-like projections. Achenial morphology is helpful in delimiting species of Calycera. The length, as well as number, of expanded calycine lobes on achenes, are useful for identifying species.

Taxonomy

Calycera Cav., Ic. et Descrip. PI. 4: 34-35. 1797. TYPE

SPECIES: Calycera herbacea Cav. 150 Discophytum Miers ex Lindl., Veg. Kingd. 701. 1847.--

TYPE SPECIES: Discophytum pulvinatum Miers ex Lindl. =

Calycera pulvinata Remy.

Anomocarpus Miers, Ann. Mag. Nat. Hist. 3: 350. 1850.-- TYPE SPECIES (HERE CHOSEN): Anomocarpus axillaris

Miers. = Calycera sessiliflora F.Phil.

Gymnocaulos F.Phil., Linnaea 28: 706. 1856.-- TYPE

SPECIES: Gymnocaulos viridiflorus F.Phil. = Calycera herbacea Cav.

Leucocera Turcz., Bull. Soc. Nat. Mosc. 21: 582. 1848.--

TYPE SPECIES: Leucocera annua Turcz. = Calycera

leucanthema (Poepp. ex Less.) Kuntze.

Annual to perennial herbs or rosette growth forms (C. crenata and C. pulvinata). Stems erect, decumbent, hollow; surface finely ribbed. Leaves simple or pinnately dissected, exstipulate; blades spathulate to elliptical; upper surfaces glabrous, dull; lower surfaces dull, glabrous, paler; margins entire, or dentate-lobed with mucronate tips; leaf bases attenuate; apices obtuse, pungent, retuse, or subulate, usually mucronate. Heads terminal or axillary; composed of cymose subunits.

Involucre composed of 4-6 bracts; broadly triangular or 151 linear-lanceolate. Receptacle low-convex or globular, 0.5-5 cm diam. Pales lanceolate to linear, light-green. Florets usually 70-120 (between 9-25 in C. sessiliflora), perfect, epigynous, persistent on achenes; corolla funnelform, open or tightly constricted (C. calcitrapa, C. herbacea, C. involucrata, C. spinulosa), with lobes 4-5. Stamens 4-5; filaments free below anther bases (FS section); filamental nectaries, ovate, yellow-brown, located at point where stamens separate from corolla tube; anther bases sagittate

(C. coronata, C. eryngioides, C. leucanthema, C. pulvinata) or rounded; apical appendages present (C. coronata, C. eryngioides, C. leucanthema, C. pulvinata, C. sessiliflora, and C. sympaganthera) or absent. Ovary inferior, 2- carpellate with a single locule and one basal ovule; style slender, terete, glabrous; stigma capitate. Fruit a 4-5- lobed achene, with two size classes of achenes produced within heads; outer layer of fruit accessory being derived from calycine tissue with lignified exterior and parenchymous center; calycine lobes short (less than 4 mm) or long (greater than 8 mm), enrolled and spine-like.

Key to the species of Calycera

A. Involucral bracts morphologically similar to uppermost

cauline leaves; leaf margins dentate with mucronate

apices; anther bases rounded; apices truncate. 152

Subgenus Calycera.

B. Heads convex; involucral bracts ovate to

elliptical with dentate margins; achenes with

elongated calycine lobes located toward the

periphery of capitula...... 8. C. calcitrapa

B. Heads globular; involucral bracts lanceolate; achenes with elongated calycine lobes interspersed

throughout capitula.

C. Calycine lobes 4 cm long, enrolled, acicular.

D. Corolla green; involucral bracts not extending beyond capitula.

...... 11. C. herbacea

D. Corolla white; involucral bracts

extending at least a 1 cm beyond

capitula 9. C. involucrata

C. Calycine lobes less than 2 cm long, only

slightly enrolled with wing-like bases.

...... 10. C. spinulosa 153

A. Involucral bracts triangular-lanceolate and distinct in

morphology from uppermost cauline leaves; leaf margins entire; anther bases usually sagittate; anther apices with appendages. Subgenus Leucocera.

E. Upper stems with dense, villous hairs; florets

all tetramerous...... 3. C. leucanthema

E. Upper stems glabrous; majority of florets pentamerous.

F. Leaf margins broadly dentate or pinnatifid.

G. Leaf margins broadly dentate; involucral bracts villous at base.

...... 2. C. sympaganthera

G. Leaf margins lacerate-pinnatifid;

involucral bracts glabrous at base.

...... 1. C. coronata

F. Leaf margins entire or

crenulate.

H. Involucre cup-shaped; involucral bracts

fused more than half their length. 154

I. Growth form rosette; most capitula

separate, but with central capitula fused to each other...... 6. C. pulvinata

I. Growth form sprawling; capitula all

separate and distinct......

...... 5. C. sessiliflora

H. Involucre flat; involucral bracts fused

only at base.

J. Involucral bracts broadly

triangular, 2.5 cm diam. at base; outer achenes with calycine lobes 2 cm or more in length......

...... 4. C. eryngioides

J. Involucral bracts narrowly

triangular, 0.8-1.5 cm diam. at

base; outer achenes with calycine lobes 0.7-1.5 cm long.

...... 7. C. crenata 155

Subgenus Leucocera (Turczaninow) Reiche, Bot. Jahrb. Syst.

29; 107-119. 1900. Leucocera. Turczaninow, Bull. Soc.

Nat. Mosc. 21: 582. 1848.-- TYPE SPECIES: Leucocera annua Turczaninow. = Calycera leucanthema (Poeppig ex Lessing) 0. Kuntze.

Perennials or annuals. Leaves spathulate, orbicular, or pinnate-pinnatifid. Heads usually convex (subglobose in

Calycera coronata and C. sympaganthera). Involucral bracts 5 in number, rarely 6 or 7, lanceolate or deltoid, morphologically distinct from upper cauline leaves.

Receptacle usually convex (subglobose in C. coronata and C. sympaganthera), 0.5-2 cm diam; central tissue solid-slightly parenchymous. Florets tetramerous or pentamerous; 9-120; corolla 2-9 mm long, 1-2 mm diam, funnelform-campanulate with base not tightly constricted. Anthers with connective bases, tails (except for Calycera pulvinata and C. sessiliflora), and apical appendages.

1. Calycera coronata DeVore, spec, nov. TYPE: CHILE, REGION VII (Region del Maule), El Picazo, 35° 28'S, 71° 17'W, 28 Dec 1936, Barros 129 (holotype, GHI). Figure

8 . 156

Herbae; rami teretes, decumbentes. Folia spathulato-

lanceolata, 1.5-12 cm longa, 0.2-1.2 cm lata, margine pinnatifido vel bipinnatifido. Flores capituloriam 80-110.

Corollae 6.5-9.0 mm longae; limbus 1.0-1.5 mm longus, 0.5- 0.7 mm latus; tubus 5-8 mm longus. Antherae 1.4-1.6 mm

longae, sagittatae. Laciniae calycis persistentis 1.0-1.6 cm longae.

Herbaceous perennials arising from a somewhat woody root base. Stems 0.3-0.6 m long, 0.2-1.2 cm diam, ascending to decumbent, greenish-yellow, glabrous. Leaves spathulate- lanceolate, pinnatifid to bipinnatifid, 1.5-12 cm long, 0.2-

1.2 cm wide; base attenuate; apex of blade and pinnatifid segments mucronulate; base attenuate; margins denticulate.

Heads 5-12, subglobose, 1.5-3.0 cm diam. Involucral bracts

5-7, 0.4-1 cm diam; 5-7; apices acuminate; bases fused, expanded, and lobed; margin entire-denticulate. Pales lanceolate to linear; 3.0-7.5 mm long, 0.2-0.7 mm wide; base attenuate; apex acute; base attenuate; margin scarious.

Florets pentamerous; 80-110; corollas 6.5-9.0 mm long, 0.5-2 mm diam, white, sometimes green at base; limb 1-1.5 mm long,

0.5-0.7 mm wide; tube 5-8 mm long, 0.7-1.5 mm. Filaments 2-

4 mm FCA; 1.0-1.5 mm FC, 0.6-1 mm FS. Anthers 1.4-1.6 mm long, sagittate. Achenes 3-5 sided, 0.3-1 cm long; short calycine lobes 0.2-0.4 mm long; long calycine lobes 0.8-1.3 mm long, throughout head. Chromosome number unknown.

Phenology. Flowering in December. 157

Distribution. Endemic to Chile, Central Valley and foothills of Andes between 35-39° S; 100-800 m. Figure 10.

Specimens Examined. Chile. REGION VII (Region del Biobio): Santa Julia, Neger s.n. (CONC) . REGION VIII (Region del Temuco) : Road to Laguna Malleco, 38° 11' S, 72° 19' W., Pfister s.n. (CONC).

Specimens of C. coronata have been frequently identified as C. herbacea and C. sympaganthera. Calycera herbacea, unlike C. coronata, has simple anthers without sagittate tails and not apical appendage. Achenes with distinct, elongated lobes are present within all portions of the head. Calycera sympaganthera differs from C. coronata by its broadly dentate leaves, more campanulate corolla, and villous hairs present on the nodes, involucre, and margins of the pales. No known specimens of Calycera coronata have been collected in over 40 years and all of these have been found only in the Central Valley and foothills of the Andes in

Chile. Specific localities for Calycera coronata have suffered from extreme habitat destruction caused by deforestation, establishment of Eucalyptus and Pinus plantations, and agriculture (DeVore, pers. observ.;

Ashmann, 1991). It is highly probable that C. coronata is extinct. Figure 8. Calycera coronata. a. Habit, b. Detail of capitulum. c. Corolla, limbs positioned down to show filaments, d-e. Filaments and anthers, f. Cauline leaf.

158 159

m m 1 0 cm

Figure 8 Figure 9. Distribution of Calycera coronata (squares), C. leucanthema (circles), and C. syjnpaganthera (triangles) in

south-central Chile.

160 161

40

Figure 9 162

2. Calycera sympagantbera (Ruiz & Pav.) Kuntze,

Revis, gen. pi. 3, pt. 2:217. 1898. Scabiosa

sympagantbera Ruiz & Pav., FI. Peruv. 1. 49. 1798. Boopis balsamitaefolia (Ruiz & Pav.) Jussieu, Ann. Mus.

Nat. Hist. 2: 38. 1803. Nom, illeait.. improper

combination based on S. sympagantbera. Calycera

balsamitaefolia (Ruiz & Pav.) Rich., Mem. Mus. Hist.

Nat. 6: 77. 1820. Nom, illeait. TYPE: "Peru", no

specific locality indicated [Ruiz and Pav. collected in the vicinity of Arauco 24 February - 1 March 1786 and

may have obtained the type in this region; Ruiz, 1931],

Ruiz & Pavon s.n. (holotype MA, photo MO!; isotype SCO!).

Calycera sympagantbera (Ruiz & Pav.) Kuntze var. pectinata

Kuntze, Revis, gen. pi. 3, pt. 2:217. 1898.-- TYPE:

Angol, Kuntz (holotype, NY; not located)

Calycera sympagantbera (Ruiz & Pav.) Kuntze var. pinnatifida

Kuntze, Revis, gen. pi. 3, pt. 2:217. 1898.-- TYPE:

Angol, Kuntze (holotype, NY; not located).

Herbaceous perennials arising from a somewhat woody root base. Stems decumbent, occasionaly ascending, 10-30 cm long, 0.8-1.6 cm diam, yellow to green, glabrous at base. 163 villous just below heads. Leaves oblanceolate to lanceolate; 1.5-7.0 cm long, 0.5-1.4 cm wide; base attenuate; apex mucronulate; margins broadly dentate. Heads

6-14, subglobose, 1-2 cm diam. Involucral bracts 5-7 in number, triangular-lanceolate, 0.5-1.2 cm long, 0.2-0.5 cm wide; base fused near midsections; apex attenuate; margins entire with villous hairs. Pales oblanceolate to lanceolate, 4-7 mm long, 1.5-1.75 mm; bases attenuate; apex acute; margins entire, upper one-third of margin villous- ciliate. Florets pentamerous; 90-110; corolla 5.5-7.0 mm long, 0.5-1.2 mm diam, white; corolla limb 1.25-1.8 mm long,

0.4-0.6 mm wide; tube 4.20-6.25 long, 0.8-1.2 mm diam.

Filaments 2-4 mm long at FCA; 1-1.5 mm long at FC; 0.5-1 mm at FS. Anthers 0.8-1.2 mm long, sagittate. Achenes with 3-

5 sided; short calycine lobes 0.2-0.5 cm long; long calycine lobes 1-1.5 cm long, throughout head. Chromosome number n = 13 (DeVore, Chapter II). Phenology. Flowering December-March.

Distribution. Coastal Range of Chile, Nahuelbuta;

500-1100 m. Figure 10.

