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Systematics of the genusMulinum pers. (Apiaceae, Hydrocotyloideae, Mulineae)

Zech, James Craig, Ph.D.

The Ohio State University, 1992

UMI 300 N. ZeebRd. Ann Aibor, MI 48106 SYSTEMATICS OF THE GENUS MULINUM PERS. (APIACEAE, HYDROCOTYLOIDEAE, MULINEAE)

DISSERTATION

Presented in Partial Fulfillment of the Requirements for

the Degree Doctor of Philosophy in the Graduate

School of The Ohio State University

By

James Craig Zech, A.A., B.S., M.S.

******

The Ohio State University

1992

Dissertation Committee: Approved by

Daniel J. Crawford

Fred Sack

Tod F. Stuessy Departmen lant Biology My Parents ACKNOWLEDGEMENTS

I would like to thank Dr. Tod F. Stuessy for his help and

direction not only with this dissertation but also during my

years at Ohio State. T thank Dr. Daniel J. Crawford for

serving on my committees, for allowing me to work in his

laboratory, and also for his advice and guidance. In

addition, I thank my remaining committee members, Drs. Fred

Sack (Advisory and Dissertation Committees) and Gary L. Floyd

(Advisory Committee) for their advice and support.

Without the participation of many people this work would

not have been possible. I thank Dr. Lincoln Constance for his

continued interest and encouragement and Dr. Thomas Duncan and

the University of California, Berkeley Herbarium staff for

their kindness and assistance. Because my field collecting would have been impossible without their support and direction, I thank Dr. Jorge V. Crisci at the Museo de La

Plata, Argentina and Dr. Oscar O. Parra and Professor

Clodomiro Marticorena at the Universidad de Concepcion Chile.

I thank Maria Marta Cigliano, Claudio Contogiorgakis, Mario

Gentilli, Patricia Gentilli, Liliana Katinas, Alegandra

Moschetti, Alberto Ripalta, Fidel Alegandro Roig, Tod F.

Stuessy, Ana Maria Zavattieri, and especially Juan Jose

iii Morrone for help while in the field. Many thanks go to the

directors, curators, and staff of the following herbaria: B,

BM, CM, CONC, CORD, E, F, G, GH, GOET, K, L, LAU, LP, LY, MA,

MO, NY, OS, OXF, P, PR, PRC, S, SGO, SI, UC, UPS, US, and W.

Special thanks go to Dr. John J. Fur low (OS) for loan

assistance, Maryan Benkowski (OS) for help with annotation

labels, Mauricio Velayos (MA), Melica Mufioz Schick (SGO), and

Esther G. de Mautino (CORD) for sending additional type

material information, and Harald Riedl (W) for his efforts in

type location.

A very special thanks go to my colleagues, and friends,

Jorge Arriagada, Maryan Benkowski, Marybeth Cosner, Patty

Pacheco, Hugo Valdebenito, Loni Walker, and Jun Wen for hours of talks and constant support. I thank Sabine Zech for German translation, and Dr. R. L. Stuckey for the use of his computer. Finally, I thank my parents, A1 and Gerry Zech who have, more than anyone, supported me unconditonally during the bad times as well as the good times.

iv VITA

December 22, 1960 ..... Born - Grand Rapids, Michigan

1 9 8 1 ...... A.A., Grand Rapids Junior College, Grand Rapids, Michigan

1983 ...... B.S., Central Michigan University, Mt. Pleasant, Michigan

1984-1986 ...... Teaching Assistant, Central Michigan University, Mt. Pleasant, Michigan.

1986 ...... M.S., Central Michigan University, Mt. Pleasant, Michigan

1986-1991 ...... Teaching Associate, The Ohio State University, Columbus, Ohio

1988 ...... Research Assistant, Harvard University, Cambridge, Massachusetts.

1989-1991 ...... Lecturer, The Ohio State University, Columbus, Ohio

1 9 9 1 ...... Lecturer and Laboratory Instructor, Central Ohio Technical College, Newark, Ohio

1991-1992 ...... Lecturer and Laboratory Instructor, The Ohio State University - Newark Campus, Newark, Ohio

PUBLICATIONS

J.C. Zech. 1987. Micromorphological Characteristics of the Tegumen Layer Among Three Michigan Species of Luzula (Juncaceae). Abstract. Ohio J. Sci. 87: 5. . 1988. The Juncus effusus Complex: The Latest Look. Abstract. Ohio J. Sci. 88: 12.

. 1989. Flavonoid Survey of Four Species Within The Tribe Mulineae (Hydrocotyloideae, Apiaceae). Abstract. Ohio J. Sci. 89: 7.

. 1989. A Preliminary Flavonoid Survey of the Genus Mulinum Pers. As Well As Other Taxa of the Tribe Mulineae (Hydrocotyloideae, Apiaceae). Abstract. Amer. J. Bot. 76: 285.

. 1990. Phenetic Analysis of Fruit Characters of the Genus Mulinum Pers. (Mulineae, Hydrocotyloideae, Apiaceae). Abstract. Ohio J. Sci. 90: 6.

. 1990. Phenetic Analysis of Fruit Characters of the Genus Mulinum Pers. As Well As Other Genera Within the Mulineae (Hydrocotyloideae, Apiaceae). Abstract. Amer. J. Bot. 77: 437.

. 1991. The Cushion Habit and the Genus Mulinum Pers. (Apiaceae). Abstract. Ohio J. Sci. 91: 27.

. 1991. Mulinum Pers. (Apiaceae) and the Cushion Habit: The Environmental Factors Which Cause Cushion Formation and Relationships to the Biogeography and Phylogeny of the Genus. Abstract. Amer. J. Bot. 78: 231.

. 1992. Classical Taxonomic Problems Encountered During the Revision of the Genus Mulinum Pers. Abstract. Ohio J. Sci. 92: 17.

Zech, J. C. and D. E. Wujek. 1986. The Micromorphological Characteristics of the Tegumen Layer Between and Among Species of Juncus and Luzula including Varieties of the Juncus effusus Complex. Abstract. Pap. Michigan Acad. Sci.

. 1990. Scanning Electron Microscopy of Seeds in the Taxonomy of Michigan Juncus. Michigan Bot. 29: 3-18.

FIELDS OF STUDY

Major Field: Plant Biology

Studies in Plant Systematics with Professor Tod F. Stuessy

vi TABLE OF CONTENTS

DEDICATION ...... ii

ACKNOWLEDGEMENTS ...... iii

VITA ...... v

LIST OF TABLES ...... ix

LIST OF FIGURES ...... xi

CHAPTER PAGE

I. REVISION OF THE GENUS MULINUM PERS. (APIACEAE, HYDROCOTYLOIDEAE, MULINEAE) ...... 1

Introduction ...... 2 Taxonomic History ...... 3 Generic Relationships ...... 8 Habitat and Distribution ...... 11 Reproductive Morphology ...... 12 Chromosome Number ...... 15 Phylogenetic Analysis of Mulinum ...... 16 Biogeography ...... 24 Taxonomic Categories ...... 28 Morphology and Taxonomic Criteria ...... 28 Taxonomy ...... 33 Key to Species of M ulinum...... 36 Excluded and Doubtful Names ...... 118 Acknowledgements ...... 125 Literature Cited ...... 126 Numerical List of Taxa ...... 142 Index to Numbered Collections Examined ... 143

vii II. EVOLUTION OF CUSHION HABIT IN MULINUM (APIACEAE) ...... 155

Introduction ...... 156 General Factors Affecting Cushion Formation...... 160 Factors Affecting Mulinum Cushion Formation ...... 166 Evolution of the Cushion Habit in Mulinum ...... 172 Literature Cited...... 178

III. FLAVONOIDS OF MULINUM (APIACEAE, HYDROCOTYLOIDEAE, MULINEAE) ...... 187

Introduction ...... 187 Materials and Methods...... 191 Acquisition of Plant Material ...... 191 Extraction, Isolation, and Identification of Flavonoids ...... 191 Results ...... 193 Discussion ...... 196 Subfamilial and Tribal Flavonoid Variation ...... 196 Generic and Infrageneric Flavonoid Variation inMulinum ...... 199 Evolution of the Flavonoid System in Mulinum ...... 201 Literature Cited ...... 2 05

LIST OF REFERENCES ...... 210

viii LIST OF TABLES

TABLE PAGE

1. Chromosome numbers for species of Mulinum .... 17

2. Characters and character states used in the cladistic analysis of Mulinum. 0 = primitive; 1,2 = derived ...... 18

3. Character state matrix for phylogenetic reconstruction of Mulinum. 0 = primitive; 1, 2, = derived. See Table 2 for description of characters and states. Azorella (A. spinosa; AZO) was used as outgroup (see text). Order of Mulinum taxa is that used in taxonomic treatment ...... 19

4. Rauh's (1939) categories of cushion plants ... 157

5. Suggested environmental factors contributing to the habit of Mulinum cushion species ..... 167

6. Climatic descriptions for selected cities of Argentina (Rudloff 1981). City location corresponds to Mulinum1s distribution; * corresponds to Mulinum cushion species distribution ...... 169

7. Winds of Argentina (Rudloff 1981), which correspond to Mulinum's distribution ...... 170

8. Flavonoid composition of the Apiaceae following the classification of Drude (1897). * = tribes where flavones are absent or rare (Harborne, 1971) ...... 190

9. Collections of Mulinum species examined for flavonoid compounds. Collection numbers are those of Stuessy et al. (S), Zech & Contogiorgakis (ZC), or Zech et al. (Z) unless noted otherwise. A = Argentina, C = Chile; * = samples from herbarium material ... 192

ix 10. Identities of flavonoids isolated from Mulinum ...... 194

11. Distribution of flavonoid compounds in Mulinum ...... 195

12. Distribution of flavones and flavonols in some genera of the Mulineae. Collection numbers are those of Stuessy et al. (S), Zech & Contogiorgakis (ZC), or Zech et al. (Z) unless noted otherwise. A = Argentina, C = Chile ...... 197

x LIST OF FIGURES

FIGURES PAGE

1. Distribution of Mulinum Pers. in southern South America ...... 13

2. Cladogram of taxa of Mulinum. The derived character states are indicated by numbers (cf. Tables 2 and 3) and single bars. Double bars and asterisks indicate parallelisms and reversals, respectively. The names of taxa are represented by the first three letters of the specific epithets ...... 22

3. Consensus Cladograms of taxa of Mulinum. A, Strict. B, Semistrict. C, 50% Majority Rule. D, Adams. The names of taxa are represented by the first three letters of the specific epithets ...... 23

4. Cross section of fruit. A. Mulinum spinosum (based on Boelcke et al. 10309, UC). B. Mulinum leptacanthum (based on Correa et al. 5855, UC) ...... 43

5. Distribution of Mulinum spinosum in Argentina and Chile. • = M. spinosum var. spinosum; ▲ = M. spinosum var. minus; m = M. spinosum var. atacamense; ★ = M. spinosum var. proliferum...... 44

6. Distribution of Mulinum leptacanthum in Argentina and Chile ...... 68

7. Cross section of fruit. A. Mulinum crassifolium (based on Pfister 9374, CONC). B. Mulinum ulicinum (based on Ruthsatz 166, UC) 73

xi 8. Distribution of Mulinum crassifolium in Chile, Regiones II and III, and Argentina, Provinces Jujuy and Salta ...... 75

9. Distribution of Mulinum ulicinum in Argentina and Bolivia. C = Catamarca; LR = La Rioja; M = Mendoza; J = Jujuy; S = Salta; SJ = San Juan; T = Tucuman. • = M. ulicinum var. ulicinum; a = M. ulicinum var. multiflorum ...... 79

10. Cross section of fruit. A. Mulinum albovaginatum (based on Ruiz L. 7183, UC) . B. Mulinum valentini (based on Grondona 2202, UC) ...... 89

11. Distribution of Mulinum albovaginatum in Argentina, Provinces Mendoza and Neuquen, Chile, Region VII. • = M. albovaginatum var. albovaginatum; a = M. albovaginaatum var. pauciflorum...... 90

12. Distribution of Mulinum valentini in Argentina and Chile, Region XII ...... 100

13. Cross section of fruit. A. Mulinum hallei (based on O fDonell 3663, UC). B. Mulinum microphyllum (based on Correa & Constance 4085/3838, UC) ...... 106

14. Distribution of Mulinum hallei in Argentina ...... 107

15. Distribution of Mulinum microphyllum in Argentina ...... „..... 113

16. Distribution of Mulinum cushion species within southern South America. • = M. albovaginatum. a = M. hallei. ■ = M. microphyllum. ★ = M. valentini ...... 159

17. Suggested evolution of the cushion habit for Mulinum. A. Shrub; B. Subshrub; C. Cushion ...... 173

18. Cladogram of taxa of Mulinum, showing the cushion clade. The names of taxa are represented by the first three letters of the specific epithets ...... 175

xii 19. Distribution of the genus Mulinum in southern South America, showing the Chubutan corridor ...... 176

20. Cladogram of taxa of Mulinum. The names of taxa are represented by the first three letters of the specific epithets. Numbers correspond to the flavonoids listed in Table 3 ...... 202

xiii CHAPTER I

REVISION OF THE GENUS MULINUM PERS.

(APIACEAE, HYDROCOTYLOIDEAE, MULINEAE)

ABSTRACT. Mulinum (Apiaceae, Hydrocotyloideae, Mulineae) is a genus of shrubs, subshrubs, and cushions endemic to southern South America with concentrations along the southern coast of Argentina and the Andes of both Chile and Argentina.

The genus is most closely related to Azorella (especially A. spinosa) . Species recognition has been difficult in the genus due to morphological character variability within species and reproductive character similarity between species. This study is based on comparative morphology, including field and herbarium studies, cladistic studies, and phytochemistry. The reproductive morphology of Mulinum. controversial in previous studies, is shown to be hermaphroditic with perfect flowers.

Morphological data, combined with cladistic studies, have led to a taxonomic treatment including eight species and eight varieties. These taxa are grouped into two newly-described sections, one with two subsections, and one of these subsections with two series, i.e., Section Mulinum. sect.

Leptacantha [subsect. Leptacantha. subsect. Crassifolia (ser.

1 Crassifolia, ser. Azorellopsis)]. One variety is described as

new, M. spinosum var. atacamense, and new combinations are

made for M. albovaginatum var. pauciflomim, M . spinosum var.

minus, and M. ulicinum var. multiflorum. The polyploid origin

of M. microphyllum is discussed, with evidence from

morphology, distribution, flavonoid chemistry, and chromosome

size suggests that it may be an allopolyploid between diploid

species M. hallei and M. valentini.

INTRODUCTION

Mulinum Pers. is one of the most taxonomically difficult

genera of the family Apiaceae (Mathias 1971; Constance, in

litt.). Taxa display an overall uniformity of reproductive

structures and a general plasticity of vegetative features,

especially leaf morphology, which complicate attempts at

classification and identification.

Mulinum is one of the last genera of Apiaceae in South

America to be studied systematically (others include: Apium L.

and Eryngium L.), and the first to be studied using modern techniques. Those studied previously include: Asteriscium

Cham. & Schlechtend., Domeykoa Philippi, Eremocharis Philippi,

Gymnophyton Clos, and Pozoa Lagasca (Mathias & Constance

1962). Also, the combination of shrub, subshrub, and cushion habits, secondary compound chemistry, and adaptive morphology make Mulinum one of the most interesting genera of the family. Some species of Mulinum appear to play a significant role as pioneers, especially as cushion taxa (Rauh 1939, 1940) within barren or disturbed areas, and may be the dominant component of particular landscapes (e.g., M. spinosum).

The genus Mulinum has a rich taxonomic history. During its 187-year history there have been over forty names associated with Mulinum at the generic, sectional, specific, and infraspecific levels. It is linked with the early exploration of South America and was originally studied primarily by European botanists who understood little about it's natural history. Mulinum has been associated with a number of other genera of Umbelliferae (e.g., Azorella Lam.,

Gymnophyton) and a synthetic treatment of its Chilean,

Argentinian, and Bolivian taxa has never been produced.

Currently eight species and eight varieties are recognized, some of which are highly polymorphic.

TAXONOMIC HISTORY

Mulinum was created from the genus Selinum L. by Persoon in 1805 by the transferring of four species: M . acaule (Cav.)

Pers., M. microphyllum (Cav.) Pers., M. proliferum (Cav.)

Pers., and M. spinosum (Cav.) Pers. Selinum is a primarily

European genus which appears to have served at the time as a convenient taxon for the placement of new Umbelliferae species. Cavanilles (1799) originally placed these four species within Selinum, and not within a genus of their own, perhaps because of the general lack of knowledge about South

American Umbelliferae. In addition to Mulinum, species of

Selinum have been associated with several other genera, such as Conium L. (Selinum conium E. H. L. Krause = C. maculatum

L.) , Coriandrum L. (Selinum coriandrum E. H. L. Krause = C. sativum L.), and Cuminum L. (Selinum cuminum E. H. L. Krause

= C. cyminum L.), which were included within Selinum by Krause in 1904.

After its conception, Mulinum was treated 25 years later by Candolle (1830) who recognized eight species (including

Persoon's original four) divided into three sections: "I",

"II", and "Dipterygia". The first contained four species mirroring modern-day Mulinum, and the second had two species which in the future would be placed within other genera [i.e.

M. acaule (Cav.) Pers. within Laretia Gillies & Hook, and M. angulatum DC. within Pozoa]. Candolle created a third section, Dipterygia, by transferring two of K. Presl's species of the genus Dipterygia K. Presl into Mulinum. Candolle

(1830: 80) relayed his doubts, which were later justified, about the transfer of the two Dipterygia species into Mulinum by going as far as to ask, "An genus proprium?". In 1847, both Clos and Turczaninow independently removed section

Dipterygia from Mulinum elevating it to generic status as

Gymnophyton and Tritaenicurn Turcz., respectively [Tritaenicum was latter combined with Gymnophyton, as was the genus Dipterygia, by Mathias & Constance (1962)]. In addition, Clos

included Mulinum in his tribe "Mulineas" and stated that

Mulinum was better represented by Candolle's first section.

In 1867, Bentham placed Mulinum within his first "series"

Heterosciadiae (= subfamily) and second tribe Mulineae for the

treatment of the Umbelliferae in Bentham and Hooker's Genera

Plantarum. Although not named specifically, Bentham included

four species within the genus.

Mulinum was placed within Baillon's fifth tribe

Hydrocotyleae in his (1879) treatment of the Ombelliferes in

Histoire des plantes, He presented further generic relationships of Mulinum comparing it with Gymnophyton, and other Hydrocotyleae genera such as Asteriscium, Diposis DC., and Huanaca Cav. Baillon regarded the fruit of Asteriscium as similar to Mulinum, and Mulinum's flowers as similar to those of Laretia, which he subsequently compared to those of

Azorella.

Drude (1897) included Mulinum in his treatment of the

Umbelliferae for Engler and Prantl's Die natiirlichen

Pflamentamilien. Although Drude did not treat Mulinum infragenerically, he placed the genus within subfamily

Hydrocotyloideae and tribe Mulineae. He divided Mulineae into two subtribes, Azorellinae and Asteriscinae placing Mulinum within the latter. In addition, Drude included Asteriscium

(Gymnophyton was placed as a subgenus of Asteriscium),

Diposis, Hermas L., and Laretia and by doing so first eluded to the possible evolutionary relationships among these genera.

It should be noted however, that Drude excluded both Azorella and Pozoa from the Asteriscinae, placing them within his second subtribe Azorellinae. These genera have been linked both taxonomically and evolutionarily not only with Mulinum but also with other genera of the Asteriscinae. Drude's intrafamilial classification is still widely recognized today,

(e.g, Constance 1971).

In 1898 Kuntze included four species and four varieties within Mulinum as part of his second entry of Umbelliferae for his Revisio Generum Plantarum. Kuntze was the first to recognize infraspecific taxa for Mulinum (i.e., M. spinosum).

The only work on Mulinum completed in North America was

Macloskie's (1905) , based upon collections from the University of Princeton's expeditions to Patagonia during 1896-1899.

Although he included 17 species within Mulinum, he only named ten species encountered within Patagonia.

Through the 19th century, Mulinum has been treated by

North American and European botanists, who had scanty material at their disposition. As a result of new and more ample collections, many of the species originally placed within

Mulinum have since been transfered to other genera (e.g., M. acaule to Laretia).

It wasn't until the late 19th and early 20th centuries that comprehensive classifications of Mulinum produced from field study began appearing. This reduced the problem of classifications by botanists not familiar with the natural history of the group, but it further increased another— nomenclatural problems due to a lack of communication between South American, North American, and European workers and between workers within neighboring Chile and Argentina.

Although R. A. Philippi described a total of seven species for the genus, more than any other author, it wasn't until 1899 that an in-depth treatment of Mulinum was produced by Reiche, as part of his ongoing publication of the Flora de

Chile, the first completed by a South American. Reiche followed Drude's (1897) earlier intrafamilial classification insofar as including Mulinum within subfamily

Hydrocotyloideae. He recognized seven species within the genus, and submerged four Mulinum species within M. spinosum at unspecified infraspecific rank, perhaps to illustrate M. spinosum's polymorphism. Reiche (1899: 799) indicated

Mulinum's affinities to Azorella by stating, "Sin tener a la vista el fruto, este jenero se separa dificilmente de

Azorella". For the first time Mulinum's ties to Azorella were documented and the general problematic nature of the genus presented.

Within neighboring Argentina, no treatment of Mulinum has been completed for the entire country. Botanists such as

Hieronymus (1881) and Italian-born Argentine Spegazzini (1899,

1902) described a number of taxa but never considered Mulinum as a whole. This only added to the growing taxonomic disarray of the genus, which remained treated primarily by European and

Chilean botanists.

Two recent treatments of Mulinum are Hiroe's (1979), for his Umbelliferae of World and Constance's (1988) for Flora

Patagonica. Hiroe based his work solely on herbarium collections (AHU, GH, KYO, PC, TI, TNS, TUF, UC, US), excluding material housed in South American herbaria and excluding the examination of type material. Hiroe reduced thirty species to five, combining species which clearly have little or no affinity (e.g., M. leoninum with M. microphyllum). Constance conservatively treated seven

Patagonian species. Although his was not a complete evaluation of the genus, Constance described Mulinum's current condition: "No ha sido nunca revisado sistematicamente y estd en un estado algo confuso” (1988: 357).

GENERIC RELATIONSHIPS

Drude (1897) includes Mulinum in his subtribe

Asteriscinae along with four other genera: Asteriscium

(including Gymnophyton), Diposis, Hermas, and Laretia.

While Mulinum and Asteriscium have characteristics in common (e.g., perennial, winged fruit, subshrub habit, ternately compound leaves), these features represent part of a wide range of variation found within Asteriscium (Mathias &

Constance 1962). Asteriscium differs further from Mulinum by chromosome number, n = 10 vs. n = 8, 16, and 24 (Bell &

Constance 1957; Constance et al. 1971, 1976), basal leaves

(Mathias & Constance 1962) vs. cauline, and the possession of a median abaxial gland on each petal (Mathias & Constance

1962) vs. glands absent.

Mulinum shares several characteristics with Gymnophyton.

Both have winged fruit, pollen similar in size and structure

(e.g., thick exine; Heusser 1971), and a similar flavonoid compound profile (see Chapter 3). However Gymnophyton differs from Mulinum in chromosome number, n = 10 vs. n = 8, 16 and 24

(Bell & Constance 1966; Constance et al. 1971, 1976), primarily basal leaves with multiple lobes (Mathias &

Constance 1962) vs. cauline leaves with primarily 3 lobes, and shrubs (Mathias & Constance 1962) vs. shrubs, subshrubs, and cushions.

Diposis are succulent, glabrous perennials, with few cauline leaves, and white flowers (Constance 1988), differing significantly from Mulinum which is dry, generally pubescent, with only cauline leaves, and yellow flowers. The two genera are similar in having winged fruit and simple umbels.

Hermas also differs from Mulinum by the production of compound umbels vs. simple umbels, narrow leaves which are similar in size to the calyx lobes vs. leaves much larger than the calyx lobes, and chromosome numbers, n = 7 vs. n = 8, 16, and 24 (Constance et al. 1971, 1976).

While taxa of Laretia produce winged fruit, and possess 10

a compact habit, the leaves are uniformly entire, differing

from the lobed leaves of Mulinum.

Although Drude (1897) placed Azorella in his subtribe

Azorellinae, the genus is vegetatively similar to some species

of Mulinum. Azorella and Mulinum share the same chromosome

number, n = 8 (Bell & Constance 1960, 1966; Constance et al.

1971, 1976) and some taxa are similar by having cauline leaves

with 3-5, often 3, lobes (e.g., A. spinosa Ruiz Lopez & Pav6n)

and the cushion habit (e.g., A. compacta Philippi). Azorella

is clearly defined and separated by the presence of wingless

fruit and pollen which is smaller with a thinner exine

(Heusser 1971; Markgraf & D'Antoni 1978).

Wolff (1924) described Azorellopsis (including

Azorellopsis trisecta H. Wolff) based upon material collected by Bridges. Previous (Constance 1965) and current searches

(Riedl, pers. comm.; Hiepko, in litt.) failed to locate the holotype or a specimen annotated by Wolff. While differences exist in Wolff's descriptions when compared to Mulinum (i.e., winged peduncles and compound umbels, both absent in Mulinum) ,

Azorellopsis is combined with Mulinum and A. trisecta with M. pauciflorum Reiche (= M. albovaginatum var. pauciflorum), a combination first made by Constance (1965), and until further evidence exists to the contrary, the placement of these taxa in Mulinum will remain.

Conflicting data make selection of a close extant relative to Mulinum difficult. Of the seven genera discussed, 11

Gymnophyton and Azorella are the most likely. Winged fruit is the strongest morphological character which links Gymnophyton and Mulinum. While both produce winged fruit, the degree of wing development is variable in Gymnophyton, ranging from minimally to well developed (Mathias & Constance 1962), and probably represents an independent parallel evolution of this character.

Azorella is most closely related to Mulinum, based strongly on chromosome number. Because Azorella as a whole displays extreme variability in habit and morphology, a single species, A. spinosa, which has been previously placed within

Mulinum (see Excluded Names), is considered most morphologically similar to Mulinum. The habit (mats), leaf arrangment (cauline), and lobing (3-5, primarily 3) are so similar to Mulinum that A. spinosa is often difficult to identify without the fruit (Martinez 1989). Therefore,

Azorella spinosa is identified as the sister group of Mulinum and is treated as outgroup in the cladistic analysis.

HABITAT AND DISTRIBUTION

Species of Mulinum occur in extreme habitats with open, full-sun, strong winds, and poor soils (sandy, gravely, or rocky). Altitudinally, species of Mulinum occur from sea level to 3500 m. All species are found in very dry areas with dramatic fluctuations in diurnal temperatures. Taxa of 12

Mulinum are subjected to intense heat and radiation during the day and cool (often freezing) temperatures or blanketed in fog at night. Some species of Mulinum (e.g., M. spinosum) are often the dominant component of a particular landscape and appear to play a significant role as pioneers, especially cushion taxa (Rauh 1939, 1940), within barren or disturbed areas.

Mulinum is endemic to Southern South America, primarily concentrated in Chile and Argentina (Fig. 1). A single species, M. ulicinum, extends its range across the northern border of Argentina into Southern Bolivia. In Chile, Mulinum occurs throughout the entire country with the exception of

Region I and the southern portion of Region X, especially along the coast (Fig. 1). In Argentina, Mulinum is distributed in the west, particularly along the cordillera, and along the southeastern coast. The genus is absent from northeastern and central Argentina with the exception of a single corridor which traverses Central Patagonia (Fig. 1).

REPRODUCTIVE MORPHOLOGY

Reproductive structure (i.e., the presence/absence of carpels and stamens) of taxa of Mulinum has been unclear. It has either not been addressed (Cavanilles 1799; Hooker 1830;

Philippi 1860, 1894; Spegazini 1899; Chodat & Wilczek

1902; Constance 1988) or, most taxa, have been described 13

FIG. 1. Distribution of Mulinum Pers. in southern South America. 14 simply as dioecious (Reiche 1899; Speggazini 1899; Wolff

1921), sometimes with hesitation (Skottsberg 1916; Constance

1988). Perfect flowers have been described as occurring in umbels of dioecious plants (e.g., M. microphyHum, M. valentinij Constance 1988). Individual specimens have been annotated as andromonoecious (e.g., by G. Webb, Correa &

Constance 4202/3846).

Examination of both alcohol preserved and herbarium material, showed that perfect flowers exist for Mulinum.

Single flowers with a fully developed stylopodium and stamens with dehisced anthers as well as fruit with stamens attached, have both been observed (e.g., M. albovaginatum, Ricardi S.

2180). In some cases, the majority of flowers are perfect, but some flowers do appear to be something other than complete. Flowers with stamens have been observed on plant material which bore fruit (e.g., M. crassifolium, Cabrera

8790), suggesting the plant to be monoecious or subgynoecious

(Richards 1986). Flowers with mature stamens and poorly developed or degraded stylopodia are also present (e.g., M. albovaginatum, Boelcke 10042), giving the appearance of staminate flowers. Flowers with a well developed stylopodium and no stamens exist (e.g., M. leptacanthum, Boelcke et al.

10970), giving the appearance of carpellate flowers. Flowers and fruit also exist with well developed, but with undehisced anthers (e.g., M. valentini, Soriano 2103).

Most members of the Apiaceae are protandrous (Bell 1971). 15

Protandry coupled with deciduous stamens may suggest the presence of either carpellate or staminate flowers depending

upon the stage of developement observed, and make

identification of the breeding system difficult. Observed

degradation of initially fleshy stylopodia may be the result

of a lack of fertilization.

Protandry coupled With deciduous stamens is considered

likely for Mulinum as evidenced by the large number of flowers

examined which lack stamens. In addition, after detaching

Mulinum stamens leave behind a marginal groove in the fleshy

stylopodium. These grooves may be easily obstructed during drying, giving the impression of a carpellate flower.

Based upon existing evidence, the flowers of Mulinum are herein interpreted as perfect with a combination of protandry, stamens which are deciduous, and variation in style size and development, all of which have been responsible for misleading floral descriptions in the past.

CHROMOSOME NUMBER

Chromosome numbers range consecutively from n = 4 to 13 in the Apiaceae and, based on predominance, x = 8 and x = 11 have been suggested as the base numbers for the family (Moore

1971). Of Drude's (1897) 12 tribes, the two Hydrocotyloideae tribes are delimited by chromosome number, with Hydrocotyleae n = 11 and 12 and Mulineae n = 8 and 10. Within tribe 16

Mulineae haploid chromosome numbers are n = 5, 8, 9, 10, and

16 with n = 8 representing almost half of all counts (Moore

1971).

Chromosome numbers have been counted for six species of

Mulinum (Table 1). The number n = 8 is prominent within the

genus with counts of n = 16 and n = 24 also reported.

Polyploidy appears present in M. spinosum with counts of n =

8, 16, and a questionable 24 recorded (Constance et al. 1971) .

These data clearly reveal an x = 8 base for the genus.

PHYLOGENETIC ANALYSIS OF MULINUM

A phylogenetic analysis of Mulinum, using morphological

data, was undertaken to provide insights into ancestor-

descendent relationships and the recognition of natural

subgeneric groupings. In particular, the origin of the

cushion habit, relationships among species, and character state evolution are addressed.

Materials and Methods. Fifteen morphological characters encoding 29 character states were used in the phylogenetic analysis. Tables 2, 3). Several aspects of the selection and use of characters require explanation. Some taxa (e.g., M. spinosum and M. ulicinum) exhibit a broad range of morphological variation, especially in the leaves. This kind of variability, which is common in Mulinum, causes difficulty in delimiting characters and states. The problem was resolved 17

Table 1. Chromosome numbers for species of Mulinum.

Taxon n Reference

M. hallei 8 Constance et al. 1971.

M. leptacanthum 8 Constance et al. 1976.

M. microphyllum 16 Constance et al. 1971.

M. spinosum 8 (as M. echinus)

Constance et al. 1971.

M. spinosum 8, 16, 24 Constance et al. 1971. 18

Table 2. Characters and character states used in the cladistic analysis of Mulinum. 0 = primitive; 1,2 = derived.

1, Plant habit: shrub or subshrub (0), cushion (1); 2, Leaf lobe number: variable (0), 3 (1); 3, Lobe apex: glabrous (0) , with hair (1); 4, Sheath: open (0), closed (1); 5, Number of flowers per umbel: 15 or more (0), 7-14 (1), 1-6 (2); 6, Umbel pubescence: scarce or absent (0), lanate (1); 7, Umbel position: peduncled (0), sessile (1); 8, Involucral bract number: 6 or more (0), 2-5 (1); 9, Involucral bract composition: Leaf-like (0), membranous (1); 10, Involucral whorls: 2 (0), 1(1); 11, Calyx development: lobed (0), with teeth (1); 12, Fruit: wings (1), wingless (0); 13, Carpophore composition: lobed (0), entire (1); 14, Blade surface: smooth

(0), sulcate (1); 15, Lobe sinus: shallow (0), extending to petiole (1). 19

Table 3. Character state matrix for phylogenetic reconstruction of Mulinum. 0 = primitive; l, 2, = derived. See

Table 2 for description of characters and states. Azorella (A. spinosa; AZO) is used as outgroup (see text) . Order of Mulinum taxa is that used in taxonomic treatment.

Taxa Characters

1 2 3 4 5 6 7 8 9 10 11 12 13 14 15

AZO 0 0 0 0 1 0 0 0 1 1 1 0 1 0 1

SPI 0 0 0 1 0 0 0 0 0 0 0 1 0 0 1

LEP 0 0 0 0 0 0 0 0 1 1 1 1 0 0 0

CRA 0 1 0 0 1 0 1 1 1 1 1 1 1 0 1

ULI 0 1 0 1 1 0 1 1 0 1 1 1 0 0 1

ALB 1 1 0 0 2 0 0 1 1 1 1 1 0 1 1

VAL 1 0 0 0 1 0 0 1 1 1 1 1 1 1 1

HAL 1 0 0 0 1 1 0 0 1 1 1 1 0 0 1

MIC 1 0 1 0 0 1 0 0 1 1 1 1 0 0 1 by eliminating these characters, or, in some cases (e.g.,

habit, and umbel position), by coding the most prevalent

condition. Three features of the involucral bracts

(characters 7-9, Table 2) were included. This places more

weight on aspects of involucral bracts, but it is believed

justified because these variation are believed to be

taxonomically very important. Autapomorphic characters were

included for better definition of groups and to aid in

ranking. The origin of the cushion habit is believed to be an

important adaptive event within Mulinum (see Chapter 2) .

Because of this importance habit (character 2, Table 2) was

weighted twice the other characters. However, in order to

establish a stronger hypothesis for cushion origin, an

analysis with habit weighted equally was also included.

The branch and bound option of PAUP (Phylogenetic

Analysis Using Parsimony) Version 3.0s (Swofford 1991) was

used. Outgroup comparison was used to root trees and polarize

character states, using A. spinosa (see Generic

Relationships). The use of A. spinosa as outgroup is not

definative. Evolutionary relationships have not been

previously discussed (e.g., Martinez 1989) for Azorella,

making identification of A. spinosa as primitive difficult.

Mulinum coming out of Azorella or Mulinum giving rise to

Azorella are also two prospects. Because these problems and

alternatives exist, a hypothetical ancester, encoded entirely with primitive states, was also included. Consensus 21

cladograms were formed using strict, semistrict, Adams, and

50% majority rule options (Adams 1972, 1986; Margush &

McMorris 1981; Sokal & Rohlf 1981; Bremer 1990; Wiley et al.

1991).

Results and Discussion. The results of the phylogenetic

analysis (Fig. 2) indicate that two main lines exist within

, Mulinum, recognized taxonomically as sections. Two further

lines exist, which are recognized as subsections. Two final

lines exist and are recognized taxonomically as series.

Use of A. spinosa as an outgroup produced 5 most

parsimonious trees of 25 steps, while the hypothetical

outgroup produced a single tree of 21 steps. consistency

indices of 0.680 and 0.708, and homoplasy indices of 0.320 and

0.292 were found using A. spinosa and the hypothetical

outgroup, respectively. Figure 2 is produced in both analyses

using different outgroups and is believed to best represent

the phylogeny of the species of Mulinum. For the

consensus tree analysis (Fig. 3), branches A, B, and C (Fig.

2) were supported 100% of the time in both the semistrict and

majority-rule consensus, and branches D and F supported 60%

and 80% respectively in the majority-rule consensus. Varied

degrees of collapse occurred within the cushion clade for all

analyses (Fig. 3).

A single cushion habit origin for M. albovaginatum, M.

hallei, M. microphyllum, and M. valentini is supported by all

analyses. However, the degree is weak, with only a single AZO

42- CRA 4-4 44- ULI B 4 4 4-4 ALB 1.4 E 4-B- - 4 VAL JjLA. HAL 4 4 MIC 5 13 44- -I- I LEP

4 4 101:1t—t- SPI

FIG. 2. Cladogram of taxa of Mulinum. The derived character states are indicated by numbers (cf. Tables 2 and 3) and single bars. Double bars and asterisks indicate parallelisms and reversals, respectively. The names of taxa are represented by the first three letters of the specific epithets.

w PO Strict Semistrict HYP HYP CRA 100 CRA ULI ULI ALB too ALB VAL VAL HAL too HAL MIC MIC LEP LEP SPI SPI

Majority rule A dam s HYP HYP CRA CRA

100 ULI ULI ALB ALB 60 VAL VAL 100 HAL HAL MIC MIC LEP LEP SPI SPI

FIG. 3. Consensus Cladograms of taxa of Mulinum. A, Strict. B, Semistrict. C, 50% Majority Rule. D, Adams. The names of taxa are represented by the first three letters of the specific epithets. 24

character supporting the cushion clade. Development of the

cushion habit is considered an important process in Mulinum's

evolution and reflects a strong adaptation to extreme

environments (see Chapter 2).

A few species as well as characters deserve further

discussion. The position of M. leptacanthum makes it the

sister species to M. spinosum and the clade which contains the

cushion taxa (Fig. 2). Mulinum leptacanthum is similar to M.

spinosum in fruit size, and leaf morphology. Mulinum spinosum

is variable in habit with shrubs, subshrubs, and mats all

represented. Individuals of M. leptacanthum are intermediate

between mats and cushions, corresponding to the intermediate

placement of this taxon between M. spinosum and the cushion

clade.

The clade exhibiting the most collapse and containing

branches supported the least during consensus (Fig. 3) and nonconsensus tree analyses includes the 4 cushion taxa (Fig.

2). The collapse and decreased support of the cushion clade

is believed a reflection of the characters used and not necessarily the clade and/or taxa.

