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By Craig M. Johnson

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Scholarly Press

Taxonomic Revision of the 

Chiliotrichum Group 

sensu stricto

(Compositae: Astereae)

José Mauricio Bonifacino

s m i t h s o n i a n c o n t r i b u t i o n s t o b o t a n y • n u m b e r 9 2
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smithsonian contributions to botany • number 92

Taxonomic Revision of 

the Chiliotrichum Group 

sensu stricto

(Compositae: Astereae)

José Mauricio Bonifacino

WASHINGTON D.C.

2009
ABSTRACT

Bonifacino, José Mauricio. Taxonomic Revision of the Chiliotrichum Group sensu stricto (Compositae: Astereae). Smithsonian Contributions to Botany, number 

92, vi + 119 pages, 62 fi gures, 1 table, 2009.—The Chiliotrichum Group sensu lato (Compositae: Astereae) comprises 11 genera and 31 species distributed from 

Colombia to Tierra del Fuego, and it is composed of four main clades: the Chiliotrichum clade (composed of Aylacophora, Chiliophyllum, Chiliotrichum, and 

Lepidophyllum), the Haroldia clade (composed of Haroldia), the Llerasia clade (composed of Llerasia and Ocyroe), and the Nardophyllum clade (composed of 

Cabreraea, Chiliotrichiopsis, Katinasia, and Nardophyllum). A taxonomic revision is presented for Chiliotrichum, Haroldia, and Nardophyllum clades, which are 

referred to here as Chiliotrichum Group sensu stricto. The Chiliotrichum Group sensu stricto, composed of 9 genera and 16 species, constitutes the most diverse 

assemblage of taxa in the Chiliotrichum Group sensu lato. All species in the Chiliotrichum Group sensu stricto were taxonomically evaluated and circumscribed 

based on analysis of the protologues, laboratory studies, and careful study of nearly all type material, dried specimen collections, and extensive fi eld observations. 

Keys for genera in the Chiliotrichum Group sensu lato and species for the genera treated are provided. Also provided for these species are complete synonymies, 

detailed descriptions, line drawings, photographs of live specimens and their habitat, and distribution maps. A list of all specimens examined is presented for 

each species. Three genera are described herein as new: Cabreraea, Katinasia, and Haroldia; and four new combinations are presented: Chiliotrichum fuegianum

(O. Hoffm.) Bonif., Cabreraea andina (Cabrera) Bonif., Katinasia cabrerae (Bonif.) Bonif., and Haroldia mendocina (Cabrera) Bonif. One species name is neotypifi ed (Barnadesia lanata Meyen), and six synonyms of accepted taxa are lectotypifi ed. One species, Chiliotrichum rosmarinifolium, is reduced to the synonymy 

of Chiliotrichum diffusum. KEY WORDS: Astereae, Argentina, Aylacophora, Bolivia, Cabreraea gen. nov., Chile, Chiliophyllum, Chiliotrichiopsis, Chiliotrichum, Chiliotrichum Group sensu lato, Chiliotrichum Group sensu stricto, Compositae, Haroldia gen. nov., Katinasia gen. nov., Lepidophyllum, Llerasia, 

morphology, Nardophyllum, nomenclature, Ocyroe, Peru, taxonomy.

RESUMEN Bonifacino, José Mauricio. Revisión taxonómica del Grupo Chiliotrichum sensu stricto (Compositae: Astereae). Smithsonian Contributions 

to Botany, número 92, vi + 119 páginas, 62 fi guras, 1 tabla, 2009.—El Grupo Chiliotrichum sensu lato (Compositae: Astereae) se compone de 11 géneros y 31 

especies, distribuídas desde Colombia a Tierra del Fuego y esta constituído por cuatro clados principales: el clado Chiliotrichum (integrado por Aylacophora, 

Chiliophyllum, Chiliotrichum y Lepidophyllum), el clado Haroldia (integrado por Haroldia), el clado Llerasia (integrado por Llerasia y Ocyroe) y el clado 

Nardophyllum (integrado por Cabreraea, Chiliotrichiopsis, Katinasia y Nardophyllum). Se presenta aquí la revisión taxonómica de los clados Chiliotrichum, 

Haroldia, y Nardophyllum, que en conjunto se designan Grupo Chiliotrichum sensu stricto. El Grupo Chiliotrichum sensu stricto, compuesto por 9 géneros y 

16 especies, presenta la mayor diversidad morfológica dentro del Grupo Chiliotrichum sensu lato. Se evaluaron taxonómicamente todas las especies del Grupo 