Specimens examined. Chile. REGION VIII (Region del Biobio): MALLECO: Parque Nacional de Nahuelbuta, 500 m W along road leading to Piedra del Aquila, DeVore & Baeza 1611

(CONC, OS, OSH); Cordillera de Nahuelbuta, Alto de La Cueva,

Quezada 216 (CONC); Parque Nacional de Nahuelbuta, between 164 the center of the park and the entrance to the road to Pichinahuel, Ricardi et al. (CONC); Parque Nacional de

Nahuelbuta, Ricardi 5358 (CONC); Parque Nacional de

Nahuelbuta, side of road at Piedra del Aquila, Rodriguez & Torres s.n. (CONC); Cordillera de Nahuelbuta, O. Zollner 13426 (MO).

Calycera sympagantbera is easily distinguished from other Chilean species by its broadly dentate leaves and pales with scarious margins. The species differs from

Calycera leucanthema in the following features: (1) overall size (C. sympagantbera has stems 0.8-1.6 cm diam; C. leucanthema 0.2-0.6 cm diam); (2) degree of pubescence

(glabrous upper stem segments vs. densely pubescent ones);

(3) number of floral parts (pentamerous vs. tetramerous); and leaf morphology (leaves with dentate margins vs. deeply pinnatifid leaves). Calycera sympagantbera is commonly found in small canopy gaps and granite balds within Araucaria-Notbofagus forests. Surprisingly, the species has never been collected in equivalent plant communities within the Southern Andean

Range. A possible explanation for the isolation of C. sympagantbera from other Calycera species may be the formation of The Central Valley. This geomorphic feature is a large, extensive graben separating the Coastal Andean

Cordilleras. It has been estimated that the Central Valley formed prior to the Pliocene (Aguirre, 1985). Possibly C. 165 sympagantbera may have evolved in isolation from other species. Calycera sympagantbera, with its large chromosomes, exhibits a karyotype quite different from the

Andean species (see Fig. 1-7, Chapter 2).

3. Calycera leucanthema (Poepp. ex Less.) Kuntze, Revis, gen. pi. 3, pt. 2: 127. 1898. Boopis leucanthema

Poepp. ex Less., Nov. Gen. 1: 21. 1835. Anomocarpus

leucantbemus (Poepp. ex Less.) Miers, Ann. Mag. Nat.

Hist. Ser. 3, 6: 350. 1850.-- TYPE: CHILE, Andes of Antuco, 1828 [Poeppig, 1960], Poeppig 774 (holotype W, photo FI).

Calycera integrifolia (F.Phil.) Reiche, Bot. Jahrb. 29: 116.

1901. Boopis integrifolia F.Phil., Anal. Univ. Chile

85: 814.-- TYPE: CHILE: Linares, hot springs near

Longavi, Jan 1888, Scboenemann s.n. (holotype SCOI, photo GH!).

Anomocarpus tenuifolius F.Phil. ex Miers, Contr. Bot. 2:

28. 1860. Calycera tenuifolia F.Phil. ex Miers,

Contr. Bot. 2: 28. 1860. Pro, svn. TYPE: Chile,

Region VIII, Cordillera near Talca, 1854 [Papavero, 1971], Germain s.n. (holotype P, isotype MOl). 166 Leucocera annua Turcz., Bull. Soc. Nat. Mosc. 21: 582.

1848.-- TYPE: CHILE, Cordillera near Linares, 1829-44

[Lanjouw, 1945], Bridges s.n. (holotype B, isotype CW, not located).

Annuals, with shallow roots. Stems 2-23 cm long, 2-7 mm diam, decumbent, yellowish-green tinged purple, densly villous at nodes and distal portions beneath heads. Leaves pinnatifid, occasionally linear; 0.5-4.5 cm long, 0.5 to 2.5 cm wide; base attenuate; apex mucronate. Heads 7-16, convex, 0.4-1.3 cm diam. Involucral bracts 0.5-7 mm long, 0.8—2-1 mm wide; bases truncate, fused; apexes attenuate; margins entire. Pales 1-3.8 mm long, 0.1-0.85 mm wide; base attenuate; apex mucronate; border scarious, sparsely vilous towards base. Florets tetramerous; 20-80; corollas 2.2-4.5 mm long, 0.3-1 mm diam, white; limb 1.2-1.6 mm long; tube

1.8-2.30 mm long, 0.2-0.8 mm diam. Filaments 1.75-2.1 mm FCA; 0.2-0.6 mm FC; 0.2-0.3 mm FS. Anthers 0.6-.85 mm long, sagittate. Achenes 4 sided, 1-4.5 mm long; short calycine lobes 0.3-1.5 mm long; long calycine lobes 2-5 mm long, throughout head. Chromosome number n = 17 (DeVore, Chapter

III) .

Phenology. Flowering December-March. Distribution. Populations of C. leucanthema often

invade grazed lands, gardens, and other disturbed habitats within the western foothills of the Andes; 600-2500 m. 167

Figure 10.

Representative specimens. Chile. REGION VII (Region del Maule): CURICO: Los Quenes, Barros s.n. (CONC), Milner s.n.

(CONC); Valley of Rio Teno, Grandjot s.n. (CONC); Upeo,

Zollner 8555 (CONC, MO) ; LINARES: sandy bank along Canal de Melado, Zollner 10482 (CONC); Robleria, road to El Melado, Ricardi 2764 (CONC); Cord. Villagran near Bullileo,

Villagran s.n. (CONC); TALCA: El Picazo, Barros s.n. (G) ;

Laguana del Maule, Behn s.n. (CONC); 10 km S of Vilches

Alto, DeVore 1486 (CONC, OSH); Las 7 Tazas del Rio Clare,

near El Radal, Garaventa 4160 (CONC); Monte Grande, Vilches Alto, Zollner 2544 & 10482 (CONC); Laguna del Maule, Pfister

509 (CONC); Monte Grande, Werdermann 552 (CONC, F, G, NY) . REGION VIII (Region del BioBio): BIOBIO: Chilpa, Neger s.n.;

NUBLE: Tennas de Chilian, Behn s.n. (CONC); Village of

Recinto, DeVore 1486 (CONC, OSH); road to Termas de Chilian

near Pinto, Garaventa 4711 (CONC); road to Termas de Chilian, Matthei s.n. (CONC); San Fabian de Alice, near El

Palo, Matthei s.n. (CONC); San Miguel, near San Pedro,

Matthei s.n. (CONC, OS); 10 km outside of Yungay, Matthei

s.n. (CONC); Atacalco, Pfister 80 & 4988 (CONC); El

Castillo, near Recinto, Rodriguez 363 (CONC); Atacalco,

along Rio Diguillin, Rodriguez & Torres s.n. (CONC); outskirts of Recinto, pasture land. West 5099A (G, UC);

Recinto, road to Termas de Chilian, Zollner 6574 (CONC). 168

REGION IX (Region de la Araucania): CAUTIN: Cunco, Barros (CONC); Pailahuegue, Pirion 403 (G); MALLECO: Victoria, Gunckel 20963 (CONC); Pelluhue, Quiros s.n. (CONC).

Calycera leucanthema is easily distinguished by its pubescent stems, pinnatifid leaves, and tetramerous florets.

This species is also the shortest member of the genus, rarely reaching over 20 cm.

The presence of only tetramerous florets in Calycera leucanthema is unique within the genus. Other Calycera species (i.e., C. pulvinata and C. sessiliflora) have occasional tetramerous florets dispersed within the capitula. The only other genus within Calyceraceae that contains members with tetramerous florets is Acicarpha.

Based on available data, it appears that this character may have evolved independently within in the family (DeVore,

Chapter III).

4 Calycera eryngioides Remy, in Gay, FI. chil. 3: 254. 1848. Anomocarpus eryngioides (Remy) Miers, Ann. Mag.

Nat. Hist. Ser. 3: 351. 1850.-- TYPE: CHILE, locality

uncertain, but Gay made collections in the provinces of

Coquimbo in 1836 and Santiago in 1830-32, 1834, 1838,

and 1840-41 [Munoz P., 1944], Gay s.n. (holotype P,

photo FI; isotypes F! GH! US!). 169

Annuals with shallow roots. Stems 5.0-40 cm long, 1-4 cm wide, ascending to procumbent, yellow to brownish green, glabrous. Leaves spathulate to lanceolate, 0.7-6 cm long,

0.3-1.8 cm wide; tip acute; base attenuate, slightly expanded towards the node; margin entire to crenulate.

Heads 5-16, convex, 2-5 cm in diam. Involucral bracts 5, broadly triangular, fused to the lower 1/3-1/2 of receptacle, 1.8-3 cm long, 0.5-2.2 cm wide; bases connate with a circular orifice forming at maturity around the point of attachment with stem; tips acute; margins entire. Pales linear to lanceolate-spathulate, 3.0-12 mm long, 0.1-2 mm wide; tip acute; base attenuate; margin entire. Florets pentamerous, 40-80; 6.0-8.0 mm long, 0.5-1.3 mm diam, white; limb 1.8-2.2 mm long, 0.4-0.7 wide; tube 4.5-6 mm long, 1-2 mm diam. Filaments 4.5-6.0 mm FCA; 0.6-0.8 FC; 0.2-0.4 FS.

Anthers 1.5-2.4 mm long, sagittate. Achenes dimorphic to trimorphic, yellow. Achenes with long calycine lobes develop towards the outside of the head and fuse with receptacle, 6-9 in number; calycine lobes 3-4 in number, 3-5 cm long, 0.2-0.4 mm wide at base. Achenes 3-5 sided, 4-7 mm long; short calycine lobes 0.4-2 mm long; long calycine lobes 1-6 cm long, only present on outer edge of head.

Chromosome number n = 21-22 (DeVore, Chapter II).

Phenology. Flowering December-March.

Distribution. Open pastures, ski slopes, roadsides and other disturbed areas in the Andean Cordillera of Chile 30- 170 34° S; 1900-3300 m. Figure 10.

Representative specimens. Chile. REGION IV (Region de Coquimbo): LIMARI: La Hualtata, Jiles 1558 (CONC). REGION V

(Region de Valpraiso): ACONCAGUA: Las Lagunillas, Brunner s.n. (CONC); Putaendo, Laguna del Copin, Parra 44 (CONC). REGION VI (Region del Libertador General Bernardo

0'Higgins): SANTIAGO: Laguna Negra, Barros s.n. (CONC);

Cordillera of Potrero Grands, Behn s.n. (CONC); Valle Los Tres Esteros, Grandjot s.n. (CONC, G, MO); San Gabriel,

Cajon del Maipo, Gunckel s.n. (CONC); Rio Colorado, Hastings

473 (US); Potrero Grande, Zollner 1457 (CONC); FIG. 10. Distribution of Calycera. eryngioides (squares) and C. sessiliflora (circles) in central Chile.

171 172

30

Figure 10 173

Lagunillas, Cajon del San Jose, Pena 6 (CONC); Refugio

Lagunillas, Cerro La Cruz, F. Schlegel 5035 (CONC); Puente

Alto, Lagunillas, Schlegel 5914 (CONC).

Calycera eryngioides is easily recognized by its sprawling growth form, five broad (2-3.5 cm wide), green, fleshy involucral bracts, and achenes with long calycine lobes at the periphery of the heads. The means of achene dispersal are also distinctive within the subgenus. The small achenes are freely dispersed, whereas achenes with long calycine lobes become fused with the involucre and dispersed with it. A distinctive character of the species is prominent apical anther appendages, clearly visible at 10

X.

Calycera calcitrapa has been suggested as the closest species to C. eryngioides (Pontiroli, 1963). Both species have large, triangular involucral bracts and long calycine lobes. Despite these apparent similarities in capitular morphology, Calycera calcitrapa is distinct from C. eryngioides. Unlike C. eryngioides, C. calcitrapa has a more robust habit and leaves with dentate, mucronate margins. Also, the triangular involucral bracts of C. calcitrapa are similar to the upper cauline leaves. Calycera eryngioides possesses five involucral bracts that become fused to one another to form a cup-like involucre extending around the lower half of the receptacle. Anther 174 microcharacters also differ. Calycera calcitrapa lacks anther tails and apical appendages, while C. eryngioides exhibits both these features.

Of all species of Calycera, C. eryngioides has the largest calycine lobes (2-5 cm long) and involucral bracts

(2-3.5 cm long, 3.5-2.2 cm wide). Since the species possesses a few, small leaves, one may suspect that the large involucral bracts might be responsible for supplying necessary photosynthate for production of achenes.

5. Calycera sessiliflora F.Phil., Linnaea 28: 706. 1856.

Aimocarpus subsessiliflorus (F.Phil.) Miers, Contr. Bot

2: 29. 1860.-- TYPE: CHILE. REGION VI (Region del

Libertador General Bernardo O'Higgins) Santiago: Cerro

Bravo, 1852-1854, Philippi s.n. (holotype SGO!; isotype

P, photo MO!).