BIOGEOGRAPHY

Present biotic distributions in South America can be deduced from patterns of speciation and phylogeny, geologic and paleobotanic data, and Quaternary paleoecological 25

information. Studies of speciation patterns of high Andean

plant and avian taxa (B. S. Vuilleumier 1970; F. Vuilleumier

1968, 1969a, 1969b, 1970a, 1970b; Smith & Vuilleumier 1971)

have led to the conclusion that Pleistocene climatic events

were factors that ultimately shaped the patterns now observed

in the paramo-puna and the related Patagonian flora and fauna.

The uplift of the Andes at the end of the Tertiary

automatically limits the age of the high Andean habitats and

their biotas to the Quaternary. Within this period, the

number of ecological fluctuations caused by the glaciations

could easily have provided the mechanism to produce the

patterns now present in these habitats (B. S. Vuilleumier

1971) .

Current distribution of Mulinum (Fig. 1) shows that

species are absent from southern-central Patagonia and the

lower portion of the lake region of southern Chile. During

the Pleistocene all of the land west of the Andes and south of

44° to 45° S was covered by ice, with the major part of

Patagonia appearing to have been a bogland dotted with glacial

lakes within and along the Atlantic coast (Auer 1958, 1960).

Glaciers on the eastern Andean slopes were restricted to the

base of the Andes which underwent a series of limited advances

and retreats (B. S. Vuilleumier 1971). The central Andes,

including northwestern Argentina and adjacent Chile, during the Pleistocene consisted of several parallel mountain chains.

These chains did not consist of continuous ice as was seen in 26

the South but were periodically interrupted by glacial lakes

and patches of ice.

Several studies support a general drying period or the

existence of arid to semi-arid climatic conditions during the

Pleistocene (Sanguinetti de Bormida & Borreo 1983; Tonni &

Fidalgo 1983). These arid and semi-arid conditions were

probably accompanied by a general lowering in mean annual

temperatures, which at lowermost levels favored the expansion

of Patagonian faunas and at highermost levels induced the

distribution of Central and Pampean faunas, with the latter

being quite similar to the present one (Tonni & Fidalgo 1983).

The present distribution of Mulinum is believed to have

been shaped during the Pleistocene. The open habitats (Tonni

& Fidalgo 1983; Villagr&n 1988a, 1988b), arid and semi-arid

conditions (Montero & Silveira 1983; Pearson & Pearson 1980;

Sanguinetti de Bdrmida & Borreo 1983; Tonni & Fidalgo 1983),

and a lowering in mean annual temperatures (Tonni & Fidalgo

1983) described for the Pleistocene and early Holocene

correlate well with Mulinum's current growing conditions and

distribution.

Mulinum may have originated within the puna grassland of

northern Argentina. Migration to the north along the Andes would have been restricted by a large glacial formation

(during Wlirm glaciation) therefore directing dispersal south,

resulting in Mulinum's southern South American concentration

seen today. Opportunities for southern and higher altitude migrations would have occurred during the numerous glacial

advances and retreats (Auer 1960; Mercer 1972; Mercer &

Laugenie 1973; Clapperton & Sugden 1988), some of which are

still occurring today (Clapperton & Sugden 1988). Once

Mulinum occupied higher elevations, species would have had the

opportunity to migrate from mountain to mountain along a

north-south axis, which has been the suggested scenario for

other organisms that exist in high altitudes (B. S.

Vuilleumier 1970; F. Vuilleumier 1968, 1969a, 1969b, 1970a).

Throughout this period, the opportunity for speciation events

to occur would have been possible, especially if correlated with repeated cycles of glacial advances and retreats followed

by geographic isolation. As the glaciers retreated on the

Chilean side, approximately 36,000 years ago (Mercer 1972;

Mercer & Laugenie 1973), new habitats to the west would have been available to Mulinum. As the southern glaciers retreated and Patagonia became dryer, Mulinum would have been able to continue it's southern migration. In the Rio Gallegos valley

(Province Santa Cruz) Sanguinetti de Bdrmida and Borreo (1983) suggest that a general trend towards a drier landscape between the Late Pleistocene and Early Holocene has occurred. This trend would therefore limit the southern migration of Mulinum species until during or after this time. During the periods of arid glacial retreats Mulinum probably adapted to drier habitats. After the retreat of the northern glaciers,

Mulinum's distribution continued to be restricted to the north 28

by increasing temperatures and humidity.

TAXONOMIC CATEGORIES

The following definitions were used for the categories

included within this revision. A species is defined as a

group of individuals that share a set of morphological

features different from those of other such groups (Cronquist

1978; Stuessy 1990). While this study dictates a

morphological species definition, it is assumed that the

species recognized here reflect biological species and

reproductive boundaries (Mayr 1969, 1992).

Because species of Mulinum are variable vegetatively and

because this variation sometimes has a geographic component,

it is beneficial to recognize formally these patterns of

variation as varieties (Stuessy 1990). Variety was chosen

over subspecies because of its historical use (Hieronymus

1876; Kuntze 1898; Hicken 1923).

MORPHOLOGY AND TAXONOMIC CRITERIA

The present classification of Mulinum is based primarily upon comparative morphology. Definitions of qualitative characters and states follow Radford (1986) and stearn (1986) .

Measurements of quantitative characters were made from pressed and dried material. In the case of reproductive structures 29 and small leaves, plant material was rehydrated before measuring.

Intergradation of leaf morphology among some Mulinum taxa

(e.g., the varieties of M. spinosum and M. ulicinum) make specific delimitations and identification somewhat difficult.

Taxa can often be recognized only by a combination of character states.

This study is in agreement with previous works on Mulinum which have emphasized fruit characters (Reiche 1899). General habit was useful in delimiting sections, subsections, and series, while both qualitative and quantatative characteristics of the fruit, peduncle, involucral bracts, pubescence, leaves, and distribution were useful in delimiting sections, subsections, species and varieties.

In general, floral characters (e.g., petals and stamens) in Mulinum are conservative while vegetative characters (e.g., leaf size) are variable. The variablity of vegetative characters, often within the same species, is believed to correlate with generally unstable environmental conditions in which the taxa occur. The conservation of floral characters is consistent with a relatively high degree of floral uniformity found throughout the Umbelliferae (Bell 1971). The uniformity has been suggested as representing an ancient adaptive peak, or specialization, for pollination by a large number of unspecialized pollinators (Bell 1971).

Habit. The taxa vary from large shrubs, to subshrubs, to 30 small compact cushions, with habit being useful for recognizing sections, subsections, and series. Section

Mulinum contains shrubs, subshrubs, and mats, while section

Leptacantha has subshrubs, mats, and cushions. Subsection

Leptacantha is defined by mats, while subsection Crassifolia has subshrubs and cushions. Series Azorellopsis is the most strongly defined by habit, with only cushions, and series

Crassifolia contains subshrubs, with mats occurring rarely.

Although variable in size, all species share an extensive branching pattern from a central tap root resulting in dome­ like shrubs, low mats, and compact cushions.

Leaves. While leaf morphology is extremely variable, most leaves are thrice divided with strongly sheathing leaf bases.

In some species (e.g., M. crassifolium) leaf size and blade development is constant, while in others (e.g., M. spinosum and M. ulicinum) it is not, ranging from large leaves with spine-like lobes to small leaves with broader blades.

Although variation exists, in general leaf morphology for

Mulinum is unique compared to other genera, and specific for most species, and especially varieties.

Stems & Roots. Because of the strongly sheathing leaf bases, the stems are often hidden or appear to be lacking.

Well developed central taproots with or without branch roots are common to all taxa, with neither stems or roots providing diagnostic characters.

Inflorescence. A simple umbel is common to all taxa with 31 few exceptions, making the characteristic unique in comparison to some, but not all, genera of the tribe. While compound umbels have been depicted for Mulinum (Cavanilles 1799: t.

486, f . 1), these inflorescences are better described as two simple umbels on a single axis and are considered an anomaly.

Although the umbel is characteristic for Mulinum, some individual inflorescences have been reduced to a single flower

(e.g., M. albovaginatum).

Peduncle length varies between species, with some umbels appearing sessile, making the characteristic significant for species recognition (e.g., M. crassifolium vs. M. ulicinum).

Involucre. Involucral bracts differ in number (2-12), size (0.5-9 mm long) , and qualitative composition (e.g., lobed vs. entire; green vs. membranous) among taxa, representing one of a few floral characters which displays diversity and is useful in species delimitation.

Flowers. The flowers are typical of the Apiaceae with a

5-merous perianth and androecium. Floral size and general morphology is extremely uniform for Mulinum, with floral parts in general offering few diagnostic characters.

Calyx. The sepals are fused into a shallow or narrowly shallow cup. The free portions are considerably small, often reduced to teeth or appearing absent, except in M. spinosum where the calyx in general is larger and more developed.

Corolla. The petals are of uniform size and shape, even within the same umbel. Most petals are yellow with the 32

exception of some which are tinged with red (e.g., M.

spinosum). Mulinum taxa lack a median dorsal petal gland

which was suggested as characteristic by Mathias and Constance

(1962) for several additional genera of Mulineae (Asteriscium,

Domeykoa, Eremocharis, Gymnophyton, and Pozoa).

Androecium. The stamens are uniform in size and number,

with only the filaments of M. hallei, which are wider at the

base, displaying variation. The anthers are dorsifixed and

versatile and open longitudinally.

Pollen. The pollen grains are uniform in size and shape

among Mulinum species. The grains were first described by

Heusser (1971) and latter by Markgraf and D'Antoni (1978) as

isopolar, radiosymmetric monads being tricolporate with long,

straight, narrow colpi. The exine is tectate, appearing

finely perforate, with distinct columellae (Heusser 1971) .

Gynoecium. The stylopodium is ± fleshy and convex to

shortly conical. The styles are short, elongating to ± 1 mm

after fertilization. While these characters are constant for

Mulinum species, and in general within the tribe, they do aid

in distinguishing Mulinum from other genera of Mulineae (e.g.,

Eremocharis with styles longer than the stylopodium).

Fruit. Of all the organs, the fruit contains characters most diagnostic for the genus (Drude 1897; Reiche 1899; Hiroe

1979; Constance 1988). The fruit of Mulinum is winged, either

appearing 2-winged or 4-winged in cross-section, and dorsally compressed. The inner fruit wall is somewhat woody. 33

Previously referred to as a woody endocarp (Drude 1897;

Constance 1988), the stucture is better described as the inner

fruit wall because of the inferior ovary and epigynous flower.

Carpophore. The carpophore is free with the degree of

bifurcation often being diagnostic at the species level (e.g.,

M. leptacanthum).

Cotyledons. Observation of a single seedling of M.

spinosum (Stuessy et al. 11069) in the Ohio State University

greenhouse, showed the cotyledons to be linear, and the first

leaf 3-lobed, with the lobes spine-like. This differs from

several individuals of Azorella triacanthum (Stuessy et al.

11068) and one individual of Gymnophyton spinosisimum (Stuessy

& Marticorena 9825) whose cotyledons are spatulate and first

leaves with the lobes denticulate and sinuses shallow.

TAXONOMY

Mulinum Pers. Syn. pi. Is 309. 1805.— LECTOTYPE, designated by

Hiroe, 1979: Mulinum spinosum (Cav.) Pers.

Azorellopsis H. Wolff, Repert. Nov. Sp. 19: 312. 1924.—

TYPE: Azorellopsis trisecta H. Wolff [= Mulinum

albovaginatum var. pauciflorum (Reiche) Zech].

Woody perennials as shrubs, subshrubs, mats, and cushions, 0.01- 0.8 m tall, 0.01-3 m diameter; hermaphroditic.

Stems prostrate or erect, extended or reduced, 1-7 mm diameter, concealed by sheathing leaf bases and decaying

leaves, leaf bases and/or leaf skeletons, brown to black.

Leaves thick to thin, fleshy to dry, stiff to flexible, easily

separating when dry or remaining attached, 2-44 mm long, 1-43 mm wide, sheathing, sessile or petiolate, strongly to loosely

imbricated; phylotaxy 5/5, alternate spiral; blade triangular

to rhomboidal, smooth to sulcate to ± inflated, glabrous or pubescent, 1-25 mm long, 1-43 mm wide, 3-veined to venation extensive, at apex ± 900 to obtuse# with margin entire; lobes 1-

8, with the division again parted or lobed, 0.5-25 mm long,

0.1-3.25 mm wide, wide to lanceolate to spine-like, reflexed or spreading, equal or with center lobe larger, sinusus shallow or deep, at apex finely acute to acute to mucronate; with constriction between blade and petiole or blade continuous; leaf bases strongly sheathing; petiole to 18 mm long, 0.5-5 mm wide, at base dilated or uniform; sheath green to membranous, open or closed, 3-7 nerved, 1.8-7 mm long, 1.5-

6 mm wide or 3 mm diameter, with margin entire or pubescent, hairs while, simple, straight to curled, to 8 mm long; stipules 3-4 mm long, 1.75 mm wide, or as hairs, 1-5 mm long, or absent. Umbel simple, 2-20 mm diameter, 1-37 flowered, ± globose to spreading, axillary among the upper leaves, extending beyond them or contained within them, sessile or peduncled; peduncle yellow to brown, smooth to ribbed to angled, to 7.5 cm long, 0.5-2 mm diameter, glabrous or lanate, hairs white, simple, at apex uniform to enlarged and constricted. Involucre glabrous or lanate, hairs white,

simple; involucral bracts 2-12, equal or not, in a single whorl, rarely 2, 0.5-9 mm long, 0.5-1 mm wide, green to membranous, with a single green to red-brown rib, linear-

lanceolate to lanceolate, at apex narrowly linear to rounded,

or lobed, with single, simple white apical hair, to 0.5 mm

long or entire, at base ± free or fused into an elongated cup, to 3 mm long, with margin entire or pubescent, hairs white, simple, straight to curly, to 1.25 mm long. Rays 1-16 mm

long, 0.25-1 mm diameter, smooth to ribbed; receptacle enlarged and constricted or uniform. Flowers complete, yellow, actinomorphic, 5-merous, 2.5-5 mm diameter; sepals 5, green to yellow-green, with median rib brown or same as blade, distinct or reduced to teeth, 0.25-2 mm long, 0.5-1 mm wide, at apex acute to mucronate to shortly acuminate to ± 90°, at base connate, forming a shallow stellate disc or narrow conical cup; petals 5, yellow or yellow-red, separate, with median rib darker, without median dorsal gland, 1.25-2 mm long, 0.5-2 mm wide, oval to elliptic, curled adaxially or straight or curled abaxially, reflexed abaxially or planar, at apex mucronate to acute to rounded, at base truncate; stamens

5, yellow, deciduous or persistent, with filaments 0.5-1.5 mm long, at base umiformly wide or flared, with anthers oval,

0.3-0.75 mm long, 0.2-0.5 mm wide, dorsifixed and versatile, dehiscing longitudinally; carpel entire or grooved; stylopodium yellow, fleshy, convex to shortly conical, 1-2 mm 36 diameter, with marginal filament grooves, evident especially after stamens have detached, with styles small, +0.1-0.5 mm long, elongating to 1.25 mm and reflexing after fertilization.

Fruit oblong to circular, yellow, yellow-purple, or purple, 2 or 4-winged in cross-section, dorsally compressed, surface granular without ornamentation, body inflated or uniform, squarish to rectangular in cross-section, 3-7 mm long, 2-8 mm wide, at apex acute to rounded, at base rounded to cordate; wings reniform to lunate, 4-7 mm long, 1-3 mm wide, lateral wings as long or longer and narrower or broader than or equal to the fruit body, straight or revolute; mericarps linear to rectangular or lunate to triangular in cross-section, separating easily or with difficulty at maturity, abaxial face angular to slightly concave to planar, commissural face slightly concave to planar to convex, to angular, ribs 5-7, 1-

3 abaxial, 2 marginal, and 2 adaxial, prominent or obscure, equal, filiform or variable; oil tubes 5, associated with ribs, round to deltoid, or irregular 0.25-0.75 mm diameter; commissure planar, scar linear; carpophore free, entire, shortly 2-cleft, or equally bilobed; stylopodium persistent; odor or absent. Chromosome numbers: n = 8, 16, 24.

Key to Species of Mulinum

1. Shrubs, subshrubs, or mats.

2. Involucre 1-2-whorled; involucral bracts green with single red-brown rib; sepals distinct, to 2-ram long.

1. M. spinosum.

2. Involucre 1-whorled; involucral bracts membranous;

sepals as teeth, 0.5-1 mm long.

3. Mats, often creeping; leaf lobe number 3-5; umbels

peduncled; involucral bracts 6 or more.

2.M. leptacanthum.

3. Subshrubs; leaf lobes 3; umbels sessile; involucral

bracts 2-5.

4. Leaves thick; leaf sheath open; umbels 2-11-

flowered; fruit 3-4 mm long, 3-4 mm diameter;

carpophore entire. 3. M. crassifolium.

4. Leaves mostly with spine-like lobes; leaf sheath

closed; umbels 3-14-flowered; fruit 4-5 mm long,

1-2 mm diameter; carpophore bilobed or shortly 2-

cleft. 4. M. ulicinum.

Cushions.

5. Involucre lanate; involucral bracts 6 or more; sepal

with median rib brown.

6. Leaf lobe apex entire, mucronate, or rarely

with hairs; umbels 7-14-flowered. 7. M. hallei.

6. Leaf lobe apex with simple white hair; umbels 15+

flowered. 8. M. microphyllum.

5. Involucre glabrous; involucral bracts 2-5; sepal with

median rib yellow.

7. Cushions 2-10 cm tall, 4-15 cm diameter; leaves with 38

3 lobes; umbels 1-6-flowered; petals reflexed

abaxially, 1.25-2 mm long; fruit 5-7 mm long, 4-5 mm

diameter; mericarps appearing lunate in cross-

section; carpophore shortly 2-cleft, rarely entire.

5. M. albovaginatum.

7. Cushions 1-2 cm tall, 3-33 cm diameter; leaves with

3-5 lobes; umbels 2-12-flowered; petals planar, 1.25

mm long; fruit 5 mm long, 4-6 mm diameter; mericarps

appearing triangular in cross-section; carpophore

entire. 6. M. valentini.

For type citations, collection location, date, collector, and number, were taken from the protologue. Additional information from type specimen labels and/or additional publications is given in brackets.

Specimens cited for each taxon are arranged alphabetically first by country and then by division, and finally by collector. Both Regiones (in Chile only) and individual collector numbers are arranged in ascending order.

Argentinean and Bolivian specimens are grouped first by

Province then by Department. Chilean specimens are grouped first by Regidn and then by Province. Because of their north to south arrangement, Regiones are included as an aid to

Province location. Number and names of Regions as well as the

Provinces included within follow Romero et al. (1983): I

(Region de Tarapacd): Arica, Iguique, Parinacota; II (Region de Antofogasta): Antofagasta, Loa, Tocopilla; III (Region de

Atacama): Chanaral, Copiapo, Huasco; IV (Region de Coguimbo):

Elqui, Limari, Choapa; V (Region de Valparaiso): Isla de

Pascua, Los Andes, Petorca, Quillota, San Antonio, San Felipe de Aconcagua, Valparaiso, (Regidn Metropolitana de Santiago)

Chacabuco, Cordillera, Maipo, Melipilla, Santiago, Talagante;

VI (Regidn del Libertador General Bernardo 0 'Higgins):

Cachapoal, Cardenal Caro, Colchaqua; VII (Region del Maule):

Cauquenes, Curicd, Linares, Talca; VIII (Regidn del Bio-Bio):

Arauco, Bio-Bio, Concepcion, Nuble; IX (Region de La

Araucania): Cautln, Malleco; X (Regidn de Los Lagos): Chiloe,

Llanquihue, Osorno, Palena, Valdivia; XI (Regidn Aisen del

General Carlos Ibdnez del Campo): Aisen, CapitSn Prat,

Coihaique, General Carrera; XII (Regidn de Magallanes y de la

Antdrtica Chilena): AntSrtica Chilena, Magallanes, Tierra del

Fuego, Ultima Esperanza.

Herbarium acronyms follow Index Herbariorum (Holmgren et al. 1991). Author citations are according to Draft Index of

Author Abbreviations (Halliday et al. 1980). Book abbreviations comply with Taxonomic Literature II (Stafleu &

Cowan 1976-1988). Journal abbreviations follow BPH (Lawrence et al. 1968). Chromosome numbers (Table 1) are from Constance et al. (1971, 1976).

I. Mulinum section Mulinum 40

Shrubs, subshrubs, or mats; involucre l-2-whorled;

involucral bracts green with single red-brown rib; sepals

distinct, to 2 mm long.

This section is defined by the same characters as its

single member, M. spinosum. The shrubby habit, green

involucral bracts arranged in 1-2 whorls, and elongated,

separate sepals makes this section, as well as M. spinosum,

unique as compared to other taxa.

1. Mulinum spinosum (Cav.) Pers., Syn. pi. 1. 309. 1805.

Selinum spinosum Cav. [nec Selinum spinosum (L.) E. H.

L. Krause, Deutschl. FI., ed. 2, 12: 81. 1904], Icon.

PI. 5: 59, t. 487, f. 1. 1799. Bolax spinosus (Cav.)

Spreng., Syst. Veg. 6: 362. 1820. Mulinum spinosum

typicum Reiche, Anal. Univ. Chile, 104: 801.

1899.— TYPE: Chile. Regidn V. "Cordillera del Planchon

et del Portillo versus Mendozae urbem", Jan-Feb 1789-

94, L. Wee 1801 (holotype: MA, photo: OS!; isotype: F!

MA!) .

Shrubs, subshrubs, or mats, 0.2-0.8 m tall, 0.3-3 m diameter; loosely to densly branched. Stems, erect to lax, 2-

4 mm diameter. Leaves variable, 1-5 cm long, 0.6-4.3 cm wide; blade rhomboidal, glabrous, at apex acute to obtuse, with margin entire; 2-6 lobes, generally 3 or 5, spine-like or with broad blades, 0.2-2.5 cm long, 0.5-3.25 mm wide, at apex acute to shortly acuminate; constriction absent, blade continous with petiole; petiole 1-1.8 cm long, 1.8-5 mm wide, with recessed, yellow-gold veins, base dilated; sheath green to white, closed, 5 mm long, 3 mm diameter, with margin entire or pubescent, hairs few to many, straight, or curled, 8 mm long or less; stipules absent. Umbel 0.7-2 cm diameter, 5-37 flowered, ± globose, maturation indeterminant, extending beyond upper leaves; peduncle yellow to brown, ribbed, 1.3-6 cm, 2 mm diameter. Involucre glabrous; involucral bracts 5-

12, equal, green with red-brown rib, in 1 or 2 whorls, with the outer whorl containing the majority of bracts, 2-9 mm long, 1 mm wide, linear-lanceolate to lanceolate, at apex acute, at base connate into a narrow cup, with margin entire or pubescent, hairs few. Rays 1.25-16 mm long, 0.33 mm diameter, green to brown; receptacle enlarged and constricted.

Flowers 5 mm diameter; sepals green, distinct, alternating with petals and opposite the stamens, to 2 mm long, 0.75 mm wide, at apex acute, at base a shallow cup; petals yellow, 2 mm long, 1.25 mm wide, curled adaxially, planar, elliptic to ovate, at apex rounded; stamens with filament 1.5 mm long, 0.1 mm wide, at base uniformly wide, with anthers 0.5 mm long, 0.3 mm wide; carpel entire; stylopodium 1-2 mm diameter, with styles ± 0.1 mm long, elongating to 1.25 mm; fruit development determinant. Fruit widely ovate, yellow, yellow/purple, or purple (etruscan red), slightly 4-winged in cross-section, linear to rectangular in cross-section, 6-7 mm long, 7-8 mm 42 wide, at apex acute, at base cordate; wings widely lunate to reniform, 6-7 mm long, 2-3 mm wide, lateral wings longer and broader than the fruit body, straight; mericarps linear to lunate in cross-section, separating easily, abaxial face planar to concave, commissural face planar to convex, ribs 5,

1 abaxial, prominent or obscure, equal, filiform; oil tubes round to irregular; carpophore bilobed to shortly 2-cleft; odor absent or coconut-like. Chromosome numbers: n = 8, 16,

24. Fig. 4.

Phenology. Flowering mid-November through early March; fruit maturing mid-December through April.

Distribution (Fig. 5) : In Chile with the exception of the the southern portion of Region X. In Argentina from Salta and

Buenos Aires Provinces south. Absent in northeast and central

Argentina with the exception of a corridor traversing Chubut

Province; 10-3500 m.

Common Names: "Coirdn" (Cavanilles 1799); "Yerba Negra",

"Chinchimari" "Chinchimal" (Kurtz 1894); "Matabarrosa" (Montes

& Wilkomirsky 1985); "Herba Negra", "Dichillo", "Hierba de la

Culebra" (Baeza R. 1921; Montes & Wilkomirsky 1985); "El

Spina", "Neneo", "El Pescado" (Carminatti s.n., LP) .

Hooker (1830) treated Fragosa spinosa Ruiz Lopez & Pavon, based on a different type, as a synonym of M. spinosum. He conveyed doubts about this inclusion (by "?") and pointed out the different structure of the leaf and the larger fruit 43

a ?

A.

B.

FIG. 4. Cross section of fruit. A. Mulinum spinsoum (based on Boelcke et al. 10309, UC). B. Mulinum leptacanthum (based on Correa et al. 5855, UC). 44

FIG. 5 Distribution of Mulinum spinosum in Argentina and Chile. • = M. spinosum var. spinosum; ▲ = M. spinosum var. minus; ■ = M. spinosum var. atacamense; ★ = M. spinosum var. proliferum. 45

as compared to M. spinosum. Hooker was correct in questioning

the placement of F. spinosa, which has since been

better combined as Azorella spinosa (Reiche 1899; this

treatment).

"Peru" is listed on the type label of M. spinosum. In

addition, "Cordillera del Planchon y Portillo" are included,

which clearly places the type locality in Chile. Nee

completed extensive trips, including collecting in Peru, and

this probably represents an error in labeling.

Krause (1904) incorrectly combined Echinophora spinosa L.

as Selinum spinosum (L.) Krause creating a later homonym.

Selinum spinosum (L.) Krause is in no way similar to S.

spinosum Cav. which was the basis for Persoon's Mulinum

spinosum. While the leaves of E. spinosa resemble those of

Mulinum, the flowers and fruit are dramatically different

(Candolle 1829, t. 16; Drude 1897, f. 57), as supported by

Drude (1897) who placed Echinophora within a completely different subfamily, the Apioideae.

Polyploidy is present in M. spinosum with n = 8, 16, and

24 (Constance et al. 1971). Little evidence exists to designate the polyploids as auto- or allopolyploids. Limited morphological variation exists among the tetraploids (n = 16), hexaploid (n = 24), and diploid (n = 8). In general the leaf lobes of the hexaploid (Correa & Constance 4181/3844) and tetraploids (Correa & Constance 4065/3835, 4096/3840,

4136/3841) are wider (2 mm wide) compared to one diploid (0.5 46

mm wide; Correa & Constance 3936/3824). The other diploid

(Correa & Constance 4202/3846) has leaves which are as wide as

the polyploids. In addition, a slight increase in leaf

pubescence is present in the polyploids compared to both

diploids. The chromosome figures (Constance et al. 1971,

their Figs. 324a, 324b) provide little insight; chromosomes of

the tetraploid are uniform in size and smaller in comparison

to the diploid. The uniformity of the tetraploid's

chromosomes and general morphological similarity between the

diploid and polyploids, may suggest autopolyploidy.

Previously, no observations have been made concerning

pollinators of Mulinum. The bee fly, Anthrax sp.

(Bombyliidae), was observed, generally in swarms, visiting

large, low, mats of M. spinosum var. spinosum. Bee flies are

more common in arid areas, corresponding to the habitat of

several Mulinum species, and are found on flowers, feeding on

pollen or nectar, or hovering over or resting on the ground in

open sunny places (Curran 1934; Bland 1978; Borror et al.

1981; Pena 1987).

The stems and leaves of M. spinosum have been screened

for anticancer activity, and have shown antitumor activity

against P-388 lymphocytic leukemia (Bhakuni et al. 1976).

Livestock which eat the species acquire a bad taste (Mario

Gentili, pers. comm.) . The species has also been used to numb teeth nerves (Montes & Wilkomirsky 1985).

Mulinum spinosum is vegetatively variable, especially the 47 composition of the leaf and in particular the leaf lobes. The majority of leaves display three lobes which is characteristic for Mulinum in general. However, anywhere from 2 to 5 lobes, with the division again lobed, have been observed. Two lobes are rare and often coincide with what appear to be juvenile leaves. Four and five lobes are somewhat more common, being the distinguishing characteristic of M. spinosum var. spinosum. The degree of leaf blade is also variable ranging from virtually bladeless, or the lobes being extremely spine­ like, to containing a distinct blade below, independent of the lobes, and lobes with wide blades.

Mulinum spinosum is an extremely variable species, especially vegetatively, here interpreted to comprise four varieties, most of which have a geographic basis.

Key to the Varieties of Mulinum spinosum

1. Leaf blades pubescent. lc. M. spinosum var. atacamense.

1. Leaf blades glabrous.

2. Leaf lobes 2-5 mm long, 0.5-0.75 mm wide; petiole

0.75-1 mm wide. lb. M. spinosum var. minus.

2. Leaf lobes 0.07-2.5 cm long, 0.5-3.25 mm wide;

petiole 1.8-5 mm diameter.

3. Leaf lobes 3, spine-like, 0.5 mm wide.

Id. M. spinsoum var. proliferum.

3. Lobes 3-5, blade-like, 3.25 wide. 48

la. M. spinosum var. spinosum.

la. Mulinum spinosum var. spinosum

Mulinum laxum Philippi, Linnaea 33: 90. 1864-5.— TYPE:

Chile. "In Andibus prope Antuco", Jan 1839, C. Gay

1513 (holotype: SGO!, photo: F!).

Mulinum chillanese Philippi, Anal. Univ. Chile, 85: 707.

1894.— TYPE: Chile. "In jugo Andium inter thermas

Chillanenses et Vallem Nebularum" [=Nieblas, fide

Munoz P., 1960], Jan 1877, R. A. Philippi s.n.

(lectotype, designated by Constance, 1954: SGO!,

photo: F!; isolectotype: SGO!).

Mulinum hirsutum Philippi, Anal. Univ. Chile, 85: 708.

1894.— TYPE: Chile. "In Andibus prov. Colchaqua",

[Cord, de San Fernando], Jan 1884, A. Hirth s.n.

(lectotype, here designated: SGO!; isolectotypes: B

BM! K!, photo: F! G! GH! NY!).

Mulinum spinosum var. leoninum Kuntze, Revis. gen. pi.

3(2): 114. 1898. Mulinum spinosum var. leoninum f.

Toscae Kuntze, Revis. gen. pi. 3(2): 114. 1898.

Mulinum leoninum Lorentz, Exped. Rio Negro, Bot. 222,

t. VI, pi. 1, f. 1-5. 1881.— TYPE: Argentina. "En una

colina de tosca, cerca de Leones, entre el Fuerte

Argentino y Nueva Roma" [Rio Sauce chico], 4 May 1879,

P. G. Lorentz s.n. (holotype: B, destroyed; isotype:

photo: OS!). 49

Mulinum spinosum var. quinquepartitum Kuntze, Revis.

gen. pi. 3(2): 114. 1898.— TYPE: Chile. "Paso Cruz [de

la Piedra], 1500 m, [34°, Cordillera], Jan 1892 O.

Kuntze s.n. (holotype: NYl).

Mulinum spinosum var. longeinvolucrata Hicken, Darwiniana

1: 61. 1923.— TYPE: Argentina. "Arroyo Blanco (rio

Atuel)", Dec 1912, C. M. Hicken 58 (holotype: SI!).

Leaves with 2-6 lobes, generally 3 or 5; lobes wide to

spine-like. Peduncles long.

Phenology. Flowering December through early March; fruit

maturing early December through mid-April.

Distribution (Fig. 5). Primarily along the Andes of Chile

from Regidnes V to IX, with concentrations near Santiago,

Curicd, and Termas de Chilian. In Argentina in the Provinces

of Buenos Aires and Salta south to Santa Cruz, primarily along

the western cordillera; 450-3200 m.

Representative Specimens. Argentina. Buenos Aires.

Tornquist: Cerro Tres Picos, faldeos inferiores, Boelcke et a l . 13042 (MO), Sierra de la Ventana, cerro de la Ventana,

Cabrera 5767 (LP) , Choique, Cabrera 6720 (NY) , Sierra de La

Ventana, Cerro Tres Picos, laderas, Fabris & P&rez M. 6741

(LP), Cerro VacacuS, Rossi & Bachmann 482 (G, UPS), Estancia

Funke, Cerro Fundacion Funke, Ventania 248 (LP, NY). Chubut.

Rio Senquer: Rio Union, Lago la Plata, Soetbeer & Glesias 66 (LP) . Mendoza. Las Her as: Quebrada Potrero, Puerta, Semper 252

(BM, NY); Lujdn de Cuyo: Puente del Inca, Wall & Sparre 23

(S) ; Malarqiie: 78 km W de El Sosneado, Valle del Rio Atuel,

Boelcke et al. 10236 (UC), Alto Valle del Atuel, Refugio Gral.

Soler, Ruiz L. 15716 (UC), Valle de Las Lenas, Arroyo de Las

Lenas, Ruiz L. 24544 (UC), 8.8 kms W of Los Molles on road to

Las Lenas, Stuessy et al. 10276 (CONC, GH, LP, OS, UC), 7.2

kms W of Las Lenas on road to Valle Hermoso, Stuessy et al.

10296 (CONC, GH, LP, OS, UC), 2-4 kms NW of Las Lenas on ski

road, Stuessy et al. 10318 (CONC, GH, LP, OS, UC), 5 km from

Los Molles on the S side of the road to Las Lenas, Zech &

Contogiorgakis 2 (CONC, GH, LP, OS, UC) , 15 km from Los Molles

on the road to Las Lenas, Zech & Contogiorgakis 3 (CONC, GH,

LP, OS, UC), 8 km before Valle Hermoso, at the monument, Zech

& Contogiorgakis 4 (CONC, GH, LP, OS, UC), Valle de Rio

Tordillo, near the Rio Tordillo, Zech & Contogiorgakis 6

(CONC, GH, LP, OS, UC), Valle de Rio Tordillo, growing along

hillside, E of river facing N, Zech & Contogiorgakis 7 (CONC,

GH, LP, OS, UC), 15 km from the sign in Valle Hermoso on the

road to Las Lenas, Zech & Contogiorgakis 8 (CONC, GH, LP, OS,

UC). Neuquen. LScar: 15 km up form San Martin De Los Andes

toward Cerro Chapelco, Stuessy et al. 10073 (CONC, GH, LP, OS,

UC); Loncopue: Above Copahue to the NE toward snow patches,

Stuessy et al. 10216 (CONC, GH, LP, OS, UC); Los Lagos: Lago

Nahuel Huapi, Sierra Lopes, Ljungner 6 (GOET, UPS). Rio Negro.

Bariloche: Camino a Nirihuau, Cabrera & Job 384 (LP) , Cerro Otto, Corte 165 (LP); Pilcaniyeu: Ruta 23, a 5 km del cruce de ruta 237 (5 km W of Estancia Perito Merino), Correa &

Constance 3815/3822 (UC), Entre Comallo y Zimmermann, Meyer

8029 (NY). Salta. Pampa Grande, C. S. [Spegazzini] s.n. (LP) .

Santa Cruz. Lago Argentino: Lago San Martin, Hogberg 23 (BM) ;

Lago Buenos Aires: Lago Belgrano, Peninsula de las Ciervos, von Platen & Greiner 152 (BM, MO). Chile. Regidn IV. Limari:

Planta Electrica Los Molles, Bocatoma, Ricardi S. et al. 756

(CONC, OS). Regidn V. [Valparaiso] Los Andes: Entre Caracoles de Portillo y Juncal, Arroyo 81337 (CONC), Uspallata-Pass,

Juncal, Buchtien s.n. (BM, E, G, LY, PRC, S, US), Andes above

Rio Blanco 10 km E of Rio Blanco on Trans-Andean Highway,

Constance 3516 (BM, CONC, F, MO, US) , Roadside ditch, 2 km downslope from Paulina, DeVore 1309 (OS), Los Andes, Grandjot

1063 (CONC, MO), Saladillo, Rio Blanco, Silva 19694 (CONC),

Juncal, Quebrada Chepica, Ricardi S. 2967 (CONC, OS, UC) ;

Quillota: Cerro Roble, punta Imdn, Boelcke 6482 (F, MO) , La

Campana de Quillota, Constance et al. 3544 (BM, CONC, F, MO,

US), Cumbre del Cerro Campana, Limache, Garaventa H. 1399,

1447 (CONC), Limache, El Roble, Garaventa H. 2790 (CONC),

Caquis, 15 km E of Melon, Morrison 16915 (G, GH, MO, S, UC);

San Felipe de Aconcugua: Above Ramayama Copper Mine, Cerro Las

Vizcaches, Hutchison 94 (F, G, GH, UC, US) , San Felipe, Joseph

3825 (US); Valparaiso: Cerro de la Campana, Looser 361 (CONC),

Lagunillas, Cajdn del Rio Maipo, Mahu 4064 (CONC), Puente

Alto, Lagunillas, Matte 715 (CONC), Lagunillas, Cajdn del Volcdn San Jose, Pena 14 (CONC), San Jose, Reed s.n. (BM) .

[Regidn Metropolitana de Santiago]. Area de MetropoXitana de

Santiago: El Salto de San Ramon, Ball s.n. (NY, US) , Entre

Farellones y Paso de Jorquera, Garaventa H. 5344 (B, CONC),

Maitenes, Montero O. 306 (CONC), Cerro Manguehue, Schlegel

3774, 3775 (CONC), Rinconada de Lo Cerda, Quebrada de La

Plata, Cerro Buitreras, Schlegel 5905 (CONC, F), El Colorado

Ski area near La Parva, 50 km from Santiago, Skog 1104 (UC),

2 kms W of Farellones on winding road to Santiago, Stuessy et al. 11123 (OS), Perez Caldera, near Rio San Francisco, Zech et al. 29 (CONC, GH, LP, OS, UC), Maitenes near Rio Colorado,

Zollner 11121 (MO); Chacabuco: Cajdn del Maipo, Minenweg Lo

Valdes, zu den Baiios Colina, Grau 3117 (BM) , Colina, Macrae s.n. (G), Portzuelo del Copado, Pennell 12239, 12240 (F, GH

{12239), NY); Cordillera: Banos de Morales, Arriagada F. s.n.