Chiliotrichum sensu stricto y las mismas fueron defi nidas en base al análisis de los protólogos, el estudio de casi todo el material tipo, especímenes de herbario y 

extensas observaciones a campo. Se presentan claves para la identifi cación de los géneros del Grupo Chiliotrichum sensu lato y para la identifi cación de las especies 

dentro de los géneros del Grupo Chiliotrichum sensu stricto. Para cada especie tratada se presenta sinonimia completa, descripción morfológica detallada, dibujos, 

fotografías de especímenes a campo y sus ambientes, mapa de su distribución geográfi ca, y una lista de todos los especímenes examinados. Tres nuevos géneros 

son descriptos: Cabreraea, Katinasia y Haroldia. Cuatro nuevas combinaciones son presentadas: Chiliotrichum fuegianum (O. Hoffm.) Bonif., Cabreraea andina

(Cabrera) Bonif., Katinasia cabrerae (Bonif.) Bonif. y Haroldia mendocina (Cabrera) Bonif. El nombre de una especie es neotipifi cado (Barnadesia lanata Meyen) 

y seis sinónimos de nombres aceptados son lectotipifi cados. Chiliotrichum rosmarinifolium es transferido a la sinonimia de Chiliotrichum diffusum. PALABRAS 

CLAVE: Astereae, Argentina, Aylacophora, Bolivia, Cabreraea gen. nov., Chile, Chiliophyllum, Chiliotrichiopsis, Chiliotrichum, Chiliotrichum Group sensu 

lato, Chiliotrichum Group sensu stricto, Compositae, Haroldia gen. nov., Katinasia gen. nov., Lepidophyllum, Llerasia, morfología, Nardophyllum, nomenclatura, Ocyroe, Peru, taxonomía.

Published by Smithsonian Institution Scholarly Press

P.O. Box 37012 / MRC 957

Washington, D.C. 20013-7012

www.scholarlypress.si.edu

Library of Congress Cataloging-in-Publication Data

Bonfacino, José M. (José Mauricio)

 Taxonomic revision of the Chiliotrichum group sensu stricto (Compositae: Astereae) / José M. Bonifacino.

 p. cm. — (Smithsonian contributions to botany ; no. 92)

 Includes bibliographical references and index.

 1. Compositae—Classifi cation. I. Title. II. Series.

 QK495.C74B66 2009

 583'.99—dc22 2008040837

ISSN (print): 0081-024X, ISSN (online): 1938-2812

 The paper used in this publication meets the minimum requirements of the American National Standard for Permanence of Paper for Printed Library 

Materials Z39.48–1992.
Contents

LIST OF FIGURES v

INTRODUCTION 1

MATERIALS AND METHODS 3

TAXONOMIC HISTORY 4

MORPHOLOGY 5

CHROMOSOME NUMBERS 14

POLLEN 15

DISTRIBUTION AND HABITAT 15

TAXONOMIC TREATMENT 16

 Key to Genera of the Chiliotrichum Group sensu lato 

 (Compositae: Astereae) 16

Aylacophora Cabrera 17

Cabreraea Bonif. gen. nov. 20

Chiliophyllum Phil. nom. cons. 21

Chiliotrichiopsis Cabrera 27

Chiliotrichum Cass. 37

Haroldia Bonif. gen. nov. 50

Katinasia Bonif. gen. nov. 53

Lepidophyllum Cass. 56

Nardophyllum (Hook. and Arn.) Hook. and Arn. 62

SPECIES EXCLUDED FROM CHILIOTRICHUM GROUP 

 SENSU STRICTO 85

ACKNOWLEDGMENTS 111

REFERENCES 113

INDEX OF SCIENTIFIC NAMES 117

Figures

Cover images. From left to right, details from Figures 47D, 50A, and 56A: 

Chiliotrichiopsis keidelii (capitula, leaves), Chiliotrichum diffusum

(capitula), and Nardophyllum bryoides (capitula, leaves).