Amnocctrpus axillaris Miers, Contr. Bot. 2: 28-29. 1860. Calycera sessiliflora F.Phil. var. axillaris (Miers)

Reiche. 1900. Anal. Univ. Chile 106: 1045. 1900.---

TYPE: CHILE, REGION V (Region de Valparaiso),

Valparaiso, 1828-1830, [Papavero, 1971; Dance, 1980],

Cuming 664 (K, not located). 175

Annuals with shallow roots. Stems 5-33 cm long, 1-6 mm diam, ascending to decumbent, nodes of lower branches stunted, green-yellow tinged pink or brown, glabrous. Leaves ovate to spathulate, 1-8 cm long, 0.25-2.5 cm wide; base attenuate, slightly expanded at node; apex acute; margin entire to broadly serrate or slightly lobed. Heads, 3-18, 0.4-1.5 cm diam, convex, solitary at leaf axils or clustered in groups of three or more, sessile or attached to stem on a . Involucral bracts 5-6, 3.0-7.0 mm long,

0.8-3.0 mm wide; bases fused; apex mucronate; margins entire. Pales linear lanceolate, 2-4 mm long; 0.3-0.5 mm wide; base attenuate; apex acuminate; margin entire.

Florets tetramerous and pentamerous; 9-30; corolla 1.7-4.0 mm long, 0.5-0.7 mm wide, white; limb 0.5-1.25 mm long, 0.2-

0.4 mm wide; tube 0.8-2.5 mm long, 0.5-1 mm diam. Filaments

0.5-0.8 mm FCA; 0.5-0.8 mm FC; 0.5-1.1 mm FS. Anthers 0.5-

0.8 mm long with prominent connective base. Achenes 4-5 sided, 2-4 mm long; short calyxine lobes 0.5-2 mm long; long calycine lobes 7-20 mm long, only at center of head. Chromosome number unknown.

Phenology. October-December.

Distribution. Andean Cordillera of central Chile; 700- 2100 m. Figure 10. 176

Representative specimens. Chile. REGION IV (Region de

Coquimbo): CHOAPO: Cordillera de Combarbala, Hacienda Ramadilla, El Churque, Jiles 5733 (CONC); ILLAPEL; sandy banks along Rio Leiva, Jiles 4339 (CONC); Cerro Curimahuida,

10 km E of Malancilla, 15 km NE of Sanchez Mine, Worth &

Morrison 16697 (UC); LIMARI: Tulaheuen, Jiles 1011 (CONC);

El Polvo, Jiles 6369 (CONC). REGION V (Region de Valparaiso): ACONCAGUA: Jahuel (El Zaino), Barrientas 1707

(CONC); near Kaki in Valle de Las Banaderas, Bultmann s.n.

(CONC); Los Maitenes, Rio Colorado, Zollner 2835} QUILLOTA:

Ocoa, Collantes 21256 (CONC); sandy area near the foot of

Cerro El Roble, Zollner 447 (CONC). REGION V (Region de

Valparaiso), (CONC). REGION VI (Region del Libertador General Bernardo O'Higgins): SANTIAGO: Luiache, Garaventa 2760 (CONC); Las Condes, Gunckel 40801 (CONC); Macul,

Jaffuel 485 (CONC); Arriba de La Obra, Cajon de Maipo, Mahu & Stebbins 8878 (UC); Cerro Ramon, Maldonado s.n. (CONC); El

Marrsanco, Montero 8296 (CONC); Las Vizacachas, 10 km from

La Dormida, Morrison 16784 (MO, UC, SI); Cordillera de Macul, 20 km E of Santiago, Pirion 485 (G) ; 33° 50' S, 70° 30' W, Ramon 1791 (US); Villa Paulina, Uslar 51 (CONC).

Calycera sessiliflora is easily recognized by its small heads (0.4-1.5 diam), involucral bracts with prominent mucronate apices, and the presence of only one or two achenes with elongate calycine lobes per capitulum. The 177 small number of florets per each capitulum (never more than

30) is also a distinctive feature of C. sessiliflora. Calycera sessiliflora also flowers at a much earlier time of the year (October-early December) than other members of the genus.

The closest species to Calycera sessiliflora is C. pulvinata. Both species possess campanunlate-funnelform corollas, and filamental glands toward the base of the corolla tube, and have occasional tetramerous florets throughout the head.

6. Calycera pulvinata Remy, Ann. Sc. Nat. Ser. 3, 6: 352.

1846. Amnocarpus pulvinatus (Remy) Miers, Ann. Mag.

Nat. His. 3, 6: 353. 1850. TYPE: BOLIVIA, Altiplano of Bolivia, near Lagunas de Potosi, 1839-40, D'Orbigny s.n. (holotype P, isotype, F!).

Rosette perennials, 2-19 cm in diam arising from 10-30 cm long roots. Outer stems free, 0.5-2.0 cm long, 0.4-1 cm wide, tan to pink, tinged green, glabrous; central stems fused. Leaves orbicular to oblanceolate, 1.5 to 10 cm long, 0.3-2.0 cm wide; base attenuate; apex mucronate; margin entire to crenulate. Heads 6-16, convex, 1-2.2 cm diam; inner heads fused. Involucral bracts 5-7, 0.5-0.7 cm long,

0.3-1 cm wide; base fused; apex acuminate; margins entire.

Pales lanceolate, 2-4 mm long; 0.2-0.6 mm wide; base 178 attenuate; apex acute; margins entire. Florets tetramerous and pentamerous; 9-14; corolla 4-6 mm long, white; tube 2.5-

5.2 mm long, 0.4-1 mm diam; limb 0.5-1.3 mm long, 0.4-0.6 mm diam. Filaments 0.5-1.5 mm at FCA; 0.3-0.4 at FC; 1-2 mm at FS. Anthers 0.4-0.7 mm long with a prominent connective base. Achenes 4-5 sided, 4-6 mm long; short calycine lobes

0.2-1.2 mm long; long calycine lobes 0.5-2.0 cm long, throughout head.

Chromosome number unknown.

Phenology. Flowering December-March. Distribution. Altiplano of Peru and Bolivia; 3000-4000 m. Figure 11.

Representative specimens. Peru. CHUCUITO: SW shore of Lake

Titicaca, vicinity of Juli, Madison s.n. (GH); Lake Titicaca, Yunguyo, Plowman & Davis s.n. (F); MOQUIGUA: Calacoto, Cordillera E of Carumas, Asplund 5737 (US) ;

Cordillera E of Carumas, Weberbauer s.n. (NY); PACAJES:

Panaccahi, Asplund 2605 (A), 5738 (US); Pampas, Rosario,

Shepard 233 (GH). Figure 11. Distribution of Calycera pulvinata (circles) and C. crenata (squares) in northern Argentina, southern

Bolivia, and southern Peru

179 180

20

Figure il 181

Calycera pulvinata is most similar to C. sessiliflora in both floral and capitular morphology (see C. sessiliflora description for details) . There is also a tendency for some capitula of C. sessiliflora to be produced on reduced stems at the stem-root base. In fact, central portions of some

Calycera sessiliflora specimens {Barrientas 1707, Mahu & Stebbins 8878, Zollner 447) approach a rosette growth form.

On the basis of growth form, capitular morphology, and floral morphology, C. sessiliflora and C. pulvinata appear to share a common ancestry.

Calycera pulvinata and C. crenata both have compact, rosette growth forms. In C. pulvinata, the central portion of the rosette is composed of fused stems and no additional parenchyma tissue. Unlike C. pulvinata, the central portion of the rosette in C. crenata consists of parenchymous tissue at the stem bases. Derivation of the rosette habit via different organ modifications suggests that the rosette growth form has evolved twice within Calycera.

7. Calycera crenata R.E.Fr., Nova Acta Regiae Soc. Sci. Upsal. ser. 4, 1; 6. 1905. Calycera pulvinata f.

crenata Hicken, Primera Reunion Nac. Soc. Argent. Cien.

Nat. 246. 1918-19.-- TYPE: ARGENTINA, Jujuy, Nevado de Chaûa, March 1903, Fries 861 (holotype S!, isotype SI!). 182

Calycera pulvinata Remy f. cauligera (R.E.Fr.) Hicken, Primera Reunion Nac. Soc. Argent. Cien. Nat. 246. 1918-

1919. TYPE: ARGENTINA, Cumbre de Malamala, 3400 m, Lillo

4894, Dec 1910 (holotype LP!)

Perennials, with rosette growth form or herbs arising from a 7-12 cm long tap root. Stems variable, reduced (rosette growth form) to 20 cm, 0.5-2 cm wide (decumbent growth forms); green-tan, glabrous. Leaves spathulate to orbiculate, 2-8 cm long, 0.3-4 cm wide; bases attenuate; tips obtuse; margins entire, serrate-crenulate. Heads 10-

22, convex, 2-4 cm in diam; outer heads usually free; inner heads may be fused. Involucral bracts 5-6 in number, 1.5-

2.5 cm long, 0.8-1.3 cm wide; base fused; apex acuminate, margins entire. Pales lanceolate, 2-5 mm long, 0.3-1.1 mm wide; base attenuate; apex acute; margins entire. Florets pentamerous, 60-95, white; corolla 3-6 mm long, 0.4-1.3 mm diam; tube 2.5-5 mm long, 0.4-1.4 mm diam; limb 0.7-1.7 mm long, 0.3-0.6 mm wide. Filaments 0.8-1 mm a FCA; 1-1.5 mm

FC; 0.5-0.8 FS. Anthers 0.8-1.2 mm long, shortly sagittate.

Achenes 3, 4, or 5 sided, 4-7 mm long; short calycine lobes

0.5-1.8 mm long; long calycine lobes 0.5-2.5 cm long.

Chromosome number unknown.

Phenology. Flowering November-March.

Distribution. Sandy open habitats, Argentina, Eastern 183 Cordillera of the Central Andes; 2000-5000 m. Figure 11.

Specimens examined. Argentina. JUJUY; Abra Pampa, Cerro Huancai, Cabrera 15426 (LP, SI); Tres Cruces, Cabrera et al.

17715 (LP, SI); Puente del Diablo, Tres Cruces, Fabris &

Zhloaga 7811 (LP) ; Mina Aguilar, Fernandez 2024 (LP) ,

Fernandez 2036 (LP), Frangi & Kiesling 21 (LP, SI) ; Mina

Pirquita, Schwabe 652 (LP), Schwabe 1114 (LP); Mina Aguilar, Scott et al. s.n. (LP).

Calycera crenata exhibits a number of growth forms, making delimitation of the species difficult. Individuals may have a rosette or decumbent growth form, and vary in height (or diam. in the case of rosette forms) from 5 to 30 cm. Examination of herbarium sheets indicates that vegetative and achenial morphology is variable in individuals, as well as within and among populations.

Floral characters are consistent among the various growth forms. Also consistent within populations is the tendency of heads within the central portion of the plant to be produced on shortened stems (seen clearly in Fabris &

Zhloaga 7811; Frangi & Kiesling 21; Schwabe 652).

Calycera subgenus Calycera

Calycera Cav., Ic. et Descrip. PI. 4: 34-35. 1797. TYPE

SPECIES: Calycera herbacea. 184

Robust perennials with thick tap roots. Leaves spathulate or elliptical; margins with mucronate teeth.

Heads usually globular or subglobular (Calycera calcitrapa). Involucral bracts 5-6; and similar in appearance to uppermost cauline leaves; spathulate or elliptical.

Receptacle 2-4 cm diam; central tissue parenchymous. Pales lanceolate-linear, 3-6 mm long, 0.5-1.5 mm wide; bases attenuate; apices attenuate; margins entire. Florets pentamerous; 80-140; corolla 6-8 mm long, 1-3 mm diam, funnelform with a tightly constricted base; limbs 2-3 mm long, 1-1.5 mm diam; tube 4-6 mm long. Filaments 4-6 mm long at FCA; 1-1.5 mm long at FC; 1-2 mm long at FS.

Anthers simple and lacking connective bases, tails, or apical appendages. Stigmas capitate, at least twice the diameter of style, twice the diam of style) with numerous long papillae. Species 8-11.

8. Calycera calcitrapa Griseb., Ph. Lorentz 115. 1874.---

TYPE: ARGENTINA, Catamarca, between Nacimientos and

Laguna Blanca, 1872, Grisebach 432 (holotype GOETTl).