(CONC), Cajdn del Maipo, Barros 21339 (CONC), El Volcdn,

Crisci 408 (LP), Quebrada El Morado, Refugio Aleman, Rio El

Volcdn, Frodin 401 (UPS), Valle del Diablo, Grandjot 3876

(GH), Cajdn del Rio Volc&n, Gunckel L. 46512 (CONC), Canelo,

Valle del Rio Maipo, Looser 855 (UC), Lo Valdes, Marticorena

& Matthei 821 (OS), San Gabriel, Montero O. 530 (MO), Santa

Rosa de los Andes to Uspallato Pass, Moseley s.n. (BM), Lo

Valdes, Cordillera de Santiago, Pfister 176 (CONC), Rio

Colorado, Cordillera de San Rosa, Poeppig 530 (G), Valle del

Colorado, San Ramon, Cubre Grat., Schlegel 2477 (CONC),

Formacion Lo Valdes, Cerro Catedral, Weldt 712 (CONC, OS). Region VI. Cachapoal: Cajon de los Helados, Ricardi S. s.n.

(CONC); Colchagua: Termas del Flaco, NE of Los Banos, Aravena

33334 (GH, MO, UC), El Huemul, Barrientos 1881 (CONC), Vegas

del Flaco, E de la Quebrada de Los Rios, Mahu 9816 (UC) , Banos

El Flaco, Cajon de Las Danas, Milner s.n. (CONC), San

Fernando, Vegas del Flaco, Ricardi S. 3168 (CONC, OS), Junta

del Rio Tinguiririca y Rio del Azufre, Villagriin & Arroyo s.n.

(CONC). Regidn VII. Cauquenes: Cordillera Cauquenes, Ball s.n.

(US); Curicd: Termas del Flaco (Andes de Colchaqua), summits

NE of Los Banos, Aravena 33323 (GH, MO, UC) , Sumit of the

Andes near Volcano of Peteroa, Bridges 1189 (BM, E, OXF) ,

Abfahrt vom Embalse de Planchdn zum Tal des Rio Vergara, Grau

& Grau 1582 (MO), Curicd, Joseph 5271 (US), Camino

Internacional a Paso Vergara, 2 km, Marticorena & Mattei 937

(CONC, OS), Paso Vergara en las laderas de los cerros,

Marticorena & Matthei 1042 (CONC) , Camino de Curicd a la

Laguna de Teno, 1 km antes de la laguna, Marticorena et al. 64

(CONC), Hacienda Monte Grande, Werdermann 554 (B, BM, E, F, G,

K, MO, NY, S, UC, US), Valley of the Rio Teno, Zollner 5200

(UC), Near Lago Planchdn, near Rio Teno, in the high

cordillera, Zollner 5209 (UC), On the margins of Laguna

Planchdn, Zollner 6139 (UC); Talca: Cordillera, Barros 3 (GH),

Vilches Alto, Open slopes above W end of Laguna del Maule,

Constance 3523 (BM, CONC, GH, MO, UC, US), Parque Nacional,

DeVore 1196, Gardner & Knees 4452 (OS). Regidn VIII. Bio-Bio:

Las Cuevas, extreme S of La Laguna Laja, Behn s.n. (CONC), Lolco to Vilucura, Elwes s.n. (K), Volc&n Antuco, Fabris &

Crisci 7595 & 7624 (LP), Antuco, en la cumbre del Mirador,

Junge s.n. (CONC), Cerca de los Banos de Nitrao, Marticorena

& Quezada 9585 (CONC, OS), Valle Vilucura, Neger s.n. (CONC);

Nuble: Termas de Chilian, Cerro Pirigallo, Cabrera 3660 (LP),

Cordillera Chilldn, Valle de las Nieblas, Junge s.n. (CONC),

Banos de Chilian, Pennell 12367, 12450 (F, GH, NV), Termas de

Chilian, Lomas sobre las Fumarolas, Pfister 946 (CONC),

Cordillera de Chilian, R. A. Phillipi s.n. (K), Termas de

Chilian, Valle de las Nieblas, Ricardi S. 5583 (CONC),

Pemehue, Kendel 19608 (CONC). Regidn XX. Malleco: Volc&n

Lonquimay, conglomerated scoria above Nothofagus - Araucaria forest, Constance & Sparre 3579 (BM, CONC, F, GH, MO, UC, US),

South bank of Rio Picoiquen, 5 km E of Vegas Blancas,

Hutchison 305 (B, F), A orillas del Rio Lolco, 3 km antes de la conf luencia con el estero Lancia, Marticorena et al. 1348

(CONC), Lago Gulletud, coironal a orillas del lago, Matthei &

Bustos 145 (CONC), Curacautin, Termas de Rio Blanco, Montero

O. 2738 (GH), Camino a Lonquimay, cerca de la confuencia,

Pfister 7242 (CONC), Lago Gualletui, Weisser S. 1544 (CONC),

Road to Angol from Parque Naheulbuta, between road and Rio

Picoiquen before the bridge, Zech et al. 42 (CONC, GH, LP, OS,

UC) , At the ranger's house below Volcdn Lonquimay, on the E slope of the road, also to the W with Araucaria, Zech et al.

44 (CONC, GH, LP, OS, UC), On the Pass Lolco, Zollner 10249

(MO). Region XI. Coihaique: Camino de Coihaique a Balmaceda, 55

Pfister s.n. (CONC), Coihaique Ba jo, Baquedano, Santesson 1379

(S), Coihaique, cercanias Lago Seco, Schlegel 2360 (CONC), 2 km from El Blanco on the road to Balmaceda, Zech et al. 50

(CONC, GH, LP, OS, UC).

Mulinum spinosum var. spinosum represents the nucleus of the species. A core of leaf structure can be detected, 3-5 lobes, with lobes to 4.3 cm wide. However, variation from this core exists. Plants occur with: a mixture of wide lobes and thin spine-like lobes, with rarely more than 3 lobes (Zech

& Contorgiorgakis 8); consistently 3-lobed leaves, with wide lobes and a trend towards larger leaves (Stuessy et al. 10073,

10216, 10318; Zech et al. 42, 50); a mixture of juvenile leaves similar to var. spinosum and mature leaves similar to

M. spinosum var. proliferum (Zech & Contorgiorgakis 3); 3- lobed plants with a reduction in lobe width (Correa &

Constance 3815/3822); primarily 3-lobed leaves of uniform width (Ricardi S. et al. 756); primarily 3-lobed leaves, with lobes long and thin (Aravena 33323); and leaves primarily 3- lobed, with long and linear lobes (Cajbrera 6720, Corte 165) .

lb. Mulinum spinosum var. minus (Kuntze) Zech, comb, et stat.

nov., Mulinum spinosum var. triacanthum (Griseb.)

Kuntze f. minus Kuntze, Revis. gen. pi. 3(2): 114.

1898.— TYPE: Argentina. Santa Cruz. Patagonia 50/3°,

April 1882, F. P. Moreno & Torini 372 (holotype: NY!). 56

Leaves small, 3-lobed, lobes spine-like, central lobe reflexed abaxially, lateral lobes spreading adaxially.

Phenology: Flowering mid-November through February (mid-

March) ; fruit maturing late November through February.

Distribution (Fig. 5). In Argentina, Mendoza Province. In

Chile, Regidnes IV and V, with a concentration in Limari

Province; 900-4600 m.

Representative Specimens. Argentina. Mendoza. Luj&n de

Cuyo: Santa Rosa de los Andes to Uspallato Pass, Moseley s.n.

(BM). Rio Negro. Bariloche: Parque Nacional Nahuel Huapi,

Eskuche 0927a, 0927b (UC). Chile. Region IV. Choapa:

Limahuida, Salamanca, Pfister 9597 (CONC), Cordillera de

Queldn, Salamanca, Zollner 11793 (MO); Elqui: Banos del Toro,

Collantes T. 23121 (CONC); Limari: Cabreria, Morro Blanco, iTiles 1232 (CONC) , Rio Molles, en los faldeos, Jiles 1972

(CONC), Cordillera San Miguel, Jiles 3594 (CONC), Rio Torca,

Jiles 3750 (CONC), Planta electrica Los Molles, Bocatoma,

Ricardi S. et al. 758 (CONC, OS), In the interior of Ovalle,

Zollner 5650 (UC). Regidn V. San Felipe de Aconcaqua:

Uspallata-Pass, Juncal, Buchtien 1185 (BM, E, GH, LY, S, US),

Valle del Alte los Leones, near Rio Blanco, high cordillera,

Zollner 2081 (UC).

This variety represents a small group of plants whose leaves are small, consistantly 3-lobed, with narrow (0.25-0.5 57

mm wide) spine-like lobes.

lc. Mulinum spinosum var. atacamense Zech, var. nov.— TYPE:

Chile. Region III. Huasco, Quebrada Yerba Buena [entre

Portezuelo Yerba Buena y Rio de Valeriano], 27 Jan

1983, C. Marticorena et al. 83613 (holotype: CONCl).

Folia crassus, trilobus, pubescentia, luteolus.

Leaves thick, 3-lobed, pubescent, light yellow.

Phenology: Flowering in January; fruit maturing in late

January (based upon little material). The fruit observed for

27 January appears immature, therefore fruit maturing through

February seems reasonable.

Distribution (Fig. 5): In northern Chile, Region III,

near Vallenar; 2200-3950 m.

Specimens Examined: Chile. Region III. Huasco: Rio de

Valeriano at Las Placetas, Johnston 6026 (S, US), Vallenar,

Rio Laguna Chica, Werdermann 268 (BM, E, F, G, GH, K, MO, S,

UC) .

The leaf compsition of M. spinosum var. atacamense is uniquely thick, with shallow lobes, and the material has been suggested by several annotators as a new species. However, the long peduncled umbels and general habit clearly place this new taxon in M. spinosum, but as a distinct variety. 58

Id. Mulinum spinosum var. proliferum (Cav.) Kuntze, Revis.

gen. pi. 3(2): 114. 1898. Mulinum proliferum (Cav.)

Pers., Syn. pi. 1. 309. 1805. Bolax prolifera (Cav.)

Spreng., Syst. Veg. 6: 361. 1820. Selinum proliferum

Cav., Icon. PI., 5: 58, t. 486, /. 1. 1799.— TYPE:

Argentina. Santa Cruz. Puerto Deseado, Dec, L. Nee

s.n. (holotype: MA, photo: OS!).

Mulinum echinus (K. Presl) DC., Prod. 4: 79. 1830. Bolax

echinus K. Presl, Prod. 4: 79. 1830.— TYPE: Chile. T.

P. X. Haenke s.n. (holotype: PR, not available).

Mulinum ovalleanum Philippi, Anal. Univ. Chile, 85: 709.

1894.— TYPE: Chile. "In Andibus departamenti Ovalle

provinciae Coquimbo et quidem en alto monte Huatulame"

[Rio Grande), Jan 1890, G. Geisse s.n. (lectotype,

designated by Constance, 1954: SGO!; isolectotypes:

SGO[2]!) .

Mulinum spinosum f. humilis Kurtz, Verh. Bot. Vereins

Prov. Brandenburg 34: 111. 1893.— TYPE: Argentina.

Mendoza. Los Molles, Rio Salado, 7 Jan 1893, Kurtz

7505 (lectotype, here designated: NY!).

Mulinum spinosum var. trispinesens Kuntze, Revis. gen.

pi. 3(2): 114. 1898.— TYPE: Argentina. Santa Cruz.

Patagonia 50/3°, April 1882, F. P. Moreno & Tonini 371

(holotype: NY!).

Mulinum patagonicum Gand. [nec M. patagonicum Speg.

Anales Soc. Ci. Argent. 48: 1899.], Bull. Soc. Bot. 59

France. 59: 710. 1912. Nom. illegit. Mulinum gandogeri

(Gand.) Hiroe, Umbelliferae of World. 1732. 1979. Nom.

illegit.— TYPE: Argentina. Santa Cruz. Patagonia,

prope Lago San Martin, 9 Jan 1909, C. J. F. Skottsberg

706 (holotype: LY!; isotype: SGO!).

Leaves large; lobes 3, narrow, 0.5 mm wide, and spine­

like.

Phenology: Flowering mid-November through February; fruit

maturing late November through March.

Distribution (Fig. 5): Throughout Argentina from San Juan

south to Santa Cruz Province. In Chile from Regidn IV to

Regidn XII, with the exception of Regidn X; 300-2500 m.

Representative Specimens. Argentina. Chubut. Cushamen: 11 km despuds de Leleque, 12 km N, toward El Maitdn, Correa &

Constance 4181/3844 (UC) , El Guri, Spasando Leleque, Isabel et al. 655 (LP), Colonia Cushamen, Rio Chubut, Meyer 8018 (NY);

Escalante: 14 km S del Comodoro Rivadavia, Atlantic Coast toward Caleta Olivia, Correa & Constance 4036/3834 (G, GH,

UC), El Trebol, De Marco de Kreibohm 119 (LP, MA, US);

Florentino Ameghino: 35 km S de Garayalde, Correa & Constance

3972/3827 (UC); Futaleufti: Tecka, Meyer 9633 (BM, F) , Reserva

Nacional de los Alerces at Lago Futalaufquen, Pedersen 293

(US) , Monte Nahuel Pan, ad est di Esquel, Pichi Sermolli &

Bizzarri 7372 (K), 22.7 km ESE of Esquel on Route 40, Stuessy et al. 6819 (OS); Languineo: Las Henicos, Castellanos s.n.

(NY); Rawson: ± 20 km camino a Cabo Raso, Correa et al. 4897

(UC); Rio Senguerr: Canadon de las Manos Pintadas, al SE de

Facundo, Llera & Torres 1870-2 (G); Sarmiento: 25 km del cruce

de Sarmiento hacia la Pampa de Marla Santisima, Correa &

Constance 4096/3840 (GH, MO, UC), Lago Musters, Correa et al.

4096 (UC) ; Telson: 10 km de Telsen hacia Gan Gan, Correa &

Constance 3893/3823 (UC). Mendoza. Las Heras: Path leading up ridge from Hotel at Villavicencio, Bartlett 19397 (GH, US) ,

Uspallata, Los Hornillos, Fabris 1209, 1213 (LP), Cerro

Melocoton, Ruiz L. 4029 (BM), 2 kms E of customs checkpoint

(Punta De Vacas) on road toward Mendoza, Stuessy et al. 10420

(CONC, GH, LP, OS, UC) ; Malargiie: Olla Colorada, Fiqueran 2834

(G), 33.5 kms N of Las Barrancas, 2 kms S of jet. road to

Camuco and Bardas Blancas, Stuessy et al. 10264 (CONC, GH, LP,

OS, UC) , 6 kms W of Los Molles, Ruta 222, Stuessy et al. 10269

(CONC, GH, LP, OS, UC), El Sosneado, up hill W from bridge over Rio Atuel, Stuessy et al. 10354 (CONC, GH, LP, OS, UC) ;

San Carlos: On road to Laguna Diamante before first army station, Zech & Contogiorgakis 10 (CONC, GH, LP, OS, UC), Road to Laguna Diamante, 4 kms before first army station, Zech &

Contogiorgakis 11 (CONC, GH, LP, OS, UC), On road to Laguna

Diamante, before the Valle Diamante, Zech & Contogiorgakis 12

(CONC, GH, LP, OS, UC); San Rafael: Sa. del Nevado, pampa entre arroya Aqua del Guacho y el zanjdn del Plateado al SSE de los Cos. Morados, Boelcke et al. 15747 (UC) , Laguna La Nina encantada, Dawson 1032 (LP) , Entre Malalhue & Ranquil-Co, Ruiz

L. 7674/23 (UC, US) , Valle de Rio Atuel, Wilczek & Lorentz 106

(G) ; Tunuyan: Cerca Puerto del Maurano, Ruiz L. 1107, 1724

(BM) , Los Arboles, Ruiz L. 2021 (BM) ; Tupungato: Valle de Rio

Tupungato, Hauman 252 (G). Neuquen. Alumine: 16 kms N of

Alumine, Ruta 23, Stuessy et al. 10158 (CONC, GH, LP, OS, UC) ;

Anelo: R. P. F., Med&n & Alvarez 292 (MO); Collon-Curd: Piedra

del Aquila, Correa & Constance 4206/3847 (G), 15 km N de Paso

Limay, Fabris 2251 (LP); Confluencia: Plaza Huincul, Plotnick

49 (LP); Huiliches: 13 km S de Junin de los Andes, near Rio

Quilquihue, Correa & Constance 4202/3846 (UC), 11 kms SW of

Junin De Los Andes on paved road to San Martin De Los Andes,

Stuessy et al. 10152 (CONC, GH, LP, OS, UC), 31.5 kms N of

Junin de Los Andes on Ruta 23, Stuessy et al. 10154 (CONC, GH,

LP, OS, UC); Lac&r: AlicurA, GonzAlez 210 (LP), Volcdn Lanin,

Joseph 5557 (CONC, S, US), 6.8 km N of San Martin De Los Andes on road to Parque Nacional Lanin, Stuessy et al. 10116A,

10116B, 10116C (CONC, GH, LP, OS, UC); Las Lagos: Estancia

Fortin Chacabuco, Hager 1044 (B), Along road from Bariloche to

Villa La Angostura, Landrum 3309 (LP); Loncopue: 44 kms S of

Loncopue, Ruta 231, Stuessy et al. 10187 (CONC, GH, LP, OS,

UC) , 36.3 kms N of Loncopue, Ruta 231, Stuessy et al. 10236

(CONC, GH, LP, OS, UC); Norquin: Termas de Copahue, Cabrera

6177 (LP), Andacollo, Mina Guaraco, Cabrera 11132 (LP),

Laderas orientales del Valle Chacaycd, Chicchi 72 (LP),

Trolope, O'Donell 2728 (S), 40 kms S Chos Malal, Ruta 40, Stuessy et al. 10252 (CONC, GH, LP, OS, UC) , 7.2 kms N of Chos

Malal on dirt road toward Tricao Malal, Stuessy et al. 10256

(CONC, GH, LP, OS, UC); Pehuenches: 10 km N of Buta Ranguil,

Ruta 40 toward Las Barrancas, Stuessy et al. 10260 (CONC, GH,

LP, OS, UC); Picdn-Leufu: Rio Limay, Roth s.n. (F) ; Picunches:

Pino Hachado, Cabrera & Crisci 19112 (LP); Rio Senguerr:

Vuelta del Senguerr, Canadon Grande, Cabrera 33 (LP); Zapala:

Zapala, Comber 193 (£) . Rio Negro. Bariloche: San Carlos de

Bariloche, en la estepa al E del pueblo, Burkart 6213 (F, US) ,

Arroyo Castillo, Cordini 229 (US), Rio Nirihuau, De Barba 1470

(MO) , Parque Nacional Nahuel Huapi, Lago Martin, Fabris &

Solbrig 1130 (LP), Arroyo Nireco, Meyer 8052 (BM, S) , Cerro

Leones, Perez M. s.n. (UC), Rio Limay, cerca del Lago Nahuel

Huapi, Teague s.n. (K) , El Bolson, Walter & Walter 543 (B) ,

Banks of Rio Limay, below Nahuel Huapi, West 4743 (UC) ;

General Roca: Entre Tricacd, Cabrera 18663 (LP, UC), Vicinity of General Roca, Fischer 189 (CM, F, MO, NY, US) ; Pichi-

Mahuida: Pichi Mahuida, Meyer 7577 (NY); Pilcaniyeu: Cuadro La

Figura, Estancia Pilcaneu, Carminatti s.n. (LP), NE de

Comallo, Klein 0160 (G); San Antonio: Ruta 3, 1272 km, Sierra

Grande, Vervoorst 5577 (UC); 25 de Mayo: Cerro Abanico,

Cabrera & Job 14 (NY), Ingeniero Jacobacci, Canadon La

Angostura, Estanica Abi-Saad, Carminatti s.n. (LP). San Juan.

Calingasta: Manantiales, Fabris & Zuloago 8428 (LP),

Manantiales, Zardini 101 (LP). Santa Cruz. Deseado:

Tehuelches, Donat 38 (GOET) ; Giier Aike: Rio Gallegos, Estancia Stag River, Tweedie 251 (K), Lago Argentino: Calafate, cercanas al Lago Argentino, Morales & Martin 236RM (MA) .

Chile. Regidn IV. Limari: Combarbald, Potrero Grande, Jiles

4564 (CONC). Regidn V. [Regidn Metropolitana de Santiago].

Area de Metropolitana de Santiago: El Rio de los Ojos de Aqua, cordillera, Gillies 561 (OXF). Regidn VI. Colchaqua: 10 km before Termas del Flaco, Zech et al. 33 (CONC, GH, LP, OS,

UC). Regidn VII. Linares: Valle Gualquivilo, Los Cipreses,

Schlegel 3592 (CONC) . Regidn VIII. Concepcidn: Camino entre el longitudinal y Pangal del Laja, cerca de El Guape, Marticorena

& Rodriguez 8407 (B, CONC), Concepcidn, Marticorena et al. 107

(OS), Pangal del Laja, al oeste de Yungay, Marticorena et al.

25287 (CONC), Camino de Estacion Umbel a Salto del Laja,

Pfister 7132 (CONC). Regidn IX. Malleco: Lonquimay,

Hollermayer 438, 438a, 438b (CONC, NY (438)), Malalcahuello,

Marticorena et al. 1248 (B). Regidn XI. Aisen: 43.9 kms E of

Cisne Medio on gravel rd. to La Tapera, Stuessy et al. 7527

(CONC, OS); Coihaique: Balmaceda, Magens 42 (B). Regidn XII.

Magallanes: Puerto San Jose, DusSn 5249 (PRC, S); Ultima

Esperanza: Bories, Elliot 334 (E, K), Salto Grande del Payne,

Pisano-Valdez 2372 (CONC), Road from Puerto Natales towards

Torres Del Paine, before Laguna Figueroa, Zech et al. 55

(CONC, GH, LP, OS, UC); Tierra del Fuego: Casa Tweedie,

Nordenskiold 32 (UPS).

While the type of M. echinus was requested from both PR 64 and PRC it has not yet been received, and at this writing is assumed unavailable for study. This taxon is placed in M. spinosum var. proliferum based upon the number (7-9) and nature of the involucral bracts, "foliolis", and the leaves being described as "3-fidis", and the divisions "spinoso- subulatis" (Candolle 1830: 79).

The leaves of this variety are consistantly 3-lobed and large, representing the larger individuals of M . spinosum in general.

II. Mulinum section Leptacantha Zech, sect. nov.— TYPE:

Mulinum leptacanthum Philippi.

Subshrubs, mats, or cushions; involucre 1-whorled; involucral bracts membranous; sepals dentate, 0.5-1 mm long.

This section is based upon the absence of the shrub and presence of the cushion habit in addition to subshrubs. In addition the taxa included within this section produce a single whorl of membranous involucral bracts, and sepals which are reduced to teeth.

IIA. Mulinum subsection Leptacantha— TYPE: Mulinum

leptacanthum Philippi.

This subsection is distinguished by the characteristics of its single member, M. leptacanthum (e.g., low, creeping, 65

habit; leaves with closed sheaths; peduncled umbels; umbels 15

or more flowered; and 6 or more involucral bracts).

2. Mulinum leptacanthum Philippi, Anal. Univ. Chile 85: 709.

1894.— TYPE: Chile. "Prope Nitrito en Araucania", 30

Jan 1887, C. Rahmer s.n. (holotype: SGO!, fragment:

UC!; isotype: SGO!).

Azorella nivalis Philippi, Anal. Univ. Chile 85: 703.

1894. Mulinum nivale Constance ex Crowden,

Phytochemistry 8: 1963-1984. 1969. Norn. invalidum,

basionym not given.--TYPE: Chile. "In Andibus

provinciae Valdiviae supra nivem pertetuam loco dicto

Huahuim" [en las nieves eternas, por Huahuin, fide

Munoz P. 1960], Jan 1887, O. Philippi s.n. (holotype:

SGO!, photo: F!, photocopy: OS!; isotype: SGO!).

Mulinum patagonicum Speg. Anales Soc. Ci. Argent. 48: 56.

1899 [nec M. patagonicum Gand., Bull. Soc. Bot.

France. 59: 710. 1912. ] .— TYPE: Argentina. "In

rupestribus montanis prope Lago Nauel-Huapi", Dec

1897, C. Speggazini s.n. (holotype: LP, photocopy:

OS!) .

Low, creeping, compact to spreading perennials, 4-6 cm tall, 8-18 cm diameter. Stems compact, 2 mm diameter, covered with decaying brown leaves and/or leaf skeletons. Leaves thick, 0.6-2(-3) cm long, 2.5-15 mm wide; blade triangular, smooth to sulcate, venation extensive; lobes 3, rarely 4-6, 2-

8 mm long, 0.3-2 mm wide, lanceolate, sinusus not extending to

petiole, at apex finely acute to mucronate; constriction

absent, blade continuous with petiole; petiole 0.5-5 mm long,

0.2-2 mm wide, at base dilated; sheath ± membranous, open, 5-7

nerved, 4 mm long, 4-6 mm wide, with margin shortly cilliate,

hairs straight; stipules absent. Umbel 8-13 mm diameter, 6-31

flowered, spreading, rarely extending beyond upper leaves;

peduncle 2-5(9) mm long, 1 mm diameter, uniform. Involucre

glabrous; involucral bracts 5(-7), equal, membranous, 1.5-2.5

mm long, 0.5-1.25 mm wide, linear-lanceolate, at apex narrowly

acute, at base fused into an elongated cup, 3 mm long, with margin entire or shortly lobed, sparsely to evenly cilliate, hairs short. Rays 2-3 mm long, 0.5 mm diameter, at apex receptacle enlarged and constricted or uniform. Flowers 3 mm diameter; sepals as teeth, green, at apex acute to shortly acuminate, at base forming a shallowly lobed stellate disc,

0.5 mm long, 1 mm wide; petals yellow, 1.5 mm long, 0.5 mm wide, planar, elliptic to obovate, at apex acute to rounded; stamens deciduous, with filament 1 mm long, 0.1 mm wide, at base uniformly wide, with anthers 0.75 mm long, 0.5 mm wide; carpels entire; stylopodium 1-1.25 mm diameter, with styles

0.1 mm long, elongating to 1 mm. Fruit widely elliptic to circular, yellow, 4-winged in cross-section, body deflated, rectangular in cross-section, 6 mm long, 6 mm wide, at apex , at base ; wings reniform, 6 mm long, 3 mm wide, lateral wings 67

longer and broader than the fruit body, straight; mericarps

linear in cross-section, separating easily, abaxial face

slightly concave, commissural face convex, ribs 5, 1 abaxial,

prominent and equal; oil tubes round to oval; carpophore

equally bilobed; odor absent. Chromosome number: n = 8. Fig.

4.

Phenology: Flowering the end of December through February

[with the exception of a single specimen dated 24 April (Diem

3349, UC)]; fruits maturing mid-January through late April.

Distribution (Fig. 6) . Primarily in Neuquen province,

Argentina, and adjacent in Regidn IX, Chile, with an extension south into Province Rio Negro, Argentina, and extensions north and south into Regidnes VIII and X, Chile; 1300-2700 m.

Representative Specimens. Argentina. Neuquen. Alumine:

Pass near Quillen [Paso de Afiiheraqui], Elwes s.n. (K); Chos

Malal: extremo NW Pampa Ferraina, Boelcke et al. 11378 (UC);

Ldcar: Lago Huechulafauen, Dawson & Schwabe 2633 (UC), Parque

Nacional Lanin, cerro Bolsico del Diablo, Eskuche 0928, 0929

(UC) , Hua Hum, Co. Malo, Hunziker 6993 (UC), Near Chapelco, 22 kms SE of San Martin De Los Andes, Up from Refugio Federico

Graef, Stuessy et al. 10017, 10047, 10048, 10049, 10072 (CONC,

GH, LP, OS, UC), Upper reaches of Arroyo Chapelco Grande, up

N fork to Lago La Kika, Stuessy et al. 10129 (CONC, GH, LP,

05, UC) ; Los Lagos: Puerto Manzano, Lago Nahuelhuapi, Diem

3349 (UC); Minas: Lagunas Epu-lauquen, cerro al NW de las 68

FIG. 6. Distribution of Mulinum leptacanthum in Argentina and Chile. lagunas, Boelcke et al. 10970 (UC), Valle superior del Atreuco en pedregal, Boelcke et al. 11500 (UC), Cordillera del Viento, cruzada de Tricao Malal al Cajon de Butalo, Boelcke et al.

11613 (UC), Paso del Macho, Boeleke et al. 13897 (UC), Sierra de Cochico, cajon de la Botica, Boeleke et al. 14121 (UC) ,

Cajon del Portillo, Boeleke et al. 14179 (UC); Norquin: Termas de Copahue, Cabrera 6215, 6223 (LP) , 1 km S of Copahue just below road to Las Maquinitas, Stuessy et al. 10198 (CONC, GH,

LP, OS, UC); Pehuenches: Pampa de Lonco Luan, semidesertico,

Ruiz L. 25807 (UC) , Paso Pino Hachado, N of the

Chilean-Argentinean border, Stuessy et al. 10162 (CONC, GH,

LP, OS, UC) . Rio Negro. Bariloche: Parque Nacional Nahuel

Huapi, Filo cerro Colorado, Boeleke & Correa 6985, 6986, 6987,

6988, 6989, 6990 (UC), Cerro Gutierriz, Buchtien 43 CK TO SEE

IF ALL ARE EQUAL (LY, US) , Parque Nacional Nahuel Huapi, cerro

Catedral, Eskuche 01448, 299-1 (UC), Parque Nacional Nahuel

Huapi, Catedral, Nido de Condores, Perez M. 49907 (UC), Parque

Nacional Nahuel Huapi, cerro Nireco, Perez M. 50370 (UC) ,

Parque Nacional Nahuel Huapi, Cordon del Catedral, Piedra del

Codon, Perez M . 50371 (UC), Gathered in Bariliche, Zollner

5208 (UC) . Chile. Region VIII. Nuble: San CcLrlos, Cordillera de Los Sauces, Barros 1171 (CONC). Region IX. Cautin: Andes,

Villarrica, Reger 1897 (W) ; Malleco: Lonquimay, Joseph 5489

(US) , 1.7 km W del Paso Pino Hachado, Keever & Greer 1 (CONC) ,

7 km from Liucura on the road to Paso Pino Hachado, Zech et al. 46 (CONC, GH, LP, OS, UC). Region X. Valdivia: Valdivia, 70

Hollermayer 489 (W).

Some specimens of M. leptacanthum (e.g., Stuessy et al.

10162) have features which are similar to M. spinosum var. spinosum. The mat-like habit is similar, along with larger leaves (to 30 mm long, 15 mm wide), and leaf lobes which are

3-5 in number and somewhat wider (to 2 mm).

When fruits of M. leptacanthum (Diem 3349) were cultivated in the University of California Botanical Garden by

Constance for chromosome studies (Voucher C-762) , this species lost its generally compact nature, suggesting an environmental influence on compactness.

The name Mulinum nivale was found on several sheets of material from K, L, and UC determined by Lincoln Constance in the 1960s. Constance suggested that Azorella nivalis Philippi be combined with Mulinum and went so far as to write "Mulinum nivale (Phil.)" on his annotation labels. However, Constance

(in litt.) never authored the transfer nor published the new combination. Crowden et al. (1969) used material of this same taxon for flavonoid analysis and published the name in their list of vouchers. Constance (1988) later combined A. nivalis with M. leptacanthum, a combination that is supported by this present study.

IIB. Mulinum subsection Crassifolia Zech, subsect. nov.—

TYPE: Mulinum crassifolium Philippi. 71

Subshrubs or cushions; leaf sheaths open or closed;

umbels 1-14 flowered, rarely 15 or more, peduncled or sessile;

involucral bracts 2-5, rarely 6 or more.

This subsection is defined by a combination of flower and

involucral bract number, and to a lesser extent by habit, leaf

sheath closure, and umbel position.

IIB1. Mulinum series Crassifolia— TYPE: Mulinum crassifolium

Philippi.

This series is defined by sessile umbels, closed leaf

sheaths, and a more northerly distribution.

3. Mulinum crassifolium Philippi, Florula Atacamensis, In

Viage al Desierto de Atacama 198. 1860.— TYPE: Chile,

"in monte Alto de Puquios 23° 52/H, 12500 m, Feb 1854,

R. A. Philippi s.n. (holotype: SGO!, photo: F!;

isotype: B, photo: F! G! GH! NY!).

Caespitose shrub, 5-10 cm tall, 8-30 cm diameter; resinous. Stems dense to spreading, 2-7 mm diameter, covered with old leaf skeletons. Leaves thick and fleshy, especially the new ones, sessile, 5-15 mm long, 1-5 mm wide; blade rhomboidal, glabrous, 4 mm long, 2-5 mm wide, 3-veined, at apex obtuse; lobes 3, incurved, 0.5-4 mm long, 1-2.5 mm wide, center lobe larger than the laterals, apex rounded to acute and mucronate; constriction absent, blade continuous with

petiole; petiole 1-4 mm long, 1.25-1.75 mm wide, base dilated;

sheath open, membranous, 4-5 mm long, 3-6 mm wide, with margin

entire or serrate; stipule absent. Umbel 2-5 mm diameter, 2-

11 flowered, spreading, not extending beyond leaves, sessile.

Involucre glabrous; involucral bracts 2-5, equal or unequal,

membranous, 2-3 mm long, 0.5-1 mm wide, linear-lanceolate, at

apex acute, at base connate, into a cylindrical cup, with

margin entire; rays 1-4 mm long, 0.25 mm diameter. Flowers 2-

3 mm diameter; sepals as teeth, green, at apex acute, at base

forming a stellate disc, 0.5-0.6 mm long, 0.5-1 mm wide;

petals yellow, 1-2 mm long, 1 mm wide, planar, elliptic to

obovate, at apex rounded to acute; stamens deciduous, with

filament 0.75-1.25 mm long, 0.2 mm wide, at base uniformly wide, with anthers 0.3-0.5 mm long, 0.2 mm wide; stylopodium

1.25 mm diameter, with styles ± 0.1 mm long, elongating to 1 mm. Fruit yellow, elliptic to widely elliptic, 4-winged in cross-section, body deflated, rectangular in cross-section, 3-

4 mm long, 3-4 mm wide; wings lunate, 4 mm long, 1 mm wide,

lateral wings longer and broader than the fruit body, revolute; mericarps lunate in cross-section, separating with difficulty, abaxial face concave, commissural face slightly convex to planar, ribs 5, 1 abaxial, prominent and equal; oil tubes oval to deltoid, adaxials small, abaxial and marginal large; carpophore entire; odor absent. Fig. 7.

Phenology: Flowering late September through February; 73

A.

B.

FIG. 7. Cross section of fruit. A. Mulinum crassifolium (based on Pfister 9374, CONC). B. Mulinum ulicinum (based on Ruthsatz 166, UC). 74

fruit maturing January through mid-February. This is based

upon a small number of collections (7 each for flowers and

fruit) which were mostly sterile. In general fruit was

absent. The fruit maturation time seems narrow, and probably

begins earlier, to coincide with flowering and may extend

beyond February.

Distribution (Fig. 8). This species occurs in northern

Chile within Regidnes II and III, and also in Argentina in the northern Provinces of Salta and Jujuy; 2300-4500 m.

Common Names: "Zucurco" ["Sucurco"] by the Atacamian

Indians (Philippi 1860); "Sucurco", "Epinilla", "Chuquicdn"

(Baeza R. 1921; Montes & Wilkomirsky 1985); "Chuquicana", or

"Chuchican" (MuRoz P. 3770, 3800, 3807)

Specimens Examined. Argentina. Jujuy. Moreno in campo,

Fries 840 (UPS). Salta. Camino a Mina La Julia, Corrida de

Cori, Cabrera 8790 (LP), Quebrada del Ajua, Lecoi & Montana

5321 (LP). Chile. Regidn II. Antofagasta: Cospa, Castro s.n.

(CONC), Salar de Tara, Hoffmann s.n. (CONC) , Locaire, Munizaga

A. s.n. (CONC), Faldeos del VolcSn Llullaillaco, camino a la

Mina Esperanto, MuHoz P. 3800, 3807 (UC), Laguna Meniques,

Niemeyer F. s.n. (CONC), Camino Chindor, Novas B. 2097 (CONC),

Desertum Atacama, R. A. Philippi s.n. (K) , San Pedro de

Atacama, Casablanca, Ricardi S. 2986 (CONC, OS) , Taltal, Cord.

Volcdn Llullailaco, Werdermann 1009 (B, BM, CONC, E, F, G, GH,

MO, NY, S, UC, US); Loa: Laguna Lejia, Behn s.n. (CONC), El 75

FIG. 8. Distribution of Mulinum crassifolium in Chile, Regi6nes II and III, and Argentina, Provinces Jujuy and Salta. 76

Tatio, Castillo s.n. (CONC), De Talabre a la Launa Lejla, Mahu

6261 (UC), Pampa Choculagui, entre Socaire & Laguna Miscanti,

Niemeyer F. s.n. (CONC), Cerro Le6n Chuquicamata, Perry s.n.

(CONC), Camino a El Tatio, Pfister s.n. (CONC), Quebrada de

Amincha, al pie del cerro Aucanquilcha, Pisano-Valdez &

Venturelli E. 1978 (UC), Toconce, Schlegel 2653 (CONC),

Alrededores de Toconce, El Tatio, llaretales cerca de las

fumarolas, Schlegel 5124 (CONC), Villagriin 1334 (CONC),

Monturaqui, Zollner 1280 (CONC), Volcdn Tatio, Zollner 3517

(UC). Regidn III. Copiapd: Monte Altos de Puquios, Munoz P.

3770 (UC).

Mulinum crassifolium differs from other species in its

unique leaf structure and distribution. More resinous than

any other species, all parts contain an amber-colored resin

which may be an adaptation against water loss, heat abortion,

or herbivory. Although deterioration of the leaves is present

in other Mulinum taxa, it is most striking in M. crassifolium.

The initally fleshy and thick leaf tissue deteriorates

revealing a spine-like skeleton of vascular traces below (5 mm

long, 0.2-0.3 mm wide), appearing in stages similar to the

flesh of a mumified corpse.

Mulinum crassifolium has been suggested as medicinal for

respiratory ailments, diabetes, cold prevention, indigestion, and against coughing (Montes & Wilkomirsky 1985). 77

4. Mulinum ulicinum Gillies & Hook., In Hooker's Bot. Misc. 1.

328, t. 64, 1830.— TYPE: Argentina. "Prope 'La Cienega

de Bonillo,' in summum fere jugum montium

Uspalatensium", 9500 ft, 1820-1828, J. Gillies s.n.

(holotype: K!; isotypes: E[2]!.

Subshrub 5-13 cm tall, 1.6-15 cm diameter; resinous.