 1. Stem anatomy of Chiliotrichum Group sensu stricto 

representative taxa. 7

 2. Leaf morphology of Chiliotrichum Group sensu stricto species. 8

 3. Leaf anatomy of Chiliotrichum Group sensu stricto 

representative taxa. 10

 4. Trichomes in Chiliotrichum Group sensu stricto. 11

 5. Morphology of reproductive features Chiliotrichum Group 

sensu stricto. 12

 6. Geographic distribution of Chiliotrichum Group sensu stricto 

(Compositae: Astereae). 15

 7. Aylacophora deserticola Cabrera. 18

 8. Geographic distribution of Aylacophora deserticola Cabrera. 19

 9. Cabreraea andina (Cabrera) Bonif. 22

10. Geographic distribution of Cabreraea andina (Cabrera) Bonif. 23

11. Chiliophyllum densifolium Phil. 25

12. Geographic distribution of Chiliophyllum densifolium Phil. 26

13. Chiliotrichiopsis keidelii Cabrera. 29

14. Geographic distribution of Chiliotrichiopsis keidelii Cabrera. 30

15. Chiliotrichiopsis ledifolia (Griseb.) Cabrera. 33

16. Geographic distribution of Chiliotrichiopsis ledifolia (Griseb.) 

Cabrera. 34

17. Chiliotrichiopsis peruviana G. L. Nesom, H. Rob. and 

A. Granda. 36

18. Geographic distribution of Chiliotrichiopsis peruviana

G. L. Nesom, H. Rob. and A. Granda. 37

19. Chiliotrichum diffusum (G. Forst.) Kuntze. 41

20. Geographic distribution of Chiliotrichum diffusum

(G. Forst.) Kuntze. 42

21. Chiliotrichum fuegianum (O. Hoffm.) Bonif. 48
22. Geographic distribution of Chiliotrichum fuegianum

(O. Hoffm.) Bonif. 49

23. Haroldia mendocina (Cabrera) Bonif. 51

24. Geographic distribution of Haroldia mendocina (Cabrera) Bonif. 52

25. Katinasia cabrerae (Bonif.) Bonif. 55

26. Geographic distribution of Katinasia cabrerae (Bonif.) Bonif. 56

27. Lepidophyllum cupressiforme (Lam.) Cass. 59

28. Geographic distribution of Lepidophyllum cupressiforme

(Lam.) Cass. 60

29. Nardophyllum bryoides (Lam.) Cabrera. 66

30. Geographic distribution of Nardophyllum bryoides

(Lam.) Cabrera. 67

31. Nardophyllum chiliotrichioides (J. Rémy) A. Gray. 72

32. Geographic distribution of Nardophyllum chiliotrichioides

(J. Rémy) A. Gray. 74

33. Nardophyllum genistoides (Phil.) A. Gray. 77

34. Geographic distribution of Nardophyllum genistoides (Phil.) 

A. Gray. 78

35. Nardophyllum lanatum (Meyen) Cabrera. 80

36. Geographic distribution of Nardophyllum lanatum

(Meyen) Cabrera. 81

37. Nardophyllum patagonicum (Cabrera) G. L. Nesom. 84

38. Geographic distribution of Nardophyllum patagonicum

(Cabrera) G. L. Nesom. 85

39. Stem and leaf anatomy of Chiliotrichum Group 

representative taxa. 87

40. Pollen of Chiliotrichum Group. 88

41. Pollen of Chiliotrichum Group (cont.). 89

42. Pollen of Chiliotrichum Group (cont.). 90

43. Pollen of Chiliotrichum Group (cont.). 91

44. Aylacophora deserticola Cabrera. 92

45. Cabreraea andina (Cabrera) Bonif. 93

46. Chiliophyllum densifolium Phil. 94

47. Chiliotrichiopsis keidelii Cabrera. 95

48. Chiliotrichiopsis ledifolia (Griseb.) Cabrera. 96

49. Chiliotrichiopsis peruviana G. L. Nesom, H. Rob. and 

A. Granda. 97

50. Chiliotrichum diffusum (G. Forst.) Kuntze. 98

51. Chiliotrichum diffusum (G. Forst.) Kuntze (cont.). 99

52. Chiliotrichum fuegianum (O. Hoffm.) Bonif. 100

53. Haroldia mendocina (Cabrera) Bonif. 101

54. Katinasia cabrerae (Bonif.) Bonif. 102

55. Lepidophyllum cupressiforme (Lam.) Cass. 103

56. Nardophyllum bryoides (Lam.) Cabrera. 104

57. Nardophyllum bryoides (Lam.) Cabrera (cont.). 105

58. Nardophyllum chiliotrichioides (J. Rémy) A. Gray. 106

59. Nardophyllum chiliotrichioides (J. Rémy) A. Gray (cont.). 107

60. Nardophyllum genistoides (Phil.) A. Gray. 108

61. Nardophyllum lanatum (Meyen) Cabrera. 109

62. Nardophyllum patagonicum (Cabrera) G. L. Nesom. 110
Taxonomic Revision of 

the Chiliotrichum Group

sensu stricto (Compositae: Astereae)

INTRODUCTION

The Chiliotrichum Group (Compositae: Astereae Cass.) consists of shrubby 

daisies that grow mainly in the Andean and Patagonian regions of South America. It was defi ned by Cabrera (1954a) and later modifi ed by Bremer (1994), 

Nesom (1994), and Bonifacino and Sancho (2004). The group is characterized 

by its shrubby habit, dense to more rarely laxly disposed coriaceous leaves that 

are densely tomentose underneath, radiate or discoid capitula, paleate receptacles that are smooth and not alveolate, ray fl orets with staminodes, tubular 

corollas, slightly sagittate anther bases, and pappus elements largely composed 

of fl attened bristles or scales.