Perennials, woody at base, usually many branched, with taproots 3-10 cm long, 0.5-1.5 cm diam. Stems 12-25 in number, erect to procumbent, 5 cm (young plants) to 100 cm long, 0.3-1.5 cm diam, procumbent, yellow to green-pink in color, weakly ribbed. Leaves elliptic, occasionaly ovate. 185

1-4.5 cm long, 0.4-2.5 cm wide; bases rounded to cordate, commonly clasping; apex acute; margins dentate. Heads subglobose, numerous, 12-25 on axillary branches 1-13 cm long. Involucral bracts 5, 0.5-2.0 cm long, 0.7-2.0 cm wide, triangular; base rounded, fused; apex acuminate; margin dentate. Florets 80-110; corolla white. Achenes 4-5 sided, 3-7 mm long, short calycine lobes 1-2 mm long; long calycine lobes up to 5 mm long, towards outside of head. Chromosome number n = 21 (Turner, 1973).

Phenology. Flowering October through April.

Distribution. Catamarca, La Rioja, and San Juan

Provinces, Argentina; usually found in arenosic (quartz- rich) sandy areas; 500-1500 m. Figure 12. Figure 12. Distribution of Calycera. calcitrapa (squares) ,

C. herbacea (circles), and C. involucrata (triangles)

186 187

30

Figure 12 188

Specimens examined. Argentina. CATAMARCA: Rio Zapata, NE of Tinogasta, Ruta 40, km 763, Bocher et al. 2297 (LP); Pie del Medano, Cabrera et al. 18119 (LP) ; Telarito, Cabrera &.

Kiesling 25266 (LP); 20 km S of Andalgala, Cantlno 399 (BHO,

GH), Cantino 548 (GH); La Crenaga, D'Antoni et al. 23 (LP) ; Andalgala, Jorgensen 1843 (US); km 1537 between Andalgala and Belén, Turner 9192 (LL); LA RIOJA: Vinchina, Covas 1230

(NY); km 147 about 55 km SE of Villa Union, Gibson 3050 & Hunziker (NY, UC) ; SALTA: Vina, Alemania, Abbiatti & Claps

711 (F), Abbiatti & Claps 717 (LP); Cafayate Medano, Cabrera

& Marchionni 13083 (LP); Medanos de Cafaya, Cardenas 4213

(GH); Cafayate, dunes, Humbert 21056 (MICH, NY); 10 km E of

Cafayate along Highway 68 toward Salta, Lavin 5783 (UT) ;

Guayrpa, 15 km S of Patquiâ, Lourteig 1195 (SI); SAN JUAN: Between Cuacete and Difunta Correa, Cabrera et al. 29572 (F,

SI); San Augustin, Haene 475 (SI); Between Estaclon and

Digue de Ullûn, Kiesling 4366 (SI); Caucete, Pie de Palo,

Mezzo 1054 (US); TUCUMAN: Tinogasta, on road between

Fiambala and Chaschuil, km 98, O 'Donell-Meyer 5126 (US) ;

Quebrada de Belen, Sleumer & Vervoorst 2362 (US); Tafi, Sierra del Cajon, Venturi 4220 (F, GH, LP).

Calycera calcitrapa is easily recognized by its many branched stems, dentate leaf margins with mucronate teeth, and wide (greater than 1.5 cm) bracts with dentate margins. 189

Another distinctive feature is the distribution of achenes with elongated calycine lobes towards the periphery of the

capitula.

Calycera involucrata and C. eryngioides are the two species within the genus that may be confused with C.

calcitrapa. Calycera involucrata has white corollas and

long achenial lobes, as is found in C. calcitrapa, but the

latter have spathulate-shaped leaves with dissected margins

and spathulate-linear involucral bracts. Calycera eryngioides has leaves with entire margins and lobes, as well as triangular involucral bracts with entire margins. In contrast, Calycera calcitrapa has leaves and triangular bracts with dentate margins.

9. Calycera involucrata F.Phil., Anal. Univ. Chile 36: 174.

1870. TYPE: ARGENTINA, Mendoza, Portezuelo de Mendoza, 1860 [Papavero, 1971], Philippi s.n. (holotype

SGO, not located).

Calycera horrida Hicken, Physis 1: 129. 1912. TYPE:

ARGENTINA: Neuquen, Arroyo Huingaco, Cordillera del

Viento, Jan 1909, Pastore 76 (holotype SII, isotype SI!) . 190 Perennial herbs, rosette (vegetative individuals), becoming many branched (reproductive individuals) with taproots 4-10 cm long, 0.5-2.5 cm diam. Stems 13-21 in number, 10-30 cm long, 0.3-2 cm diam, decumbent, light green, weakly ribbed. Leaves spathulate to pinnatifid- pinnate, 2.5-9 cm long, 1.5-4 cm wide; base attenuate; apex obtuse; margins denticulate. Heads 12-22, convex on axillary branches 2-20 cm long. Involucral bracts 5-6, lanceolate- oblanceolate, 2-4 cm long, 0.7-1.5 cm wide; base attenuate; apex acuminate; margins denticulate. Florets 70-140; corollas white. Achenes 3-5 sided, 5-13 mm long; short calycine lobes 0.3-0.5 cm long; long calycine lobes 1-4 cm, throughout head. Chromosome number unknown. Phenology. Flowering from December through early March.

Distribution. Back-arc folded region (just E of high central region) of the Andes between 31-39° S; commonly found in arenosic (quartz-rich) sands; 1800-3000 m. Figure

12.

Specimens examined. Argentina. MENDOZA: El Portillo,

DeVore 1738 (OS, OSH) ; Rincon Colorado, Ruiz Leal 1311 (RL) ;

NEUQUEN: Arroyo Huingaco, Cordillera del Viento, Pastore 76 (SI); SAN JUAN: Las Minitas, Kiesling 6856. 191

Calycera involucrata is most similar to C. herbacea.

Both species have large, globular receptacles (up to 5 cm diam.) and long, lobed achenes to 4 cm long. Calycera involucrata can be distinguished from C. herbacea by habit

(many branched stems in C. involucrata, 1-5 branched in C. herbacea), leaf shape (pinnatifid vs. spathulate with incised margins), involucral bracts (extending 1.5-2 cm beyond the receptacle vs. 0.2-0.5 cm), and corolla color

(white vs. green).

A personal search for type material of Calycera involucrata at SGO was not successful. The description includes references to a plant which resembles Calycera herbacea and has long involucral bracts and white corollas.

The name can be applied based on these distinct features

Philippi included in the description.

10. Calycera spinulosa Gillies ex Miers, Ann. Mag. Nat.

Hist. Ser. 3, 6: 400. 1850.--- TYPE: ARGENTINA,

Mendoza, Los Arboles, 1835 [Lanjouw, 1945], Gillies

s.n. (holotype K, not seen)

Calycera foliosa J.Phil., Anal. Univ. Chile 106: 1046. 1860.-- TYPE: ARGENTINA, Andes near Mendoza, Valle del

Tunuyan, 1859, Philippi s.n. (holotype SGO!). 192 Calycera crassifolia (Miers) Hicken, Physis 2: 117. 1916.

Boopis crassifolia (Miers) A.Gray, Proc. Amer. Acad. 5: 321. 1861. Acicarpa crassifolia Miers, Ann. Mag.

Nat. Hist. Ser. 3, 6: 400. 1850.-- TYPE: ARGENTINA,

Buenos Aires, Maldronado, 1819, Miers s.n. (holotype K, not located).

Calycera crassifolia (Miers) Hicken var. spinuligera Hicken,

Anal. Soc. Cien. Arg. 48: 174. 1901.--- TYPE:

ARGENTINA, Rio Negro, Dec 1893, Fisher s.n. (holotype SI!) .

Perennial herbs, rosette (vegetative individuals), becoming few branched (reproductive individuals) with taproots 5-20 cm long, 0.5-3.0 cm in diam. Stems 1-6 in number, erect or procumbent, 8-40 cm long, 0.7-2.5 cm in diam, strongly ribbed, green-pink in color. Leaves spathulate to lanceolate, 2-8 cm long, 1.5-5 cm wide; base obtuse to truncate; apex acute; margins dentate; abaxial and adaxial surfaces often appearing waxy. Heads 2-12, globose, usually terminal on branches. Involucral bracts 5-7 in number, lanceolate-spathulate, 0.5-2.5 cm long, 0.2-1.5 diam; base attenuate to obtuse; apex obtuse; margin dentate to entire. Florets 80-110; corolla white. Achenes 3-5 sided, 0.7-2.2 cm long; calycine lobes not clearly 193 differentiate into long and short size classes, 0.3-2.0 cm long. Chromosome number n = 21 (DeVore, Chapter 2).

Phenology. December-February. Distribution. In arenosic or volcanic sands from

Uruguay south to Chubut Province, Argentina; Atlantic coast to fold-thrust belt (region east of high crystalline zone) of the southern Andean Range. Figure 13.

Specimens examined. Argentina. BUENOS AIRES: Villa Gesell,

Boelcke 14586 (MO), Burkart 22386 (US [2]); San Clemente,

Cabrera 4259, 4923 (GH); Juancho, Cabrera 2681 (NY [2] );

Pinamar, Cabrera 10670 (F, US); Monte Hermosa, Carette 157; Punta Negra, 15 km W of Necochea, Eyerdan et al. 23713; Mar del Plata, Hicken s.n. (US); Miramar, Rodrigues 809 (US);

Pehuén-Cô, Villamil et al. 3443 (NY); Miramar, Scala 13

(NY); Necochea, Las Grutas, Villamil et al. 3560 (NY); San

Antonio, Wetmore 778 (US); CHUBUT: Valdez Peninsula, Bahia

Riacho, Frick 24 (GH); LA PAMPA: 5 km S of Cerro Centinela, Pedersen s.n. (NY); MENDOZA: Atuel Valley near El Sosneado,

Bocher, Hjerting & Rahn 999 (MO); St. Raphael, Hauman 1928

(US [2]); NEUQUEN: Colunco, Ammann 12 (F); Along Route 40,

1-2 km N of Barrancas, King & Heinz 9421 (UC); Paso Rio

Salado, Ruta 40, Senn 4324 (F); RIO NEGRO: Chelforo, Cabrera et al. 32822 (SI); Vicinity of General Roca, Fischer 32 (F, MO, NY, US). Uruguay. MONTEVIDEO: Playa Carrasco, Osten

21852 (GH); Canelones, La Floresta, Rosengurtt B1582 (US) ; 194

SAN JOSE: Estancia Pascual, Herter 6707 (GH,NY,ÜS).

Calycera spinulosa is easily distinguished from other

Argentinean Calycera species by the absence of long (over 2 cm) achenial lobes. Unlike other species of Calycera which produce two distinct classes of achenes, C. spinulosa is characterized by achenial lobes which vary continuously within a shorter range fashion.

The last treatment of Calycera (Pontiroli, 1963) recognized C. crassifolia (C. crassifolia vars. crassifolia and spinuligera) and C. spinulosa as separate species.

Characters used to distinguish C. crassifolia and C. spinulosa (degree of branching, presence or absence of mucronate teeth on leaves and involucral bracts) vary both within and among populations. Both herbarium and field observations for the present treatment support combining these species into C. spinulosa. Figure 13. Distribution of Calycera spinulosa.

195 0 0*

SCALE

100 200 )00 200 «OOUiLCt

200 400 MO #00 KILOUdlRI

SINUSOIDAL PROJECTION

H U> 0» Figure 13 197

11. Calycera. herbacea Cav., Ic. et Descrip. PI. 4: 34-35. 1797. Calycera canvanillesi Rich., nom, superf 1. Mém. Mus. Hist. Nat. 6: 34. 1820.-- TYPE: CHILE. REGION VI

(Region del Libertador General Bernardo O'Higgins),

Portillo, 1790, Née s.n.(holotype M, photo OS!).

Calycera nudicaulis F.Phil. ex Burkhill, Fitzgerald's

Exped. 361. 1899.--- TYPE: CHILE, REGION VII (Region del Maule), Cordillera of Linares, 1854, [Papavero

1971], Philippi s.n. (holotype G, photo MO!).

Calycera sinuata Miers, Contr. Bot. 2: 35. 1860. Calycera

viridiflora (F.Phil.) Miers var. sinuata Hicken, Primera Reunion Nac. Soc. Argent. Cien. Nat. 247. 1918-19. Calycera herbacea Cav. var. sinuata (Miers)

Pontiroli, Revist. Mus. La Plata 9, 41: 191. 1963.---

TYPE: ARGENTINA, Mendoza, Puente del Inca, date and

collector unknown (holotype K, not seen).

Calycera squarrosa Miers, Contr. Bot. 2: 35. 1860. Calycera viridiflora F.Phil. (Miers) f. squarrosa

Hicken, Primera Reunion Nac. Soc. Argent. Cien. Nat.