Stems dense to spreading, 3-4 mm diameter, covered with old

leaf skeletons. Leaves thick to narrow, stiff, easily seperating when dry, 8-44 mm long, 3-30 mm wide, axillary leaf production copious, sessile or petioled; blade glabrous to rarely pubescent, hairs short, at apex; 3-lobed, spreading, 3-

15 mm long, 0.5-1 mm wide, center lobe larger than the laterals or lobes equal, center lobe strongly reflexed abaxially, lateral lobes reflexed adaxially, at apex acute to shortly mucronate; constriction absent, blade continous with petiole; petiole 9.5 mm long or less, 1.5 mm wide, base dilated; sheath closed, 3-7 mm long, 3.5-10 mm wide, 1-7 veined, with margin entire or with 3 to 4 hairs, simple, white, 2.5 mm long or less; stipules absent. Umbel 4-7 mm diameter, 3-14 flowered, spreading, not extending beyond leaves, sessile to shortly peduncled; peduncle 0.5-4 mm long,

0.3-0.5 mm wide, at apex uniform. Involucre glabrous; involucral bracts 3-5, equal, green to membranous, 2-5 mm long, 1 mm wide, lanceolate to linear-lanceolate, at apex acute to shortly mucronate, at base connate into a cylindrical cup, with margin entire. Rays 2-4 mm long, 1 mm diameter at

apex receptacle enlarged and constricted. Flowers 2.5-3 mm

diameter; sepals as teeth, at apex acute, at base forming a

stellate cup, 0.5-1 mm long, 0.75 mm wide; petals yellow, 1-

1.75 mm long, 1-1.25 mm wide, planar, ovate to obovate, at

apex acute to rounded; stamens deciduous, with filament 0.5-1 mm long, 0.2 mm wide, at base uniformly wide; carpels entire;

stylopodium 1.5 mm diameter, with styles ± 0.1 mm long,

elongating to 1 mm. Fruit oblong to widely elliptic, 4-winged

in cross-section, body deflated, squarish to rectangular in cross-section, 4-6 mm long, 2-5 mm wide, at apex rounded to acute, at base rounded to cordate; wings lunate to widely

lunate, 4-5 mm long, 1-2 mm wide, lateral wings longer and broader than the fruit body, straight or revolute; mericarps lunate, linear, or shallowly triangular in cross-section, separating easily, abaxial face concave, commissural face planar to convex, ribs 5, 1 abaxial, prominent, equal; oil tubes round, oval, or irregular; carpophore entire, shortly

2-cleft, or bilobed; odor absent or coconut-like. Fig. 7.

Phenology: Flowering mid-December through mid-March; fruit maturing middle to late December through early April.

Distribution (Fig. 9). Primarily in Argentina from

Mendoza Province north to Jujuy Province with a limited extension over the border into Southern Bolivia; 1000-4950 m.

Common Names: "Espinillo"(Rodriguez 293), "Chuchicana"

(Cabrera 8353), and "Espina Amarilla" (Dell'Arciprete s.n., 79

FIG. 9. Distribution of Mulinum ulicinum in Argentina and Bolivia. C = Catamarca; LR = La Rioja; M = Mendoza; J = Jujuy; S = Salta; SJ = San Juan; T = Tucuman. • = M. ulicinum var. ulicinum; a - M. ulicinum var. multiflorum. 80

NY) .

Mulinum ulicinum differs from all other Mulinum taxa by the consistant production of axilllary, ± sessile umbels.

Mulinum ulicinum is closest to M. crassifolium. The 2 species share a northernly distribution as well as leaves with spine­ like lobes.

Weddell (1857) treated M. ulicinum Gillies & Hook, in

Mulinum for Chloris Andina, but his illustration is different from the illustration and type of M. ulicinum Gillies & Hook.

Because Weddell's plate has caused some confusion, it is useful to indicate that Weddell's illustration is based upon

F . Pafaltique 360 (P) , which is closer in appearance to M. spinosum var. proliferum.

Mulinum ulicinum has been described as being used against cough and the leaves as aromatic (Jorgensen 1692, UC).

This species is variable in leaf size and composition and in peduncle length and therefore is considered to comprise two varieties, both of which have a geographic basis.

Key to the Varieties of Mulinum ulicinum

1. Leaves small, 7-15 mm long, 3-10 mm wide.

4a. M. ulicinum var. ulicinum.

1. Leaves large, 19-44 ram long, 13-30 mm wide.

4b. M. ulicinum var. multiflorum. 4a. Mulinum ulicinum var. ulicinum

Mulinum echegarayi Hieron., Bol. Acad. Nac. Ci. 4: 27-28.

1881.— TYPE: Argentina. San Juan, "pie del Tontal y en

la Quebrada del Paramillo", Jan 1876, D. S. Echegaray

s.n. (holotype: CORD, photo: OS!; isotype: B, photo:

F! G! GH! NY!).

Most leaves small, 7-15 mm long, 3-10 mm wide.

Involucral bracts 2 mm long, lanceolate. Fruit oblong, appearing squarish in cross-section, 4 mm long, 3-4 mm wide, at apex acute, at base cordate; wings lunate, 1.75-2 mm wide,

4-4.5 mm long, revolute; mericarps lunate in cross-section, commissural face convex; oil tubes oval; carpophore free, shortly 2-cleft or bilobed; odor absent.

Phenology: Flowering mid-December through February; fruit maturing late December through late March.

Distribution (Fig. 9): In Argentina from Jujuy south to

Mendoza Province; 2700-4950 m.

Specimens Examined: Argentina. Catamarca. Cerro Negro,

Chiquerito, Briicher s.n. (S) , Altura del Espiritu Santo del

Cerro Negro, Sierra Famatina, Hieronymus & Niederlein 819 (G) ;

Santa Maria: La Hoyack, Quebrada del Zarzo, Sierra Aconquya,

CKPeirano s.n. (S), Cordillera del Queouquija, Venturi 4785

(CM, GH, US). Jujuy. Moreno in campo arenoso ricco, Fries 840

(S, UPS); Humahuaca: Mina Aguilar, Cabrera 9214 (LP), Mina Aguilar, Cabrera & Constance 18964 (LP), Cordillera Oriental,

Tres Cruces Mts., Colquiri, Carter s.n. (F), Foot of Cerro

Tres Cruces enroute to Tres Cruces, Constance & Cabrera 3854

(US), above main settlement of La Hina Aguilar, Constance &

Cabrera 3856 (CM, UC) , Mina Aguilar, Frangi & Kiesling 55

(LP), Toreia de aqua de la Veta, Ruthsatz 4 (UC) , Mina

Aguilar, Veta, Ruthsatz 7 (UC), Mina Aguilar, Rio Despensas,

Ruthsatz 44 (UC), Mina Aguilar, Toria de aqua de Illolivo,

Ruthsatz 57 (UC), Mina Aguilar, Espinazo del Diablo, Ruthsatz

275 (UC), Mina Aguilar, Ruthsatz 336 (UC); Susques: Sey,

Cabrera 9069 (UC), Sey, Ruthsatz 204 (UC), Abra de Chorrillos,

Ruthsatz 210 (UC), N de la Salina Olaroz, Ruthsatz 225 (UC),

Humahuaca, Ruthsatz 507 (UC); Tumbaya: Abra de Pives, Ruthsatz

126 (UC) , E de la Abra de Pives, Ruthsatz 135, 136 (UC) .

Mendoza. Las Heras: 24 km NE Uspallata camino a Villavicencio,

Boelcke et al. 9940 (UC), Cordillera del Paramillo de

Uspallata, Las Minas del Paramillo, Kurtz 9463 (UC), Paramillo de Uspallata, Roig & Ruiz L. 20166 (UC), Arroyo Tambillo,

Semper 10091 (UC) ; Luj&n de Cuyo: Los Vallecitos, Ruiz L.

13400 (UC) ; Malargiie: Paso de la Sierra Bomilla, s.c. 83 (LP) .

Salta. Los Andes: Angosto de Tocomar, Cabrera et al. 22540

(UC); San Antonio de los Cobres: Quebrada de Pohorillos,

Cabrera 8353 (GH); San Carlos: Cerro de Cachi, Venturi 6977,

7275 (US). San Juan. Calingasta: Paso del Espinacito,

Castellanos 15501 (US), Rio del Palque, Talguana, Fabris &

Marchionni 2374 (LP), Entre Quebrada Honda y Las Lenas, Paso 83 del Espinacito, Valle de los Patos Perez M. s.n. (UC), Las

Frias, Roig s.n. (UC); Tafi: Cumbres Calchaquies, Burkart 5344

(UC) , Calchaquies, Quebrada del Matadero, de la Sota 2725

(LP), Cumbres Calchaquies, Lillo 5509 (G).

While Mulinum ulicinum var. ulicinum partially overlaps

in distribution with and has been suggested as similar to M. albovaginatum, this variety differs in smaller leaves (8 mm long, 3-4 mm wide) and a generally less compact nature. The compactness of var. ulicinum does vary with some plants (e.g.,

P6rez M. s.n., UC) approaching M. albovaginatum.

Mulinum ulicinum var. ulicinum differs from var. multiflorum by producing smaller leaves, mm long, mm wide, with broad lobes, mm wide. However, some plants of var. ulicinum (e.g., Fries 840) produce both broader lobed and leaves with spine-like lobes which are similar to the leaves of var. ulicinum.

4b. Mulinum ulicinum var. multiflorum (Hieron.) Zech, comb.

nov. Mulinum triacanthum var. multiflorum Hieron.,

Bol. Acad. Nac. Ci. 4: 27. 1881.— TYPE: Argentina. San

Juan. Feb 1876, D. S. Echegaray s.n. (holotype: CORD,

photo: OS!).

Mulinum axilliflorum Griseb., Abh. Konigl. Ges. Wiss.

Gottingen 19: 106. 1874.— TYPE: Argentina. Catamarca.

"In convallibus arenosis exceisis inter Nacimientos et 84

Laguna blanca", 9-10,000 ft, Jan 1872, P. G. Lorentz

430 (holotype: GOETi).

Mulinum spinosum var. triacanthum (Griesb.) Kuntze, Revis.

gen. pi. 3(2): 114. 1898. Mulinum tricanthum Griseb.,

Abh. Konigl. Ges. Wiss. Gottingen 19: 106.

1874.— TYPE: Argentina. Catamarca. "In alpinis Vayas

altas pr, Belen", Jan 1872, P. G. Lorentz 586

(holotype: GOET!; isotype: B, photos: F! G! GH! NY!).

Most leaves large, 19-44 mm long, 13-30 mm wide.

Involucral bracts 4-5 mm long, linear-lanceolate. Fruit

widely elliptic, appearing rectangular in cross-section, 4-6

mm long, 4-5 mm wide, at apex acute, at base cordate; wings

widely lunate, 5 mm long, 1.5 mm wide, straight; mericarps

shallowly triangular to linear in cross-section, commissural

face convex to plane; oil tubes oval, round, or irregular;

carpophore free, entire or shortly 2-cleft; odor absent or

coconut-like.

Phenology: Flowering [December] January through mid-

March; fruit maturing mid-December through early April.

Distribution (Fig. 9) : Primarily within Argentina with an extension over the northern border of Argentina into southern

Bolivia; 1000-4400 m.

Specimens Examined: Argentina. Catamarca. Cuesta del

Abajo, Cerro de las Capillitas, Schickendantz 334 (G) ; Andalgalci: Cerro Medanitos, Jorgensen 1692 (GH, MO, UC, US);

Belen: Faldeos N del Portezuelo del Blanco, arriba del

Granadillas, Sleumer & Vervoorst 2578 (UC, US); Santa Marla:

Sierra del Queouquija, Venturi 6450 (US). Jujuy. Cochinoca:

camino de Cochinoca a Abra Pampa, que pasa por I NT A, 10 km de

Abra Pampa, Dell'Arciprete s.n. (NY), Laguna de Quayalayoc,

Ruthsatz 166 (UC), Abra Pampa, Ruthsatz 241 (UC); Humahuaca:

Mina Aguilar, Bajada al Rio Despensa, Ancibor 32 (UC), Mina

Aguilar, Cabrera 9221 (LP), Esquinas Blancas, Cabrera 17734

(LP, UC), Foot of Cerro Tres Cruces enroute to Tres Cruces,

Constance & Cabrera 3854 (GH, UC), Tres Cruces, Cabrera &

Constance 18947 (LP, UC), Mina Aguilar, Cabrera & Constance

18964 (UC), Tres Cruces, Cabrera et al. 21433 (LP), Azul

Pampa, Claren s.n. (S), Esquinas Blancas, Ruthsatz 35 (UC),

Mina Aguilar, Espinazo del Diableo, Ruthsatz 40, 47 (UC);

Tilcara: Tilcara, Venturi 6451 (US); Yaui: Camino de Elku a

Caugregillos, Ruthsatz 363 (UC). La Rioja. Chilecito: sierra

Famatina, Hieronymus & Niederlein 361 (G), Mina Yareta, Sierra

Famatina, Hieronymus & Niederlein 808 (G), Sierra de Famatina,

Quebrada de La Cunsche, Krapovickas 6250 (UC), Famatina,

Stuckert 15140 (G), Famatina, Stuckert 15125 (UC) ;

Chicligasta: Pueblo Viejo, Venturi 6449 (US), Estancia Santa

Rosa, Venturi 9042 (US); Sarmiento: Cortaderas, precordillera,

Hunziker 2099 (UC), Quebrada del Rio Salado, Hunziker & Caso

4145 (UC); Vinchina: Cordillera d los Andes Venturi 10279 (BM,

GH, MO, S, US) . Mendoza. Las Heras: Rio Vacas, NE of Aconcaqua, Miche 171/80/24.4, 171/80/25.4 (UC); San Carlos:

Pampa de Las Osamentas, Estancla Llancha, Ruiz L. 7182 (UC);

Tunuyan: Camino al Paso del Portillo Mendocino, Cuesta de Los

Afligidos, Ruiz L. 1855 (BM, UC) , Valle del Alto Tunuyan, Ruiz

L. 2135 (BM), Precordillera, cerca del macimiento del Arroyo de Las Cuevas, Ruiz L. 3126 (BM, UC) . Salta. San Carlos: Cerro de Cachi, C. S. [Speggazini] s.n. (LP), Ambiaio, Cachi Pampa,

C. S. [Speggazini] s.n. (LP), Cerro de Cachi, Venturi 7276,

7277 (GH (7276), US). Tucuman. Lara, Rodriguez 293 (BM, UC) ;

Tafi: Sierra del Cajun, Venturi 10277 (US). Bolivia. Puna

Palanca, Fiebrig 2601 (B, BM, F, G, GH, MA, MO, S, UPS, US).

Pacajes. La Paz: Ulloma, Asplund 2536 (S, UPS).

Mulinum ulicinum var. multiflorum differs from var. ulicinum by the consistant production of larger leaves (19-44 mm long, 13-30 mm wide) with spine-like lobes. The plants of var. multiflorum are large (7-18 mm tall) subshrubs with some exceptions. Few plants are smaller (3-4 mm tall) and more mat-like (e.g., Ruiz L. 1855, Venturi 6450).

IIB2. Mulinum series Azorellopsis (H. Wolff) Zech, stat. et

comb. nov.— TYPE: Mulinum albovaginatum var.

pauciflorum (Reiche) Zech

Pulvini; umbellae sessilis.

Cushions; umbels sessile. 87

This series is characterized by habit and sessile umbels.

5. Mulinum albovaginatum Gillies & Hook., In Hook. Bot. Misc.

1. 328-329. 1830.— TYPE: Argentina. "'Cerro de la

Polcura', in Andibus versus Mendozam", 1820-1828, J.

Gillies s.n. (holotype: K!; isotypes: E[2]!).

Caespitose perennials, creeping, often covering rocks

forming small, compact, cushions, 2-10 cm tall, 4-15 cm diameter. Stems erect and compact, 1-2 mm diameter, covered with dead leaves. Leaves sessile or petiolated, strongly imbricated, the upper leaves rossette-like, 4-17 mm long, 1.5-

8 mm wide; blade rhomboidal, sulcate, granular, glabrous to rarely pubescent, 1-6 mm long, 1.5-8 mm wide, at apex ± 90° to obtuse; lobes 3, chunky to slender, 1-4 mm long, 0.5-2 mm wide, at apex acute to 90° to mucronate; constriction absent, blade continous with petiole; petiole 0.5-2 mm long, 0.5-2 mm wide, 3-5-veined, generally 3, at base dilated; sheath open,

1.8-6 mm long, 1.5-4 mm wide, with margin entire or rarely pubescent, hairs straight, 0.4 mm long. Umbel 1-6 flowered, spreading, often not extending beyond leaves, sessile or peduncled; peduncle ribbed, 3-4 mm long, 0.75 mm diameter.

Involucre glabrous; involucral bracts 5, equal, membranous,

0.5-3 mm long, 0.5-1 mm wide, ± linear-lanceolate, at apex narrowly acuminate, at base connate, into cylindrical cup, with basal margin entire or fringed, hairs 1-4, ± 0.1 mm long; rays ribbed, yellow to membranous, 3-7 mm long, 0.25-0.5

diameter, at apex receptacle enlarged and constricted.

Flowers 3-4 mm diameter; sepals as teeth, at apex ± 90°, at

base forming a stellate disc, 0.5-1 mm long, 1 mm wide; petals

yellow, reflexed abaxially, 1.25-2 mm long, 0.5-1.25 wide,

ovate to obovate to oblong, at apex acute to mucronate;

stamens deciduous, with filament 1.25 mm long, 0.2 mm

diameter, at base uniformily wide, with anthers 0.75 mm long,

0.5 mm wide; stylopodium 1-2 mm diameter, with styles ± 0.1 mm

long, elongating to 1 mm. Fruit yellow, elliptic, 4-winged in

cross-section, body deflated, rectangular in cross-section, 5-

7 mm long, 4-5 mm wide, at apex acute, at base cordate; wings

lunate, 5-6.25 mm long, 1.25-2 mm wide, lateral wings as long

or longer and broader than the fruit body, straight or

revolute; mericarps lunate in cross-section, separating easily, abaxial face concave, commissural face convex, ribs 5,

1 abaxial, prominent and ± equal; oil tubes oval, 0.25-0.75 mm diameter; carpophore entire or shortly 2-cleft; odor celery-like or absent. Fig. 10.

Phenology: Flowering mid-November through late February; fruits maturing December through early April,

Distribution (Fig. 11). In the Andes of Central

Argentina, with a concentration in Mendoza Province. This species does extend southward into Neuquen and northward into

San Juan Provinces, Argentina, and crosses into Chile around

VolcSn Peteroa and Cordillera de Curicd; 1700-3700 m. 89

FIG. 10. Cross section of fruit. A. Mulinum albovaginatum (based on Ruiz L . 7183, UC). B. Mulinum valentini (based on Grondona 2202, UC). 90

FIG. 11. Distribution of Mulinum albovaginatum in Argentina, Provinces Mendoza and Neuqu§n, and Chile, Regidn VII. • = M. albovaginatum var. albovaginatum; ▲ = M, albovaginatum var. pauciflorum. 91

Constance (1988) questioned the application of the name of this taxon because it was based on sterile material.

However, reference to the fruit was made by Hooker (1830) and loose fruit was found within the folder of the type from K.

While the type material is scanty at best, and appears similar to the type material of M. ulicinum var. ulicinum (e.g., leaf size and sulcate blades), the original descriptions and type

localities differ enough to continue the use of this epithet.

Mulinum albovaginatum has been regarded as similar to several other species. Hooker (1830: 329) described M. albovaginatum as "very distinct" with "few and very indifferent specimens" among Gillies' collection. Hooker compared the fruit to M. ulicinum Gillies & Hook, and the branches and leaves to Selinum microphyllum [= M. microphyllum] which he further stated as differing in peduncle length. Reiche (1899) included M. albovaginatum under the category of "Especie problem^tica" and with a question mark compared it to Azorella spinosa (Reiche 1899: 804). A single leaf upon the type is similar to A. spinosa, but A. spinosa's wingless fruit makes it clearly different from Mulinum.

In my opinion, M. albovaginatum is most similar to M. valentini. The two species are virtually identical in leaf morphology (e.g., leaves mm long, mm wide, and sulcate blades) and share a cushion habit. The two differ most dramatically in cushion size (2-10 cm tall, 4-15 cm diameter in M. albovaginatum vs. 1-2 cm tall, 3-33 cm diameter in M. valentini), petal structure and size (reflexed abaxially,

1.25-2 mm long, 0.5-1.25 mm wide in M. albovaginatum vs. planar, 1.25 mm long, 0.75-1 mm wide in M. valentini), and distribution, with M. albovaginatum restricted to the higher elevations of the central Andes and M. valentini concentrated along the southern coast, and less distinctly in flower number

(1-6 in M. albovaginatum vs. 2-12 in M. valentini), involucral bract number (in M. albovaginatum vs. in M. valentini), fruit size (5-7 mm long, 4-5 mm wide in M. albovaginatum vs. 5 mm long, 4-6 mm wide in M. valentini), and carpophore structure

(shortly 2-cleft, rarely entire in M. albovaginatum vs. entire in M. valentini).

Mulinum albovaginatum is vegetatively uniform with the exception of pubescence and a few unusual collections. Kuntze

32 contains a larger plant, with larger internode lengths, while still displaying the typical dense cushion nature.

Bridges s.n. is more lax with elongated internodes. This same elongatedness has been observed on the peripheral edge of more typical cushions, as well as on the holotype and isotypes.

The plants of Salinas 2180 and Semper 11855 are extremely small and have been annotated by Constance as possibly representing a distinct variety. The small stature, however, falls within the range of other collections and the difference is believed to be a result of environmental pressures such as constant wind and temperature, and/or how the plant was originally collected, dried, and mounted. 93 Mulinum albovaginatum is somewhat variable in leaf and

fruit composition and especially variable in leaf pubescence,

both of which have a geographic basis, and therefore two

varieties are recognized.

Key to the Varieties of Mulinum albovaginatum

1. Leaves glabrous; fruit 5 mm long, 4 mm wide.

5a. M. albovaginatum var. albovaginatum.

1. Leaves pubescent; fruit 6.25-7 mm long, 4-5 mm wide.

5b. M. albovaginatum var. pauciflorum.

5a. Mulinum albovaginatum var. albovaginatum.

Mulinum cryptanthum Clos var. pulvinaris Chodat & E.

Wilcz. Bull. Herb. Boissier 2: 525. 1902.— TYPE:

Argentina. "Cajon del Burro, Vallee du Rio Tordillo",

2900 m, Jan-Feb 1897, E. Wilczek 65 (holotype: LAU!;

isotypes: G [2]!).

Leaves glabrous. Fruit 5 mm long, 4 mm wide, at base cordate; wings 5 mm long, 2 mm wide, lateral wings broader than and as long as the fruit body, straight or revolute; oil tubes 0.25-0.75 mm in diameter; carpophore free, entire; odor celery-like.

Phenology: Flowering mid-November through late February; fruits maturing December through early April. Distribution (Fig. 10). In the Andes of Central

Argentina, primarily in Mendoza Province with an extension south into Neuqu&n; 1700-3700 m.

Specimens Examined. Argentina. Mendoza. Lujdn de Cuyo:

Lujdn del Cuyo del Piata, Piedra Grande, Heinziman 21045 (UC),

Quebrada Mulas, Sierra del Plata, King 114 (BM), Nacimiento del Arroyo de Las Cuevas, Ruiz L. 3155 (BM, UC); Malarqiie:

Choique, near Ruta 40 (km 470), Bocher et al. 1546 (S),

Portezuedo del Choique, Boelcke 4217 (UC), 15 kms NE Valle

Hermosa, camino a Los Molles, Boelcke 10322.5 (UC), Calmuco,

Covas 10913 (UC), Portezuelo del Choique, Hunziker 6379 (UC),

Portezuelo del Choique, Ruiz L. 21439 (UC), Alto Valle del

Atuel, Ruiz L. & Roig 15654 (UC), Alto Valle del Atuel, Ruiz

L. & Roig 15763 (UC), 2-4 kms NW of Las Lenas on ski road,

Stuessy et al. 10333 (CONC, GH, LP, OS, UC), 12 km from Las

Lenas on the road to the Valle Hermosa, Zech & Contogiorgakis

5 (CONC, GH, LP, OS, UC) ; San Carlos: Camino a Laguna

Diamonte, Boelcke 4624 (UC), Laguna Diamonte ribera oeste,

Boelcke 10033 (UC), Laguna Diamonte ribera W, Boelcke 10042

(UC), Laguna Diamonte 16 km. E. prox. Vegas Yaucha, Boelcke

10078 (UC), Laguna Diamonte bajada de El Paramillo, Boelcke et al. 10064 (UC), Road to Laguna del Diamante, Cuezzo & Barkley

20MZ383 (UC) , Paso Cruz, Kuntze 32 (NY, US), Pampa de Los

Osamentas (Estancia LLancha), Ruiz L. 7183 (UC), Quebrada del

Paso de La Cruz de Piedra, Ruiz L. 11682 (UC), Quebrada de La 95

Cruz de Piedra, Ruiz L. 14578 (UC), Laguna Diamonte, Zech &

Contogiorgakis 9 (CONC, GH, LP, OS, UC); San Rafael: Sa. del

Nevado, Boelcke 15927 (UC); Tunuy&n: Camino al Paso del

Portillomendocino, Ruiz L. 1983 (BM, UC), Rincones del Cerro

Morado, Ruiz L. 3169 (BM, UC); Tupungato: Estancia Silva, Loma

Larga, Ricardi S. 2180 (BM, UC), Las Tres Quebradas, Ruiz L.

3619 (BM, UC) , Arroyo La Horqueta, Semper 11855 (UC) . Neuquen.

Chos Malal: 25 km a 1 NW de Chos Malal, Ancibor et al. 90214

(UC), Caj6n del A del Cruce, Boelcke 11228 (UC) , 33 km de

Tricao Malal camino a Mina de Azufre, Boelcke 11633 (UC),

Minas, Cordillera del Viento, Boelcke et al. 11610 (UC) ,

Parque Nacional Lanin, Correa 5815 (UC), Huiliches, Parque

Nacional Lanin, Correa 5817 (UC).

Mulinum albovaginatum var. albovaginatum represents the majority of individuals of M. albovaginatum.

5b. Mulinum albovaginatum var. pauciflorum (Reiche) Zech,

comb, et stat. nov. Mulinum pauciflorum Reiche, Anal.

Univ. Chile 104: 803. 1899. Mulinum reichei H. Wolff,

Repert. Spec. Nov. Regni Veg., 17: 441-442. 1921. Mom.

superfl., based on type of M. pauciflorum

Reiche.— TYPE: Chile. Regidn VII. "En terrenos de

arena volcanica [Peteroa] de la cordillera de Curico",

2500 m, Jan, K. Reiche s.n. (holotype: SGO!).

Azorellopsis trisecta H. Wolff, Repert. Nov. Sp. 19: 312. 96

1924.— TYPE: Bolivia, "in regione alpina andina loco

non indicato", Bridges (holotype: W, not located).

Leaves pubescent. Fruit 6.25-7 mm long, 4-5 mm wide, at

base cordate; wings 6.75 mm long, 1.25 mm wide, lateral wings

broader than and longer than the fruit body, straight; oil

tubes small, 0.25 mm in diameter; carpophore free, entire to

shortly 2-cleft; odor absent.

Phenology: Flowering in late December; fruit maturing in

January. A single specimen (Reiche s.n.) is dated 15 October

and contains mature fruit. Flowering would have had to have

occurred in September or before which seems unlikely given the

climate and elevation. Therefore these fruits are believed to

be still attached from the previous year. Because this is

based on a very limited amount of material, the phenology

probably is closer to that of variety albovaginatum.

Distribution: (Fig. 10) . In Chile around Volccin Peteroa

and Cordillera de Curicd, the type locality; 2000-2800 m.

Specimens Examined. Chile. Regidn VI. Colchagua: Bridges

s.n. (BM). Regidn VII. Curicd: Summit of Andes near Volcano of

Peteroa, Bridges 1188 (BM, E, G, K, OXF, W), Cordillera de

Curico, Reiche s.n. (GOET), Cordillera de Volcdn Peteroa,

Werdermann 613 (E, F, G, GH, MO, S, UC, US), Upper valley of the Rio Teno, near the limit, Zollner 5950 (L, UC). 97

The leaves of M. pauciflorum are shortly pubescent on all

surfaces, differing from the glabrous M. albovaginatum. Leaf

pubescence along with a Chilean distribution are the basis for

the recognition of M. albovaginatum var. pauciflorum.

6. Mulinum valentini Speg., Anales Soc. Ci. Argent. 48: 57.

1899.— TYPE: Argentina. Chubut. "In aridis saxosis

prope Trelew", Nov 1898 [1897], G. Valentin 66

(holotype: LP!, photo: F! GH!).

Mulinum lycopodiopsis Speg. Anal. Soc. Ci. Argent. 48: 55-

56. 1899.— TYPE: Argentina. Patagonia, "Pan de Aztacar

secus Rio Chico," Dec 1897, C. Ameghino s.n.

(lectotype here designated: LP, photocopy: OS!;

isolectotype: LP, photocopy: OS!).

Azorella concolor Rendle, J. Bot. 42: 368. 1904.— TYPE:

Argentina. Santa Cruz. Patagonia, Mountain Tops,

Burmeister Peninsula [Lake Argentino], 1900-1901, H.

H. Prichard s.n. (holotype: BM!).

Compact cushions, 1-2 cm tall, 3-33 cm diameter. Stems, erect, dense, 1-4 mm diameter, covered with brown-black decaying leaves and/or leaf skeletons. Leaves sessile or petiolated, imbricate, strongly reflexed, 4-15 mm long, 1-8 mm wide; blade glabrous, strongly to weakly sulcate, 4.5-6 mm long, 1-8 mm wide, with margin entire; lobes 3, rarely 4, 0.5-

4 mm long, 0.1-1.5 mm wide, central lobe reflexed abaxially, generally larger, lateral lobes reflexed adaxially, generally smaller, at apex mucronate to finely acuminate; constriction absent, blade continuous with petiole; petiole to 6 mm long,

0.5-3 mm wide; sheath closed, 3-5 nerved, dilated or width uniform, 2-7 mm long, 1.5-4 mm wide, with margin entire or rarely pubescent, hairs straight, to 2 mm long; stipules absent. Umbel 5-7 mm diameter, 2-12 flowered, spreading, often not extending beyond upper leaves, sessile or peduncled; peduncle to 3 mm long, 1.5 mm wide. Involucre glabrous, involucral bracts 3, rarely 4-5, equal, membranous, 0.5-2.5 mm long, 0.75-1.25 wide, linear-lanceolate, at apex acuminate, at base connate into a cylindrical cup, with basal margin fringed; rays membranous, 2-4 mm long, 0.25-0.75 mm diameter, at apex receptacle enlarged and constricted. Flowers 3-5 mm diameter; sepals as teeth, green, at apex ± 90fi, at base forming a shallowly stellate disc, 0.25 mm long, 0.5 mm wide; petals, yellow, 1.25 mm long, 0.75-1 mm wide, planar, elliptic to obovate, at apex rounded to acute; stamens deciduous or persistant on fruit, with filament 0.75-1 mm long, 0.2 mm wide, at base uniformly wide, with anthers 0.5-0.75 mm long,

0.5 mm wide; carpel entire; stylopodium 1.25 mm diameter, with styles to 0.5 mm long, elongating to 1 mm. Fruit elliptic to widely ovate, yellow, strongly 4-winged in cross-section, body inflated, squarish to rectangular in cross-section, 5 mm long,

4-6 mm wide, at apex acute, at base cordate; wings widely lunate to widely lunate, 5 mm long, 2 mm wide, lateral wings 99

longer and broader than or equal to the fruit body, straight;

mericarps triangular in cross-section, separating easily,

abaxial face concave to angular, commissural face convex, ribs

5, 1 abaxial, prominent and equal; oil tubes round; carpophore

entire; odor weakly celery-like or coconut-like. Chromosome

number: n = 8. Fig. 10.

Phenology: Flowering November through late January;

fruits maturing mid-November through early February.

Distribution (Fig. 12). This species occurs in Southern

Argentina within the provinces of Chubut and Santa Cruz. In

Santa Cruz M. valentini grows along the coast and at higher

elevations near the Andes and is lacking within the central

portion of the province. In Chubut M. valentini is

concentrated primarily along the Andes with a small coastal

extension extending over the border of Santa Cruz. This

species also has a limited distribution in Southern Chile

between Parque Nacional Torres del Paine and Puerto Natales.

A single disjunct collection exists near La Plata, Argentina,

suggesting a more extensive northern distribution in the past;

130-1600 m.

Common Name: "Neneo" (Soriano 2193).

Specimens Examined. Argentina. Buenos Aires. Buenos

Aires: Chascomds, in der Nahe der Estancia Vitel [Lauquen],

Berg 67 (LP). Chubut. Cushamen: top of Mt. Quichaura, 100 km

S of Esquel, Beetle & Soriano 509 (UC). Escalante: Pampa del 100

'Santa Cru

FIG. 12. Distribution of Mulinum valentini in Argentina and Chile, Regidn XII. 101

Castillo, Castellanos 6117, 6119 (UC), 30 km al NW de Comodoro

Rivadavia, Campo Diadema Norte, Ruiz L. 14784 (UC), Estancia

Oriental, Soriano 2103 (UC). 85 km E of Sarmiento on Ruta 26,

Zech & Contogiorgakis 24 (CONC, GH, LP, OS, UC) 25 (OS);

Lanquineo: Tecka, Estancia "La Carlota", Grondona 2202 (UC);

Rio Senguerr: Valle de La Laguna Blanca,

Koslowsky s.n. (BM, UC), Valle del Lago Blanco, Koslowsky 1314

(UC); Sarmiento: Bosque petrif icado de Maria Santisima a 60 km

sw de Sarmiento, Correa & Constance 4090/3839 (UC);

Tehuelches: Estancia El Cherque 30 km E de Rio Pico, Soriano

2160 (UC), Arroyo Temenhuau, Soriano 2193 (UC) , Entre Rio Pico

and Lago Vintter, Soriano 3072 (UC). Santa Cruz. Deseado:

Colonia Las Heras, Ancibor 119 (UC), Tehuelches, Donat 60, 61

(GH, GOET, UC), 60 km S of Puerto Deseado, road to San Julian,

Eyerdam et al. 23876 (G, MO, UC); Giier Aike: Rio Gallegos,

Estancia La Carlota, Arroyo et al. 358 (UC) , Estancia La

Verdadera Argentina Co de la Virgen, en ladera, parte

superior, Arroyo et al. 2252 (UC), Estancia Las Viscachas,

Lomada a ± 2 km de Laguna Viscachas, Arroyo et al. 2384 (UC),

Estancia Las Viscachas, Co Las Viscachas, laderas inferiores,

Arroyo et al. 2437 (UC), Estancia Las Viscachas, cono volc£nica Pan de Azticar, Arroyo et al. 2566 (UC), Estancia Las

Viscachas, Laguna Viscachas, ladera SSE de la Laguna, Arroyo et al. 2610 (CONC, UC), Estancia Las Viscachas, Co Sin Nombre, ladera N, Arroyo et al. 2650 (UC) , Estancia La Angelina, Faggi

& Lopez 4416 (UC), Meseta De Las Vizcachas, Estancia de Las Vizcachas, Ruiz L. 27020 (UC), Rio Gallegos, cerro N, Sleumer

842 (G, US), Rio Gallegos, cerro Tres Hermanos, Sleumer 927

(G, US), 7 km S of Hotel Lemarchand on Ruta 3, Zech &

Contogiorgakis 18 (CONC, GH, LP, OS, UC) ; Lago Argentino: Lago

San Martin, Dusen 5931 (F), Cerro El Calafate, Lago Argentino,

Sleumer 1178 (G, S, US), Margen de la Meseta [del Quemado] arroba de la Perez, Rio de las Vueltes, Sleumer 1358 (G, S,

US). Lago Buenos Aires: 20 km de Bajo Caracoles to Perito

Moreno, Boelcke 12912 (UC), Lago Posadas, Ruiz L. 24225 (UC),

10 km de Estancia La Vizcaina, Ruiz L. 26423 (UC) , En Puerto

Lebrum, Ruiz L. 26434 (UC) , Laguna El Sello, Ruiz L. 26451

(UC), Fenix-dalen, Rio Fenix, N Lago Buenos Aires, Skottsberg

619 (LY, S, UC, UPS) , 88 km de Perito Moreno, Soriano 4795

(UC), Perito Moreno, Vidal 24 (LP). Chile. Regidn XII. Ultima

Esperanza: Sierra de Los Baguales, cerro Santa Lucia, Arroyo et al. 841002 (CONC), Cerro Donoso, Sector Rio de las Chinas,

Arroyo et al. 870276 (CONC), Sierra Contreras, Eruperaire

1395, 1455 (B), Sierra de Los Baguales, Estancia La Cumbre,

Frontera Chile-Argentina, Paso Baguales IV, Caravana Estancia

La Cumbre, Landero 643 (CONC), Sierra de Los Baguales,

Estancia La Cumbre, Campo de La Tropilla, Landero 699 (CONC),

Estancia "Cerro Castillo" cumbre de la Sierra del Cazador,

Magens 918 (B, CONC, UC).

Mulinum valentini is similar to M. albovaginatum. As discussed earlier, M. valentini differs most in cushion size, 103 petal structure and size, and distribution, and to a lesser extent in flower and involucral bract number, fruit size, and carpophore structure.

Although most plants are uniformly small-leaved, compact cushions, there does exist some variation. Cushions exist with elongated, almost viney, peripheral stems and/or larger leaves (e.g., Anclbor 119, Donat 61, Euperaire 1455, Faggi &

Ldpez 4416). This elongatedness may be a result of relaxed environmental pressures, which are believed responsible for maintaining cushion compactness (see Chapter 2).

7. Mulinum hallei Skottsb., Kgl. Sv. Vet. Akad. Handl. 56(5):

278-279. 1916. PI. 20, fig. 6,7.— TYPE: Argentina.

Santa Cruz. "Patagonische Kiiste: Puerto Mazaredo", 17

Dec 1904, P. Dus&n 5304 (lectotype, here designated:

UPS!; isolectotypes: K! S!).

Perennial forming small cushions, 2-8 cm tall, 2.5-12 cm diameter. Stems erect rarely lax, compact rarely spreading,

1-2 mm diameter, covered with brown decaying leaf bases.