In certain areas, especially in southern South America, some species—e.g., 

Chiliotrichum diffusum (G. Forst.) Kuntze, Lepidophyllum cupressiforme 

(Lam.) Cass., Nardophyllum bryoides (Lam.) Cabrera—can form dense stands 

constituting the dominant shrub in the Patagonian scrub; they often extend over 

a considerable area. As for habitat, the Chiliotrichum Group grows from sea 

level up to 4,200 m, and from extremely dry conditions in eastern Patagonia to 

considerably humid areas such as the southern Andean subantarctic forest and 

sub-Paramo vegetation. Most species belonging to Chiliotrichum Group usually 

grow in wide open areas or mountain slopes, although Chiliotrichum diffusum

is also found as an understory element in the subantarctic forests dominated 

by Nothofagus spp., particularly towards the southernmost end of its distribution. Llerasia Triana ocurrs in the low forest that is so characteristic of the subParamo vegetation along the northwestern Andes from Colombia to Bolivia.

Up to the present, there has been no agreement among authors regarding the 

exact composition of the group (Bremer, 1994; Nesom, 1994; Bonifacino and 

Sancho, 2004). However, a recent effort (J. M. Bonifacino and V. Funk, Smithsonian Institution, National Museum of Natural History, Department of Botany, 

unpublished phylogenetic data) to understand the group from a molecular perspective has shed some light on this problem as well as helped to clarify the phylogenetic relationships among its constituent genera and species (see Table 1).

According to Cabrera’s (1954a) original circumscription, nine genera were included in Chiliotrichum Group: Aylacophora Cabrera, Chiliophyllum Phil., Chiliotrichiopsis Cabrera, Chiliotrichum Cass., Lepidophyllum Cass., Nardophyllum

José Mauricio Bonifacino, Laboratorio de 

Botánica, Facultad de Agronomia, Casilla de 

Correos 1238, Montevideo, Uruguay. Email: 

bonifacinoj@fagro.edu.uy. (This work was prepared while the author was a Postdoctoral Fellow 

at the Department of Botany, National Museum 

of Natural History, Smithsonian Institution, 

Washington, D.C., USA.) Manuscript received 

18 January 2008; accepted 19 May 2008.
2

• SMITHSONIAN CONTRIBUTIONS TO BOTANY TABLE 1. Taxonomic history of the Chiliotrichum Group (Compositae: Astereae). The X’s indicate genera considered to be similar or related to each other by different authors. Genera in boldface are described herein as new. Abbreviations: CG = Chiliotrichum Group, s.l. = sensu lato, s.s. = sensu stricto. Authors Genera Cassini (1816) Hook. f. (1847) Remy (1849) Weddell (1856) Gray (1862) Benth. & Hook. f. (1873) Cabrera (1954 & 1969) Blake (1930) Nesom (1991) Zhang & Bremer (1993) Nesom (1993) Nesom (1994) Nesom et al. (2001) Bonif. & Sancho (2004) Karaman (2006) This study CG s.l. CG s.s. Aylacophora X X X XXX X X X Baccharis X Cabreraea X X X XXX X X X X Chiliophyllum a X X X XXX X X X Chiliotrichiopsis b X X X XXX X X X X Chiliotrichum X X X X X X XXX X X X X Diplostephium X XX

Ericameria XX XHaroldia X X X XXX X X X

Hinterhubera XKatinasia X XLepidophyllum X X X X X X XX X X X X

Llerasia XX X X

Nardophyllum c X X X X X X X X XXX X X X X

Ocyroe X X X XXX X X

Oritrophium X XPaleaepappus d X X XXX X X X

Parastrephia X X X X X XX X XPteronia X X X 

a In this study, Chiliophyllum includes only Chiliophyllum densifolium.b In this study Chiliotrichiopsis mendocina is excluded from Chiliotrichiopsis. c In this study Nardophyllum cabrerae and Nardophyllum armatum are excluded from Nardophyllum.d In this study, Paleaepappus is included in Nardophyllum.
number 92 • 3

(Hook. and Arn.) Hook. and Arn., Parastrephia Nutt. emend 

Cabrera and Pteronia L., all of them from southern South 

America, except for Pteronia, which is from South Africa. 