247. 1918-19.-- TYPE: ARGENTINA, Mendoza, date and

collector unknown (holotype K, not seen). 198

Calycera. viridi flora (F.Phil.) Miers, Contr. Bot. 2; 36.

1860. Gymnocaulos viridiflorus F.Phil. Linnaea 28;

706. 1856. Calycera herbacea Cav. var. viridi flora (F.Phil.) Pontiroli, Revist. Mus. La Plata 9, 41: 191.

1963.-- TYPE: CHILE, REGION VII (Region del Maule),

Cordillera of Linares, 1854, Germain s.n. [SGO 57270; Munoz P, 1960] (holotype SGO, not located).

Perennial herbs, rosettes (vegetative individuals) becoming few to many branched (reproductive individuals) with taproots 4-12 cm long, 1-4 cm diam. Stems 2-12 in number, erect, 8-60 cm long, 1.5-5 cm diam, green, ribbed.

Leaves 2-10 cm long, 0.5-4.5 cm wide, spathulate; base attenuate; apex obtuse to retuse; margins entire-dentate

(juvenile) to denticulate (mature). Heads globose, terminal on branches. Involucral bracts 5-6 in number, oblanceolate to lanceolate; 0.5-2.0 cm long, 0.5-1.5 cm wide; base truncate; apex obtuse to attenuate; margins denticulate, rarely entire. Florets 90-150; corollas green. Achenes 0.3-

1 cm long; short calyxine lobes 0.4-0.8 cm long; long calyxine lobes 2-6 cm long, throughout head. Chromosome number n = 21 (DeVore, Chapter 2). Phenology: Flowering December-February.

Distribution: Main Andean Cordillera of Argentina and Chile between 28-40°; 2000-3500 m. Figure 12. 199

Specimens examined. Argentina. MENDOZA: Las Heras, 24 km

NE of Uspallata road to Villavicencio, Boelcke 9929 et al. (LP); Rio Salado Valley, Bocher, Hjerting & Rahn 1014 (LP) ;

2 km E of Puente del Inca, DeVore 1721 (LP, OS, OSH) ; Puente del Inca, DeVore 1722 (LP, OS, OSH); Maripotal-Santiago, Gabriel s.n. (SI); Puente del Inca, Rio Mendoza, Hauman s.n.

(SI); San Rafael, Alrededores Hotel el Sosueado, Lagiglia &

D^Antoni 1200 (LP); Las Heras, Puente del Inca, Lourteig 659

(AH); Quebrada de Vargas, Miehe 48 (UC); Las Heras,

Caracoles Villavicencio, Rotman et al. s.n. (SI); SAN JUAN: Calingasta, Hanantiales, Volponi 33 (LP); Volponi 166 (LP). Chile. LINARES: Cajon Troncoso, Schlegel 3700 (CONC);

TALCA: Laguna de Maule, Behn s.n. (CONC); SANTIAGO: Maipo, Grandjot 3588 (GH); Los Andes in Maitenes near Rio Colorado,

Zollner 11807 (MO).

Calycera herbacea is easily recognized by its large

(3.5-5 cm wide) globular heads, bright green flowers, and long (1-4 cm) calycine lobes. Individuals of the species, with heads bearing numerous spine-like achenal lobes, have a mace-like appearance.

There is a considerable of variation within and among populations of Calycera herbacea. In the past, this species has been taxonomically treated in various ways. Miers

(1860) described two new species (C. sinuata, C. squarrosa) and made a new combination [C. viridiflora (F.Phil.) Miers]. 200 Later workers (Hicken, 1918-19; Pontiroli, 1963) reduced these species to varieties. This treatment will recognize no infraspecific taxa within Calycera herbacea.

Examinations of populations in Argentina and Chile reveal no correlation between morphologic variation and geographic distribution. Field observations found that the overall size, as well as the number of branches produced, seem to correlate with soil moisture. Populations, or portions of populations, located on dry microhabitats (e.g., DeVore

1721) consisted of short individuals (8-20 cm in height) with few branches (3-4). Populations located along meltwater streams or in wet habitats (e.g.,DeVore 1722) were robust (40-80 cm in height) with numerous branches. Based on these observations, variation within Calycera herbacea appears to be related to microhabitat differences.

Field observations have also been helpful in elucidating means of achene dispersal within C. herbacea. Populations of this species observed are located in open, sandy regions near meltwater streams and talus slopes.

Means of dispersal to new open habitats apparently occur via the entire capitulum. At maturity, the stem tissue dies and the capitula loosen and fall from the . The capitula from Calycera herbacea have been observed being transported by wind (DeVore, pers. observ.) 201

DOUBTFUL AND EXCLUDED NAMES

Calycera boopidea Hicken, Pr. Reun. Nac. Soc. Arg. Cienc. Nat. 251. 1918-19. TYPE: ARGENTINA: Neuquen, Confluencia del Limay, 1912 Hicken 184 (holotype SIl).

= Boopis gracilis F.Phil. The holotype clearly has

pinnatifid leaves, short corolla limbs (in relation to

corolla length, outer filamental glands positioned at the base of the corolla, and ribbed achenes. All of

the above characters make it identifiable as Boopis

gracilis.

Calycera castilloni Hicken, Physis l: 386. 1914. = Boopis

castilloni (Hicken) Pontiroli, Revist. Mus. La Plata 9,

41: 191. 1963.--- TYPE: Catamarca, El Rodeo, 15 Jan

1911, Castilloni 2053 (holotype LIL).

Calycera intermedia F.Phil., Anal. Univ. Chile 36: 174.

1870.-- TYPE: ARGENTINA, Mendoza, Portezuelo de

Mendoza, 1860 [Papavero, 1971], Philippi s.n. (holotype

SGO, not located). A personal search for the holotype at SGO was not successful. Based on the description

alone it is impossible to determine if the name is a 202 synonym of C. involucrata F.Phil. or C. spinulosa

Gillies ex. Miers.

Calycera neuquenensis Suess., Rep. Spec. Nov. 51: 197.

1942.-- TYPE: Argentina, Neuquen, exact locality, date, and collector, unknown (holotype M). Based on

the description it is impossible to tell if the name is

a synonym of C. herbacea Cav. or C. involucrata F.Phil. 203

References

Aguirre, L. 1985. The Southern Andes. Pp. 265-376 in The Ocean Basins and Margins, vol. 5, eds. A. E. M. Nairn,

F. G. Stehli, and S. Uyeda. New York: Plenum Press.

Aschmann, H. 1991. Human impact on the biota of

mediterranean-climate regions of Chile and California.

Pp. 33-49 in Biogeography of Mediterranean Invasions, eds. R. H. Groves and F. Di Castri. Cambridge:

Cambridge Press.

Bentham, G. 1873. Calyceraceae. Pp 161-163 in Genera

plantarum, vol. 2, eds. A. Engler and K. Prantl.

Leipzig: W. Engelmann.

Brown, R. 1817. Some observations on the natural family of

plants called Compositae. Transactions of the Linnean

Society of London 12:76-142.

Brummitt, R. K. 1994. Report of the Committee for Spermaphyta: 40. Taxon 43:113-126.

Dance, S. P. 1980. Hugh Cuming (1791-1865) Prince of

collectors. Journal of the Society for the

Bibliography of Natural History 9:477-501. 204

Davis, P. H. and V. H. Heywood. 1963. Principles of

Angiosperm Taxonomy. Princeton, NJ: Van Nostrand.

Fries, R. 1905. Zur Kenntnis der alpinen Flora im nordlichen Argentinian. Nova Acta Regiae Societatis

Scientiarum Upsaliensis (series) 4:1-205.

Grisebach, A. 1874. Plantae Lorentzianae. Bearbeitung der

ersten und zweiten Sammlung argentinischer Pflanzen des

Professor Lorentz zu Cordoba. Abhandlungen der Koniglichen Gesellschaft der Wissenschaften zu

Gottingen 19:49-279.

Hicken, C. 1919. Catalogo de las Caliceraceas argentinas.

Primera Reunion Nacional de Ciencas Naturales. Pp. 238- 253.

Hansen, H. 1992. Studies in the Calyceraceae with a

discussion of its relationship to Compositae. Nordic

Journal of Botany 12:63-75.

Hicken, C. 1919. Catalogo de las Caliceraceas argentinas.

Primera Reunion Nacional de Ciencas Naturales. Pp. 238-

253. 205

Hock, F. 1894. Calyceraceae. In A. Engler and K. Prantl

(editors). Die natiirlichen Pflanzenfamilien, tell 4,

abt. 5, eds. W. Engelman, Leipzig.

Jussieu, A. L. 1803. Sur I'Acicarpha et le Boopis, deux

generes noveaux de plantes de la famille de

Cinarocephales. Annales du Museum D' Histoire Naturelle 2:345-350.

Lanjouw, J. 1945. The location of Central and South American botanical collections. Pp. 225-235 in Plants

and Plant Science in Latin America, ed. F. Verdoorn.

New York: Roland Press.

Miers, J. 1860. On the Calyceraceae. Contributions to

Botany 2:1-42.

Munoz P, C. 1944. El itinerario de Don Claudio Gay.

Boletin de Historia Natural 22:27-45.

Munoz P, C. 1960. Las Especies de Plantas Descritas par R.

A. Philippi in el siglo XIX. Santiago: Ediciones de la Universidad de Chile. 206

Papavero, N. 1971. Essays on the History of Neotropical Dipterology. Sac Paulo: Emprêsa Grafica da Revista dos

Tribunals.

Phillipi, R. 1870. Sertum mendocinum alterum, o sea, catalogo de las plantas recojidas cerca de Mendoza i en

los caminos que conducen de Chile a esa ciudad. Anales

de la Universidad de Chile 36:159-212.

Poeppig, E. 1960 (transi. C. R. Keller). Un Testigo en la

Alborada de Chile (1826-1829) por Eduard Poeppig. Santiago, Chile: Zig-Zag.

Pontiroli, A. 1963. Flora Argentina, Calyceraceae. Revista

Museo de La Plata 9:175-241.

Reiche, K. 1900. Beitrage zur Systematik der Calyceraceen.

Botanischer Jahresbericht 29:107-119.

Remy, J. 1846. Analecta boliviana, seu nova genera et

species plantarum in Bolivia crescentium. Annales des

Sciences Naturelles Botanique 6:345-357.

. 1848. Calicéreas. in Gay, Flora of Chile 3: 246-

256. 207 Reveal, J. L. and R. D. Hoagland. Proposal 1010. 1991.

Taxon 40:651-652.

Richard, M. 1820. Mémoire sur une famille de plantes dites

les Calycérées. Mémoires d'Histoire Naturelle Botanique 6:28-82.

Ruiz, H. (transi. B. E. Dahlgren). 1931. Relaciôn del Viaje Hecho a los Reynos del Perû y Chile par los

Botânicos y Dihuxantes Enviados para Aquella

Expediciôn, Extractado de los Diaros per el Orden que Llevô en estos su Autor. Mandrid: Huelves y Compaha.

Stuessy, T. F. 1990. Plant Taxonomy. New York: Columbia University Press. APPENDIX A THE DISTRIBUTION OF ACICARPHA TRIBÜLOIDES IN EASTERN NORTH AMERICA

208 209

RHODORA, VoL 93, No. 873, pp. 26-35, 1991

THE OCCURRENCE OF ACICARPHA TRIBÜLOIDES (CALYCERACEAE) IN EASTERN NORTH AMERICA

M el a n i e L. D e V o r e

ABSTRACT

Acicarpha tiibuloides (Calyceraceae) ivsi. has a natuial distribution in soutbem Brazil, Uruguay, Paraguay, and northeastern-central Argentina. This species has also been collected in the United States, and Small (1933) reported it as being naturalized in northern Florida. Recent herbarium studies reveal thatAcicarpha tribuloides occurred as a ballast plant in Pennsylvania, New Jersey, Alabama and Florida. Small's account was based on two specimens collected by Curtiss in 1885; no specimenso f Acicarpha tribuloides collected after 1888 are Imown.Acicarpha tribuloides appears to be a well adapted weed in South America based on its abundance, multiple adaptations for disposal, and on its ability to invade dis­ turbed habitats and cultivated fields. This species may have ^ e d to become naturalized in the United States due to: (1) dimatic and edaphic barriers; (2) inability to compete with native and naturalized species; (3) low reproductive capacity.