Leaves petiolated, rarely sessile, 2-15 mm long, 1.2-5 mm wide; blade shape, ± inflated to strongly sulcate, glabrous; lobes 3, rarely 2, 4-6, narrow to slightly obovate, 5-11 mm long, 0.5-1 mm wide, ± equal or center lobe larger, at apex rounded to mucronate, rarely pubescent with 1 white, simple, apical hair, to 0.5 mm long; constricted at petiole; petiole base dilated; sheath open, membranous, 3-nerved, 2-4 mm wide, with margin ciliate to frimbriate, sparse or dense, up to 20 hairs per margin, hairs straight, to 5 mm long; stipules present as hairs or absent; petiole 2-5 mm long, 0.5-1 mm wide. Umbel 4-12 mm diameter, (1)5-14 flowered, spreading, extending beyond upper leaves; peduncle (1)8-30 mm long, 0.75-

0.8 mm diameter, 4-5 angled, at apex receptacle enlarged and constricted. Involucre lanate; involucral bracts 3-8, equal, green to membranous, 2-3.5 mm long, 0.5-1 mm wide, linear- lanceolate, at apex entire or 2-3 lobed, lobes shortly bifurcated, to 1.25 mm long, at base ± free or connate, into a shallow cup, to 1 mm long, with margin entire, shortly ciliate, and/or frimbriate, hairs simple, white, curly, to

1.25 mm long; rays yellow, 1.5-3 mm long, 0.3 mm diameter, ribbed, relaxed, but erect with fruit, at apex receptacle weakly constricted. Flowers 4 mm diameter; sepals as teeth, yellow-green with median rib brown, 0.5-1 mm long, 0.75 mm wide, at apex acute to narrowly acute, at base forming a stellate disc; petals yellow or yellow-red, reflexed and curled abaxially, 1.5 mm long, 1 mm wide, ovate to obovate, at apex acute to mucronate; stamens weakly deciduous, with filament 0.5-1.25 mm long, 0.1 mm wide, at base flared, with anthers 0.5 mm long, 0.5 mm wide; carpel entire; stylopodium mm diameter, with styles to 0.5 mm, elongating to 1 mm. Fruit widely elliptic, yellow, 2-winged to slightly 4-winged in cross-section, body deflated, rectangular in cross-section, 105

5.5-6 mm long, 5 mm; wings lunate, 6 mm long, 1-1.25 mm wide,

lateral wings as long and narrower than the fruit body,

slightly revolute; mericarps lunate in cross-section,

separating easily, abaxial face slightly concave, commissural

face slightly convex, ribs 7, 3 abaxial, prominent and equal;

oil tubes irregular; carpophore shortly 2-cleft; odor absent.

Chromosome number: n = 8. Fig. 13.

Phenology: Flowering late November through February

[March]; fruit maturing middle to late November through March.

Distribution (Fig. 14). Endemic within southern

Argentina, primarily in Chubut and Santa Cruz Provinces with

sparse extensions north into Neuquen and Rio Negro Provinces;

60-1200 m.

Specimens Examined. Argentina. Chubut. Escalante: Golfo

San Jorge, Ameghino s.n. (LP), Manantiales Behr, Castellanos

s.n. (UC), 35 km cruce Ruta 3, camino Pampa del Castillo, near

Comodoro Rivadavia, Correa & Constance 3836 (UC), 35 km N of

Comodoro Rivadavia, Canaddn Ferrais, Correa & Constance

3985/3829 (G, GH, MO, UC) , 35 km W of Cruce de Rutas 3 & 277,

Correa et al. 4063 (UC), Pampa del Castillo, Ferruglio s.n.

(UC), Cafiaddn Ferrais, Ferruglio s.n. (UC) ; Languifieo:

Quichaura, Tecka, Soriano 2257 (UC), Tecka, Soriano 2426 (UC) ;

Rio Senguerr: Canaddn de las Manos Pintadas al SE de Facundo,

Eskuche 1868-7f 1921-11 (UC), Valle del Lago Blanco, Koslowsky

33 (UC); Sarmiento: N-NW Lago Musters, Feruglio s.n. (UC) . 106

A.

B.

FIG. 13. Cross section of fruit. A. Mulinum hallei (based on O'Donell 3663, UC) . B. Mulinum microphyllum (based on Correa & Constance 4085/3838, UC). 107

<, Rpo Negro Chubut

FIG. 14. Distribution of Mulinum hallei in Argentina. Neuquen. PicCin Leufti: PicCin Leufti, Ruiz L. 23988 (UC) ; Zapala:

Entre Zapala and Laguna Blanca, Ruiz L. 26772 (UC). Rio Negro.

Norquincd: Cango, W Rio Chico, Hager 1003 (B) . Santa Cruz.

Corpen Aike: Rio Santa Cruz, Burmeister s.n. (UC), Comandante

Luis Piedrdbuena, O fDonell 3799 (UC), 160 km from Rio

Gallegos, Ruiz L. 26905 (UC), Santa Cruz, Santesson 230, 225

(K(225), S), Colonia Las Heras, Colonia Pastoril Presidente

Carlos Pellegrini, Estancia La Flora, von Thiingen 14, 72, 97,

103 (LP) , 155 km N of Rio Gallegos, Ruta 3 to San JuliSn, Zech

& Contogiorgakis 21 (CONC, GH, LP, OS, UC); Deseado: 6 km N of

Puerto Deseado, Correa & Constance 4011/3832 (G, UC), Puerto

Deseado, 3 km de Cabo Blance a Tellier, Correa & Nicora 3335

(UC), Bahia Uruguay, 30 km W of Puerto Deseado, Correa et al.

2586 (UC), Camino Deseado, Cabo Blanco, Correa et al. 2598

(UC), Puerto Mazanedo in campo, Camino Tellier - Paso Gob.

Gregores, Ruta 282, Correa et al. 2651 (UC) , Tehuelches, Donat

60, 61 (BM, G, GH, F, GOET, MO, NY, S(61), UC(60)), Camino

Deseado, Cabo Blanco, Dus&n 5304 (S), 15-20 km from Estancia

La Aurora, Ruiz L. 24082 (UC); Lago Buenos Aires: 55 km S Rio

Mayo, Boelcke et al. 16082 (UC), Entre Arroyo Teka & Perito

Moreno Ruiz L. 24218 (UC), Meseta del Lago Buenos Aires, a 10 km de Estancia La Vizcaena a Puesto Lebrum, Ruiz L. 26425

(UC) , Lago Ghlo, Skottsberg 857 (UPS); Magallanes: Puerto San

Julian, Blake 45 (BM, K), San Julian, Blake 160, 167 (K,

M A (160)), Tres Cerros, Estancia La Lomita pie del co. i 3 km ruta 3, Boelcke et al. 15368 (CONC, UC) , 50 km N of San 109

Julian, road to Puerto Deseado, Eyerdam et al. 23963 (GH, UC),

San Julidn, O'Donell 3663 (UC) ; Rio Chico: 10 km de Las

Horguetas hacia Bajo Caracoles, Boelcke et al. 12809 (UC),

Tres Cerritos & Lago Cardiel, Ruiz L. 24109 (UC), Canadon

Le6n, Soriano 5076, 5098, 5099 (UC), Estancia Rio Lista, von

Platen & Greiner 153 (BM).

Skottsberg (1916) described this species based upon

Dusen's collection at Puerto Mazaredo, Departamento Deseado,

and his own collections from the Rio Fenix Valley and at the

east end of Lago Buenos Aires, both in Departamento Lago

Buenos Aires and both Skottsberg 855. Skottsberg includes the

locations for his collections within the protologue of M.

hallei, but does not refer to either specimen specifically by

collector or number (Skottsberg 855). He does include the

location and specific collection number for Dusen's material.

Because it is unclear which of the two specimens represent the

holotype, Dusen 5304 is here designated the lectotype based

upon its explicit reference within Skottsberg's protologue

(1916).

Mulinum hallei is distinct from all other Mulinum

species, with the exection of M. microphyllum, based upon

copious lanate hairs, especially within the involucre, peduncle length (to 30 mm long), lobed involucral bracts, and

its small size (2-8 cm tall, 2.5-12 cm diameter). However,

M. hallei is extremely similar morphologically to M. 110

microphyllum. The two species differ by terminal simple hairs

in M. microphyllum, which occur at the apex of the leaf lobes

versus entire or shortly mucronate apices in M. hallei.

Although they are partially sympatric in distribution, M. hallei has a broader distribution along the southern coast of

Argentina in the provinces of Chubut and Santa Cruz. The

species do differ in chromosome number. Mulinum hallei is n =

8 whereas M. microphyllum is n = 16 and is believed to be

involved in the origin of the polyploid M. microphyllum.

8. Mulinum microphyllum (Cav.) Pers., Syn. pi. 1. 309. 1805.

Selinum microphyllum Cav., Icon. PI. 5: 59, t. 486, f.

2. 1799. Mulinum microphylli (Cav.) Pers. in

Spegazzini, Anales Mus. Nac. Hist. Nat. Buenos Aires.

7(Ser. 2a, t. 4): 295. 1902. Orthogr. var.— TYPE:

Argentina. Santa Cruz. Puerto Deseado, L. N&e s.n.

(holotype: MA, photo: OS!).

Mulinum morenonis (Kuntze) Speg., Anales Mus. Nac. Hist.

Nat. Buenos Aires, 7(Ser. 2a, t. 4): 295. 1902.

Huanaca morenonis Kuntze, Rev. gen. pi. 3(2): 113.

1898. Diplaspis morenonis (Kuntze) K. Schum. Just's

Bot. Jahresber. 26(2): 367. 1900. Azorella morenonis

(Kuntze) Macloskie, Rep. Princeton Univ. Exp.

Patagonia, Botany 8: 629. 1905.— TYPE: Argentina.

"Patagonia" 50/3°, April 1882, E. P. Moreno & Torrini

368 (holotype: NY!). Perennials forming small compact to lax cushions, 1.4-9

cm tall, 1-15 cm diameter; branches erect, spreading, or

trailing. Stems, erect to lax, dense to spreading, 2-3 mm

diameter, covered with brown decaying leaf bases. Leaves 4-30

mm long, 1.5-13 mm wide; blade shape, inflated, sulcate, or

flat, glabrous or abaxially and/or adaxially pubescent; lobes

3-8, variable, sinus length equal or unequal, 1-10 mm long,

0.5-1.25 mm wide, longly obovate, elliptic, or linear-

lanceolate, at apex round or acute with single, simple, white

hair, 3 mm long or less; lobes constricted at petiole or

constriction absent, constriction glabrous or pubescent,

sparse to dense, hairs white, simple, curled; petioles 1-7 mm

long, 0.5 mm wide, triangular in cross-section, base dilated;

sheath, thick, membranous, open, 3-nerved, 2-7 mm long, 1.75-3 mm wide, with margin ciliate to frimbriate, up to 20 hairs per margin, hairs straight, 5 mm long or less; stipules 3-4 mm

long, 0.75 mm wide, at apex acute, with margin fringed, hairs mm long, or stipules reduced to hairs, 1-5 mm long. Umbel rarely with 2 simple umbels along a single axis, 6-15 mm diameter, 3-33 flowered, ± globose, extending beyond the upper

leaves; peduncle (0.1-0.3)1-7.5 cm long, 0.5 mm wide, 3 to 4 to 6 angled, glabrous to lanate, at apex enlarged and constricted. Involucre lanate; involucral bracts 4-7, equal, green, thick, 2-3 mm long, 0.5-1 mm wide, linear to lanceolate, at apex 2-lobed, rounded, or acute, with single, simple, white, apical hair, 0.5 mm long or less, at base connate, into a shallow cup, with margin entire, shortly

ciliate, and/or frimbriate, hairs simple, white, curly, to

1.25 mm long; rays 2-5 mm long, 0.5 mm wide, yellow; receptacle enlarged and constricted. Flowers 4.5 mm diameter;

sepals as teeth, yellow-green with median rib brown, at apex

acute to mucronate, at base forming a conical cup, 0.5 mm

long, 0.5 mm wide; petals yellow or yellow-red, 1.5-2 mm long,

1-1.5 mm wide, planar, ovate to narrowly ovate, at apex acute

to rounded; stamens weakly deciduous, with filament 1.25 mm

long, 0.1 mm wide, at base uniformly wide, with anthers 0.5 mm

long, 0.5 mm wide; carpels grooved, easily separating in the

flower; stylopodium with styles to 0.5 mm, elongating to 1 mm.

Fruit elliptic, 4-winged in cross-section, body deflated, rectangular in cross-section, 6-7 mm long, 4-5 mm wide; wings lunate, 6-7 mm long, 1-2 mm wide, lateral wings longer and broader than the fruit body, slightly revolute; mericarps lunate to rectangular in cross-section, separating easily, abaxial face concave, commissural face slightly concave to planar to squared, ribs 5, 1 abaxial, equal and filiform; oil tubes round to oblong; carpophore shortly 2-cleft; odor weak.

Chromosome number: n = 16. Fig. 13.

Phenology: Flowering late October through mid-March; fruit maturing late November through early March.

Distribution (Fig. 15). Endemic to southern Argentina, mainly at lower elevations of the eastern cordillera within

Neuquen, Rio Negro, Chubut, and Santa Cruz Provinces and Rio Negro Chubut

FIG. 15. Distribution of Mulinum microphyllum in Argentina. 114

sparsely along the coast of Chubut and Santa Cruz; 770-1500 m.

Specimens Examined. Argentina. Chubut. Cushamen: Epuyen,

Perez M. 18663 (UC) ; Escalante: 35 km al N de Comodoro

Rivadavia, Pampa de Salamanca, directly across highway from

Pico Salamanca, Correa & Constance 3997/3831 (UC), 10 km al W de Hermitte, Correa et al. 4085 (UC), Comodoro Rivadavia,

O'Donell 3363 (GH, NY, S), Estancia Begona, 30 km al NW de

Comodoro Rivadavia, Soriano 2025 (UC); Florentino Ameghino:

Camarones, Aurelius 16 (S), Lochiel, 30 km al W de Camarones,

Soriano 1986 (UC); Futaleuffi: Corcovado, Illin s.n. (LP),

Futalaufguen, Lahitte s.n. (UC), Futaleufd, Skottsberg 856 (S,

UPS), Cerro Cuche, Estancia Pampa Chica, Soriano 2508 (UC) ;

Gaiman: Ruta 3, 21 km al S de Garayalde, Correa & Constance

3970/3826 (G, UC); Languifieo: Rio Corcovado, Illin 257 (UC) ,

Cerro de la Zeta, Kiihnemann 471 (UC), Esguel, campo de aterrizaje, Kiihnemann 552 (UC), Esquel, Soriano 3788 (UC); Rio

Senguerr: Lago Blanco, Koslowsky s.n. (UC); Sarmiento: 10 km

W de Hermitte a Sarmiento, Correa & Constance 4085/3838 (G,

UC), 50 kms antes de llegar a Sarmiento, Ricardi S. & Matthei

282 (CONC); Tehuelches: a 30 km de Gobernador Costa hacia

Tecka, Ruta 40, Correa & Constance 4152/3843 (UC). Neuquen.

Alumine: Puesto Piedra, camino al Paso Tromen, Eskuche 0926

(UC) ; Ldcar: San Martin de los Andes, Bridarolli 2063 (LP) ,

San Martin de los Andes, Cabrera 11210 (LP, UC) , Piedra

Pintada, Castagnet 127 (LP), Vega Lolog, Comber 823 (K), San Martin de los Andes, cerros al N de la cindas, Dawson 1271

(CONC, UC) , Circa Lago San Martin, Uogberg 142 (BM) , San

Martin de Los Andes en laderas de los cerros y claros del bosgue de Nothofagus, Ruiz L. 20244 (UC); Los Lagos: Sobre costa del Lago Huemul, cerca del arroyo Quintinquenco, Boelcke

& Hunziker 3484 (UC), Island near Correntoso, Edwards s.n.

(BM), Estancia Fortin Chacabuco, Hager 993 (B) , Rio Huemul delta, Ljungner 756 (GOET), Lake Nahuelhuapi, Rio Huemul in direct part of the delta, Ljungner 773 (NY), Santa Maria,

Ljungner 794 (GOET), Lago Nahuel Huapi, Huemul N side,

Ljungner 799 (GOET). Rio Negro. Bariloche: Front range of the

Andes, SW of Bariloche on road to Cerro Otto, Arnow 3780 (UC),

Cerro Otto, Boelcke 1728, 1760 (UC), Pargue Nacional Nahuel

Huapi, Estancia Fortin Chacabuco, Boelcke & Hunziker 3512

(UC), CK THESE cerro Gutierrez, Buchtien 42 (US), Lago

Nahuelhupi, cerro Gutierrez, Buchtien 1350 (E, GH, LY, S),

Cerro Leones, Burkart 6232 (F, US), Cerro Otto, Burkart 6642

(UC), Lago Nahuel Huapi, Cabrera & Job 39 (LP, NY), Puerto

Sabana, Cordini 143, 223 (US), Lago Naheul Huapi, S side of the Huemul Bay, Ljungner 509 (BM), Cercana al Lago Nahuel

Huapi, Parodi 11859 (GH), Los Repollos por el camino a Piedra

Pintadas, P&rez M. 45363 (UC), Cerro Otto, P6rez M. 48694,

(UC), Colania Suiza, Perez M. 48695 (UC), Cerro Otto, Perez M.

49142, (UC), Cerro Otto, Soriano 1320 (LP), Pargue Nacional

Nahuel Huapi, Cerro Otto, Teague s.n. (K), Cerro del Lion, E end of Lago Nahuel Huapi, West 4757 (GH, UC) ; 9 de Julio: 116

Meseta de Somuncura, Laguna Paraguay, Ruiz L. 25591 (UC);

Norquincd: Cango W Rio Chico, Hager 1002 (B) ; Pilcaniyeu:

Comallo, Cabrera 4283 or 4823 CK THIS (LP, UC) , Ruta 23 a 5 km del cruce de ruta 237 (5 km W of Estancia Perito Merino,

Correa & Constance 3817/3820 (UC), Los Juncos, Perez M. 48689

(UC). Santa Cruz. Deseado: Puerto Mazaredo in campo, Dusen

5301 (K), Puerto Deseado, O'Donell 3560 (UC), Puerto Deseado,

Rodriguez s.n. (F).

Mulinum microphyllum is morphologically similar to M. hallei. Several morphological characters are amplified within

M. microphyllum. Mulinum microphyllum has thicker, more leaf-

like involucral bracts, thicker leaf sheaths, more leaf lobes

(to 8), an increase in leaf lobe variation (e.g., more secondary lobes), larger leaves (to 30 mm long, 13 mm wide), and an increase in flower number per umbel (to 33) as compared with M. hallei (involucral bracts membranous; leaf sheaths thin; leaf lobes generally 3; leaves to 15 mm long, 5 mm wide; to 14 flowers per umbel). A single specimen (Burkart 6232) was observed with 2 simple umbels on a single axis, and in general the carpels are grooved and/or separate easily in the flower. Most striking is a dramatic increase in overall pubescence. The peduncles and leaf blades of M. microphyllum are often covered with dense curly white hairs and the apices of the involucral bracts and especially the leaf lobes are tipped with single simple white hairs up to 3 mm long. Apical 117

leaf lobe hairs and increased production of these hairs is the most constant character amplified within M. microphyllum and the character which best delimits this species. While M. hallei is also pubescent, the amount and consistency of pubescence among leaves is considerably less than that found

in M. microphyllum. The two species are partially sympatric

in distribution, with M. microphyllum more abundant along the eastern cordillera and extending more northernly from Neuquen to Santa Cruz.

The preceeding evidence coupled with a doubled chromosome number suggests that M. microphyllum is a polyploid derivative from M. hallei. Whether M. microphyllum represents an autopolyploid or an allopolyploid is unclear, and further study is encouraged. Examples of allopolyploids resembling one parent or the other exist (Clausen et al. 1945; Grant

1981), but close morphological resemblance alone between a polyploid and diploid is no longer considered a valid criterion of autopolyploidy (Grant 1981). Mulinum valentini is partially sympatric with both M . hallei and M. microphyllum and has a chromosome number of n = 8. Mulinum valentini is similar to M. microphyllum morphologically in fruit shape and, based upon a limited sample, the number of flavonoids synthesized (see Chapter 3) . The chromosome figures of M. microphyllum, M. hallei, and M. valentini (Constance et al.

1971, their Figs. 323, 322, 325, respectively) seem to support

M. microphyllum being an allopolyploid. The chromosomes of M. 118 hallei appear larger than the chromosomes of M. valentini.

Both large and small chromosomes are apparent in M. microphyllum. While the designation of some of M. microphyllum's chromosomes as large or small is debatable and chromosomes were counted at different stages, diakinesis (M. hallei), metaphase I (M . microphyllum), and metaphase II (M. valentini), the mixture of chromosome size may support an allopolyploid origin.

While most of Donat 60 & 61 is M. hallei, some sheets are mixtures of M. hallei and M. valentini [Donat 60 (GH, GOET),

Donat 61 (GH) ]. This may suggest that M . microphyllum orignated at the location of Donat 60 & 61, near Tehuelches,

Chubut Province, Argentina. Further support is added by the fairly extensive occurance of M. microphyllum in Chubut, including Tehuelches (Correa & Constance 4152/3843) .

EXCLUDED AND DOUBTFUL NAMES

Mulinum acaule (Cav.) Pers., Syn. pi. 1. 309. 1805. Selinum

acaule Cav., Icon. PI. 5: 59-60, t. 487, f. 2.

1799.— TYPE: Argentina. Santa Cruz, "portu Deseado", ±

1789-94, L. Nee s.n. (holotype: MA, photo: OS!). =

Laretia acaulis (Cav.) Gillies & Hook., In Hook. Bot.

Misc. 1. 329, t. 65. 1830. This taxon represents one of

several of Cavanilles' species of Selinum which Persoon

included in his original concept of Mulinum. The type 119

photograph clearly shows the whorls of entire, spatulate

leaves and cushion habit of Laretia and supports Gillies

and Hooker's (1830) placement.

Mulinum ? angulatum (K. Presl) DC., Prod. 4: 80. 1830. Bolax

angulatus K. Presl in DC., Prodr. 4: 80. 1830.— TYPE:

Chile. Cordillera de Chile, T. P. X. Haenke s.n.

(holotype: PR!; isotype: PRC!) = Pozoa coriacea Lagasca,

Gen. & Sp. Nov. 13. 1816. Pozoa has a long history of

being associated with a number of widely different genera

(Mathias & Constance 1962), Mulinum being one of them.

However, Pozoa*s conspicuously united involucre,

depressed habit, and terete, unwinged fruit makes it

quite distinctive from any species of Mulinum. Candolle

(1830) displayed doubts about this species by including

a question mark between the genus and specific epithet,

and rightly so. Mathias and Constance (1962) correctly

placed this species in synonymy within their

comprehensive treatment of Pozoa.

Mulinum clandestinum Philippi, Anal. Univ. Chile, 85: 708-709.

1894.— TYPE: Chile, "in montibus Araucaniae Nahuelbuta",

Jan 1877, R. Philippi 2014 (holotype: SGO!, photo: F! CK

IF EQUAL) = Azorella spinosa (Ruiz Lopez & Pav6n) Pers.

Syn. pi. 1. 303. 1805. This is one of two species of

Mulinum which are clearly Azorella spinosa. Philippi 120

(1894) included M. clandestinum within his list of new

plants for Chile ("Plantas Nuevas Chilenas") without any

referral to A. spinosa which Persoon (1805) had

transferred earlier from Fragosa Ruiz Lopez & Pav6n.

Examination of the type material of M. clandestinum

clearly places it within Azorella based on wingless,

terete fruit.

Mulinum cryptanthum Clos, In C. Gay, FI. Chil. 3: 90-91.

1847.— TYPE: Chile, "las cordilleras de Ovalle en la

provincia de Coquimbo", 11,000 ft, C. Gay 515 (holotype:

P, photo: F!; isotype: P!). = Azorella cryptantha (Clos)

Reiche, Bot. jahrb. 28: 16. 1899. This species was

recently included within Martinez's (1989) treatment of

Argentine Azorella. Martinez follows Reiche's (1899)

original treatment of this species as A. cryptantha.

Because of its superficial resemblance to Mulinum, it is

understandable that Clos originally associated this

species with the genus. However, upon closer

examination, the wingless, terete fruit, and entire or

more rarely trilobed leaves are better associated with

Azorella, supporting Reiche's transfer, original

synonymy, and Martinez's current treatment.

Mulinum cuneatum Hook. & Arn., Bot. Beech. Voy. 26.

1841.--TYPE: Chile. "Valparaiso", [1825-1830, fide 121

Lanjouw & Stafleu, 1954], A. Cruckshanks 186 (holotype:

K!). = Azorella spinosa (Ruiz Lopez & Pavon) Pers. Syn.

pi. 1. 303. 1805. This is the second species of Mulinum

which is definitely Azorella spinosa. In Hooker and

Arnott's description of M . cuneatum, they indicated some

doubt by questioning that this species may or may not be

either Ruiz Lopez and Pavdn's Fragosa spinosa or

Persoon's A. spinosa, and referred it to Mulinum on

account of its habit. Their recognition of M. cuneatum

as a new species was somewhat justified because these

collections represented the first time plants had been

collected within Chile. Examination of the type of M.

cuneatum confirms the doubts of Hooker and Arnott; it is

clearly A. spinosa.

Mulinum dipterygia (K. Presl in DC.) DC., Prod. 4: 80. 1830.

Diptergia capitata K. Presl in DC., Prodr. 4: 80.

1830.— TYPE: Chile. "In Andibus Chilensibus", T. P. X.

Haenke s.n. (holotype: PR!; isotype: PRC!). = Gymnophyton

polycephalon (Gillies & Hook.) Clos in Gay, FI. Chil. 3:

103. 1847. The virtually leafless stems and 10-25-

flowered umbels of the type of M. dipterygia clearly

places it in G. polycephalon, agreeing with Mathias and

Constance (1962).

Mulinum famatinense Wolff, Fedde Repert. Spec. Nov. Regni Veg. 17: 441-442. 1921.— TYPE: Argentina. Sierra Famatina,

Cueva de Perez, Jan 1879, G. Hieronymus & G. Niederlein

364 (holotype: B, presumably destroyed). = Azorella.

This species is the least well known of Mulinum taxa, and

there is at present insufficient material on which to

base a final decision. In addition, the holotype was

presumably destroyed, with no isotypes or material

annotated by Wolff located. The collection Hieronymus &

Niederlein 362 however, has been seen, and comes from the

same locality. The abaxially reflected leaf lobes refers

this specimen to M. ulicinum. The fruit, however, is not

that of Mulinum. While some fruit appears dorsally

compressed, others are terete, and all the fruit examined

lacks prominent wings, which are common in Mulinum. In

addition, the elevation at which this taxon was

collected, over 4000 m, is beyond where Mulinum usually

occurs. Until further material can be obtained for study

and/or type material located, M. famatinense is

tentatively excluded to Azorella.

Mulinum integrifolium Hieron., Bol. Acad. Nac. Ci. 4: 28-29.

1881.— TYPE: Argentina, "en las cercanias de Las Cuevas,

Jan 1876, D. S. Echegaray s.n. (holotype: CORD). =

Azorella cryptantha (Clos) Reiche, Bot. Jahrb. 28: 17.

1899. No reference to Clos' (1847) M. cryptanthum was

made by Hieronymus when he described M. integrifolium nor 123

did Reiche (1899) place it into synonymy with M.

cryptanthum when he transferred M. cryptanthum into

Azorella. However, Martinez (1989) has included both

Mulinum species as synonyms of A. cryptantha in her

treatment of Argentinean Azorella.

Mulinum isatidicarpum K. Presl ex DC., Prodr. 4: 80. 1830,

"isaticarpum". Dipterygia isatidicarpa K. Presl in DC.,

Prodr. 4: 80. 1830.— TYPE: Chile. "In Andibus

Chi lens ibus", T. P. X. Haenke s.n. (holotype: PR!;

isotype: PRC!). = Gymnophyton isatidicarpum (K. Presl ex

DC.) Math. & Const., Unv. Calif. Publ. Bot. 33(2):

145-147, pi. 144. 1962. Mathias and Constance (1962)

referred M. isatidicarpum to Gymnophyton, and I agree

with this placement. Although the fruit characters are

comparable to Mulinum, this species is vegetatively

similar to and possesses a median dorsal petal gland

characteristic of Gymnophyton.

Mulinum saniculaefolium (Lam.) Spreng., PI. umbell. prodr. 15.

1813. Hydrocotyle saniculaefolium Lam., Encycl. 3: 154.

1789.— TYPE: Uruguay. "Monte-Video", [1766-1769, fide

Lanjouw & Stafleu, 1954], P. Commerson s.n. (holotype: P;

isotype: P!). = Diposis saniculaefolia (Lam.) DC., Coll.

Mdm. 5: 33. 1829. Originally .described by Lamarck (1789)

under the genus Hydrocotyle, Sprengel transferred the 124

species to Mulinum in 1813. Sprengel may have based his

decision on Lamarck's comment on the isotype label,

"Confer Selinum", Selinum being the genus from which

Mulinum emerged. Examination of the type material

clearly excludes this species from Mulinum. The broad,

three-parted, basal whorls of leaves and generally

leafless scape terminated by a compound umbel, suggest a

greater affinity to the genus Diposis and support

Candolle's 1829 transfer. Candolle (1829) does

acknowledge a similarity between the fruit of Mulinum and

Diposis but adds that the two genera generally can be

distinguished by a single inflorescence in the latter.

Mulinum spinosum var. laciniis ciliatis Hieron. in Lorentz,

Exped. Rio Negro, Bot. 222. 1881, nomen nudum. Lorentz

(1881) first published this name by including it in his

discussion of M. leoninum. Lorentz (1881) states that

based upon his [Lorentz] diagnosis, Hieronymus was of the

opinion that M. leoninum was the same as his [Hieronymus]

M. spinosum var. laciniis ciliatis [written with two

epithets and clearly referred to by Lorentz (1881) as

variety]. Mulinum spinosum var. laciniis ciliatis was

not published in Hieronymus (1881) and it is beleived

that the variety has never been published by Hieronymus

or formly described. 125

Mulinum Section Dipterygia DC., Prodr. 4: 80. 1830. =

Gymnophyton. Candolle (1830) included section Dipterygia

in Mulinum. The section, along with both of its species

(M. dipterygia and M. isatidicarpum), has since been

combined with Gymnophyton (Mathias & Constnace 1962).

ACKNOWLEDGMENTS

I would like to thank Dr. Tod F. Stuessy for his help and direction with this research. I thank Drs. Daniel J. Crawford and Fred Sack for their advice and support. I thank Dr.

Lincoln Constance for his continued interest and encouragement of this research and Dr. Thomas Duncan and the University of

California, Berkeley Herbarium staff for their kindness and assistance.

Because my field collecting would have been impossible without the support and direction of several individuals, I thank: Dr. Jorge V. Crisci at the Museo de La Plata,

Argentina; Dr. Oscar O. Parra and Professor Clodomiro

Marticorena at the Universidad de Concepcidn, Chile; Maria

Marta Cigliano, Claudio Contogiorgakis, Mario Gentilli,

Patricia Gentilli, Liliana Katinas, Alegandra Moschetti,

Alberto Ripalta, Fidel Alegandro Roig, Tod F. Stuessy, Ana

Marid Zavattieri, and especially Juan Jose Morrone for help while in the field; the directors, curators, and staff of the following herbaria: B, BM, CM, CONC, CORD, E, F, G, GH, GOET, 126

K, L, LAU, LP, LY, MA, MO, NY, OS, OXF, P, PR, PRC, S, SGO,

SI, UC, UPS, US, and W. Special thanks go to Dr. John J.

Furlow (OS) for loan assistance, Maryan Benkowski (OS) for

help with annotation labels, Mauricio Velayos (MA), Melica

Munoz Schick (SGO) , and Esther G.de Mautino (CORD) for sending

additional type material information, and Harald Riedl (W) for

his efforts in type location.

This paper is part of a Thesis presented in partial

fulfillment of the requirements for the Ph.D. at The Ohio

State University. The work was supported by: An OSU Tinker

Foundation Field Research Grant; a John Ayers Herbarium Travel

Award from the American Society of Plant Taxonomists; and

National Science Foundation Doctoral Dissertation Research

Grant, BSR-8815311.

LITERATURE CITED

Adams, E. N. 1972. Consensus techniques and the comparison tof

taxonomic trees. Syst. Zool. 21: 390-397.

. 1986. N-trees as nestings: complexity, similarity, and

consensus. Journal of classification 3: 299-317.

Auer, V. 1958. Ann. Acad. Sci. Fenn. Ser. A, III. 50: 1.

. 1960. The Quaternary history of Fuego-Patagonia. Proc. 127

Roy. Soc. London, Ser. B, Biol. Sci. 152: 507-516.

Baeza R., V. M. 1921. Los nombres vulgares de las plantas

silvestres de Chile i su concordancia con los nombres

cientificos: Con observaciones sobre la aplicacion t&nica

i medicinal de algunas especies, Santiago: Sociedad Imp.

y Lit. Universo.

Baillon, H. E. 1880 [1879]. Ombellifdres. In Histoire des

plantes, ed. H. E. Baillon, 7: 84-256. Paris: Hachette &

Cie.

Bell, C. R. 1971. Breeding systems and floral biology of the

Umbelliferae or evidence for specialization in

unspecialized flowers. In The Biology and Chemistry of

the Umbelliferae, ed. V. H. Heywood, 93-107. London:

Academic Press.

Bell, C. R. & L. Constance. 1957. Chromosome numbers in

Umbelliferae. II. Amer. J. Bot. 44: 565-572.

. 1960. Chromosome numbers in Umbelliferae. II. Amer. J.

Bot. 47: 24-32.

. 1966. Chromosome numbers in Umbelliferae. III. Amer. J. 128 Bot. 53: 512-520.

Bentham, G. 1867. In Genera plantarum, eds. G. Bentham & J. D.

Hooker, 1: 859-931, 1008-1009. London: Lovell Reeve &

Co., Williams & Norgate.

Bhakuni, D. S., M. Bittner, C. Marticorena, M. Silva, E.

Weldt, & M. Hoeneisen. 1976. Screening of Chilean plants

for anticancer activity. I. Lloydia 39: 225-243.

Bland, R. G. 1978. How to know the insects, ed. 3, i-409.

Dubuque: Wm. C. Brown Co.

Borror, D. J., D. M. De Long, & C. A. Triplehorn. 1981. An

introduction to the study of insects, ed. 5, i-827.

Philadelphia: Saunders College Publishing.

Bremer, K. 1990. Combinable component consensus. Cladistics 6:

369-372.

Candolle, A. P. de. 1829. Observations sur la famille des

Ombelliferes. Collection de M&moires 5: 1-84. Paris:

Treuttel et Wttrtz.

. 1830. Umbelliferae. In Prodromus systematis naturalis

regni vegetabilis, ed. A. P. de Candolle, 4: 55-250. 129

Paris: Treuttel et Wiirtz.

Cavanilles, A. J. 1799. Icones et descriptiones plantarum 5:

58-59. Madrid: Typographia regia.

Chodat, R. H. & E. Wilczek. 1902. Contributions a la Flora la

Republique Argentine: Ombelliferes. Bull. Herb. Boissier.

2: 525-528.

Clapperton, C. M., & D. E. Sugden. 1988. Holocene glacier

flucturations in South America and Antarctica. Quaternary

Science Reviews &: 185-198.

Clausen, J., D. D. Keck, & W. M. Hiesey. 1945. Experimental

studies on the nature of species. II. Plant evolution

through amphiploidy and autoploidy, with examples from

the Madiinae. Carnegie Institue: Washington Publishers.

564: 1-174.

Clos, D. 1847. Umbeliferas. In Historia fisica y politica de

Chile, Botanica: Flora chilena, ed. C. Gay, 3: 61-145.

Santiago: Museo historia natural.

Constance, L. 1965. Some subtractions from the Umbelliferae of

South America. Ann. Missouri Bot. Gard. 52: 274-280. 130

. 1971. History of the classification of Umbelliferae

(Apiaceae). In The Biology and Chemistry of the

Umbelliferae, ed. V. H. Heywood, 1-11. London: Academic

Press.

. 1988. Umbellifereae. In Flora Patagonica. Parte V:

Dicotiledoneas Dialipetalas (Oxalidaceae a Cornaceae),

ed. M. N. Correa, 310-379. Buenos Aires: Instituto

Nacional De Technologia Agropecuraria.

Constance, L., T-I. Chuang, & C. R. Bell. 1971. Chromosome

numbers in Umbelliferae. IV. Amer. J. Bot. 58: 577-587.

. 1976. Chromosome numbers in Umbelliferae. V. Amer. J.

Bot. 63: 608-625.

Cronquist, A. 1978. Once again, what is a species? In

Biosystematics in Agriculture, ed. L. V. Knutson, 3-20.

Montclair, NJ: Allenheld Osmun.

Crowden, R. K., J. B. Harborne, & V. H. Heywood. 1969.

Chemosystematics of the Umbelliferae: A general survey.

Phytochemistry, 8: 1963-1984.

Curran, C. H. 1934. The families and genera of North American

Diptera, 1-512. New York: Ballou Press. 131

Drude, C. G. 0. 1897-1898. Umbelliferae. In Die natiirlichen

Pflanzenfamilien, eds. A. Engler & K. Prantl, 3(8):

63-250. Leipzig: Wilhelm Engelmann.

Fries, R. E. 1905. Zur Kenntnis Der Alpinen Flora Im

Nordlichen Argentinien. Nova Acta Regiae Soc. Sci. Upsal.

Ser. IV, 1(1): 1-205.

Gandoger, M. M. 1912. Manipulus plantarum novarum praecipue

Americae austral ior is. Bull. Soc. Bot. France. 59:

704-710.

Grant, V. 1981. Plant Speciation, ed. 2. New York:

Columbia University Press.

Grisebach, A. H. R. 1874. Plantae Lorentzianae: Umbelliferae.

Abh. Konigl. Ges. Wiss. Gottingen. 19: 105-108.

Halliday, P., R. D. Meikle, J. Story, & H. Wilkinson. 1980.

Draft Index of Author Abbreviations: Compiled at The

Herbarium Royal Botanic Gardens, Kew. Basildon: Her

Majesty' Stationary Office.

Harborne, J. B. 1971. Flavonoid and phenylpropanoid patterns

in the Umbelliferae. In The Biology and Chemistry of the

Umbelliferae, ed. V. H. Heywood, 293-314. London: 132

Academic Press.

Heusser, C. J. 1971. Umbelliferae. In Pollen and Spores of

Chile. 59-60, pi. 56. Tucson: The University of Arizona

Press.

Hicken, C. M. 1922. Sertularium Andinum: Umbelliferae.

Darwiniana 1: 61-62.

Hieronymus, G. H. E. W. 1881. Sertum Sanjuaninum: Umbeliferas.

Bol. Acad. Nac. ci. 4: 26-29.

Hiroe, M. 1979. Umbelliferae of World. Tokyo City: Ariake Book

Company.

Holmgren, P. K., N. H. Holmgren, & L. C. Barnett. 1990. Index

Herbariorum. Part I: The Herbaria of the World, ed. 8.

New York: New York Botanical Garden.

Hooker, W. J. 1830. On the plants of the natural order

Umbelliferae, detected by Dr. Gillies in the

extratropical parts of South America. Bot. Misc. 1:

323-335.

Hooker, W. J. & G. A. W. Arnott. 1841. Chili: Umbelliferae. In

The Botany of Captain Beechey's Voyage, eds. W. J. Hooker 133

& G. A. W. Arnott, 25-27. London: Henry G. Bohn.

Johnston, I. M. 1929. I. Papers on the flora of northern

Chile. II. Some undescribed species from Peru. Contr.