More recent authors (Cabrera, 1969; Bremer, 1994; Nesom, 

1994) added other genera to Chiliotrichum Group: Diplostephium Kunth., Ericameria Nutt., Llerasia, Oritrophium 

(Kunth.) Cuatrec., and Paleaepappus Cabrera, all of them 

from South America, except for Ericameria, which is native 

to North America.

Nesom (1994) placed all these genera in the subtribe 

Hinterhuberinae Cuatrec. However, morphology as well 

as molecular data place an important hiatus between Chiliotrichum Group sensu lato (Bonifacino and Funk, unpublished data) and the typical elements of the subtribe 

Hinterhuberineae (e.g., Diplostephium, Hinterhubera Sch. 

Bip. ex Wedd., Parastrephia). According to the molecular 

studies of Bonifacino and Funk (unpublished data), the 

genera Celmisia Cass., Oritrophium, and Olearia Moench, 

are among the closest relatives to the Chiliotrichum Group. 

As a result of these studies, based on both nuclear and 

chloroplast markers, but also on a careful study of a wide 

array of morphological characters, the coalescence of the 

group originally proposed by Cabrera (1954a) was confi rmed, with the important addition of Llerasia, an idea 

suggested by Nesom (1994) and more recently by Karaman 

(2006). Based on the above-mentioned studies, it was evident that some taxon circumscriptions had to be reconsidered. Consequently, several signifi cant taxonomic changes 

are proposed here, including the description of new genera, 

designation of new combinations, establishment of new 

synonyms, and the exclusion of some species.

The Chiliotrichum Group sensu lato, as redefi ned by 

Bonifacino and Funk (unpublished data), comprises 11 

genera (three described herein as new) and 31 species: Aylacophora, Cabreraea Bonif., Chiliotrichiopsis, Chiliophyllum, Chiliotrichum, Haroldia Bonif., Katinasia Bonif., 

Lepidophyllum, Llerasia, Nardophyllum, and Ocyroe Phil.

Aylacophora, Cabreraea, Chiliophyllum, Haroldia, 

Katinasia, Lepidophyllum, and Ocyroe are monotypic, 

Chiliotrichum comprises two species, Chiliotrichiopsis

three species, Llerasia ~14 species, and Nardophyllum fi ve 

species.

The genera in the Chiliotrichum Group sensu lato are 

arranged in four strongly supported clades (Bonifacino 

and Funk, unpublished data): Chiliotrichum clade (composed of Aylacophora, Chiliophyllum, Chiliotrichum, and 

Lepidophyllum), Haroldia clade (composed by Haroldia), 

Llerasia clade (composed of Llerasia and Ocyroe), and 

Nardophyllum clade (composed of Cabreraea, Chiliotrichiopsis, Katinasia, and Nardophyllum). Nested within 

the Chiliotrichum Group sensu lato there is a monophyletic group that is here referred to as Chiliotrichum Group 

sensu stricto, and which contains the clades Chiliotrichum, 

Haroldia, and Nardophyllum. Future studies will cover 

the description of the Chiliotrichum Group sensu lato, to 

be completed in two other contributions: (1) a revision 

of Ocyroe, a recently reinstated genus that now holds a 

species formerly assigned to Nardophyllum (Nardophyllum armatum (Wedd.) Reiche) (Bonifacino, 2008); and 

(2) a taxonomic revision of Llerasia, which was recently 

confi rmed as part of the Chiliotrichum Group (Karaman 

2006; Bonifacino and Funk, unpublished data).

MATERIALS AND METHODS

This study was based on observation of approximately 

50 type specimens and more than 1,200 herbarium specimens belonging to 19 institutions, which are abbreviated 

as follows for subsequent mention in the text:

BAA Herbario Gaspar Xuárez, Facultad de 

Agronomía, Universidad de Buenos Aires, 

Buenos Aires, Argentina.

BAB Herbario Instituto de Recursos Biológicos, 

Centro de Recursos Naturales, Instituto 

Nacional de Tecnología Agropecuaria, 

Castelar, Buenos Aires, Argentina.

BAF Herbario Facultad de Farmacia y 

Bioquímica, Museo de Farmacobotánica 

Juan A. Domínguez, Universidad de Buenos 

Aires, Buenos Aires, Argentina.

BM Herbarium, Department of Botany, The 

Natural History Museum, London, UK.