Key Words:Acicarpha tribuloides. ballast plants, Calyceraceae, distribution. United States

INTRODUCTION Acicarpha tribuloides Juss. (Hgure 1) is one of five species of the South American genus Acicarpha, and occurs in Southern Brazil, Uruguay, Paraguay, Bolivia, Peru, and northeastern-cen­ tral Argentina (Figure 2). The epithet “tribuloides” no doubt re­ fers to the five persistent, spine-like calyx lobes protruding fi-om the center of each achene, much in the same fashion as the genus Tribulus ^ygophyllaceae). BothAdcarpha and Tribidus exhibit fiuits which in shape resemble a Roman tribulus (a device once used to impede calvary). The first known specimens ofAdcarpha tribuloides fi’om the United States were collected sometime between 1867 and 1876 (Burk, 1877; Table 1). Additional specimens firom the Eastern United States were collected between 1885 and 1888. Small (1933) reported the species firom northern Rorida, and Shetler and Skog (1978) listedÆtribuloides as an introduceda n n u a l. These reports provided a stimulus to investigate the distributional history and current status ofAdcarpha tribuloides in the United States. 210 19911 DéVora—Acicarpha

mm

cm

Rgurc I. Aàcarpha tribtdoides Juss. a, capitulum; h, flower; c, habit. From Micrs (I860: Plate 51).

NATURAL HABITAT AND MEANS OF DISPERSAL Cabrera and Tardîni (1978) noted thatAdcarpha tribuloides commonly inhabits grasslands of deltas, river banks, and sandy ravines, and is an invader of cultivated fields. Flowers are borne in capitula (0.3-0.8 cm in diameter) attached to slight peduncles. At maturity, the calyx lobes of the peripheral achenes (central flowers are female-sterile) persist, and are adnate to the achene. 211

Rhodora [VoL93

?

Figure 2. showing the natural distributionof Adcarpha tribuloides Juss. in southern South America. 212

1991] T>éVoxQ—Adcarpha

Table I. Collections ofAdcarpha tribtdoides Juss. from the eastern United States. Number o f Location of Speci­ Voucher Collection Site Collector" mens Year Specimens Philadelphia, PA Burk 3 1867-75 F (310981, 319640). GH Camden, NJ Parker 1 1887 NY North Carolina McCarthy 2 1888 US (42101, 53061) South Carolina McCarthy 1 1888 GH Walton County, FL Curtiss I 1885 GH Pensacola, FL Curtiss I 1885 GH M obile, AL Mohr 1 1888 US (53144) * No collection numbers are recorded for these voucher specimens.

The achenes and receptacle fuse together and form an ovate, rigid, spiny, disseminule that detaches &om the spindly peduncle (Rg- ure I). Ridley (1930) hypothesized that plants with adherent calyces are almost certainly dispersed by wandering mammals. The rip­ ened capitula o î Adcarpha tribuloides may break off and become attached to a passing animal. Since the species is common along river banks, die spiny capitula may be transported by animals using these sites as watering places. Disseminules may also be wind-blown. Annual herbs inhabiting deserts, , or praires often are dispersed when their infforescenses become detached and are blown across open areas (Ridley, 1930). Morphologically, the detached capitula oî Adcarpha could easily be tumbled across a sandy deltaic plain or river bank by wind. Many specimens ofAdcarpha tribtdoides examined have been collected from river banks or ravines. In aU likelihood, capitula are transported downstream and deposited on sandy river banks and bars. Species ofAdcarpha appear to be well a^pted weeds in South America based on their abundance, multiple adaptations for dispersal, and their ability to invade disturbed habitats and cultivated fields.

SITES OF INTRODUCTION IN THE UNITED STATES The earliest known occurrence ofAdcarpha tribuloides in the United States is near the Philadelphia area in 1867 (Table 1). 2 1 3

Rhodora [VoL 93

Burk collected plants from a ballast dump at Kaighn’s Point which he identified as Calycera balsamatifolia (Burk, 1877). Careful examination of Burk’s collections reveals that all three specimens are A. tribuloides. Two specimens ofAdcarpha tribtdoides collected in the late 1880’s firom ballast sites inCamden, New Jersey, and Mobile, Alabama serve as evidence of later introductions of the species. Other specimens fi’om North Carolina (1888) and Walton County, Florida (1885), lack label information regarding the specific lo­ cality (Hgure 3).

MEANS OF INTRODUCTION Ridley (1930) described six ways plants can be introduced: (1) as impurities with cereals, vegetables, and bird seeds; (2) as at­ tachments to fleeces and hides of domesticated animals; (3) in animal fodder; (4) in packing materials; (5) as escapes fi-om cul­ tivation; and (6) in sHps’ ballast and exported soiL During the late 1880’s ballast disposal was a common means by which alien plants were introduced. Ships usually took sand and gravel fi-om a near-shore area at the be^nning of a voyage, and discharged it at the port of destination. This practice continued until the early 20th century when water became the material used to weight ships. Burk (1877) noted that “improvements” made by the Penn­ sylvania Railroad and the American Steamship Company in­ creased the number of vessels entering Philadelphia to export produce and merchandise. Marshland surrounding the harbor was covered with mud and sand dredgings and ballast was constantly added. Brown (1879) watched vessels dumping ballast day and night on Gowanus Creek in New York City. Late 19th century botanists not only observed ballast dumpings, but also made care­ ful observations o f the plants growing in the sites and traced their origins. A large portion of ballast dumped in Philadelphia was oolite or chalk, materials indigenous to the British Coast. Many plants reported fi-om the Kaighn’s Point site were native to the British Isles. Others were South American or Southern European in or­ igin, and such plants captured the imagination of Burk “either fi-om their rarity or the place of nati-vity.” The circumstances possibly responsible for the introductionAdcarpha of tribuloides are intriguing. 214

1991] Adcarpha V

Figure 3. Sites of introductica ofAcicarpha tribuloides Juss. in eastern North America.

On 26 December 1864 the Paraguayan dictator Francisco Sala- no Lopez invaded Brazil. Brazil formeid a coalition with Argentina and Uruguay against Paraguay that lasted undl the Brazilian lanc­ ers killed Lopez in the mid-1870's (Mooney, 1981). During the beginning of the Paraguay-Brazil conflict, the United States and the Confederacy were engaged in the Civil War. When defeat was inevitable, some Confederates formed colonization societies to promote emigration from North America. One popular choice of a new rebel homeland was Brazil, a country glamorized in books Southerners read before the war (Harter, 1985). Some colonists 2 1 5

Rhodora [VoL 93

failed to adapt to Brazil and were returned to the United States by vessels belonging to the Brazilian Squadron. Such ships as the Kansas. Portsmouth, and Quinnebaug protected United States citizens and economic interests during the Paraguay-Brazil War (Harter, 1985). Two ships belonging to this war squadron com­ monly traveled in the region of South America whereAcicarpha tribuloides occurs naturally. The Wasp was a small iron-hulled sidewheel steamer that joined the Brazilian Squadron after she was repaired in Philadelphia in 1867. She cruised the Plata and Un^uay Rivers during the war and remained in Uruguay after the war transporting diplomats and “ guarding American interests.” A second vessel, theJuani- ata, patrolled the coast of Brazil as far south as Buenos Aires. In 1867this steam sloop-of-war returned toPhiladelphia and docked with monitors and ironclads in the Old Navy Shipyard (Coletta and Bauer, 1985; M ooney,1981). The fact that Burk collected Acicarpha tribtdoides in a Philadelphia ballast dump between 1867 and 1876 suggests that the plant may have been introduced via a naval vessel. The circumstances responsible for later introductions ofAci­ carpha tribtdoides are very sketchy. One possible means o f intro­ duction may have been by ships transporting farm equipment The Aurgentinian economy boomed during the 1880’s as a result of wheat cultivation. Lands previously utilized as sheep and cattle range lands were allocated for wheat A great need for quality farm equipment resulted in strong trade relationships between the United States and Argentina and an increase in shipping (Koe- bel, 1912; Ross and McGann, 1982; Williams, 1975).

CURRENT STATUS No specimens ofAcicarpha tribtdoides collected after 1888 are known. Burk (1877) observed a great variety of ballast plants, most of which survived a single growing season. Futhermore, herbarium studies reveal that Small’s account was based on two specimens collected by Curtiss in 1885 (Table 1). Dr. Robert Godfrey (pers. comm.) has never discovered a population A.of tribuloides in his 35 years of botanizing in northern Florida. It appears that this species never became naturalized in the United States. 2 1 6

1991] DéVoTQ— Acicarpha

Some taxa have become naturalized after being introduced in ships’ ballast. Apium leptophyllum (Apiaceae), native to Florida, Texas, and South America, became naturalized in Europe, West Africa, China, Japan, Australia, New Zealand and Polynesia.Cak~ He maritima (Brassicaceae), Glaucium leuzeum (Papaveraceae), and Plaraago coronopm (Plantaginaceae) were probably intro­ duced to Southern Australia in ballast (Ridley, 1930). It is sur­ prising that Acicarpha tribuloides never became established in Alabama or Florida. The species appears to be an abundant weed in regions of South America that are physiographically similar to southeastern United States. Acicarpha tribtdoides probably failed to become naturalized for a number of reasons. Any plant native to South America is likely to encounter some climatic and edaphic barriers to establishment. Seedlings are especially affected by these two factors since they lack the tolerance and vigor of a mature plant (Smith, 1978). If the seedling survives it is forced to compete with native and naturalized individuals. Ross and Harper (1972) suggested that an individual’s ability to capture resources is restricted by the number and proximity of neighboring plants. Even if the founder is capable of scattering seed or reproducing vegetatively, the re­ sulting population may be small and vulnerable due to low re­ productive capacity and slow population growth (Smith, 1978). A small population ofA. tribuloides growing in a constantly dis­ turbed ballast dump site would be an excellent candidate for eradication. Acicarpha tribuloides, as well as other members o f Calyceraceae, possess capitula. The family bears a striking morphological re­ semblance to Asteraceae. The aggregation of flowers into a ca­ pitulum is believed to be the result of selection acting on the flowering phase of the plant. During the firuiting phase, the ca­ pitulum is vulnerable to herbivory (Bunt, 1978). To date, the only secondary compounds known to exist in Calyceraceae are monoterpenoid cyclopentanoid lactones called kidoids (Jensen et al., 1975). Cronquist (1988) speculates that the lack of diversifi­ cation in Calyceraceae is not due to floral or vegetative mor­ phology, but tolimited chemical defenses that have evolved in the family. Cronquist’s theory will be tested in the future when more is known about the biology and secondary compounds in the Calyceraceae. Currently, no members of the Calyceraceae oc- 217

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cur as weeds outside South America, even though the family includes such weedy species as Boopis gracilis Phil., Boopis an- themoides Juss., and Adcarpha tribuloides.

ACKNOWI.EDGMENTS I wish to thank the curators of F, GH, MO, MICH, NY, UC, and US for lending specimens used in this study. Robert Godfrey and Kent Perkins provided information on the statusof Adcarpha tribuloides in Florida; Ronald Stuckey made valuable si^gestions; and Tod F. Stuessy and two anonymous reviewers critically re­ viewed the manuscript

UTERATtJRE CTTED

Brown, A. 1879. Ballast plants in New Yoric Qty and its vidsity. Bull Tcrrcy BoL Q ub. 6: 353-360. B u r k , L 1877. Plants recently collected on ship’s ballast in the neighborhood of Philadelphia. Proc. Acad. Nau Sci. Philadelphia. 29:105-109. B u r t t , B . L. 1978. Aspects of diversification in the capitulum, pp. 41-59.Itv V. H. Heywood, J. B. Harbomc and B. L. Turner, Eds., The Biology and Chemistry of the Compositae. Academic Press, LondoiL C a b r e r a , A . L. A N D E.M.ZARDINI, 1978. Manual de la Flora de losAlrededores de Buenos Aires. Editorial Acme S. A. C. I., Buenos Aires. C o l e t t a , P. E. a n d K. J. B a i j e r . 1985. United States Navy and Marine Corps Bases, Domestic. Greenwood Press, Westport, CT. CRONQtnsT. A. 1988. The Evolution and Qasdfication of Flowering Plants, 2nd ed. New York Botanical Garden, Bronx, NY. HARTER, E. C. 1985. The Lost Colony of the Confederacy. University Press of Mississippi, University, MS. J e n s q t , S. R., B. J. Nielsen AND R. D a h l c r e n . 1975. Iridoid compounds, their occurrence and systematic importance in the angiosperms. BotN o l 128: 148-180. K o e b e l , W. H. 1912. Argentina Past and Presenu Kegan PauL Trench, Trubner and C o . Ltd., LondoiL M e e r s , j. 1860. On Calyceraceae: Cont.B o l 2: 1-42. M o o n e y , J. L. 1981. Dictionary of American Naval Fighting Ships, VoL VUL Naval Historical Center. Department of the Navy, Washington, DC. R i d l e y ; H .N . 1930. The Dispersal of Plants Throughout the World. L. Reeve and Co., Ltd., Ashford, Kenu Ross, M . a . a n d j . L.H a r p e r . 1972. Occupation of biological space during seedling establishmenu J. EcoL 60: 77-88. Ross, S. R. A N D T. F. M c G a n n . 1982. Buenos Aires: 400 years. University of Texas Press, Austin, TX. S h e t l e r , s . G.a n d L. E. S k o c . 1978. A provisional checklist of species for Flora North America. FL N. Amer. Rep. 84. 2 1 8

1991] D c V qt^— A cicarpha

Small,!.K. 1933. ManioloftheSoatheasttmFlora.(Facsimile)Hafiter,New York, 1972. SK na,!. M.B. 1978. Dispersai and Establishment of Plants, pp. 207-223./n: D. Walker and !. C Guppy, Eds.. Biology and Quaternary Environments. Australian Academy o f % ence, Canberra Gty. W i l l i a m s , G. 1975. Ibe Desert and the Dream. University of Wales Press, Cardifil

DEPARTMENT OF PLANT SCIENCE THE OHIO STATE UNIVERSITY 1735 NEIL AVENUE COLUMBUS, OH 43210 LIST OF REFERENCES

Aguirre, L. (1979). The Southern Andes. In A. E. Nairn, F. G. Stehli, and S. Uyeda (editors), The Southern Andes, vol. 5, pp. 265-376. Plenum Press, New York. Alberch, P., S. J. Gould, G. F. Oster, & D. B. Wake. (1979). Size and shape in ontogeny and phylogeny. Paleobiology 5: 297-317.