Gray Herb. 85: 1-180.

Krause, E. H. L. 1904. Umbelliferae. In J. Sturm's Flora von

Deutschland, ed. 2, 12: 24-164. Stuttgart: K. G. Lutz.

Kuntze, c. E. 0. 1898. Umbelliferae. In Revisio generum

plantarum, ed. C. E. 0. Kuntze, 3(2): 110-115. New York:

Gust. E. Schechert.

Kurtz, F. 1893. Bericht iiber zwei Reisen zum Gebiet des oberen

Rio Salado (Cordillera de Mendoza), ausgefiihrt in den

Jahren 1891-1892 und 1892-1893. Verh. Bot. Vereins Prov.

Brandenburg 34: 95-120.

Lamarck, J. B. A. P. M. de. 1789. Hydrocotyle. Encylopedie

Methodique, Botanique 3: 151-156. Paris: Panckoucke et

Plomtreux.

Lanjouw, J. & F. A. Stafleu. 1954. Index herbariorum. II.

Collectors. A-D. Utrecht: IAPT.

Lawrence, G. H. M., A. F. Gunther Buchheim CK, G. S. Daniels, 134

& H. Dolezal. 1968. Botanico-Periodicum-Huntianum.

Pittsburgh, PA: Hunt Botanical Library.

Lorentz, P. G. & G. Niederlien. 1881. Umbelliferae. Expedicion

al Rio Negro (Patagonia) Entrega II. - Botanica, 221-224.

Buenos Aires: Ostwald Y Martinez.

Macloskie, G. 1905. Umbelliferae. Reports of the Princeton

University expeditions to Patagonia. 1896-1899: Botany

Part V Flora Patagonica. Section 3. Cactaceae-Compositae

8: 619-643. Lancaster, PA: New Era Printing Co.

Margush, T. & R. R. McMorris. 1981. Consensus n-trees.

Bulletin of Mathematical Biology 43: 239-244.

Markgraf V. & H. L. D'Antoni. 1978. Umbelliferae. In Pollen

Flora of Argentina. 96, pi 41. Tucson: The University of

Arizona Press.

Martinez, S. 1989. El Genero Azorella

(Apiaceae-Hydrocotyloideae) en La Argentina. Darwiniana

29: 139-178.

Mathias, M. E. 1971. Systematic survey of New World

Umbelliferae. In The biology and chemistry of the

Umbelliferae, ed. V. H. Heywood, 13-29. London: Academic 135

Press.

Mathias, M. E. & L. Constance. 1962. A revision of

Asteriscium and some related Hydrocotyloid Umbelliferae.

Univ. Calif. Publ. Bot. 33: 99-184.

Mayr, E. 1969. The biological meaning of species. Biol. J.

Linn. Soc. l: 311-320.

. 1992. A local flora and the biological species concept.

Amer. J. Bot. 79: 222-238.

Mercer, J. H. 1972. Chilean glacial chronology 20,000 to

11,000 Carbon-14 years ago: Some global comparisons.

Science 176: 1118-1120.

Mercer, J. H., & C. Laugenie. 1973. Glacier in Chile ended

a major readvance about 36,000 years ago: Some global

comparisons. Science 182: 1017-1019.

Montero, E. A. C., & M. J. Silveira. 1983. Radiocarbon

chronology of a tephra layer in Rio Traful Valley,

Province of Neuquen, Argentina. In Quaternary of South

America and Antartic Peninsula 1: 135-151. Rotterdam:

Balkema. 136

Montes, M. & T. Wilkomirsky. 1985. Medicina tradicional Chile.

Concepcion: Universidad de Concepcion.

Moore, D. M. 1971. Chromosome studies in the Umbelliferae. In

The Biology and Chemistry of the Umbelliferae, ed. V. H.

Heywood, 233-255. London: Academic Press.

Munoz Pizarro, C. 1960. Las especies de plantas descritas por

R. A. Philippi in el siglo XIX, 1-189. Santiago:

Ediciones de la Universidad de Chile.

Pearson, O., & A. Pearson. 1980. Nueve mil afios de

pequenos mamiferos en el Valle Traful. IX Reunion

Argentina de Ecologia, San Carlos de Bariloche, April

1980, Abstracts.

Pena Guzman, L. E. 1987. Introduccion a los insectos de Chile,

1-256. Santiago: Editorial Universitaria.

Persoon, C. H. 1805. Synopsis Plantarum seu Enchiridium

botanicum 1: V.-546. Paris: Treuttel et Wiirtz.

Philippi, R. A. 1864-5. Linnaea. 33: 90. INTER LIB. LOAN

. 1860. Florula Atacamensis: Umbelliferae. In Viaje al

desierto de Atacama, 198-200. Santiago: Halle en Sajonia. 137

. 1894. Plantas Nuevas Chilenas: Umbelliferae. Anales Univ.

Chile 85: 507-514, 699-726.

Presl, K. B. 1825. Reliquiae Haenkeanae seu Descriptiones et

leones Plantarum. Prague: apud J.G. Calve bibliopolam.

. 1833. Repertorium Botanicae Systematicae. Prague: Typis

filiorum Theophili Haase.

Radford, A. E. 1986. Fundamentals of Plant Systematics. New

York, NY: Harper & Row, Publishers, Inc.

Rauh, W. 1939. tiber polsterformigen Wuchs. Nova Acta

Leop. 7: 265-540.

. 1940. Die Wuchsformen der Polsterpflanzen. Botanisches

Archiv. 40: 289-374, 375-462.

Reiche, K. F. 1899. Estudios Criticos Sobre la Flora de

Chile: Umbeliferas. Anales Univ. Chile 104: 767-842.

. 1899. Zur Kenntnis einiger chilenischer

Umbelliferen-Gattungen. Bot. Jahrb. Syst. 28: 1-17.

Rendle, A. B. 1904. Mr. Hesketh Prichard's Patagonian Plants. 138

J. Bot. 42: 321-334, 367-378.

Richards, A. J. 1986. Plant Breeding Systems. Cambridge, MA:

Unwin Hyman Ltd.

Romero Aravena, H., Borgel Olivares, R., & D. Vio Urrutia.

1983. Geografla de Chile. Instituto Geogrkfico Militar.

1: Fundamentos Geogrkficos del Territorio Nacional.

Santiago: Instituto GeogrSfico Militar de Chile.

Ruiz Lopez, H. & J. A. Pav6n. 1802. Flora Peruviana, et

Chilensi. In Prodomus et Flora Peruviana et Chilensis,

eds. H. Ruiz L6pez & J. A. Pavon, 3: i-95. Madrid: en la

imprenta de Sancha.

Sanguinetti de Bdrmida, A. C., & L. A. Borrero. 1983. Las

Buiterras Cave and the palaeoenvironments of the Rio

Gallegos valley, Province of Santa Cruz, Argentina. In

Quaternary of South America and Antartic Peninsula 1:

151-156. Rotterdam: Balkema.

Schumann, K. M. 1900. Neue Arten der Siphonogamen 1898. Just's

Bot. Jahersber. 26(1): 323-396.

Skottsburg, C. J. F. 1916. Botanische Ergebnisse der

schwedischen Expedition nach Patagonien und dem 139

Feuerlande 1907-1909. V. Die Vegetationsverhaltnisse

langs der Cordillea de los Andes. Kongl. Svenska

Vetenskapsakad. Handl. 56(5): 1-366.

Smith, W. J. & F. Vuilleumier. 1971. Evolutionary

relationships of some South American Ground Tyrants.

Bull. Mus. Comp. Zool. Harvard Univ. 141: 1181-268.

Sokal, R. R. & F. J. Rohlf. 1981. Taxonomic congruence in the

Leptopodomorpha re-examined. Syst. Zool. 30: 309-325.

Speggazini, C. 1899. Nova Addenda ad Floram Patagonicam: Part

I. Anales Soc. Ci. Argent. 48: 44-59.

. 1902. Nova Addenda ad Floram Patagonicam: Part IV. Anales

Mus. Nac. Hist. Nat. Buenos Aires. 7(Ser. 2a, t. 4):

203-308.

Sprengel, K. 1813. Plantarum umbelliferarum denuo

desponendarum prodromus. Halae: Hendelianis.

. 1820. Umbelliferae. In Systema vegetabilium, eds. J. J.

Roemer & J. A. Schultes, 6: 315-628. Stuttgard: sumtibus

J. G. Cottae.

Stafleu, F. A. & R. S. Cowan. 1976-1988. Taxonomic Literature, 140

2nd ed. 1-7: A-Z. Utrecht: Bohn, Scheltema & Holkema.

Stearn, W. T. 1986. Botanical Latin, 3rd. ed., revised. North

Pomfret, VT: David & Charles Inc.

Stuessy, T. F. 1990. Plant Taxonomy: The Systematic Evaluation

of Comparative Data. New York: Columbia University Press.

Swofford, D. L. 1991. PAUP: Phylogenetic Analysis Using

Parsimony. Version 3.0s. Privately published.

Tonni, E. P., & F. Fidalgo. 1983. Geology and

palaeontology of Pleistocene sediments at Punta Hermengo

area (Miramar, Province of Buenos Aires, Argentina) : Some

palaeoclimatic aspects. In Quaternary of South America

and Antartic Peninsula 1: 23-53.

Turczaninow, N. S. 1847. Decas Tertia: Nonnullarum specierum

Myrtacearum xerocarpicarum atque Umbelliferarum

imperfectarum. Bull. Soc. Imp. Naturalistes Moscou, 20

(1): 148-174.

Villagrdn, C. 1988a. Late Quaternary vegetation of Southern

Is la Gande de Chiloe, Chile. Quaternary Research 29: 294-

306. 141

. 1988b. Expansion of magellanic moorland during

the late Pleistocene: Palynological evidence from

Northern Isla de Chiloe, Chile. Quaternary Research 30:

304-314.

Vuilleumier, B. S. 1970. The systematics and evolution of

Perezia sect. Perezia (Compositae). Contr. Gray Herb.

199: 1-163.

. 1971. Pleistocene changes in the fauna and flora of

South America. Science 73: 771-780.

Vuilleumier, F. 1968. Population structure of Asthenes

flammulata superspecies (Aves: Furnariidae). Breviora

297: 1-21.

. 1969a. Systematics and evolution in Diglossa (Aves,

Coerebidae). Amer. Mus. Novit. 2381: 1-44.

. 1969b. Pleistocene speciation in birds living in the

high Andes. Nature 223: 1179-1180.

. 1970a. Generic relations and speciation patterns in the

Caracaras (Aves: Falconidae). Breviora 355: 1-29.

. 1970b. Insular biogeography in continental regions. I. 142

The Northern Andes of South America. Amer. Natur. 104:

373.

Weddell, H. A. 1857. Umbelliferae. Chloris andina 2: 186-206.

Paris: chez P. Bertrand.

Wiley, E. O., D. Siegel-Causey, D. R. Brooks, & V. A. Funk.

1991. The compleat cladist: A primer of phylogenetic

procedures. Lawrence, KS: The University of Kansas.

Wolff, K. F. A. H. 1921. Spec. Nov. Austro-Americanae. Feddes

Repert. Spec. Nov. Regni Veg. 17: 441-442.

. 1924. Azorellopsis, genus novum Umbelliferarum

Bolivianum. Feddes Repert. Spec. Nov. Regni Veg. 19: 312.

NUMERICAL LIST OF TAXA

1. M. spinosum (Cav.) Pers. la. M. spinosum var. spinosum lb. M. spinosum var. minus (Kuntze) Zech lc. M. spinosum var. atacamense Zech

Id. M. spinosum var. proliferum ((Cav.) Pers.) Zech

2. M. leptacanthum Philippi

3. M. crassifolium Philippi

4. M. ulicinum Gillies & Hook. 143

4a. M. ulicinum var. ulicinum

4b. M. ulicinum var. multiflorum (Griesb.) Zech

5. M. albovaginatum Gillies & Hook.

5a. M. albovaginatum var. albovaginatum

5b. M. albovaginatum var. pauciflorum (Reiche) Zech

6. M. valentini Speg.

7. M. hallei Skottsb.

8. M. microphyllum (Cav.) Pers.

INDEX TO NUMBERED COLLECTIONS EXAMINED

The numbers in parentheses refer to the corresponding

taxa in the text and in the Numerical List of Species

presented above.

Anclbor, E. 32 (4b); 119 (6).

Ancibor, E., et al. 90214 (5a).

Aravena, P. 33323 (la); 33334 (la).

Arnow, L. 3780 (8).

Arroyo, M. T. K. 81337 (la).

Arroyo, M. T. K., et al. 841002 (6); 870276 (6).

Arroyo, S., et al. 358 (6); 2252 (6); 2384 (6); 2437 (6); 2566

(6); 2610 (6); 2650 (6).

Asplund, E. 2536 (4b).

Aurelius, A. N. J. 16 (8).

Barrientos, R. 1818 (la). 144

Barros, E. 3 (la); 1108 (la); 1171 (2); 21339 (la).

Bartlett, H. H. 19397 (Id).

Beetle, A. A. & A. Soriano 509 (6) .

Berg, C. 67 (6).

Besser, 101 ().

Blake, A. L. 160 (7); 167 (7).

Blake, M. E. 45 (7); 105 () .

Bocher, T. W., et al. 1546 (5a).

Boelcke, O. 1728 (8); 1760 (8); 4217 (5a); 4301 (Id); 4624

(5a); 6317 (e); 6482 (la); 10033 (5a); 10042 (5a); 10078

(5a); 11228 (5a); 11633 (5a); 12912 (6); 15927 (5a).

Boelcke, O. & M. N. Correa 6985 (2); 6987 (2); 6988 (2); 6989

(2 ).

Boelcke, O. & J. H. Hunziker 3484 (8); 3507 (e); 3512 (8).

Boelcke, O., et al. 9940 (4a); 10064 (5a); 10236 (la); 10309

(la); 10970 (2); 11378 (2); 11500 (2); 11610 (2); 11613

(2); 12809 (7); 13042 (2a); 13897 (2); 14179 (2); 14121

(2); 15368 (6); 15747 (la).

Bridarolli, A. 2063 (8).

Bridges, T. 1188 (5b); 1189 (la).

Buchtien, O. 42 (8); 43 (2); 1185 (lb); 1350 (8).

Burkart, A. E. 5344 (4a); 6213 (Id); 6232 (8); 6642 (8).

Cabrera, A. L. 33 (Id); 3466 (la); 3472 (la); 3660 (la) ;

4283/4823 (8); 5767 (la); 5856 (la); 6177 (Id); 6215 (2) ;

6223 (2); 6720 (la); 8353 (4a); 8790 (3); 9069 (4a); 9214

(4a); 9221 (4b); 11031 (e) ; 11067 (Id); 11132 (Id); 11210 145

(8); 11559 (2); 17734 (4b); 18633 (Id); 18663 (Id); 18713

(Id); 21919 (2); 23027 (2); 23086 ().

Cabrera, A. L. & L. Constance 18947 (4b); 18964 (4a & 4b).

Cabrera, A. L. & J. V. Crisci 19112 (Id); 19191 (2).

Cabrera, A. L. & M. M. Job 14 (Id); 30 (Id); 39 (8); 384 (la).

Cabrera, A. L., et al. 21433 (4b); 22540 (4a); 25964 (2).

Castagnet, C. 127 (8).

Castellanos, A. 15501 (4a); 6117 (6); 6119 (6).

Chicchi, A. 72 (Id).

Cipriano, F. 2038 (la).

Collantes T., B. 23121 (lb).

Comber, H. F. 193 (s-p); 823 (8).

Constance, L. 3506 (la); 3507 (la); 3516 (la); 3523 (la).

Constance, L. & A. L. Cabrera 3854 (4a & 4b); 3856 (4a).

Constance, L. & B. Sparre 3577 (Id); 3579 (la).

Constance, L., et al. 3544 (la).

Cordini, I. R. 143 (8); 223 (8); 229 (Id).

Correa, M. N. & L. Constance 3815/3822 (la); 3817/3820 (8);

3836 (7); 3893/3823 (Id); 3970/3826 (8); 3972/3827 (Id);

3985/3829 (7); 3997/3831 (8); 4011/3832 (7); 4036/3834

(Id); 4085/3838 (8); 4090/3839 (6); 4096/3840 (Id) ;

4152/3843 (8); 4181/3844 (Id); 4202/3846 (Id); 4206/3847

(Id).

Correa, M. N. & E. Nicora 3335 (7).

Correa, M. N., et al. 2586 (7); 2598 (7); 2651 (7); 4063 (7);

4085 (8); 4096 (Id); 4202 (Id); 4206 (Id); 4897 (Id); 5815 (5a); 5817 (5a); 5853 (2); 5855 (2); 5912 (2).

Corte, A. 165 (la).

Covas, G. 10913 (5a).

Crisci, J. V. 408 (la).

Cuezzo, A. R. & F. A. Barkley 20MZ383 (5a).

Dawson, G. 1032 (s-p); 1271 (8).

Dawson, G. & H. Schwabe 2633 (2).

De Barba, R. 1470 (Id), de la Sota, E. 2725 (4a).

De Marco de Kreibohm, E. 119 (Id).

DeVore, M. L. 1196 (la); 1309 (la).

Diem, J. 3349 (2).

Donat, A. 38 (Id); 60 (6 & 7); 61 (6 & 7) .

Duek, et al. 58 (la).

Dus&n, P. K. H. 5249 (Id); 5301 (8); 5304 (7); 5931 (6).

Elliott, C. 156 (lb); 157 (la); 334 (Id).

Eruperaire 1395 (6); 1455 (6).

Eskuche, U. 299-1 (2); 599-19 (2); 0926 (8); 0927a (lb); 0927b

(lb); 0928 (2); 0929 (2); 01448 (2); 1868-7 (7); 1921-11

(7).

Eyerdam, W. J., et al. 23876 (6); 23963 (7).

Fabris, H. A. 1209 (Id); 1213 (Id); 2225 (2); 2251 (Id).

Fabris, H. A. & J. V. Crisci 7523 (Id); 7555 (la); 7595 (la) ;

7624 (la).

Fabris, H. A. & J. M. Marchionni 2374 (4a).

Fabris, H. A. & R. L. P€rez Moreau 6741 (la). 147

Fabris, H. A. & O. T. Solbrig 1130 (Id).

Fabris, H. A. & F. 0. Zuloaga 8428 (Id); 8475 (la).

Faggi, A. M. & N. Lopez 4416 (6).

Fiebrig, K. 2601 (4b).

Fiqueran, J. R. 2834 (Id).

Fischer, W. 189 (Id).

Frangi, J. & R. Kiesling 55 (4a).

Fries, R. E., 840 (); 840 (4a).

Frodin, J. 401 (la), 537 (la).

Garaventa, H., A. 1399 (la); 1447 (la); 2790 (la); 4361 (la);

5344 (la).

Gardner, M. F. & S. G. Knees 4452 (la).

Gay, C. 1513 (Id).

Gillies, J. 561 (Id).

Gonzdlez, E. 210 (Id).

Grandjot, C. 1063 (la).

Grandjot, C. & G. Grandjot 3876 (la).

Grau, J. 3117 (la).

Grau, J. & G. Grau 1582 (la).

Grondona, E. 2202 (6).

Gunckel L., H. 46512 (la).

Hager, J. 993 (8); 1002 (8); 1003 (7); 1044 (Id).

Hastings, G. T. 304 (la); 323 (la).

Hauman, L. 252 (Id).

Hicken, C. M. 58 (Id)

Heinziman, F. P. M. 21045 (5a). 148

Hieronymous, 6. & 6. Niederlein 361 (4b); 808 (4b); 819 (4a).

Hogberg, C. 23 (la); 142 (8).

Hollermayer, R. P. A. 438 (Id); 438a (Id); 438b (Id); 489 (2) ;

1724 (2).

Holway, E. W. D. & M. W. Holway 255 (la).

Hunziker, J. H. 2099 (4b); 6379 (5a); 6993 (2).

Hunziker, J. H. & O. Caso 4145 (4b).

Hutchison, P. C. 94 (la); 305 (la).

Illin, N. 257 (8).

Isabel, M., H. S. de Birabdn, & M. Biraben 655 (Id).

Jaffuel, F. 2839 (la).

Jeltor, J. C. 2038 (la).

Jiles, C. 1232 (lb); 1972 (lb); 3594 (lb); 3750; 4564 (Id).

Johnston, I. M. 6026 (lc).

Jorgensen, P. 1692 (4b).

Joseph, C. F. 557 (Id); 3825 (la); 4103 (2); 5271 (la); 5489

(2) .

Keever, J. & J. Greer 1 (2); 1287 (2).

Kendel, G. 19608 (la).

King, D. 0. 114 (5a)

Klein, R. M. 0160 (Id).

Koslowsky, J. 33 (7); 1314 (6).

Krapovickas, A. 6250 (4b).

Kiihnemann, 0. 471 (8) ; 552 (8) .

Kuntze, C. E. O. 32 () .

Kurtz, F. 7505 (Id); 9463 (4a). 149

Landero, A. 643 (6); 699 (6).

Landrum, L. R. 3309 (Id).

Lecoi, R. & C. Montana 5321 (3) .

Lillo, M. 5509 (4a).

Ljungner, E. 6 (la); 509 (8); 756 (8); 773 (8); 794 (8); 799

(8).

Llera, V. & E. Torres 1870-2 (Id).

Looser, G. 361 (la); 855 (la).

Lorentz, P. G. 430 (); 586 ().

Magens, 0. 42 (Id).

Mahu, M. 4064 (la); 6261 (3); 9816 (la).

Marticorena, C. & O. Matthei 629 (lb); 821 (la); 937 (la);

1042 (la).

Marticorena, C. & M. Quezada 9585 (la).

Marticorena, C. & R. Rodriguez 8407 (Id) .

Marticorena, C., et al. 64 (la); 107 (Id); 1248 (Id); 1348

(la); 25287 (Id); 83613 (lc).

Matte, V. 715 (la).

Matthei, O. & N. Bustos 145 (la).

Med&n, D. & M. C. Alvarez 292 (Id).

Meyer, F. 8029 (la).

Meyer, T. 7577 (Id); 8018 (Id); 8052 (Id); 9633 (Id).

Miche, G. 171/80/24.4 (4b); 171/80/25.4 (4b).

Montero, 0., G. 306 (la); 530 (la); 2738 (la); 5945 (la); 7080

(la).

Morales, R. & T. Martin 236RM (Id). 150

Moreno, E. P. & Torrini 368 (8).

Morrison, J. L. 16915 (la).

Munoz Pizzaro, C. 3770 (3); 3800 (3); 3807 (3).

Nee, L. 1801 (Id); 4601 (Id).

Neger, F. W. 1897 (2).

Nordenskiold, O. 32 (Id).

Novas B., E. 2097 (3).

O'Donell, C. H. 1995 (Id); 2049 (2); 2305 (Id); 2728 (Id);

3363 (8); 3560 (8); 3663 (7); 3799 (7).

Parodi, L. R. 11859 (8).

Pedersen, T. M. 293 (Id).

Pena, L. E. 14 (la).

Pennell, F. W. 12239 (la); 12240 (la); 12367 (la) 12450 (la);

12998 (la).

Perez Moreau, R. 18663 (8); 45363 (8); 45379 (2); 45386 (2);

48689 (8); 48694 (8); 48695 (8); 49140 (2); 49142 (8);

49902 (2); 49907 (2); 50370 (2); 50371 (2); 55087 (5a).

Pfister, A. 176 (la); 844 (2); 946 (la); 7132 (Id); 7242 (la);

7867 (la); 8074 (2); 9597 (lb).

Philippi, R. A. 155 (3).

Pichi Sermolli, R. E. G. & P. Bizzarri 7372 (Id).

Pisano-Valdez, E. 2372 (Id).

Pisano-Valdez, E. & J. Venturelli E. 1978 (3).

Plotnick, R. 49 (Id).

Poeppig, E. F. 530 (la).

Puentes, J. 5172 (la). 151

Rahmer, C. 18 OR 2049 (2- see isotypes).

Ricardi Salinas, M. 2420 (la); 2967 (la); 2986 (3); 3168 (la);

5583 (la).

Ricardi Salinas, M. & C. Marticorena 5060/1444 (2).

Ricardi Salinas, M. & 0. Matthei 282 (8).

Ricardi Salinas, M., et al. 756 (la); 758 (lb); 821 (la).

Rodriguez, 293 (4b).

Roig, F. A. & R. A. Ruiz Leal 20166 (4a).

Rossi & Bachmann 482 (la) .

Ruiz Leal, R. A. 1107 (Id); 1724 (Id); 1855 (4b); 1983 (5a);

2021 (Id); 2135 (4b); 3126 (4b); 3155 (5a); 3169 (5a);

3619 (5a); 4029 (Id); 7182 (4b); 7183 (5a); 11682 (5a);

13400 (4a); 14578 (5a); 14784 (6); 15716 (la); 20244 (8);

21439 (5a); 23988 (7); 24082 (7); 24109 (7); 24218 (7);

24225 (6); 24544 (la); 25591 (8); 26423 (6); 26425 (7);

26434 (6); 26451 (6); 26772 (7); 26905 (7); 27020 (6).

Ruiz Leal, R. A. & F. Roig 15654 (5a); 15763 (5a) .

Ruthsatz, B. 4 (4a); 7 (4a); 35 (4b); 40 (4b); 44 (4a); 47

(4b); 57 (4a); 126 (4a); 135 (4a); 136 (4a); 166 (4b) ;

204 (4a); 210 (4a); 225 (4a); 241 (4b); 275 (4a); 336

(4a); 363 (4b); 507 (4b) .

Salinas, J. V. 2180 (5a) .

Santesson, R. 225 (7); 320 (7); 1379 (la).

Schickendantz, F. 334 (4b).

Schlegel, F. 2360 (la); 2476 (la); 2653 (3); 3592 (Id); 3774

(la); 3775 (la); 5124 (3); 5905 (la). 152

Schwabe, H. W. Ch-103 (la).

Semper, J. 252 (la); 1185 (5a); 10091 (4a).

Silva, M. 19694 (la).

Simon, J. P. 274 (lb).

Skog, L. E. 1104 (la).

Skottsberg, C. J. F. 619 (6); 706 (Id); 855 (7); 856 (8); 857

(7). Sleumer, H. 575 (la); 842 (6); 927 (6); 1178 (6); 1358 (6).

Sleumer, H. & F. Vervoorst 2578 (4b).

Soetbeer, M. & A. M. Glesias 66 (la).

Soriano, A. 1320 (8); 1986 (8); 2025 (8); 2103 (6); 2160 (6);

2193 (6); 2257 (7); 2426 (7); 2508 (8); 3072 (6); 3788

(8); 4795 (6); 5076 (7); 5098 (7); 5099 (7).

Sparre, B. 1745 (la).

Sparre, B. & L. Constance 10906 (la).

Stuckert, T. 1513 (); 7505 (Id); 15125 (4b); 15140 (4b).

Stuessy, T., et al. 6819 (Id); 7527 (Id); 10000 (2); 10017

(2); 10047 (2); 10048 (2); 10049 (2); 10072 (2); 10073

(la); 10116A (Id); 10116B (Id); 10116C (Id); 10129 (2);

10152 (Id); 10154 (Id); 10158 (Id); 10162 (2); 10187

(Id); 10198 (2); 10216 (la); 10236 (Id); 10252 (Id);

10256 (Id); 10260 (Id); 10264 (Id); 10269 (Id); 10275 () ;

10276 (la); 10276A (); 10277 (); 10296 (la); 10318 (la);

10333 (5a); 10354 (Id); 10420 (Id); 11123 (la).

Valentine, G. 66 (6).

Ventania, Proyecto 248 (la) . 153

Venturi, S. 4785 (4a); 4785 () ; 6449 (4b); 6450 (4b); 6451 () ;

6977 (); 6977 (4a); 7275 (4a); 7276 (4b); 7277 (4b); 9042

(4b); 10277 (4b); 10279 (4b).

Vervoorst, F. 5577 (Id).

Vidal, M. I. 24 (6).

VillagrSn, C. 1334 (3). von Platen, L. & U. Greiner 152 (la); 153 (7). von Thtingen, J. 14 (7); 72 (7); 97 (7); 103 (7).

Tweedie, R. 251 (Id).

Wall, E. & B. Sparre 23 (la); 23 (la); 1494 (Id).

Walter, H. & E. Walter 543 (Id).

Weisser S., P. 1544 (la).

Weldt, E. 712 (la).

Werdennann, E. 268 (lc); 554 (la); 613 (5b); 1009 (3).

West, J. 4743 (Id); 4757 (8).

Wetmore, A. 812 (Id).

Wilczek, E. 65 (5a); 81 (Id).

Wilczek, E. & P. G. Lorentz 106 (Id) .

Zardini, V. 101 (Id).

Zech, J. C. & N. C. Contogiorgakis 2 (la); 3 (la); 4 (la); 5

(5a); 6 (la); 7 (la); 8 (la); 9 (5a); 10 (Id); 11 (Id);

12 (Id); 18 (6); 21 (7); 24 (6); 25 (6).

Zech, J. C., et al. 29 (la); 33 (Id); 42 (la); 44 (la); 46

(2); 49 (Id); 50 (la); 51 (Id); 52 (Id); 55 (Id).

Zollner, O. 1280 (3); 2081 (lb); 2295 (la); 3517 (3); 5200

(la); 5209 (la); 5650 (lb); 5758 (la); 5950 (5b); 5951 154

(la); 6139 (la); 7587 (); 10249 (la); 11121 (la); II793

(lb); 13610 (la). CHAPTER II

EVOLUTION OF CUSHION HABIT IN MULINUM (APIACEAE)

ABSTRACT. The cushion habit, defined narrowly as plants with reduced internodes resulting in a compact (often small) habit, is arguably one of the world's most unique plant forms. Genetic and environmental factors, including cell size, bud protection, wind, fog, solar radiation, solifluction, microenvironment, high elevation, low moisture, and diurnal temperature fluctuation, have been proposed as responsible for the formation and maintenance of the compact habit. Because Mulinum contains both cushion and noncushion taxa, it represents a good system for studying the evolution of the cushion habit within a genus.

Both genetic and environmental factors are believed responsible for cushion formation in Mulinum with environmental factors, such as strong winds, playing a more dominant role. A single origin for the cushion habit is hypothesized within the genus, and a reductional evolutionary sequence from a shrub to subshrub to a compact cushion is believed to have occurred.

155 156

The cushion habit represents one of the most extreme

vegetative morphologies within the flowering plants. In

their 1914 survey of Angiosperm cushion plants, Schroter and

Hauri recognized 338 species, representing 78 genera and 34

families within seven geographic areas, as having this

distinctive growth form. While cushion plants dominate in

alpine communities worldwide, they are most conspicuous in

the Andes of South America, with South America containing

over 50% of all cushion taxa (Schroter and Hauri 1914).

In his comprehensive review of cushion plants, Rauh

(1939, 1940) defined cushion plants broadly by recognizing

both vascular and nonvascular (Bryophyte) taxa, classifying

them into 17 categories (Table 4). Most modern workers

(Hedberg 1964; Ruthsatz 1978; Armesto et al. 1980),

including this author, follow a more narrow definition which

corresponds to a combination of Rauh#s categories I-lc & d and I-2a & b. The habit is defined by reduced internode

length, which leads to the characteristic compact, generally

flat, growth-form. The cushion habit is most widely represented within three genera, Azorella Lam. (Apiaceae),

Saxifraga L. (Saxifragaceae), and Raoulia J. D. Hook, ex

Raoul (Asteraceae) (Rauh 1939, 1940).

According to the definition of Rauh (1939, 1940) all species of the genus Mulinum Pers. could be considered cushion plants. However following the narrower definition above, four Mulinum cushion species are currently recognized TABLE 4. Rauh's (1939) categories of cushion plants.

I. TRUE CUSHION PLANTS

1. Radially symmetrical, Spherical Cushions

a . Cushion Trees

b . Spherical Shrubs

c. Hollow Spherical Cushion

d. Solid Spherical Cushion

2. Radially Symmetrical, Flat Cushions

a. Hollow Flat Cushion

b. Solid Flat Cushion

3. Attached to the True Cushion

II. CREEPING CUSHIONS

III. GRASS-LIKE CUSHIONS

IV. ROSETTE CUSHIONS

V. SUPPORTED CUSHIONS

VI. CUSHION MOSS

VII. MONOCARPOUS CUSHIONS

Appearing As Cushions from among eight in the genus (see Chapter 1). Mulinum albovaginatum Gillies & Hook, occurs at high altitudes

(1700-3700 m) of the Andes primarily within Mendoza

Province, Argentina, with a limited extension south into

Neuquen Province and west into Chile around Volcdn Peteroa

(Fig. 16). This species forms small (4-15 cm) cushions which are often found intertwining among rock outcrops.

Mulinum valentini Speg. forms dense, flat cushions 3-33 cm in diameter, and is distributed at lower altitudes (130-

1600) primarily along the coast of Southern Argentina and to a lesser extent among the lower elevations of the Andes in the Argentinean Provinces of Chubut and Santa Cruz. In addition, M. valentini extends into Chile at a single location near the Parque Nacional Torres del Paine (Fig.

16). Mulinum hallei Skottsb. and Mulinum microphyllum

(Cav.) Pers. not only represent the smallest of the cushion species, up to 10 cm in diameter, but also the smallest taxa within the genus. The two species are extremely similar in their morphology and differ only by the presence of simple, single, terminal lobe hairs (M. microphyllum), chromosome number [M. hallei, n = 8; M. microphyllum, n = 16 (Constance et al. 1971)], and to a lesser extent distribution (Fig.

16). Like M. valentini, both M. hallei and M. microphyllum have disjuct distributions between the coast and lower elevations of the Andes within Southern Argentina. Mulinum hallei occurs primarily along the coast of Southern 159

FIG. 16. Distribution of Mulinum cushion species within southern South America. • = M. albovaginatum. ▲ = M. hallei. ■ = M. microphyllum. ★ = M. valentini. 160

Argentina with locations within the lower elevations of

Chubut and Santa Cruz Provinces (60-1200 m); whereas M.

microphyllum is distributed primarily within the lower

elevations of the Argentine Andes within the Provinces of

Neuquen, Rio Negro, and Chubut and more scarcely along the

Southern coast (90-1500 m; Fig. 16).

Because Mulinum contains both cushion and noncushion

taxa, it represents a good system for studying the evolution

of the cushion habit within a genus. The purpose of this

study is to: (1) present a review of factors which have

previously been discussed as responsible for cushion

formation and discuss these factors as they relate to

cushion formation within Mulinum; and (2) suggest hypotheses

for the evolution of the cushion habit within the genus.

GENERAL FACTORS AFFECTING CUSHION FORMATION

The cushion habit is clearly an adaptation to environmental extremes (Billings and Mooney 1968; Ralph

1978) and genetic and environmental factors have been suggested as stimuli for cushion formation. Most factors, especially environmental, have been presented alone (e.g.,

Turesson 1931; Spomer 1964) as single causes of cushion formation. However, a combination of genetic and environmental as well as a combination of several environmental factors are believed responsible not only for 161 initial cushion formation but also later cushion maintenance.

Factors— Many previous studies have addressed the genetic basis of different factors associated with a reduced and/or cushion habit (Turesson 1931; Rauh 1939, 1940;

Clausen et al. 1940, 1948; Clebsch 1960; Mooney and Billings

1961; Lona 1963; Hedberg 1964; Landolt 1967; Rochow 1967;

Korner et al. 1989). Rauh (1939, 1940) regarded the role of the environment as minimal, especially wind shearing effects, and identified the cushion habit as being

"morphogenetically determined" (Rauh 1939: 497; 1940: 446).

In a series of studies (Korner et al. 1986; Korner and

Larcher 1988; Korner et al. 1989), Korner has compared both the morphology and physiology of alpine plants, including cushion species, with their lower altitude counterparts.

Korner et al. (1989) identified both genetic and environmental factors associated with the compactness of alpine plants. While an increase in leaf thickness, inclination, and stomatal densities at higher altitudes was present, total plant dry mass and mean sum of leaves per individual remained constant between high and low altitude plants. Leaf size was significantly reduced at higher elevations, but cell dimensions did not change with altitude. Transplantation to warmer, low altitude environments showed a production of larger and thinner leaves and thinner cell walls, indicating a strong 162

environmental influence on these properties. In contrast

cell size changed little. Therefore Korner et al. (1989)

concluded that compactness at higher elevations was due to a

genetically controlled decrease in cell size.

Bud protection has been suggested as catalyst for

cushion formation (Rauh 1939, 1940; Hedberg 1964; Jolls

1975; Ruthsatz 1978; Armesto et al. 1980). Peripheral buds

are exposed to environmental fluxes in temperature and

particle degradation. A reduction in internode length

repositions the buds within the plant and more closely to

ground level where they are better protected.

Climate can be regarded as the primary expression of

the set of environmental factors impinging on an organism.