CONC Herbario Departamento de Botánica, 

Universidad de Concepción, Concepción, 

Chile.

CORD Herbario Facultad de Ciencias Exactas, 

Físicas y Naturales, Museo Botánico, 

Universidad Nacional de Córdoba, 

Córdoba, Córdoba, Argentina.

E Herbarium, Royal Botanic Garden, 

Edinburgh, Scotland, UK.

F Herbarium, Botany Department, Field 

Museum of Natural History, Chicago, USA.

G Herbarium, Conservatoire et Jardin 

botaniques de la Ville de Genève, Genève, 

Switzerland.

K Herbarium, Royal Botanic Gardens, Kew, 

England, UK.
4 • SMITHSONIAN CONTRIBUTIONS TO BOTANY

LIL Herbario Fundación Miguel Lillo, San 

Miguel de Tucumán, Argentina.

LP Herbario División Plantas Vasculares, 

Museo de La Plata, La Plata, Buenos Aires, 

Argentina.

MO Herbarium, Missouri Botanical Garden, 

Saint Louis, Missouri, USA.

MVFA Herbario Bernardo Rosengurtt, Laboratorio de 

Botánica, Facultad de Agronomía, Universidad 

de la República, Montevideo, Uruguay.

NY William and Lynda Steere Herbarium, New 

York Botanical Garden, Bronx, New York, 

USA.

P Herbier National de Paris, Département de 

Systématique et Evolution Phanérogamie, 

Muséum National d’Histoire Naturelle, 

Paris, France.

SI Herbario Instituto de Botánica Darwinion, 

San Isidro, Buenos Aires, Argentina.

SGO Herbario Sección Botánica, Museo Nacional 

de Historia Natural, Santiago, Chile.

US United States National Herbarium, 

Department of Botany, Smithsonian 

Institution, Washington, D.C., USA.

In addition, specimens belonging to Natalie Goodall (personal herbarium, Ushuaia, Tierra del Fuego, Argentina) 

were also studied.

Additionally, extensive fi eldwork was conducted 

throughout almost the whole geographic range of the 

group. For both the morphological and anatomical studies, 

the material was reconstituted through boiling and posterior bleaching with sodium hypochlorite at 50% solution 

and stained with safranin. Drawings of morphological 

details, unless otherwise noted, were made by the author 

using a Wild M5 stereoscope with camera lucida, and the 

anatomical drawings were made using an Olympus CH-2

 microscope with camera lucida. The photographs were 

shot by the author. For plane shapes, terminology follows 

that proposed by Systematics Association Committee for 

Descriptive Biological Terminology (Anonymous, 1962), 

and trichome terminology follows Ramayya (1962) and 

Freire and Katinas (1995).

TAXONOMIC HISTORY

1786. Lamark describes the fi rst two species of the group, 

Conyza cupressiformis Lam., and Conyza bryoides Lam., based on material from the Straits of 

Magellan.

1788. Forster describes the species Amellus diffusus from 

material collected by himself in Tierra del Fuego.

1816. Cassini describes Lepidophyllum to include Conyza 

cupressiformis without making the combination, 

and suggests a close relationship between Lepidophyllum and Pteronia (probably based on the 

opposite phyllotaxy of both genera).

1817. Cassini describes Chiliotrichum to include Amellus 

diffusus G. Forst. without making the combination, and coining a new (illegitimate) name, Chiliotrichum amelloideum Cass. 

1830. Cassini places Chiliotrichum inside a section of 

“true aster prototypes” together with Diplostephium and Olearia amid other miscellaneous 

Astereae genera. 

1838. Candolle describes Dolichogyne DC. with two species: Dolichogyne staehelinoides DC. and Dolichogyne gnaphalioides DC., both of which are 

currently in the synonymy of Nardophyllum lanatum (Meyen) Cabrera.

1846. Hooker describes two sections inside Chiliotrichum: Chiliotrichum section Euchiliotrichum 

Hook. f., which currently corresponds to Chiliotrichum, and Chiliotrichum section Anactinia

Hook. f. composed of species now placed in 

Nardophyllum.

1849. Rémy mentions Baccharis L., Chiliotrichum and 

Nardophyllum as the closest genera to Dolichogyne (currently Nardophyllum and Parastrephia).

1856. Weddell defi nes two sections within Dolichogyne: 

section Dolichogyne (currently Nardophyllum and 

Ocyroe) and section Tola (currently Parastrephia), 

clearly showing his view of those two taxa as 

closely related.