Alvarez, L. W., Alvarez, W., Asaro, F., and Michel, H. V., et al. (1982). Current status of the impact theory for the terminal Cretaceous extinction. Geol. Soc. Amer. Spec. Paper 190: 305-315.

Axelrod, D. I. (1937). A Pliocene flora from the Mount Eden beds, California. Publ. Carnegie Inst. Wash. 476,

. (1950). Further studies of the Mount Eden flora, southern California. Publ. Carnegie Inst. Wash. 476.

(1970). Mesozoic paleogeography and early angiosperm history. Hot Rev. 36: 277-319.

Becker, H. F. (1969) . Fossil plants of the Tertiary Beaverhead Basins in southwestern Montana. Palaeontographica 127: 1-142. Bentham, G. (1873). Notes on the classification, history, and geographic distribution of Compositae. J. Linn. Soc. Bot. 13: 335-577.

Bentham, G. (1873). Calyceraceae. Pp 161-163 in Genera plantarum, vol. 2, eds. A. Engler and K. Prantl. Leipzig: W. Engelmann.

Berry, E. W. (1924). The middle and upper Eocene floras of southeastern North America. U.S. Geol. Surv. Prof. Paper 92. Bremer, K. (1992). Ancestral areas: a cladistic reinterpretation of the center of origin concept. Syst. Biol. 41: 436-445.

and Jansen, R. K. (1992). A new subfamily of the

219 220

Asteraceae. Ann. Missouri Bot. Gard. 79: 414-415.

Brown, R. (1817). Observations on the natural family of plants called Compositae. Trans. Linn. Soc. 12: 76- 142.

Brundin, L. Z. (1966). Transantarctic relationships and their significance as evidenced by chironomid midges. K. Sven. Vetenskapsakad. Handl. 11: 1-472. Cabrera, A. L. (1959). Revision del género Dasvphvllum (Compositae). Rev. Mus. La Plata (new ser.) 9: 21- 100. (1977) . Mutisieae— systematic review. In V. H. Heywood, J. B. Harborne & B. L. Turner (editors). The Biology and Chemistry of the Compositae, pp. 1039-1066. Academic Press, London. Cande, S. C. and Mutter, J. C. (1982). A revised identification of the oldest seafloor spreading anomalies between Australia and Antarctica. Earth Planet Sci. Lett. 58: 151-160.

Candolle, A.L.P.P. de. (1836). Calyceraceae. Pp. 1-3 in Prodomus systematis naturalis regni vegetabilis, vol. 5. Paris:Treuttel et Würtz.

Carlquist, S. (1969). Wood anatomy of Goodeniaceae and the problem of insular woodiness. Ann. Missouri Bot. Gard, 56: 358-390. Carolin, R. C. (1959). Floral structure and anatomy in the family Goodeniaceae. Proc. Linn. Soc. N.S.W. 84: 242- 255.

. (1966). Seeds and fruit of the Goodeniaceae. Proc. Linn. Soc. N.S.W. 91:58-83.

_____ . (1967). The concept of the inflorescence in the order Campanulales. Proc. Linn. Soc. N.S.W. 92: 7-26.

. (1970). The trichomes of Goodeniaceae. Proc. Linn. Soc. N.S.W. 96: 8-22.

(1978). The systematic relationships of Brunonia. Brunonia 1: 9-29.

_. (1990). Nomenclatural notes, new taxa and the systematic arrangement in the genus Scaevola (Goodeniaceae) including synonyms. Telopea 3: 477-515. 221

_____, Morrison, D. A. and Rajput, T. (1992). Flora of Australia 35. Australian Government Publishing Service, Canberra. Cassini, H. (1816). Acicarpha (Bot.). Diet. Sci. Nat. 1, Suppl: 32-33. Chung, I. (1965). Revision of Barnadesia (Compositae— Mutisieae). Chicago: published by author. Cialdella , A. M. and Lopez de Riesling, A. G. (1981). Cariologîa de Schlechtendalia luzulaefolia (Compositae). Darwiniana 23:357-360.

Cockerell, T. D. A. (1933) . A fossil goldenrod. Torreya 33: 72.

Condie, K. C. (1989). Plate Tectonics and Crustal Evolution. Pergamon, New York.

Cox, A. and Hart, R. B. (1986). Plate Tectonics: How it works. Blackwell Scientific, Oxford. Craddock, C. (1982). Antarctica and Gondwanaland. In C. Craddock (editor). Antarctic Geoscience, pp. 3-13. Univ. Wisconsin Press, Madison. Crepet, W. L. (1979). Some aspects of the pollination biology of Middle Eocene angiosperms. Rev. Palaeobot. Palynol. 25: 241-252.

. (1984). Advanced (constant) insect pollination mechanisms: pattern of evolution and implications vis- a-vis angiosperm diversity. Ann. Missouri Bot. Gard. 71: 607-630. _____ and Stuessy, T. F. (1978). A reinvestigation of the fossil Vicfuiera croncmistii (Compositae). Brittonia 29: 137-153.

and Friis, E. M. (1987). The evolution of insect pollination in angiosperms. In E. M. Friis, W. G. Chaloner & P. R. Crane (editors). The Origins of Angiosperms and their Biological Consequences, pp. 181- 202. Cambridge Univ. Press, Cambridge. and Nixon, K. (1991). Fossil evidence for the evolution of biotic pollination. Phil. Trans. Roy. Soc. Lond. B33: 187-195.

Covas, G & B. Schanck, (1947). Numéro de cromosomas en antofitas de la region de Cuzco (Republica Argentina). Rev. Argent. Agron. 13: 153-166. 222 Cronquist, A. (1955). Phylogeny and taxonomy of the Compositae. Amer. Midi. Naturalist 53: 478-511. . (1977). The Compositae revisited. Brittonia 29: 137-153.

. (1981). An Integrated System of Classification of Flowering Plants. Columbia Univ. Press, New York.

Dalziel. I. W. (1983). The evolution of the Scotia Arc: A review. In R. L. Olivier, P. R. Jones & J. B. Jago (editors). Antarctic Earth Science, pp. 238-288. Cambridge Univ. Press, Cambridge. _____ and Elliot, D. H. (1973). The Scotia Arc and Antarctic margin. In A. E. M. Mairn & R. G. Stehl (editors). The Ocean Basins and Margins, Vol. 1, The South Atlantic, pp. 171-246. Plenum Press, New York.

Darlington, C. D. and E. K. Janaki Ammal. (1955). Chromosome Atlas of Flowering Plants. George Allen and Unwin, London.

Davis, P. H. & and V. H. Heywood. (1963). Principles of Angiosperm Taxonomy. Van Nostrand, Princeton. Demarcq, G., Méon-Vilain, H., Miquet, R. and Kujawski, H. (1976). Un bassin paralique Néogène: celui de Skanes- Monastir (Tunisie orientale). Notes Serv. Géol. Tunisie 42: 97-147.

DeVore, M. L. (1991). Acicarpha tribuloides in eastern North America. Rhodora 93: 26-35.

_____ . (1993). The systematics of Acicarpha (Calyceraceae). Ohio J. Sci. 92:41. Abstr. Downie, S. and Palmer, J. (1992). Restriction site mapping of the chloroplast DNA inverted repeat: a molecular phylogeny of the Asteridae. Ann. Missouri Bot. Gard. 79: 266-283.

Ezcurra, C. (1985). Revision del genéro Chucmiraoa (Mutisieae, Compositae). Darwiniana 26: 219-284.

and Crisci, J. V. (1987). Relaciones fenéticas entre las especies del género Chucmiraaa (Compositae- Mutisieae): un analisis numérico. Darwiniana 28: 219- 229.

Farris, J. S. (1989). The retention index and rescaled consistency index. Cladistics. 5: 417-419. 2 2 3

Frakes, L. A. (1979). Climates Throughout Geologic Time. Elsevier, Amsterdam. Friis, E. M. and Crepet, W. L. (1987). Time of appearance of floral features. In E. M. Friis, W. G. Chaloner & P. C. Crane (editors). The Origins of Angiosperms and their Biological Consequences, pp. 145-179. Cambridge Univ. Press, Cambridge.

Gamerro, J. C. (1986). Morfologia del polen de Huarpea y su relaciôn con Barnadesia (Mutisieae, Compositae). Darwiniana 26: 219-284.

Gould, S. J. (1977). Ontogeny and Phylogeny. Harvard University Press, Cambridge, England. Gray, J. (1964). Northwest American Tertiary palynology: the emerging picture. In L. M. Cranwell (editor). Ancient Pacific floras, pp. 21-30. Univ. Hawaii Press, Honolulu. Grunow, A. M., Dalziel, I. W. D. and Kent, D. V. (1987). Ellsworth-Whitmore Mountains crustal block, western Antarctica: New paleomagnetic results and their tectonic significance. In Gondwana Six: Structure, Tectonics, and Geophysics, pp. 161-171. Geophysical Monograph Series Vol. 40. Washington DC: American Geophysical Union.

Ha11am, A. (1980). A reassessment of the fit of Pangaea components and the time of their breakup. In D. W. Strangway (editor). The Continental Crust and its Mineral Deposits, pp. 375-387. Geol. Assoc. Canada Spec. Paper 20.

Hansen, H. V. (1991a). Phylogenetic studies in Compositae tribe Mutisieae. Opera Bot. 109:1-50.

(1991b). SEM-studies and general comments on pollen in tribe Mutisieae (Compositae) sensu Cabrera. Nord. J. Bot. 10:607-623.

(1992). Studies in the Calyceraceae with a discussion of its relationship to Compositae. Nord. J. Bot. 12: 63-75.

Hennig, W. (1966). Phylogenetic Systematics. University of Illinois Press, Urbana.

Hicken, C. (1919). Catalogo de las Caliceraceas argentinas. Primera Reunion Nacional de Ciencas Naturales. Pp. 238- 253. 224

Hochuli, P. A. (1978). Palynologische Untersuchungen im Oligozan und Untenniozan der Zentralen und Westlichen Paratethys. Beitr. Palaont. Ost. 4: 1-132. Hock, F. (1894). Calyceraceae. In A. Engler and K. Prantl (editors). Die natUrlichen Pflanzenfamilien, tell 4, abt. 5, eds. W. Engelman, Leipzig.

Howden, P.A. (1981). Zoogeography of some Australian Coleoptera as exemplified by the Scarabaeoidea. In A. Keast (editor). Ecological Biogeography of Australia, vol. 2, pp. 1007-1035. W. Junk, The Hague. Hut, P. et al. (1987). Comet showers as a cause of mass extinctions. Nature 329: 118-127.

Jamieson, B. G. M. (1981). Historical biogeography of Australian Oligochaeta. In A. Keast (editor). Ecological Biogeography of Australia, vol. 2, pp. 885- 921. W. Junk: The Hague.

Jansen, R. K. and Palmer, J. D. (1987). Chloroplast DNA from lettuce and Barnadesia (Asteraceae): structure, gene localization, and characterization of a large inversion. Curr. Genet. 11: 553-564. and _____. (1988). Phylogenetic implications of chloroplast DNA restriction site variation in the Mutisieae (Asteraceae). Amer. J. Bot. 75: 751-764.

Jones, J. B. and Fitzgerald, M. J. (1984). Extensive volcanism associated with the separation of Australia and Antarctica. Science 226: 346-348.