Climate is one of the major determinants of plant forms;

therefore, plant forms are an expression of climate (Jolls

1975).

a. Temperature— In an open vegetation of isolated

cushion plants, local temperature gradients are controlled by the distribution of soil moisture, with small differences

in microtopography making considerable differences in soil temperature and wind-chill on alpine ridges keeping maximum summer temperatures low (Billings and Mooney 1968). In general cushion plants benefit by being closer to ground level where temperatures remain higher and more constant

(Rauh 1940; Hedberg 1964; Ruthsatz 1978). Cushions are subject to less severe temperature fluxes by being exposed 163

to slower wind velocities (Jolls 1975) and often maintain

higher leaf temperatures compared to external temperature

(Gates and Janke 1965; Warren-Wilson 1959). While many

studies have indicated temperature as a contributing factor

to cushion formation (Schimper 1898; Schroeter 1908; Troll

1948, 1958, 1959; Troll 1960; Hedberg 1964; Billings and

Mooney 1968; Korner et al. 1989), Spomer (1964) suggested

that temperature was the primary factor controlling the

habit of the Rocky Mountain cushion species studied.

b. Moisture— Several studies have included moisture

conservation in connection with the cushion habit (Schroeter

1908; Warming 1909; Rauh 1939, 1940; Hedberg 1964; Billings

and Mooney 1968). Although many mountain environments are

efficient producers of precipitation, annual fluctuations in

available moisture occur. Cushion plants adapt to both

winter and summer drought stress by decreasing water

potentials, increasing soluble carbohydrate concentrations,

and closing stomates (Billings and Mooney 1968).

c. Solar Radiation— Solar radiation reaches the soil in

relatively great amounts, especially in open areas scattered with isolated cushion plants (Billings and Mooney 1968) but

is in turn lost in great amounts by reradiation to the sky

(Jolls 1975). As a result of this fluctuation in the amount of solar radiation available, cushion plants, through their form, have adapted an efficient means of heat energy conservation (Schimper 1898; Schroeter 1908; Hann and Suring 164

1939; Sauberer and Hartel 1959; Troll 1959; Hedberg 1964;

Billings and Mooney 1968; Cernusca 1976; Ralph 1978; Korner and Cochrane 1983; Korner et al. 1989).

d. Wind— Wind has been identified as being responsible

for form, particularly in cushion plants, in several studies

(Schimper 1898; Warming 1909; Neger 1913; Espinosa 1932;

Warren-Wilson 1952; Troll 1966; Billings and Mooney 1968;

Smith 1972; Ralph 1978; Van Gardingen and Grace 1991). Wind can have a great effect on plant growth and form, with windswept ridges providing one of the most severe environments in terms of temperature, drought stress, and wind abrasion (Billings and Mooney 1968). In exposed places of the tundra or alpine environments wind blasts and blowing snows limit "tree" growth to a twisted shrubby ,,Krummholz,, near the ground (Billings and Mooney 1968). Wind depresses plant growth through mechanical injury, indirect effects such as soil erosion, cooling, and desiccation (Warren-

Wilson 1959). Cushion plants are well-adapted to wind stress

(Schimper 1898; Warming 1909; Espinosa 1932; Rauh 1939;

Hedberg 1964) with wind speed increasing more or less logarithmically with increasing height above the ground's surface (Whitehead 1951; Warren-Wilson 1959; Gates and Janke

1965; Geiger 1965). Cushion plants occur in a stratum where the lowest wind speeds occur with the wind being reduced internally by the breaking action of the dense branches and foliage (Jolls 1975). Warren-Wilson (1959) reported that a 165

vegetation barrier of 50% density will decrease wind speeds

by 80% for distances of three to five times the height of

the barrier.

e. Fog— The cushion form is adapted for harvesting

moisture carried within nightly and coastal fogs (Hedberg

1964; Troll 1966; Smith 1972).

f. Elevation— Many studies have suggested increased

altitude as the primary factor behind the cushion form

(Turesson 1931; Rauh 1939, 1940; Clausen et al. 1940, 1948;

Hedberg 1964; Billings and Mooney 1968; Jolls 1975; Ralph

1978; Korner et al. 1989), with cushions becoming more

compact with increasing elevation (Spomer 1964).

g. Solifluction— Solifluction is all types of soil movements caused by seasonal or diurnal freezing and thawing

(Troll 1944; Hedberg 1964). Plants, especially seedlings,

often are uprooted by solifluction, specifically in the case

of frost. Cushion plants occur mainly in regions which have nightly frosts throughout the year (Hedberg 1964) The extensive taproots associated with most cushions have been

suggested (Troll 1944, 1947; Hedberg 1964) as a means of preventing the disruption and often destruction of these plants which occur potentially solifiable soils.

h. Microenvironment— The microenvironment hypothesis

(Geiger 1950; Griggs 1956; Hedberg 1964; Billings and Mooney

1968; Baust 1976; Schinner 1982; Korner, et al. 1989) is a synthesis of several factors which have been previously 166 presented alone. In general cushion taxa occur in environmentally extreme locations and represent islands of productivity in virtually unproductive landscapes. The compact cushion habit provides a habitat for other organisms

(Heilborn 1925; Billings and Mooney 1968), particularly microorganisms. An increase in microbial activity within

the cushion leads to an increase in humus production which

in turn contributes to both moisture (Heilborn 1925) and

temperature conservation. A lower day-time and higher night-time temperature (Warren-Wilson 1957; Hedberg 1964;

Coe 1967; Barry and Van Wie 1974) and higher moisture

concentration is internally maintained as compared to the external environment.

GENERAL FACTORS AFFECTING MULINUM CUSHION FORMATION

Although genetic and environmental factors (Table 5) are believed responsible for Mulinum cushion formation and maintenance, the environment is believed to play a more dominant role. When grown under greenhouse conditions at the University of California, Berkeley, M. leptacanthum, a noncushion species, and the cushion Azorella compacta, both generated elongated internodes and lost their general compact nature. This suggests that internode length and the compactness associated with it may be more environmentally influenced. Another possibility is that selection is acting 167

TABLE 5. Suggested environmental factors contributing

to the habit of Mulinum cushion species.

Factor albovaginatum hallei microphyl1urn valentini

DTS: + + + +

LM: + + + +

SR: + + + +

W: + + + +

F: + + + +

HE: + + + +

S: + + + +

M: + + + +

DTS = Diurnal temperature shift; LM = Low moisture; SR =

Solar radiation; W = Wind; F = Fog; HE = High elevation; S =

Solifluction; M = Microenvironment; + = always present; ± = often present. 168

upon genotypes which are phenotypically plastic (Schlichting

1986). Mulinum cushion species occur in barren habitats and

are continually subjected to environmental extremes.

a. Temperature— Cushion species of Mulinum are

subjected to annual (Table 6) as well as extreme diurnal

temperature fluctuations.

b. Moisture— Mulinum cushion species occur in dry

areas. In Mendoza Province, the primary location of M.

albovaginatum, the agricultural lands receive an average of

less than 250 mm of annual precipitation (Hoffmann 1975),

while annual potential evapotranspiration is ± 800 mm

(Burgos and Vidal 1951). Further, cloudiness which may

contribute precipitation and

snowfall are much reduced in the Andes during situations of

extreme drought (Minetti and Sierra 1989).

c. Solar Radiation— Mulinum cushion plants which occur

at high altitudes are subjected to increased solar radiation

as compared to lower elevations and Mulinum cushion species

in general occur in areas with high percentages of annual

solar radiation (Table 6).

d. Wind— All cushion species of Mulinum occur in windswept habitats (Table 6). The winds at both high and

low elevations are extremely intense, projecting a constant force on the plants they come in contact with. Of the winds which correspond to Mulinum cushion distribution (Table 7) the Zonda wind is one of the most intense, with the Zonda TABLE 6. Climatic descriptions for selected cities of Argentina (Rudloff 1981). City location corresponds to Mulinum's distribution; * corresponds to Mulinum cushion species distribution.

Avg. Yearly Avg. Yearly City Alt.(m) Temp.(°C) Precep.(mm) % Sunsh. Wind Fog

La Quiaca: 3459 9 323 76 G O Tucuman: 420 19 977 47 G O Catamarca: 547 20 370 — G O *Sarmiento: 268 11 152 — G O *Comodoro/ Rivadavia: 61 13 218 48 G 0 *Pto. Deseado: 79 10 192 — G 0 San Antonio W: 7 15 243 63 G 0 *Santa Cruz: 111 8 192 45 G 0 *Rlo Gallegos: 17 7 249 — GO La Rioja: 450 20 316 65 WGMOH *Mendoza: 827 16 196 66 WGM OH ♦Neuquen: 270 14 170 — WGMOH Chos Malal: 858 13 265 — GM WH *Bariloche: 836 8 954 58 GM WH

G = Gale force; O = Occasionally; H = High elevation; W = Winter; M = Mountains. 170

TABLE 7. Winds of Argentina (Rudloff 1981), Which

Correspond to Mulinum's Distribution.

1. Zonda: Warm dry fall-wind; blowing from the N or

NW in the Andes; famous for their biting

cold.

2. Travestas: W or NW-erly; at times stormy; winds in

Patagonia blowing from the Pacific Ocean.

3. Sudestado: Storm from the SW after a cold front

passage on the coast; often lasts for

several days.

4. Pampero: Abruptly-starting storm blowing from the S

or SW. 171 wind blowing over the low-lands of Mendoza causing very high evaporation rates (Norte 1986).

e. Fog— Mulinum cushion species do occur in areas which are blanketed with fog, both along the coast and at higher elevations in the Andes (Table 6). Cushion species of

Mulinum are probably able to take up the moisture carried by fog, and use it as a valuable water source in a generally dry habitat.

f. Elevation— Previous cushion plant studies (Armesto et al. 1980) have mainly included cushion taxa which occur exclusively at high elevations. Mulinum is one of a few unique genera (a second being Azorella) that has cushion taxa which grow at both high and low elevations. This suggests that elevation alone does not play a significant role in cushion formation and/or maintenance within Mulinum.

g. Solifluction— An extensive taproot often accompanied by a network of branch roots is common to all Mulinum cushion species and may reflect not only an adaptation for moisture conservation but also for the prevention of solifluction as well.

h. Microenvironment— It is clear that the cushion species as well as noncushion taxa of Mulinum represent adaptive microenvironments. Internally the plants are composed of decaying leaf bases which contributes to increased humus production and moisture and temperature conservation. While measurements of internal cushion and 172 external environment temperature and moisture differences have not been documented, lower daytime and higher nighttime temperatures as well as higher moisture content are predicted for Mulinum cushion species with further study suggested.

EVOLUTION OF THE CUSHION HABIT IN MULINUM

Wind is proposed as a primary influence and initial force in the evolution of the Mulinum cushion habit.

Mulinum spinosum is composed of several stems which radiate from a more or less single source forming spherical to hemispherical shrubs. Subjected to constant, strong winds, the branches of M. spinosum could have been forced to ground level and the remaining aerial branches broken off (Fig.

17). A smaller more subshrub-like habit would have been selected for. Windshear of exposed branches and buds coupled with the additional environmental pressures of water and temperature conservation could further force a reduction in form culminating in the cushion habit (Fig. 17).

Because Mulinum contains both cushion and noncushion taxa, it represents a unique system for studying the evolution of the cushion habit within a genus. Shrub, subshrub, and compact cushion habits are all represented within Mulinum. A reductional evolution of the habit is proposed for Mulinum, with the shrub habit evolving into a 173

B C

Fig. 17. Suggested evolution of the cushion habit for Mulinum. A. Shrub; B. Subshrub; C. Cushion. 174

more reduced subshrub, followed by further reduction into a

more compact cushion. The form of M. spinosum, a shrub, may

have evolved into a form similar to M. leptacanthum, a

subshrub, which in turn may have evolved into a form similar

to the cushion M. alJbovaginaturn.

Although current disjunct distributions (Fig. 16)

indicate the possibility of a multiple cushion habit origin

for Mulinum, cladistic analysis supports, by a single

character, the cushion habit originating a single time (Fig.

18). While three of the four cushion species display a disjunct distribution between the coast and the Andes, evidence exists which suggests more extensive past distributions. A distributional corridor, occupied primarily by M. spinosum, exists across Chubut Province connecting the Andes and the coast. This corridor suggests a more extensive past distribution and a possible east-west migration route (Fig. 19). The distributions of Mulinum hallei, M. valentini, and especially M. microphyllum extend from both the coast and the Andes into the corridor, suggesting a more extensive and nondisjunct past distribution. In addition, a single northern disjunct location for M, valentini exists, which further supports a more extensive past distribution. Based upon the current sympatric distribution of three of the four cushion species, an ancestral cushion taxon probably originated along the

Andes of Chubut Province, Argentina. Species then evolved HYP

CRA D

ULI

ALB A

VAL E HAL

MIC

LEP

SPI

FIG. 18. Cladogram of taxa of Mulinum, showing the cushion clade. The names of taxa are represented by the first three letters of the specific epithets. 175 176

Fig. 19. Distribution of the genus Mulinum in southern South America, showing the Chubutan corridor. and migrated. Mulinum albovaginatum advanced into higher

elevations to the north, while M. hallei, M. microphyllum,

and M. valentini migrated to the south along the Andes and to the east with M. hallei and M. valentini further

extending south along the coast (Fig. 19). A U-shaped migration south along the Andes, east across southern Santa

Cruz Province, and north along the coast seems less likely

than the proposed H-shaped route. The initial leg of the U-

shaped route along the Andes would have been repeatedly blocked by glacial advancement which was limited to just beyond the western cordillera (Vuilleuraier 1971). In addition, a current sympatric distribution along the Andes of Santa Cruz Province would be expected for the three southern cushion species.

The cladistic analysis also supports the proposed progression in cushion formation from spreading shrub to compact cushion. The branching pattern suggests a shrub­ like Ancestor A which gives rise to M. spinosum and a less shrub-like Ancestor B (Fig. 18). Subshrub-like Ancestor B, then gives rise to M, leptacanthum and Ancestor c (Fig. 18).

The subshrub-like habit is then passed from Ancestor c to

Ancestor D, the common ancestor shared by M. crassifolium and M. ulicinum (Fig. 18) both of which are interpreted as subshrubs (see Chapter 1). Ancestor C's subshrub-like habit is reduced to a cushion or cushion-like habit in Ancestor E, the common ancestor shared by the four cushion species (Fig. 178

18) .

This study represents the first attempt to study the evolution of the cushion habit within a genus. While the branching patterns are suggestive of a reductional development from shrub to cushion and a strong environmental force is probable, additional investigation is needed. The precise role of the environment and its influence on cushion formation and maintenance is unclear.

ACKNOWLEDGMENTS. The author thanks Drs. Tod F. Stuessy for his assistance with this research. Thanks also go to:

Daniel J. Crawford, and Fred Sack for helpful comments;

Maria Marta Cigliano, Claudio Contogiorgakis, Jorge Crisci,

Mario Gentili, Patricia Gentili, Liliana Katinas, Juan Jose

Morrone, Alegandra Moschetti, Alberto Ripalta, Fidel

Alegandro Roig, Tod F. Stuessy, and Ana Maria Zavattieri for field and project assistance in South America. This paper is a part of a Thesis presented in partial fulfillment of the requirements for the Ph.D. at The Ohio State University and supported by NSF grant BSR-8815311, an OSU Tinker

Foundation Field Research Grant, and a John Ayers Herbarium

Travel Award from the American Society of Plant Taxonomists.

LITERATURE CITED

Armesto, J. J., M. D. Arroyo, and C. Villagrdn. 1980. 179

Altltudinal distribution, cover and size structure of

umbelliferous cushion plants in the high Andes of

Central Chile. Acta OEcologica/OEcologia Generalis 1:

327-332.

Barry, R. G., and C. C. Van Wie, 1974. Topo- and

microclimatology in alpine areas. 73-83. In Ives, J.

D., and R. G. Barry, eds, Arctic and alpine

environments. London: Methuen and Co., Ltd.

Baust, J. G. 1976. temperature buffering in an arctic

microhabitat. Annales of the Entomological Society of

America 69: 117-119.

Billings, W. D. and H. A. Mooney. 1968. The ecology of

arctic and alpine plants. Biol. Rev. (London) 43: 481-

529.

Burgos, J. J. and A. L. Vidal, 1951. Los climas de la

Reptiblica Argentina, segdn la nueva clasifcacion de

Thornthwaite. Meteoros 1: 3-32.

Cernusca, A. 1976. Bestandesstruktur, Biokima und

Energiehaushalt von alpinen Zwergstrajichbestanden.

Oecol. PI. 11: 71-102. 180

Clausen, J., D. D. Keck, and W. M. Heisey, 1940.

Experimental studies on the nature of species. I.

effect of varied environments on western North American

plants. Publ. Carnegie Inst. Wash. 520: 1-452.

. 1948. Experimental studies on the nature of species

III. Environmental responses of climatic races of

Achillea. Publ. Carnegie Inst. Wash. 581: 1-129.

Clebsch, E. E. c. 1960. Comparative morphological and

physiological variation in arctic and alpine

populations of Trisetum spicatum. Ph.D. dissertation,

Duke University, 1-241.

Coe, M. J. 1967. Microclimate and animal life in the

equatorial mountanins. Zoolog. Africana 4: 101-128.

Constance, L., T. Chuang, and R. C. Bell. 1971. Chromosome

numbers in Umbelliferae. IV. Amer. J. Bot. 58: 577-587.

Espinosa, R. 1932. Okologische Studien iiber

Kordillerenpflanzen. Bot. Jaarb. 65: 120-211.

Geiger, R. 1950. Das Lima der bodennahen Luftschicht. Die

Wissenschaft, 78, ed. 3. Braunschweig. 181

. 1965. The climate near the ground. 4th ed. Cambridge,

MA: Harvard University Press.

Griggs, R. F. 1956. Competition and succession on a Rocky

Mountain fellfield. Ecology 37: 8-20.

Hann, J. and R. Siiring. 1939. Lehrbuch der Meteorologie, ed.

5. Leipzig:

Hedberg, O. 1964. Features of Afroalpine plant ecology. Acta

Phytogeogr. Suec. 49: 1-144.

Heilborn, O. 1925. Contributions to the ecology of the

Ecuadorian pdramos with special reference to cushion

plants and osmotic pressure. Svensk Bot. Tidskr. 19:

153-170.

Hoffmann, J. A. J. 1975. Atlas climatico de America del Sur.

Budapest: WMO-UNESCO. 71.5

Jolls, c . L. 1975. Adaptive strategy of the cushion form in

arctic and alpine environments. Senior Honors Thesis,

University of Michigan.

Kbrner, C. and P. Cochrane. 1983. Influence of plant

physiognomy on leaf temperature on clear midsummer days 182

in the Snowy Mountains, south-eastern Australia. Oecol.

PI. 4: 117-124.

, P. Bannister, and A. F. Mark. 1986. Altitudinal

variation in stomatal conductance, nitrogen content and

leaf anatomy in different plant life forms in New

Zealand. Oecologia 69: 577-588.

, M. Neumayer, S. P. Menendez-Riedl, and A. Smeets-

Scheel. 1989. Functional Morphology of Mountain Plants.

Flora 182: 353-383.

Kuschel, G. 1958. Nuevos Cyldrorhininae de la Patagonia

(Col. Curculionoidea, Aporte 18). Investigaciones

Zoologicas Chilenas 4: 231-252.

Landolt, E. 1967. Gebirgs- und Tieflandsippen von

Bltitenpflanzen im Bereich der Schweizer Alpen. Bot.

Jaarb 86: 463-480.

Lona, F. 1963. II nanismo di molte piante alpine e

essenzialmente genetico. Giorn. Bot. Ital. 70: 560-564.

Minetti, J. L. and E. M. Sierra. 1989. The influence of

general circulation patterns on humid and dry years in

the Cuyo Andean Region of Argentina. International 183

Journal of Climatology. 9: 55-68.

Mooney, H. A. and W. D. Billings. 1961. Comparative

physiological ecology of arctic and alpine populations

of Oxyria dignya. Ecol. Monogr. 31: 1-29.

Neger, F. W. 1913. Biologie der Pflanzen. Stuttgart:

Norte, F. A. 1986. Characteristicas termodinSmicas y

aerologicas fundamentales en una situacidn de Zonda

intenso. GEO ACTA 13(2): 11-24.

Ralph, c. P. 1978. Observations on Azorella compacta

(Umbelliferae), a tropical andean cushion plant.

Biotropica 10: 62-67.

Rauh, W. 1939. tiber polsterformigen Wuchs. Nova Acta

Leop. 7: 265-540.

. 1940. Die Wuchsformen der Polsterpflanzen. Bot. Arch.

40: 289-374, 375-462.

Rochow, T. F. 1967. The ecology of Thlaspi alpestre in the

Central Rocky Mountains along altitudinal gradients.

Ph.D. diss., Duke University, 1-265. 184

Rudloff, W. 1981. World-Climates. Stuttgart:

Wlssenschaftliche Verlagsgesellschaft.

Ruthsatz, B. 1978. Las plantas en cojin de los semidesiertos

andinos del noroeste Argentino. Darwiniana 21: 491-539.

Sauberer, F. and 0. Hartel. 1959. Pflanze und Strahlung.

Leipzig.

Schimper, A. F. W. 1898. Pflanzengeographie auf

physiologischer Gmndlage. Jena.

Schinner, F. 1982. C02-Freisetzung, Enzymaktivitaten und

Bakteriendichte von Boden unter Spalierstrauchern und

Polsterpflanzen in der alpinen Stufe. Oecol. PI. 3: 49-

58.

Schlichting, C. D. 1986. The evolution of phenotypic

plasticity in plants. Ann. Rev. Ecol. Syst. 17: 667-

693.

Schroeter, C. 1908. Das pflanzenleben der alpen. Zurich:

Verlag von Albert Raustein.

Schroter, E., and H. Hauri. 1914. Versuch einer Ubersicht

der angiospermen Polsterpflanzen. Bot. Jahrb. Syst. 185

Suppliment. 50:

Smith, A. P. 1972. Notes on wind related growth patterns of

paramo plants in Venezuela. Biotropica 4: 10-16.

Spomer, G. G. 1964. Physiological ecology studies of alpine

cushion plants. Physiol. PI. (Copenhagen) 17: 717-724.

Troll, C. 1944. Strukturboden, Solifluktion und Frostklimate

der Erde. Geol. Rundschau 34: 545-694.

. 1947. Die Formen der Solifuktion und die periglaziale

Bodenabtragung. Erdkunde 1, 162-175.

. 1948. Der asymmetrische Aufbau der Vegetationszonen und

Vegetationsstufen auf der Nord- und Sudhalbkugel. Ber.

geobot. Forschungsinst. Rubel 1947: 46-83. Zurich.

. 1958. Zur physiolgnomik der Tropengewachse. Jber. Ges.

Fr. Ford, rhein. Friedrich-Wilhelms Univ., Bonn.

. 1959. Die tropischen Gebirge, ihre dreidimensionale

klimatische und pflanzengeographische Zonierung. Bonner

geogr. Abh. 25.

. 1960. The relationship between the climates, ecology 186

and plant geography of the southern cold temperate zone

and of the tropical high mountains. Proc. Roy. Soc.

London, Ser. B, Biol. Sci. 152: 529-532.

. 1966. Okologische Landschaftsforschung und

vergleichende Hochgebirgsforschung. Wiesbaden: F.

Steiner.

Turesson, G. 1931. The geographical distribution of the

alpine ecotype of some Euasiatic plants. Hereditas 15:

329-346.

Van Gardingen, P. and J. Grace. 1991. Plants and Wind.

Advances Bot. Res. 18: 189-253

Warming, E. 1909. Oecology of Plants. An Introduction to the

Study of Plant-Communities. Oxford: Oxford Clarendon

Press.

Warren-Wilson, J. 1952. Vegetation patterns associated with

soil movement on Jan Mayen Island. J. Ecol. 40: 249-

264.

Whitehead, R. H. 1951. Ecology of the altiplano of Monte

Maiella, Italy. J. Ecol. 39: 330-335. CHAPTER III

FLAVONOIDS OF MULINUM

(APIACEAE, HYDROCOTYLOIDEAE, MULINEAE)

Flavonoid chemistry of the genus Mulinum and selected species of Azorella and Gvmnophvton was studied. Both flavones and flavonols were identified. The presence of both flavones and flavonols weakens previous generalizations that the Mulineae contain only flavonols and is primitive compared to other tribes of Apiaceae. In general a more homogeneous flavonoid compound composition for the Apiaceae is suggested. In addition, both carbon and oxygen glycosides were isolated from Mulinum. All species contain

6,8-di-C-glycosyl chrysoeriol (flavone) and quercetin

(flavonol). Evolution of the flavonoid system within

Mulinum was inferred based on correlation with an hypothetical phylogeny based on morphology.

Secondary plant products have continued to provide valuable data for examining a wide range of taxonomic groups and systematic questions (Giannasi and Crawford, 1986). The

187 188

Apiaceae are rich in secondary compounds and many chemical studies have been completed, especially on the furanocoumarins, terpenoids, and polyacetylenes (Crowden et al., 1969). While Crowden et al. (1969) found flavonoids to be the only secondary constituent showing significant variation within the Apiaceae, relatively few flavonoid studies have been completed for intrafamilial groups (e.g.

Caucalideae: Harborne, 1971); previous studies have primarily represented familial reviews (Von Gerichten, 1901;

Oesterle & Wander, 1925; Bate-Smith, 1962; Matsushita &

Iseda, 1965; Harborne, 1967; Crowden et al., 1969; Harborne et al. 1969; Harborne, 1971; Harborne, 1975; Saleh et al.,

1983) .

Crowden et al. (1969) and Harborne (1967, 1971, &

1975), based on a survey of leaf flavonoids, reported that nearly all Umbelliferae species have either flavonols or flavones but not both (Daucus L. and Laserpitium L. containing exceptions). Furthermore, flavones are found almost entirely in taxa generally considered to be advanced or more specialized (e.g., Daucus, Scandix L., Torilis

Adans.) while flavonols predominate in the less advanced genera (e.g., Hydrocotyle L.).

On the basis of flavonoid data, Harborne (1971) divided the tribes of Apiaceae into two broad groups: the majority of eight tribes in which flavones are rare or absent; and four tribes (Apieae, Dauceae, Laserpiteae, and Scandiceae) 189

in which flavones are common or predominant (Table 8). This

division correlates reasonably well with Drude's (1897)

subfamilial divisions (Table 8). According to Harborne

(1971: 304), the Hydrocotyloideae (which lacks flavones) is

"less typically umbelliferous" (i.e, more primitive) than

the Apioideae (which contains flavones) and the

Saniculoideae (also lacking flavones), which exhibits a

distinctive chemistry in other regards (Hiller 1971).

Further, Harborne (1971) regards the Hydrocotyloideae as

probably evolving from the rest of the family at an earlier

time.

Flavonols have been identified for the tribe Mulineae, with quercetin and kaempferol found together or separately

in seven of the eight genera studied (Crowden et al. 1969).

No flavones have been identified within the tribe with the

exception of a questionable glycoflavone in Laretia acaulis

(Crowden et al. 1969). Of the nine Mulineae genera previously studied by Crowden et al. (1969), no genus was analyzed completely in terms of including all species. A single sample of Mulinum, M. nivalis (= M. leptacanthum

Phil.; see Chapter 1), was included within Crowden et al.

(1969). Crowden et al. (1969) identified the flavonols quercetin, kaempferol, and isorhamnetin and an additional phenolic compound, aesculetin, but no flavones.

The relative lack of flavonoid data for Mulinum suggests the need for a more intensive survey in the context 190

TABLE 8. Flavonoid composition of the Apiaceae following the classification of Drude (1897). * = tribes where flavones are absent or rare (Harborne, 1971).

No. of species with

Subfamily and Tribe Flavonol Flavone

Hydrocotyloideae:

*Hydrocotyleae 16 0

♦Mulineae 7 0

Saniculoideae:

♦Saniculeae 26 0

♦Lagoecieae 2 0

Apioideae:

♦Echinophoreae 2 0

Scandiceae 3 35

♦Coriandreae 3 0

♦Smyrnieae 21 5

Apieae 79 20

♦Peucedaneae 46 1

Laserpiteae 7 11

Dauceae 5 7 191

of the new revision of the genus. The purpose of this

study, therefore, was to: (1) document the flavonoids of all

species of Mulinum; (2) compare the flavonoid composition of

Mulinum with previous generalizations made for the Mulineae,

Hydrocotyloideae, and Apiaceae; and (3) infer the evolution

of the flavonoid system in Mulinum by correlation with an hypothetical phylogeny based on morphology.

MATERIALS AND METHODS

Acquisition of plant material— Plant samples were collected during three expeditions to Argentina and three expeditions to Chile during January-February 1987-1990 by personnel from laboratories of: the Department of Plant

Biology of The Ohio State University; Museo de La Plata,

Instituto de Cientifica, Mendoza, and Instituto Patagonico de Ciencias Naturales, Argentina; and the Universidad de

Concepcion, Chile. Five species of Mulinum were sampled from field collected material (Table 9). Specimens were air-dried in the field and vouchers are on deposit at OS with duplicates at CONC, GH, LP, and UC. The remaining three species of Mulinum were sampled from herbarium material (Table 9) with permission of the curators.

i

Extraction, isolation, and identification of flavonoids— Ground leaves and stems from each species were 192

TABLE 9. Collections of Mulinum species examined for flavonoid compounds. Collection numbers are those of Stuessy et al. (S), Zech & Contogiorgakis (ZC), or Zech et al. (Z) unless noted otherwise. A = Argentina, B = Bolivia, C = Chile; * = samples from herbarium material.

M. albovaginatum: A, 12 km from Las Lenas along road to Valle Hermoso, ZC 5, Laguna Diamonte, ZC 9; *M. crassifolium: C, San Pedro de Atacama, Casa blanca, Ricardi 2986 (OS); M. hallei: A, ± 155 km N Rio Gallegos, Ruta 3, ZC 21j M. leptacanthum: A, Chapelco, S 10000, Chapelco, S 10047, Chapelco, S 10048, Chapelco, S 10049, Chapelco, 5 10072, Lago La Kika, S 10129, C, Paso Pino Hachado, S 10162, 7 km E of Liucura on road to Paso Pino Hachado, Z 46; *M. microphyllum: A, 21 km S Garayalde, Correa & Constance 3970/3826 (UC); M. spinosumt A, 11 km SW Junin de Los Andes, S 10152, 31.5 km N Junin de Los Andes, S 10154, 10 kms N AluminA, S 10158, Copahue, S 10216, 33.5 km N Las Barrancas, S 10264, 6 kms W Los Molles, S 10269, 3 kms W Los Molles, S 10275, 2-4 kms NW Las Leftas, S 10318, El Sosneado, S 10354, 2 kms E customs, Punta de Vacas, S 10420, 5 km from Los Molles on road to Las Lenas, ZC 2, 15 km from Los Molles on road to Las Lehas, ZC 3, 8 km before Valle Hermoso, at monument, ZC 4, Banks of Rio Tordillo, ZC 6, Above Rio Tordillo, ZC 7, Valle Hermoso, 15 km from Rio Tordillo Road intersection, towards Las Lefias, ZC 8, Road to Laguna Diamonte, before 1st army station, ZC 10, Road to Laguna Diamonte, 4 km before first army station, ZC 11, Road to Laguna Diamonte, before Valle Diamonte, ZC 12, LOC, ZC 16, LOC, ZC 17, LOC, ZC 19, LOC, ZC 23, 50 km W of Commodoro/Rivadivia, ZC 26, C, Perez Caldera, Rio San Francisco, Z 29, 10 km before termas del Flaco, Z 33, Road to Angol from Nahuelbuta, near Rio Picoiquen, Z 42, Ranger's house below VoleAn Lonquimay, Z 44, ± 5 km form Coihaique, on road to Balmaceda, Z 49, Ruta 45 from Balmaceda, W & S towards Villa Cerro Castillo, Z 51, Before Laguna Fiqueroa, road from Puerto Natales to Torres Del Paine, Z 55; *M. ulicinumi B, Puna Patanca, Fiebrig 2601 (MO), Tafi, Sierra del Cajun, Venturi 10277, (US); M. valentini: A, 7 km S of Hotel Lemarchand, Ruta 3, ZC 18, 85 km E of Sarmiento, Ruta 26, ZC 24, 85 km E of Sarmiento, Ruta 26, ZC 25. extracted with 85% and then 50% aqueous methanol. The combined extracts were evaporated to dryness under vacuum, taken up in methanol and applied to sheets (46 X 50) cm of

Whatman 3MM chromatographic paper. The 2D chromatograms were developed with tert-butyl alcohol: acetic acid: water

3:1:1 (TBA) and 15% acetic acid (HOAc) and examined over ultraviolet light following standard procedures (Mabry,

Markham, and Thomas, 1970). In some cases, the same chromatograms were run a second or even a third time in TBA and HOAc for maximum compound separation. Individual flavonoids were obtained by elution of compounds from paper with methanol and when necessary further purified by thin layer chromatography (TLC) using precoated polyamide DC-6

(Macherey-Nagel). The solvent systems were: 1,2 dichloro- ethane: MEK: MeOH: H20: Formic Acid (50:25:20:4:0.5) and

Chloroform: MeOH: MEK: Acetone (20:10:5:1). Identification of the flavonoids was performed by ultraviolet spectral analysis and when possible by comparison with standard Rf values (Mabry, Markham, and Thomas, 1970; Neuman,

Timmermann, and Mabry, 1979; Markham, 1982). Mono- and diglycosides were determined by chromatogram position.

RESULTS

Eleven flavonoids were isolated and identified from the

8 species of Mulinum (Tables 10, 11). The flavonoid system in TABLE 10. identities of flavonoids isolated from Mulinum.

Compound Tentative Rf UV Spectral Data designation Identity TBA/HOAc MeOH NaOMe AlCl, A1C13/HC1 NaOAc

1 6,8-di-C—glycosyl Chrysoeriol -- 345 405 395 387 348

2 Luteolin 7-O-diglycoside ------340 410 420 385 350

3 4 ' methoxy flavone ------

4 4'7-dihydroxyflavone 7-O-diglycoside 0.634/0.577 315 360 315 315 310

5 Quercetin 3-O-diglycoside 0.456/0.409? 350 405 412 390 355

6 Quercetin 3,7-O-diglycoside ------345 405 412 395 345 7 Isorhamnetin-3, 7-O-diglycoside ------350 400 445 445 350

8 Flavonol 3-0-monoglycoside ------

9 Flavonol 3-0-diglycoside ------

10 Flavonol 7-0-monoglycoside ------

11 Tamarixten aglycone ——— 360 435 435 420 415

to 4 k TABLE 11. Distribution of flavonoids in Mulinum.

Flavone Flavones Flavonols Aglycone

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

M. albovaginaturn X X

M . crassifolium X X

M. hallei X X X X X X X

M. leptacanthum X X X

M. microphyllum XX

M . spinosum XXX X X X X X

M. ulicinum X X

M. valentini X X 196

Mulinum is based on diglycosides of the flavones luteolin and chrysoeriol, monoglycosides of the flavonol

isorhamnetin, and mono- and diglycosides of the flavonol quercetin. Chrysoeriol glycosides are carbon glycosylated, while luteolin and the flavonols are oxygen glycosylated.

Flavonol glycosides are substituted at the 3 and 7 positions while flavone glycosides are substituted at the 6, 7, and 8 positions.

Two compounds, the flavone 6,8-di-C-glycosyl chrysoieriol (#1) and a derivatives of the flavonol quercetin (#5), were found in all species of Mulinum, with no differences being apparent between species. C- glycosylated chrysoeriol co-occurred with O-glycosylated derivatives of quercetin in most plants studied (Table 11).

Flavonoids isolated from other genera in the Mulineae are included in Table 12, including results obtained in the present study and from Crowden et al. (1969). Although flavonols are more common than flavones in the taxa studied, the number of flavones isolated within the Mulineae has been increased. A single questionable flavone was published previously for Laretia (Crowden et al., 1969).

DISCUSSION

Subfamilial and tribal flavonoid variation— The presence of both flavones and flavonols within Mulinum and Table 12. Distribution of flavones and flavonols in some genera of Mulineae. Collection numbers are those of Stuessy et al. (S), Zech & Contogiorgakis (ZC), or Zech et al. (Z). A = Argentina, C = Chile.

Flavonol Flavone Reference or Taxon Que Kae Iso Chr Mf Gf F70M Voucher

Asteriscium chilense Cham. & Schlect. x" Crowden et al. 1969 Asteriscium glaucum Hieron. & Wolff X A, Cerro Chapelco, S 10017. Azorella monantha Clos ex Gay X X A, 204 kms NW of Las Lenas, S 10319C, Paso Pino Hachado, S 10163B, ± 30 km from the border, toward Punta Delgada, Ruta 255, Z 53. Azorella patagonica Speg. A, 2 km from Calafate, shore of Lago Argentino, ZC 20. Azorella trifurcata (Gaertn.) Pers. A, 5 km from Los Molles, on the road to Las Lenas, ZC 1, C, Near Volcan Longuimay, Z 43, ± 20 km S of Punta Delgada on Ruta 255 to Punta Arenas, Z 54. Diposis saniculifolia Crowden et al. 1969 Table 12. (Continued).

Domevkoa amolexicaulis X Crowden et al. 1969 Gvmnoohvton isatidicaroum (Presl ex DC.) Math. & Const. X X C, Perez Caldera, Rio San Francisco, Z 30. Gvmnoohvton oolvceohalum (Gillies & Hook.) Clos in Gay X X X X A, ± 50 kms E of Puente de Inca on Ruta 7 to Mendoza, S 10377. 2 km from Hotel Termas Villavicencio on road to Uspallata, ZC 13. Gvmnoohvton soinosissumum Phil. X X C, 1.6 kms E of Puguios on rout to Tinocrasta. Stuessv & Marticorena 9825. Hermas villosa X Crowden et al. 1969 Huanaca andina X X Crowden et al. 1969 Laretia acaulis • Crowden et al. 1969 Mulinum nivale (=M. leotacanthum) XX X Crowden et al. 1969 Soananthe oaniculata XX Crowden et al. 1969 *Que = quercetin; Kae = kaempferol; Iso = isorhamnetin; Chr = 6,8-di-C-glycosyl chrysoeriol; Mf = 4'methoxy flavone; Gf = glycoflavone; F70M = flavone 7-0-monoglycoside. 199

the species of Gymnophyton and Azorella herein studied

differ from the previous generalizations (Crowden et al.

1969; Harborne, 1971) that nearly all species have either

flavonols or flavones but not both and that flavones are

found almost entirely in taxa generally considered to be

advanced or more specialized while flavonols predominated in

the less advanced taxa. While admittedly based on a small

sample, the present results suggest more flavonoid variation

than previously documented for the Mulineae, and call into

question the report that presence of flavonoids only is primitive within the family.

Generic and infrageneric flavonoid variation in

Mulinum— Mulinum appears similar to other genera of the

Mulineae (Table 12), previously studied in part by Crowden et al. (1969), in the production of the flavonol quercetin.

Kaempferol was also commonly identified in many Mulineae genera, including Mulinum (Crowden et al, 1969; Table 12).

The present study did not isolate kaempferol from Mulinum

(Table 11), including M. leptacanthum from which Crowden et al. (1969) previously obtained it (Table 12). Crowden et al. (1969) also isolated isorhamnetin from M. leptacanthum

(Table 12). Isorhamnetin was isolated from M. spinosum but was not found in M. leptacanthum or the other six remaining species (Table 11). Each species flavonoid profile contained compounds which were unidentifiable. Isorhamnetin and kaempferol may have been present in quantities too small 200

for identification, especially in the case of AT.

leptacanthum.

The flavonoid compound profiles of the eight Mulinum

species studied are simple, containing relatively few

compounds. The greatest number of compounds were isolated

from M. hallei and M. spinosum (Table 11). The larger

number of compounds isolated for M. spinosum is probably reflective of genetic variation as seen by morphology (see

Chapter 1) as well as flavonoids, while M. hallei's large number of compounds is considered more derived. Fewer compounds were isolated from M. microphyllum as compared to

M. hallei (Table 11). In addition, few compounds were isolated from M. crassifolium, M. leptacanthum, M. ulicinum, and M. valentini. Mulinum microphyllum is suggested as an allopolyploid with M. hallei and M. valentini as parents

(see Chapter 1). Mulinum hallei is clearly one parent to M. microphyllum, while the evidence supporting M. valentini as the other parent is less strong. The apparent reduction in flavonoid production in M. microphyllum and M. valentini

(Table 11), adds limited support, because of sample size, for M. valentini to be one of the parents of the polyploid

M. microphyllum (see Chapter 1).

The species of Azorella and Gymnophyton studied thus far suggest some possible generic trends. Flavonols and flavones have been isolated from both genera (Table 12).