1862. Gray mentions Ericameria and Chrysothamnus 

Nutt. as the most closely related genera to Nardophyllum and Lepidophyllum sensu lato (currently 

Lepidophyllum and Parastrephia), and shared 

Rémy’s opinion of the close relation between 

Nardophyllum and Chiliotrichum.

1873. Bentham mentions the close relationship between 

Chiliotrichum and Olearia, and also of these two 

genera with Diplostephium.

1873. Bentham and Hooker mention Nardophyllum and Ericameria as genera closely related to Lepidophyllum. 

1898. Kuntze places Pteronia close to Nardophyllum.

1905. Macloskie agrees with Kuntze on the Pteronia and 

Nardophyllum relationship. 

1930. Blake synonymizes Parastrephia with Lepidophyllum largely due to similarities in habit that he 

could infer from the scanty type material at P.
number 92 • 5

1953. Cabrera describes Aylacophora, noting its similarity 

with Nardophyllum.

1954. Cabrera (1954a) proposes Chiliotrichum Group indirectly (without naming it as such) when referring 

to the South American genera related to Nardophyllum, namely Aylacophora, Chiliophyllum,

Chiliotrichiopsis, Chiliotrichum, Lepidophyllum, 

and Parastrephia. Cabrera also pointed out Pteronia as the closest genus to Nardophyllum. 

1969. Cabrera adds the monotypic Paleaepappus from 

central Patagonia to Chiliotrichum Group.

1991. Nesom defi nes a group of closely related South 

American genera and refers to them as the Chiliophyllum Group (tribe Astereae, subtribe Solidagininae O. Hoffm.) which, according to him, 

includes Chiliophyllum, Chiliotrichiopsis, Chiliotrichum, Nardophyllum, Parastrephia, and 

probably Aylacophora. 

1993. Zhang and Bremer formally defi ne Chiliotrichum

Group as such, placing it in the tribe Astereae, subtribe Asterinae (Cass.) Dumort., and including the 

following genera: Aylacophora, Chiliophyllum, 

Chiliotrichiopsis, Chiliotrichum, Diplostephium, 

Lepidophyllum, Nardophyllum, Paleaepappus 

and Parastrephia.

1993. Nesom (1993a and 1993b) agrees to a great extent on the group composition suggested by Cabrera (1954a) and later modifi ed by Zhang and 

Bremer (1993), but placing the group inside tribe 

Astereae, subtribe Hinterhuberinae, and further expanding the group circumscription when 

including genera Hinterhubera, Llerasia, and 

Oritrophium.

1994. Bremer keeps unmodifi ed the group defi nition of 

Zhang and Bremer (1993).

1994. Nesom adds Ericameria and excludes Parastrephia

and Hinterhubera from Chiliotrichum Group, 

placing the latter two genera in the Hinterhubera Group. Nesom also defi nes subgroups inside 

Chiliotrichum Group: (a) Chiliophyllum, Chiliotrichiopsis, Ericameria, Lepidophyllum and 

Nardophyllum (including Aylacophora and Paleaepappus); (b) Chiliotrichum, Diplostephium,

and Oritrophium; (c) Llerasia, and (d) Pteronia.

2001. Bonifacino and Sancho discuss the probable generic 

affi nities of Aylacophora and Paleaepappus, indicating Chiliophyllum, Chiliotrichiopsis, Chiliotrichum, and Nardophyllum as the most closely 

related genera. 

2001. Nesom et al. consider Aylacophora, Chiliophyllum, 

Chiliotrichiopsis, Chiliotrichum, Lepidophyllum, 

Nardophyllum, and Paleaepappus as a group of 

South American related genera, more distally related to Diplostephium, Hinterhubera, Lepidophyllum, and Parastrephia, although they do not 

mention the Chiliotrichum Group explicitly.

2004. Bonifacino and Sancho, exclude Nardophyllum scoparium Phil. from Nardophyllum, and defi ne Chiliotrichum Group sensu stricto as composed by 

Aylacophora, Chiliophyllum, Chiliotrichiopsis, 

Chiliotrichum, Lepidophyllum, Nardophyllum, 

and Paleaepappus.

2006. Karaman, based on molecular data, confi rms the 

idea earlier suggested by Nesom (1993a), that 

Llerasia belong to the Chiliotrichum Group. This 

author also establishes the close relatedness of 

Lepidophyllum and Chiliotrichum, and the also 

close relationship between Chiliotrichiopsis and 

Nardophyllum. In addition, Karaman also presents molecular evidence of the polyphyletic nature 

of Chiliophyllum.