Jussieu, A. L. (1803). Mémoire sur 1^Acicarpha et le Boopis. deux genres nouveaux de plantes de la famille des Cinarocephales. Ann. Mus. Nat. Hist. 2: 345-350. Keast, A. (1981). The evolutionary biogeography of Australian birds. In A. Keast (editor). Ecological Biogeography of Australia, vol. 2, pp. 1587-1635. W. Junk, The Hague.

Kluge, A. J. and J.S. Farris. (1969). Quantitative phyletics and the evolution of anurans. Syst. Zoo. 18: 1-32.

Knowlton, F. H. (1923). Revision of the flora of the Green River Formation, with descriptions of new species. U. S. Geol. Surv. Prof. Paper 131: 133-182. Koch, L. E. (1981). The scorpions of Australia: aspects of 2 2 5

their ecology and zoogeography. In A. Keast (editor). Ecological Biogeography of Australia, vol. 2, pp. 873- 884. W. Junk, The Hague.

Krassilov, V. A. (1973). Mesozoic plants and the problem of angiosperm ancestry. Lethaia 6: 163-178. Krutzsch, W. (1970). Die stratigraphisch verwertbaren Sporen- und Pollenformen des mitteleuropaischen Alttertiars. Jahrb. Geol. Bot. 3: 309-379.

Lammers, T. G. (1988). Chromosome numbers and their systematic implications in the Hawaiian Lobelioideae (Campanulaceae). Amer. J. Bot. 75: 1130-1134.

(1992). Circumscription and phylogeny of the Campanulales. Ann. Missouri Bot. Gard. 79: 388-413.

Leins, P. and Erbar, C. (1990). On the mechanisms of secondary pollen presentation in the Campanulales- -complex. Botan. Acta 103: 87-92.

Leopold, E. B. and MacGinitie, H. D. (1972). Development and affinities of Tertiary floras in the Rocky Mountains. In A. Graham (editor). Floristics and Paleofloristics of Asia and Eastern North America, vol. 2, pp. 147-200. Elsevier, Amsterdam.

Lessing, C. F. (1831). Calyceraceae Rich. Linnaea. 6:258- 260.

MacGinitie, H. D. (1933). The Trout Creek flora of southeastern Oregon. Publ. Carnegie Inst. Wash. 416. Main, B. Y. (1981). A comparative account of the biogeography of terrestrial invertebrates in Australia: some generalizations. In A. Keast (editor). Ecological Biogeography of Australia, vol. 2, pp. 1055 1077. W. Junk, The Hague.

Marshall, L. G. (1980). Marsupial paleobiogeography. In L. L. Jacobs (editor). Aspects of Vertebrate History: Essays in Honor of Edwin Harris Colbert, pp. 345-386. Museum of Northern Arizona Press, Flagstaff. McGowran, B. (1990). Fifty million years ago. Amer. Scientist 78: 30-39.

Miers, J. (1860). On the Calyceraceae. Contr. to Botany 2: 1-42.

Menke, B. (1976). Pliozane und altestquartare Sporen- und 226

Pollenflora von Schleswig-Hollstein. Geol. Jahrb. A32: 3-197. MonasteriOf M. (1986). Adaptive strategies of Espeletia in the Andean desert pâramo. In F. Vuilleximier & M. Monasterio (eds.). High Altitude Tropical Biogeography, pp 49-80. Oxford University Press, Oxford. Moore, D. M. (1967). Further records for the vascular flora of the Falkland Islands. Bot. Notiser 120: 2-24. (1968). The vascular flora of the Falkland Islands. Brit. Antarc. Surv. Sci. Rep. 60: 1-202.

_. (1983a). The flora of the Fuego-Patagonia Cordillera: its origins and affinities. Rev. Chil. Hist. Nat. 56: 123-136.

(1983b). Flora of Tierra del Fuego. Missouri Botanical Garden, St. Louis, Missouri. Muller, J. (1981). Fossil pollen records of extant angiosperms. Bot. Rev. 47: 1-42.

(1984). Significance of fossil pollen for angiosperm history. Ann. Missouri Bot. Gard. 71: 419- 443.

Nagy, E. (1969). Palynological elaborations of the Miocene layers of the Bakony Mountains (Valpalota) of Hungary. Acta Bot. 8: 153-163. Nowicke, J. W. and Skvarla, J. J. (1974). A palynological investigation of the genus Tournefortia (Boraginaceae). Amer. J. Bot. 61: 1021-1036.

Olmstead, R. G . , Jansen, R. K., Michaels, H. J., Downie, s. R. and Palmer, J. D. (1990). Chloroplast DNA and phylogenetic studies in the Asteridae. In S. Kawano (editor). Biological Approaches and Evolutionary Trends in Plants, pp. 119-134. Academic Press, London.

, Michaels, H. J., Scott, K. M. and Palmer, J. D. (1992). Monophyly of the Asteridae and identification of their major lineages inferred from DNA sequences of rbcL. Ann. Missouri Bot. Gard. 79: 249-265.

Panero, J. L. 1992. Systematics of Pappobolus (Asteraceae- Heliantheae). Syst. Bot. Monographs 36: 1-195.

Pares Regali, M. de S., Uesugui N. and Santos, A. de S. (1974). Palinologia dos sedimentos meso-cenozoicas do 227

Brasil (I, II). Bol. Teen. Petrobras, Rio de Janeiro. 17: 177-191, 263-301.

Parra, 0. and Marticorena, C. (1971). Granos de polen de plantas chilenas. II. Compositae-Mutisieae. Gayana 21: 3-107.

Parrish, J. T. (1987). Global palaeogeography and palaeoclimate of the Late Cretaceous and Early Tertiary. In E. M. Friis, W. G. Chaloner & P. R. Crane (editors). The Origins of Angiosperms and their Biological Consequences, pp. 51-74. Cambridge University Press, Cambridge.

. (1990). Gondwanaland paleogeography and paleoclimatology. In T. N. & E. Taylor (editors). Antarctic Paleobiology, pp. 15-26. Springer-Verlag, New York.

_____ and Curtis, R. L. (1982). Atmospheric circulation, upwelling, and organic-rich rocks in the Mesozoic and Cenozoic. Palaeogeogr. Palaeoclim. Palaeoecol. 40: 31- 66. Ziegler, A. M. and Scotese, C. R. (1982). Rainfall patterns and the distribution of coals and evaporates in the Mesozoic and Cenozoic. Palaeogr. Palaeoclimat. Palaeoecol. 40: 67-101.

Patterson, C. (1980). Cladistics. Biologist. 27:234-240.

Peacock, W. J. (1963). Chromosome numbers and cytoevolution in the Goodeniaceae. Proc. Linn. Soc. New South Wales 88: 8-27. Pesacreta, T. C., Sullivan, V. I. and Hasenstein, K. H. (1993). The connective base of Cirsium horridulum (Asteraceae): description and comparison with the visoelastic filament. Amer. J. Bot. 80: 411-418.

Pesecreta, T. C., V. I. Sullivan, and M. L. DeVore. The connective base of Acicarpha tribuloides. Amer. J. Bot. 81: (in press).

Platnick, N. I. (1985). Philosophy and the transformation of cladistics. Cladistics 1: 87-94. Pontiroli, A. (1963). Flora Argentina, Calyceraceae. Rev. Mus. La Plata, sec. Bot. 9: 175-241.

Rahn, K. 1960. Chromosome numbers in some South American angiosperms. Bot. Tidsskr. 56: 117-127. 228 Raven, P. H. and Axelrod, D. I. (1974). Angiosperm biogeography and past continental movements. Ann. Missouri Bot. Gard. 61: 539-673.

Raven, P. H. (1975). The bases of angiosperm phylogeny: cytology. Ann. Missouri Bot. Gard. 62: 724-764. Richard, M. (1820). Mémoire sur une famille de plantes dites les Calycérées. Mémoires d^Histoire Naturelle Botanique. 6: 28-82.

Robinson, H. (1987). Some suggestions regarding the significance of chloroplast DNA variation in the Asteraceae. Phytologia 65:316-324.

Rodriguez, C., W. T. Ormond, and M. C. Bezerra Pinheiro. (1977). Contribucao à citologia de Acicarpha spathulata R. Brown (Calyceraceae) . Bol. Mus. Nac. Nova Ssrie 45: 1.

Rohlf, F. J. (1988). NTSYS-pc. Numerical taxonomy and multivariate analysis system. Version 1.50. Setauket, Exeter Publishing Ltd, New York. Salard-Cheboldaeff, M. (1978). Sur la palynoflore Maestrichtienne et Tertiaire du bassin sédimentaire littoral du Cameroun. Pollen et Spores 20: 215-260.

Segal, R. (1965). New fossil fruit (Compositae) from the Pliocene of western Kansas. Amer. Midi. Nat. 73: 430- 432.

Shackleton, N. J. (1986). Paleogene stable isotope events. Palaeogeog. Palaeoclimat. Palaeoecol. 57: 91-102.

Skvarla, J. J., Turner, B. L., Patel, V. C. and Tomb, A. S. (1977). Pollen morphology in the Compositae and in morphologically related families. In V. H. Heywood, J. B. Harborne & B. L. Turner (editors). The Biology and Chemistry of the Compositae, pp. 141-248. Academic Press, London. Small, J. (1919). The origin and development of the Compositae. Reprint of New Phytologist No. 11.

Solbrig, O. T. (1977). Chromosomal cytology and evolution in the family Compositae. In V. H. Heywood, J. B. Harborne & B. L. Turner (editors), The Biology and Chemistry of the Compositae, pp. 264-281. Academic Press, London.

Stebbins, G. L. (1974). Flowering Plants: Evolution Above 229

the Species Level. Harvard University Press, Cambridge.

Stebbins, G. L. (1977). Development and comparative anatomy of the Compositae. In V. H. Heywood, J. B. Harborne & B. L. Turner (editors), The Biology and Chemistry of the Compositae, pp 91-109. Academic Press, London.

Stevens, P. F. (1981). On ends and means, or how polarity criteria can be assessed. Syst. Bot. 6:186-188.

Stuessy, T. F., and Sagastegui A., A. (In press). Revision de Arnaldoa (Compositae, Barnadesioideae), género endêmico del norte del Peru. Arnaldoa.

_____ and Sang, T. (1993). Phylogeny of Barnadesioideae and early evolution of Compositae. Ohio J. Sci. 92:41. Abstr.

Swofford, D.L. (1991) . PAUP: Phylogenetic analysis using parsimony, version 3.0. Champaign: Illinois Natural History Survey.

Takhatajan, A. (1959) . Die Evolution der Angiospermen. Gustav Fischer, Jena. Thorne, R. F. (1972). Major disjunctions in the geographic ranges of seed plants. Quart. Rev. Biol. 47: 365-411.

. (1978). Plate tectonics and angiosperm distribution. Notes Royal Bot. Gard. Edinburgh 36:297- 315.

(1986). Antarctic elements in Australasian rainforests. Telopea 2:611-617.

Tiffney, B. (1984) . Seed size, dispersal syndromes, and the rise of the angiosperms: evidence and hypothesis. Ann. Missouri Bot. Gard. 71: 551-576.

Tindale, N. B. (1981). The origin of Lepidoptera relative to Australia. In A. Keast (editor). Ecological Biogeography of Australia, pp. 955-976. W. Junk, The Hague.

Turner, B. L. (1973). lOPB chromosome number reports LXII. Taxon 27: 519-535.

Turner, B. L. (1977). Fossil history and geography. In V, H. Heywood, J. B. Harborne & B. L. Turner (editors). The Biology and Chemistry of the Compositae, pp. 21-39. 230 Academic Press, London.

, B. L., J. Bacon, L. Urbatsch, and B. Simpson. 1979. Chromosome numbers in South American Compositae. Amer. J. Bot. 66; 173-178.

Weissal, J. K. and Hayes, D. E. (1972). Magnetic anomalies in the Southeastern Indian Ocean. In D. E. Hayes (editor). Antarctic Oceanography II: Australia-New Zealand Sector, pp. 234-249. Antarctic Res. Ser., Vol. 19. American Geophysical Union, Washington, D.C. and Herron, E. M. (1977). Plate tectonic synthesis; the displacements between Australia, New Zealand, and Antarctica since Late Cretaceous. Marine Geol. 25: 231- 277.

Wheeler, Q.D. 1981. The ins and outs of character analysis: A response to Crisci and Stuessy. Systematic Botany 6:297-306.

Williams, W. D. (1981). Aquatic insects: an overview. In A. Keast (editor). Ecological Biogeography of Australia, vol. 2, pp. 1213-1229. W. Junk, The Hague.

Ziegler, A. M., Scotese, C. R. and Barrett, S. F. (1983). Mesozoic and Cenozoic paleogeographic maps. In P. BrosChe & J. Sundermann (editors). Tidal Friction and the Earth's Rotation, vol. 2, pp. 240-252. Springer- Ver lag, Berlin.