Azorella may be similar to other genera of the Mulineae in 201 that both quercetin and isorhamnetin were isolated (Table

12). Azorella differs from both Mulinum (Table 11) and

Gymnophyton (Table 12) in producing more kinds of flavonols and Gymnophyton appears similar to Mulinum in the production of both chrysoeriol and quercetin as well as relatively few

compounds (Table 12).

Evolution of the flavonoid system in Mulinum— Mulinum

is considered holophyletic, with Azorella spinosa Ruiz Lopez

& Pavon considered the closest living relative of Mulinum

(see Chapter 1). Comparison of the flavonoid system of

Mulinum with that of the taxa of Gymnophyton isolated thus far (Table 12) suggests Gymnophyton as the closer relative.

However, different chromosome number, presence of a petal gland, and leaf morphology and position in Gymnophyton, suggest that Mulinum's similar flavonoid composition may have evolved in parallel and not from Gymnophyton (see

Chapter 1).

Relating the flavonoid system to an existing hypothetical phylogeny based on morphology (Fig. 20) has provided additional insights for the group or groups being studied (Pacheco et al., 1985; 1991). Mulinum spinosum is considered the most primitive species based on habit and morphological features (see Chapter 1). Eight of the 11 flavonoids are synthesized in this species (Table 11). If

M. spinosum is primitive within Mulinum, then the flavonoids found within it, 1, 2, 3, 5, 6, 7, 8, and 10 (Table 11) may HYP 1,5 CRA 1.8 ULI 1.8 ALB 1.8 VAL 1.3—,4 —,8 ,8 —,8,9 H A L 1.8 MIC i.».y LEP 1-3 ,5-8,10 SPI

Fig. 20. Cladogram of taxa of Mulinum. The names of taxa are represented by the first three letters of the specific epithets. Numbers correspond to the flavonoids listed in Table 10. 202 203

be considered primitive.

The evolution of individual flavonoid compounds appears

conservative. Flavonoids 2, 3, 6, 7, and 8 were apparently

lost relatively early within M. leptacanthum with only

flavonoids 1 and 5 remaining and 11 derived (Fig. 20).

Compounds 3, 6, and 8 were apparently lost at three

different branching points, only remaining as reversals in

M. hallei (Fig. 20). Flavonoids l and 5 are the only

primitive compounds conserved in all taxa studied (Table

11). This reduction in flavonoid production may be

correlated with a general reduction in leaf, and plant size,

culminating in the four Mulinum cushion species (Fig. 20).

The cushion taxa are considered to be derived. These

species share a compact cushion habit and general reduction

in morphological and reproductive characteristics which is

clearly adaptive to the extreme environments in which they

occur (see Chapter 2). The reduction in flavonoid profile

to two compounds is consistent for three cushion species: M.

albovaginatum, M. microphyllum, and M. valentini. However,

7 of the eleven flavonoids are synthesized in the fourth, M. hallei, with compounds 4 and 9 being derived. The greater

conservation of primitive flavonoid compounds as well as the

two derived compounds within M. hallei may indicate a greater specialization as compared to the remaining three

cushion taxa.

Advanced species within a genus have been suggested as 204 having fewer and structurally more simple compounds (Mabry,

1974; Wells & Bohm, 1980), a greater flavonoid complexity

(Whalen, 1978; Crawford and Smith, 1983a; Stuessy and

Crawford, 1983), or no trend in either direction (Crawford and Smith, 1983b; Pacheco et al., 1985), indicating that flavonoid evolution is group-specific. The same group specificity has been shown to exist in the evolution of C- glycosylated flavones (Harborne, 1977; Gornall & Bohm, 1978;

Valant-Vetschera, 1985).

C-glycosylated chrysoeriol co-occurs with O- glycosylated derivatives of quercetin in most Mulinum studied (Tables 11 & 12). The co-occurrence of C- glycosylated flavones with flavonoid O-glycosides, especially quercetin, commonly occurs in the Angiosperms, but is rare in the Apiaceae (Valant-Vetschera, 1985). The

C-glycosylation of chrysoeriol appears to be rare in general

(Valant-Vetschera, 1985). Maintenance of C-glycosyl flavone production in Mulinum, therefore, would suggest that the flavonoid composition of Mulinum is relatively primitive within the family.

ACKNOWLEDGEMENTS

The author thanks Drs. Tod F. Stuessy for his assistance with this research. Thanks also go to: Daniel J.

Crawford for the use of his lab, help in compound 205

identification, and general comments; Fred Sack for general

suggestions; Marla Marta Cigliano, Claudio Contogiorgakis,

Jorge Crisci, Mario Gentili, Patricia Gentili, Liliana

Katinas, Juan Jose Morrone, Alegandra Moschetti, Alberto

Ripalta, Fidel Alegandro Roig, Tod F. Stuessy, and Ana Marla

Zavattieri for field and project assistance in South

America. This paper is a part of a Thesis presented in

partial fulfillment of the requirements for the Ph.D. at The

Ohio State University and supported by NSF grant BSR-

8815311, an OSU Tinker Foundation Field Research Grant, and

a John Ayers Herbarium Travel Award from the American

Society of Plant Taxonomists.

LITERATURE CITED

Crawford, D. C., and E. B. Smith. 1983a. The distribution of

anthochlor floral pigments in North American Coreopsis

(Compositae): taxonomic and phyletic interpretations.

Amer. J. Bot. 70: 355-362.

. 1983b. Leaf flavonoid chemistry of North American

Coreopsis (Compositae): intra- and intersectional

variation. Bot. Gaz. 144: 577-583.

Crowden, R. K., J. B. Harborne and V. H. Heywood. 1969.

Chemosystematics of the Umbelliferae— A General Survey. 206

Phytochemistry 8: 1963-1984.

Drude, C. G. 0. 1897-1898. Umbelliferae. In Die natilrlichen

Pflanzenfamilien, eds. A. Engler & K. Prantl, 3(8): 63-

250. Leipzig: Wilhelm Engelmann.

Giannasi, D. E. and D. J. Crawford. 1986. Biochemical

Systematics II. A Reprise. Evol. Biol. 20: 25-248.

Harborne, J. B. 1967. Comparative Biochemistry of the

Flavonoids. London: Academic Press.

. 1971. Flavonoid and phenylpropanoid patterns in the

Umbelliferae. In The Biology and Chemistry of the

Umbelliferae, V. H. Heywood, ed., 293-314. New York:

Academic Press.

. 1975. Biochemical systematics of flavonoids. In The

Flavonoids, J. B. Harborne, T. J. Mabry, and H. Mabry,

eds., 1057-1095. London: Academic Press.

. 1977. Flavonoids and the evolution of the Angiosperms.

Biochemical Systematics and Ecology 5: 7-22.

Harborne, J. B., V. H. Heywood, and C. A. Williams. 1969.

Distribution of Myristian in the seeds of the 207

Umbellifereae. Phytochemistry 8: 1729-1732.

Hiller, K. 1971. Chemosystematics of the Saniculoideae. In

The Biology and Chemistry of the Umbelliferae, V. H.

Heywood, ed., 369-384. New York: Academic Press.

Mabry, T. J. 1974. Chemistry of disjunct taxa. In Chemistry

in botanical classification, G. Bendz, J. Santesson,

and V. Runnstrom-Reio, eds., 63-66. Nobel Symposium 25,

New York: Academic Press.

Mabry, T. J., K. R. Markham, and M. B. Thomas. 1970. The

Systematic Identification of Flavonoids. New York:

Springer-Verlag.

Markham, K. R. 1982. Techniques of Flavonoid Identification.

New York: Academic Press.

Mathias, M. E. and L. Constance. 1962. A revision of

Asteriscium and some related hydrocotyloid

Umbelliferae. Univ. Calif. Publ. Bot. 33: 99-184.

Matsushita, A. and S. Iseda. 1965. Nippon Nogei Kagaku

Kaishi 39:.

Neuman, P. R., B. N. Timmermann, and T. J. Mabry. 1979. 208

Laboratory Manual for the Systematic Identification of

Flavonoids. Privately Published.

Oesterle, 0. A. and G. Wander. 1925. Helv. Chim. Acta 8:

Pacheco, P., D. J. Crawford, T. F. Stuessy, and M. Silva O.

1985. Flavonoid evolution in Robinsonia (Compositae) of

the Juan Fernandez Islands. Amer. J. Bot. 72: 989-998.

. 1991. Flavonoid evolution in Dendroseris (Compositae,

Lactuceae) from the Juan Fernandez Islands, Chile.

Amer. J. Bot. 78: 534-543.

Saleh, N. A. M., S. I. El-Negoumy, M. N. El-Hadidi, and H.

A. Hosni. 1983. Comparative study of the flavonoids of

some local members of the Umbelliferae. Phytochemistry

22: 1417-1420.

Stuessy, T. F., and D. J. Crawford. 1983. Flavonoids and

phylogenetic reconstruction. Plant Systematics and

Evolution 143: 83-107.

Von Gerichen, E. 1901. Liebig's Ann 318:

Whalen, M. D. 1978. Foliar flavonoids of Solanum section

Androceras: a systematic survey. Syst. Bot. 3: 257-276. Wells, E. F., and B. A. Bohm. 1980. Chemotaxonomic studies

in the Saxifragaceae S.I. 15. The flavonoids of

subsection Villosae section Heuchera in the genus

Heuchera. Canad. J. Bot. 58: 1459-1463. LIST OF REFERENCES

Adams, E. N., III. 1972. Consensus techniques and the comparison of taxonomic trees. Syst. Zool. 21: 390-397.

. 1986. N-trees as nestings: complexity, similarity, and consensus. Journal of Classification 3: 299-317.

Armesto, J. J., M. D. Arroyo, and C. VillagrSn. 1980. Altitudinal distribution, cover and size structure of umbelliferous cushion plants in the high Andes of Central Chile. Acta OEcologica/OEcologia Generalis 1: 327-332.

Auer, V. 1958. Ann. Acad. Sci. Fenn. Ser. A, III. 50: 1.

. 1960. The quaternary history of Fuego-Patagonia. Proc. Roy. Soc. London, Ser. B, Biol. Sci. 152: 507-516.

Baeza R., V. M. 1921. Los nombres vulgares de las plantas silvestres de Chile i su concordancia con los nombres cientificos: Con observaciones sobre la aplicacion t&nica i medicinal de algunas especies. Santiago: Sociedad Imp. y Lit. Universo.

Baillon, H. E. 1880 [1879]. Ombellif&res. In Histoire des plantes, ed. H. E. Baillon, 7: 84-256. Paris: Hachette & Cie.

Barry, R. G., and C. C. Van Wie, 1974. Topo- and microclimatology in alpine areas. 73-83. In Ives, J. D., and R. G. Barry, eds, Arctic and alpine environments. London: Methuen and Co., Ltd.

Baust, J. G. 1976. temperature buffering in an arctic microhabitat. Annales of the Entomological Society of America 69: 117-119.

Bell, C. R. 1971. Breeding systems and floral biology of the Umbelliferae or evidence for specialization in unspecialized flowers. In The Biology and Chemistry of the Umbelliferae, ed. V. H. Heywood, 93-107. London: Academic Press.

210 211

Bell, C. R. & L. Constance. 1957. Chromosome numbers in Umbelliferae. II. Amer. J. Bot. 44: 565-572.

. I960. Chromosome numbers in Umbelliferae. II. Amer. J. Bot. 47: 24-32.

. 1966. Chromosome numbers in Umbelliferae. III. Amer. J. Bot. 53: 512-520.

Bentham, G. 1867. In Genera plantarum, eds. G. Bentham & J. D. Hooker, 1: 859-931, 1008-1009. London: Lovell Reeve & Co., Williams & Norgate.

Bhakuni, D. S., M. Bittner, C. Marticorena, M. Silva, E. Weldt, & H. Hoeneisen. 1976. Screening of Chilean plants for anticancer activity. I. Lloydia 39: 225-243.

Billings, W. D. and H. A. Mooney. 1968. The ecology of arctic and alpine plants. Biol. Rev. (London) 43: 481- 529.

Bland, R. G. 1978. How to know the insects, ed. 3, i-409. Dubuque: Wm. C. Brown Co.

Borror, D. J., D. M. De Long, & C. A. Triplehorn. 1981. An introduction to the study of insects, ed. 5, i-827. Philadelphia: Saunders College Publishing.

Bremer, K. 1990. Combinable component consensus. Cladistics 6: 369-372.

Burgos, J. J. and A. L. Vidal, 1951. Los climas de la Repdblica Argentina, segdn la nueva clasifcacion de Thornthwaite. Meteoros 1: 3-32.

Candolle, A. P. de. 1829. Observations sur la famille des Ombellifdres. Collection de Memoires 5: 1-84. Paris: Treuttel et Wtirtz.

. 1830. Umbelliferae. In Prodromus systematis naturalis regni vegetabilis, ed. A. P. de Candolle, 4: 55-250. Paris: Treuttel et Wtirtz.

Cavanilles, A. J. 1799. Icones et descriptiones plantarum 5: 58-59. Madrid: Typographia regia.

Cernusca, A. 1976. Bestandesstruktur, Biokima und Energiehaushalt von alpinen Zwergstrauchbestanden. Oecol. PI. 11: 71-102.

Chodat, R. H. & E. Wilczek. 1902. Contributions a la Flora la Rgpublique Argentine: Ombellifdres. Bull. Herb. 212

Boissier. 2: 525-528.

Clapperton, C. M., and D. E. Sugden. 1988. Holocene glacier flucturations in South America and Antarctica. Quaternary Science Reviews &: 185-198.

Clausen, J., D. D. Keck, and W. M. Heisey, 1940. Experimental studies on the nature of species. I. effect of varied environments on western North American plants. Publ. Carnegie Inst. Wash. 520: 1-452.

. 1945. Experimental studies on the nature of species. II. Plant evolution through amphiploidy and autoploidy, with examples from the Madiinae. Publ. Carnegie Inst. Wash. 564: 1-174.

. 1948. Experimental studies on the nature of species III. Environmental responses of climatic races of Achillea. Publ. Carnegie Inst. Wash. 581: 1-129.

Clebsch, E. E. C. 1960. Comparative morphological and physiological variation in arctic and alpine populations of Trisetum spicatum. Ph.D. dissertation, Duke University, 1-241.

Clos, D. 1847. Umbeliferas. In Historia fisica y politica de Chile, Botanica: Flora chilena, ed. C. Gay, 3: 61-145. Santiago: Museo historia natural.

Coe, M. J. 1967. Microclimate and animal life in the equatorial mountanins. Zoolog. Africana 4: 101-128.

Constance, L. 1965. Some subtractions from the Umbelliferae of South America. Ann. Missouri Bot. Gard. 52: 274-280.

. 1971. History of the classification of Umbellifereae (Apiaceae). In The biology and chemistry of the Umbelliferae, ed. V. H. Heywood, 1-12. London: Academic Press.

. 1988. Umbellifereae. In Flora Patagonica. Parte V: Dicotiledoneas Dialipetalas (Oxalidaceae a Cornaceae), ed. M. N. Correa, 310-379. Buenos Aires: Instituto Nacional De Technologia Agropecuraria.

Constance, L., T-I. Chuang, & C. R. Bell. 1971. Chromosome numbers in Umbelliferae. IV. Amer. J. Bot. 58: 577-587.

. 1976. Chromosome numbers in Umbelliferae. V. Amer. J. Bot. 63: 608-625.

Crawford, D. C., and E. B. Smith. 1983a. The distribution of 213

anthochlor floral pigments in North American Coreopsis (Compositae): taxonomic and phyletic interpretations. Amer. J. Bot. 70: 355-362.

. 1983b. Leaf flavonoid chemistry of North American Coreopsis (Compositae): intra- and intersectional variation. Bot. Gaz. (Crawfordsville) 144: 577-583.

Cronquist, A. 1978. Once again, what is a species? In Biosystematics in Agriculture, ed. L. V. Knutson, 3-20. Montclair, NJ: Allenheld Osmun.

Crowden, R. K., J. B. Harborne, & V. H. Heywood. 1969. Chemosystematics of the Umbelliferae: A general survey. Phytochemistry 8: 1963-1984.

Curran, C. H. 1934. The families and genera of North American Diptera, 1-512. New York: Ballou Press.

. 1948. Experimental studies on the nature of species III. Environmental responses of climatic races of Achillea. Publ. Carnegie Inst. Wash. 581: 1-129.

Drude, C. G. 0. 1897-1898. Umbelliferae. In Die natiirlichen Pflanzenfamilien, eds. A. Engler & K. Prantl, 3(8): 63-250. Leipzig: Wilhelm Engelmann.

Espinosa, R. 1932. Okologische Studien iiber Kordillerenpflanzen. Bot. Jaarb. 65: 120-211.

Fries, R. E. 1905. Zur Kenntnis Der Alpinen Flora Im Nordlichen Argentinien. Nova Acta Regiae Soc. Sci. Upsal. Ser. IV, 1(1): 1-205.

Gandoger, M. M. 1912. Manipulus plantarum novarum praecipue Americas australioris. Bull. Soc. Bot. France. 59: 704-710.

Geiger, R. 1950. Das Lima der bodennahen Luftschicht. Die Wissenschaft, 78, ed. 3. Braunschweig.

. 1965. The climate near the ground. 4th ed. Cambridge, MA: Harvard University Press.

Giannasi, D. E. and D. J. Crawford. 1986. Biochemical Systematics II. A Reprise. In M. H. Hecht, B. Wallace, and G. T. Prance [eds.], Evolutionary Biology 20: 25- 248.

Grant, V. 1981. Plant Speciation, ed. 2. New York: Columbia University Press. 214

Griggs, R. F. 1956. Competition and succession on a Rocky Mountain fellfield. Ecology 37: 8-20.

Grisebach, A. H. R. 1874. Plantae Lorentzianae: Umbelliferae. Abh. Konigl. Ges. Wiss. Gottingen. 19: 105-108.

Halliday, P., R. D. Meikle, J. Story, & H. Wilkinson. 1980. Draft Index of Author Abbreviations: Compiled at The Herbarium Royal Botanic Gardens, Kew. Basildon: Her Majesty' Stationary Office.

Hann, J. and R. Siiring. 1939. Lehrbuch der Meteorologie, ed. 5. Leipzig:

Harborne, J. B. 1967. Comparative Biochemistry of the Flavonoids. London: Academic Press.

. 1971. Flavonoid and phenylpropanoid patterns in the Umbelliferae. In The Biology and Chemistry of the Umbelliferae, ed. V. H. Heywood, 293-314. London: Academic Press.

. 1975. Biochemical systematics of flavonoids. In J. B. Harborne, T. J. Mabry, and H. Mabry [eds.], The Flavonoids. London: Academic Press.

. 1977. Flavonoids and the evolution of the Angiosperms. Biochemical Systematics and Ecology 5: 7-22.

Harborne, J. B., V. H. Heywood, and C. A. Williams. 1969. Distribution of Myristian in the seeds of the Umbellifereae. Phytochemistry 8: 1729-1732.

Hedberg, 0. 1964. Features of Afroalpine plant ecology. Acta Phytogeogr. Suec. 49: 1-144.

Heilborn, O. 1925. Contributions to the ecology of the Ecuadorian p£ramos with special reference to cushion plants and osmotic pressure. Svensk Bot. Tidskr. 19: 153-170.

Heusser, C. J. 1971. Umbelliferae. In Pollen and Spores of Chile. 59-60, pi. 56. Tucson: The University of Arizona Press.

Hicken, C. M. 1922. Sertularium Andinum: Umbelliferae. Darwiniana 1: 61-62.

Hieronymus, G. H. E. W. 1881. Sertum Sanjuaninum: Umbeliferas. Bol. Acad. Nac. Ci. 4: 26-29. 215

Hiller, K. 1971. Chemosystematics of the Saniculoideae. In The Biology and Chemistry of the Umbelliferae, ed. V. H. Heywood, 369-384. New York: Academic Press.

Hiroe, M. 1979. Umbelliferae of World. Tokyo City: Ariake Book Company.

Hoffmann, J. A. J. 1975. Atlas climatico de America del Sur. Budapest: WMO-UNESCO. 71.5

Holmgren, P. K., N. H. Holmgren, Si L. C. Barnett. 1990. Index Herbariorum. Part I: The Herbaria of the World, ed. 8. New York: New York Botanical Garden.

Hooker, W. J. 1830. On the plants of the natural order Umbelliferae, detected by Dr. Gillies in the extratropical parts of South America. Bot. Misc. 1: 323-335.

Hooker, W. J. & G. A. W. Arnott. 1841. Chili: Umbelliferae. In The Botany of Captain Beechey's Voyage, eds. W. J. Hooker & G. A. W. Arnott, 25-27. London: Henry G. Bohn.

Johnston, I. M. 1929. I. Papers on the flora of northern Chile. II. Some undescribed species from Peru. Contr. Gray Herb. 85: 1-180.

Jolls, C. L. 1975. Adaptive strategy of the cushion form in arctic and alpine environments. Senior Honors Thesis, University of Michigan.

Korner, C. and P. Cochrane. 1983. Influence of plant physiognomy on leaf temperature on clear midsummer days in the Snowy Mountains, south-eastern Australia. Oecol. PI. 4: 117-124.

, P. Bannister, and A. F. Mark. 1986. Altitudinal variation in stomatal conductance, nitrogen content and leaf anatomy in different plant life forms in New Zealand. Oecologia 69: 577-588.

, M. Neumayer, S. P. Menendez-Riedl, and A. Sraeets- Scheel. 1989. Functional Morphology of Mountain Plants. Flora 182: 353-383.

Krause, E. H. L. 1904. Umbelliferae. In J. Sturm/s Flora von Deutschland, ed. 2, 12: 24-164. Stuttgart: K. G. Lutz.

Kuntze, C. E. 0. 1898. Umbelliferae. In Revisio generum plantarum, ed. C. E. 0. Kuntze, 3(2): 110-115. New York: Gust. E. Schechert. 216

Kurtz, F. 1893. Bericht tiber zwei Reisen zum Gebiet des oberen Rio Salado (Cordillera de Mendoza), ausgefiihrt in den Jahren 1891-1892 und 1892-1893. Verh. Bot. Vereins Prov. Brandenburg 34: 95-120.

Kuschel, G. 1958. Nuevos Cyldrorhininae de la Patagonia (Col. Curculionoidea, Aporte 18). Investigaciones Zoologicas Chilenas 4: 231-252.

Lamarck, J. B. A. P. M. de. 1789. Hydrocotyle. Encylopkdie Meth6dique, Botanique 3: 151-156. Paris: Panckoucke et Plomtreux. t Landolt, E. 1967. Gebirgs- und Tieflandsippen von Bliitenpflanzen im Bereich der Schweizer Alpen. Bot. Jaarb 86: 463-480.

Lanjouw, J. & F. A. Stafleu. 1954. Index herbariuorum. Part II. Collectores. A-D. Utrecht: IAPT.

Lawrence, G. H. M., A. F. GUnther Buchheim CK, G. S. Daniels, & H. Dolezal. 1968. Botanico-Periodicum- Huntianum. Pittsburgh, PA: Hunt Botanical Library.

Lona, F. 1963. II nanismo di molte piante alpine e essenzialmente genetico. Giorn. Bot. Ital. 70: 560-564.

Lorentz, P. G. & G. Niederlien. 1881. Umbelliferae. Expedicion al Rio Negro (Patagonia) Entrega II. - Botknica, 221-224. Buenos Aires: Ostwald Y Martinez.

Mabry, T. J. 1974. Chemistry of disjunct taxa. In Chemistry in botanical classification, eds. G. Bendz, J. Santesson, and V. Runnstrom-Reio, 63-66. Nobel Symposium 25. New York: Academic Press.

Mabry, T. J., K. R. Markham, and M. B. Thomas. 1970. The Systematic Identification of Flavonoids. New York: Springer-Verlag.

Macloskie, G. 1905. Umbelliferae. Reports of the Princeton University expeditions to Patagonia. 1896-1899: Botany Part V Flora Patagonica. Section 3. Cactaceae-Compositae 8: 619-643. Lancaster, PA: New Era Printing Co.

Margush, T. & R. R. McMorris. 1981. Consensus n-trees. Bulletin of Mathematical Biology 43: 239-244.

Markgraf V. & H. L. D'Antoni. 1978. Umbelliferae. In Pollen Flora of Argentina. 96, pi 41. Tucson: The University of Arizona Press. 217

Markham, K. R. 1982. Techniques of Flavonoid Identification. New York: Academic Press.

Martinez, S. 1989. El Gdnero Azorella (Apiaceae-Hydrocotyloideae) en La Argentina. Darwiniana 29: 139-178.

Mathias, M. E. 1971. Systematic survey of New World Umbelliferae. In The biology and chemistry of the Umbelliferae, ed. V. H. Heywood, 13-29. London: Academic Press.

Mathias, M. E. & L. Constance. 1962. A revision of Asteriscium and some related Hydrocotyloid Umbelliferae. Univ. Calif. Publ. Bot. 33: 99-184.

Matsushita, A. and S. Iseda. 1965. Nippon Nogei Kagaku Kaishi 39:.

Mayr, E. 1969. The biological meaning of species. Biol. J. Linn. Soc. 1: 311-320.

. 1992. A local flora and the biological species concept. Amer. J. Bot. 79: 222-238.

Mercer, J. H. 1972. Chilean glacial chronology 20,000 to 11,000 Carbon-14 years ago: Some global comparisons. Science 176: 1118-1120.

Mercer, J. H., & C. Laugenie. 1973. Glacier in Chile ended a major readvance about 36,000 years ago: Some global comparisons. Science 182: 1017-1019.

Minetti, J. L. and E. M. Sierra. 1989. The influence of general circulation patterns on humid and dry years in the Cuyo Andean Region of Argentina. International Journal of Climatology. 9: 55-68.

Montero, E. A. C., & M. J. Silveira. 1983. Radiocarbon chronology of a tephra layer in Rio Traful Valley, Province of Neuqudn, Argentina. In Quaternary of South America and Antartic Peninsula. 1: 135-151. Rotterdam: Balkema.

Montes, M. & T. Wilkomirsky. 1985. Medicina tradicional Chile. Concepcidn: Universidad de Concepcidn.

Mooney, H. A. and W. D. Billings. 1961. Comparative physiological ecology of arctic and alpine populations of Oxyria dignya. Ecol. Monogr. 31: 1-29.

Moore, D. M. 1971. Chromosome studies in the Umbelliferae. 218

In The biology and chemistry of the Umbelliferae, ed. V. H. Heywood, 233-255. London: Academic Press.

Munoz Pizarro, C. I960. Las especies de plantas descritas por R. A. Philippi in el siglo XIX, 1-189. Santiago: Ediciones de la Universidad de Chile.

Neger, F. W. 1913. Biologie der Pflanzen. Stuttgart:

Neuman, P. R., B. N. Timmermann, and T. J. Mabry. 1979. Laboratory Manual for the Systematic Identification of Flavonoids.

Norte, F. A. 1986. Characteristicas termodin&micas y aeroldgicas fundamentales en una situacidn de Zonda intenso. GEO ACTA 13(2): 11-24.

Oesterle, 0. A. and G. Wander. 1925. Helv. Chim. Acta 8:

Pacheco, P., D. J. Crawford, T. F. Stuessy, and M. Silva 0. 1985. Flavonoid evolution in Robinsonia (Compositae) of the Juan Fernandez Islands. Amer. J. Bot. 72: 989-998.

. 1991. Flavonoid evolution in Dendroseris (Compositae, Lactuceae) from the Juan Fernandez Islands, Chile. Amer. J. Bot. 78: 534-543.

Pearson, 0., & A. Pearson. 1980. Nueve mil afios de pequenos mamiferos en el Valle Traful. IX Reunidn Argentina de Ecologia, San Carlos de Bariloche, April 1980, Abstracts.

Pena Guzmdn, L. E. 1987. Introduccion a los insectos de Chile. Santiago: Editorial Universitaria.

Persoon, C. H. 1805. Synopsis Plantarum seu Enchiridium botanicum 1: V.-546. Paris: Treuttel et Wtirtz.

Philippi, R. A. 1864-5. Linnaea. 33: 90.

. 1860. Florula Atacamensis: Umbelliferae. In Viaje al desierto de Atacama, 198-200. Santiago: Halle en Sajonia.

. 1894. Plantas Nuevas Chilenas: Umbelliferae. Anales Univ. Chile 85: 507-514, 699-726.

Presl, K. B. 1825. Reliquiae Haenkeanae seu Descriptiones et leones Plantarum. Prague: apud J.G. Calve bibliopolam.

. 1833. Repertorium Botanicae Systematicae. Prague: Typis filiorum Theophili Haase. 219

Radford, A. E. 1986. Fundamentals of Plant Systematics. New York, NY: Harper & Row, Publishers, Inc.

Ralph, C. P. 1978. Observations on Azorella compacts (Umbelliferae), a tropical andean cushion plant. Biotropica 10: 62-67.

Rauh, W. 1939. Ober polsterformigen Wuchs. Nova Acta Leop. 7: 265-540.

. 1940. Die Wuchsformen der Polsterpflanzen. BotanischeS Archiv. 40: 289-374, 375-462.

Reiche, K. F. 1899. Estudios Crlticos Sobre la Flora de Chile: Umbeliferas. Anales Univ. Chile 104: 767-842.

. 1899. Zur Kenntnis einiger chilenischer Umbelliferen-Gattungen. Bot. Jahrb. Syst. 28: 1-17.

Rendle, A. B. 1904. Mr. Hesketh Prichard's Patagonian Plants. J. Bot. 42: 321-334, 367-378.

Richards, A. J. 1986. Plant Breeding Systems. Cambridge, MA: Unwin Hyman Ltd.

Rochow, T. F. 1967. The ecology of Thlaspi alpestre in the Central Rocky Mountains along altitudinal gradients. Ph.D. diss., Duke University, 1-265.

Romero Aravena, H., Borgel Olivares, R., & D. Vio Urrutia. 1983. Geografia de Chile. Instituto Geogr&fico Militar. 1: Fundamentos Geogr&ficos del Territorio Nacional. Santiago: Instituto Geogr&fico Militar de Chile.

Rudloff, W. 1981. World-Climates. Stuttgart: Wissenschaftliche Verlagsgesellschaft.

Ruiz Lopez, H. & J. A. Pav6n. 1802. Flora Peruviana, et Chilensi. In Prodomus et Flora Peruviana et Chilensis, eds. H. Ruiz L6pez & J. A. Pavon, 3: i-95. Madrid: en la imprenta de Sancha.

Ruthsatz, B. 1978. Las plantas en cojin de los semidesiertos andinos del noroeste Argentino. Darwiniana 21: 491-539.

Saleh, N. A. M., S. I. El-Negoumy, M. N. El-Hadidi, and H. A. Hosni. 1983. Comparative study of the flavonoids of some local members of the Umbelliferae. Phytochemistry 22: ,1417-1420.

Sanquinetti de Bdrmida, A. C., & L. A. Borrero. 1983. Las Buiterras Cave and the palaeoenvironments of the Rio 220

Gallegos valley, Province of Santa Cruz, Argentina. In Quaternary of South America and Antartic Peninsula. 1: 151-156. Rotterdam: Balkema.

Sauberer, F. and 0. Hartel. 1959. Pflanze und Strahlung. Leipzig.

Schimper, A. F. W. 1898. Pflanzengeographie auf physiologischer Grundlage. Jena.

Schinner, F. 1982. C02-Freisetzung, Enzymaktivitaten und Bakteriendichte von Boden unter Spalierstrauchern und Polsterpflanzen in der alpinen Stufe. Oecol. PI. 3: 49- 58.

Schlichting, D. C. 1986. The evolution of phenotypic plasticity in plants. Ann. Rev. Ecol. Syst. 17: 667- 693.

Schroeter, C. 1908. Das pflanzenleben der alpen. Zurich: Verlag von Albert Raustein.

Schroter, E., and H. Hauri. 1914. Versuch einer Ubersicht der angiospermen Polsterpflanzen. Bot. Jahrb. Syst. Suppliment. 50:

Schumann, K. M. 1900. Neue Arten der Siphonogamen 1898. Just's Bot. Jahersber. 26(1): 323-396.

Skottsburg, C. J. F. 1916. Botanische Ergebnisse der schwedischen Expedition nach Patagonien und dem Feuerlande 1907-1909. V. Die Vegetationsverhaltnisse lMngs der Cordillea de los Andes. Kongl. Svenska Vetenskapsakad. Handl. 56(5): 1-366.

Smith, A. P. 1972. Notes on wind related growth patterns of paramo plants in Venezuela. Biotropica 4: 10-16.

Smith, w. J. & F. Vuilleumier. 1971. Evolutionary relationships of some South American Ground Tyrants. Bull. Mus. Comp. Zool. Harvard Univ. 141: 1181-268.

Sokal, R. R. & F. J. Rohlf. 1981. Taxonomic congruence in the Leptopodomorpha re-examined. Syst. Zool. 30: 309- 325.

Speggazini, C. 1899. Nova Addenda ad Floram Patagonicam: Part I. Anales Soc. Ci. Argent. 48: 44-59.

. 1902. Nova Addenda ad Floram Patagonicam: Part IV. Anales Mus. Nac. Hist. Nat. Buenos Aires. 7(Ser. 2a, t. 4): 203-308. 221

Spomer, G. G. 1964. Physiological ecology studies of alpine cushion plants. Physiol. PI. (Copenhagen) 17: 717-724.

Sprengel, K. 1813. Plantarum umbelliferarum denuo desponendarum prodromus. Halae: Hendelianis.

. 1820. Umbelliferae. In Systema vegetabilium, eds. J. J. Roemer & J. A. Schultes, 6: 315-628. Stuttgard: sumtibus J. G. Cottae.

Stafleu, F. A. & R. S. Cowan. 1976-1988. Taxonomic Literature, 2nd ed. 1-7: A-Z. Utrecht: Bohn, Scheltema & Holkema.

Stearn, W. T. 1986. Botanical Latin, 3rd. ed., revised. North Pomfret, VT: David & Charles Inc.

Stuessy, T. F. 1990. Plant Taxonomy: The Systematic Evaluation of Comparative Data. New York: Columbia University Press.

Stuessy, T. F., and D. J. Crawford. 1983. Flavonoids and phylogenetic reconstruction. Plant Systematics and Evolution 143: 83-107.

Swofford, D. L. 1991. PAUP: Phylogenetic Analysis Using Parsimony. Version 3.0s. Privately published.

Tonni, E. P., & F. Fidalgo. 1983. Geology and palaeontology of Pleistocene sediments at Punta Hermengo area (Miramar, Province of Buenos Aires, Argentina): Some palaeoclimatic aspects. In Quaternary of South America and Antartic Peninsula. 1: 23-52. Rotterdam: Balkema.

Troll, c. 1944. Strukturboden, Solifluktion und Frostklimate der Erde. Geol. Rundschau 34: 545-694.

. 1947. Die Formen der Solifuktion und die periglaziale Bodenabtragung. Erdkunde 1, 162-175.

. 1948. Der asymmetrische Aufbau der Vegetationszonen und Vegetationsstufen auf der Nord- und Sudhalbkugel. Ber. geobot. Forschungsinst. Rubel 1947: 46-83. Zurich.

. 1958. Zur physiolgnomik der Tropengewachse. Jber. Ges. Fr. Ford, rhein. Friedrich-Wilhelms Univ., Bonn.

. 1959. Die tropischen Gebirge, ihre dreidimensionale klimatische und pflanzengeographische Zonierung. Bonner geogr. Abh. 25. 222

. 1960. The relationship between the climates, ecology and plant geography of the southern cold temperate zone and of the tropical high mountains. Proc. Roy. Soc. London, Ser. B, Biol. Sci. 152: 529-532.

. 1966. Okologische Landschaftsforschung und vergleichende Hochgebirgsforschung. Wiesbaden: F. Steiner.

Turczaninow, N. S. 1847. Decas Tertia: Nonnullarum specierum Myrtacearum xerocarpicarum atque Umbelliferarum imperfectarum. Bull. Soc. Imp. Naturalistes Moscou, 20 (1): 148-174.

Turesson, G. 1931. The geographical distribution of the alpine ecotype of some Euasiatic plants. Hereditas 15: 329-346.

Van Gardingen, P. and J. Grace. 1991. Plants and Wind. Advances Bot. Res. 18: 189-253

VillagrSn, C. 1988a. Late Quaternary vegetation of Southern Isla Gande de Chiloe, Chile. Quaternary Research 29: 294-306.

. 1988b. Expansion of magellanic moorland during the late Pleistocene: Palynological evidence from Northern Isla de Chilo&, Chile. Quaternary Research 30: 304-314.

Von Gerichen, E. 1901. Liebig's Ann 318:

Vuilleumier, B. S. 1970. The systematics and evolution of Perezia sect. Perezia (Compositae). Contr. Gray Herb. 199: 1-163.

. 1971. Pleistocene changes in the fauna and flora of South America. Science 73: 771-780.

Vuilleumier, F. 1968. Population structure of Asthenes flammulata superspecies (Aves: Furnariidae). Breviora. 297: 1-21.

. 1969a. Systematics and evolution in Diglossa (Aves, Coerebidae). Amer. Mus. Novit. 2381: 1-44.

. 1969b. Pleistocene speciation in birds living in the high Andes. Nature 223: 1179-1180.

. 1970a. Generic relations and speciation patterns in the Caracaras (Aves: Falconidae). Breviora. 355: 1-29. 223

. 1970b. Insular biogeography in continental regions. I. The Northern Andes of South America. Amer. Natur. 104: 373.

Warming, E. 1909. Oecology of Plants. An Introduction to the Study of Plant-Communities. Oxford: Oxford Clarendon Press.

Warren-Wilson, J. 1952. Vegetation patterns associated with soil movement on Jan Mayen Island. J. Ecol. 40: 249- 264.

Weddell, H. A. 1857. Umbelliferae. Chloris andina 2: 186-206. Paris: chez P. Bertrand.

Wells, E. F., and B. A. Bohm. 1980. Chemotaxonomic studies in the Saxifragaceae S.I. 15. The flavonoids of subsection Villosae section Heuchera in the genus Heuchera. Canad. J. Bot. 58: 1459-1463.

Whalen, H. D. 1978. Foliar flavonoids of solanum section Androceras: a systematic survey. Syst. Bot. 3: 257-276.

Whitehead, R. H. 1951. Ecology of the altiplano of Monte Maiella, Italy. J. Ecol. 39: 330-335.

Wiley, E. O., D. Siegel-Causey, D. R. Brooks, & V. A. Funk. 1991. The compleat Cladist: A primer of phylogenetic procedures. 80-90. Lawrence, KS: The University of Kansas.

Wolff, K. F. A. H. 1921. Spec. Nov. Austro-Americanae. Feddes Repert. Spec. Nov. Regni Veg. 17: 441-442.

. 1924. Azorellopsis, genus novum Umbelliferarum Bolivianum. Feddes Repert. Spec. Nov. Regni Veg. 19: 312.