Finally, as a result of the present study, but also based 

on the results of Bonifacino and Funk (unpublished data), 

the Chiliotrichum Group sensu lato is considered to be 

composed of 11 genera and 31 species arranged in four 

main clades very strongly supported: Chiliotrichum clade 

(composed of Aylacophora, Chiliophyllum, Chiliotrichum, and Lepidophyllum); Haroldia clade (composed 

by Haroldia): Llerasia clade (composed of Llerasia and 

Ocyroe); and Nardophyllum clade (composed of Cabreraea, Chiliotrichiopsis, Katinasia, and Nardophyllum). 

The clades Chiliotrichum, Haroldia, and Nardophyllum form a moderately supported clade here referred to 

as Chiliotrichum Group sensu stricto, composed of nine 

genera and 16 species. Three genera are described herein 

as new (Cabreraea, Haroldia, and Katinasia); one species is transferred to the synonymy (Chiliotrichum rosmarinifolium Less.); four new combinations are coined 

(Chiliotrichum fuegianum (O. Hoffm.) Bonif., Cabreraea 

andina (Cabrera) Bonif., Haroldia mendocina (Cabrera) 

Bonif., and Katinasia cabrerae (Bonif.) Bonif.); and one 

species is excluded from Nardophyllum (Nardophyllum armatum). Monotypic Paleaepappus, in agreement 

with Nesom (1993b), is here considered as a synonym of 

Nardophyllum.

MORPHOLOGY

The descriptions that follow are based on the characteristics of the most diverse genera belonging to the 
6 • SMITHSONIAN CONTRIBUTIONS TO BOTANY

Chiliotrichum Group, an assemblage that is here referred to as the Chiliotrichum Group sensu stricto, and 

which includes: Aylacophora, Cabreraea, Chiliotrichiopsis, Chiliophyllum, Chiliotrichum, Haroldia, Katinasia, 

Lepidophyllum, and Nardophyllum. The treatment of the 

relatively uniform Llerasia is a part of an ongoing project 

(J. M. Bonifacino, Laboratorio de Botánica, Facultad de 

Agronomia, Montevideo, Uruguay, unpublished), and the 

treatment of Ocyroe appears in a separate paper (Bonifacino, 2008).

Habit. The taxa belonging to the Chiliotrichum

Group sensu stricto are small shrubs, usually globose, and 

sometimes spreading, that vary between 5 cm and 2 m, the 

branches densely or laxly disposed. Nardophyllum bryoides is a rather diverse species regarding habit, since it can 

be found as globose shrubs with laxly arranged branches or 

forming dense mats appressed to the ground. Some species 

(i.e., Nardophyllum chiliotrichioides (J. Rémy) A. Gray 

and Nardophyllum patagonicum (Cabrera) G. L. Nesom) 

present branches tapering towards the end. Growth is notably heteroblastic for Nardophyllum patagonicum, which 

presents both macro- and brachyblasts, while the rest of 

the species present only macroblasts.

Branches. The stems of the Chiliotrichum

Group sensu stricto present a transverse section with 

a vascular cylinder showing a well-marked secondary 

growth, usually presenting a sclerotized pith, and phloem 

fi bers arranged more or less regularly in bundles (Figure 

1A−D,F–I). Fibers are also important in the xylem. Additionally, those species that show notable ribs along the 

stems (i.e., Aylacophora deserticola Cabrera and Nardophyllum chiliotrichioides) or ribs more or less developed 

(i.e., Katinasia cabrerae and Nardophyllum lanatum) have 

subepidermic fi ber bundles reinforcing the ribs (Figures 

1A,B; 39A,B).

The branches in most species are covered by a dense 

tomentose pubescence formed by aseptate-fl agellate trichomes (Figure 4E) but also have an important presence 

of glandular trichomes usually concealed by the tomentose pubescence, which mainly consist of biseriate vesicular glandular trichomes (Figure 4A−C). Some species (i.e., 

Chiliotrichiopsis ledifolia (Griseb.) Cabrera, Haroldia 

mendocina, and Nardophyllum genistoides (Phil.) A. Gray, 

additionally have notable glandular trichomes (biseriate 

vesicular capitate glandular trichomes, Figure 4D) laxly 

arranged over the stems. In some species (i.e., Chiliophyllum densifolium Phil., Lepidophyllum cupressiforme), 

stems are exclusively gland-dotted, rarely with tomentose 

pubescence. Some species show a broad variation in stem 

pubescence, with individuals exclusively gland-dotted and 

others profusely tomentose and gland-dotted (Chiliotrichum diffusum, Nardophyllum bryoides). The base of 

the leaves extends along the stem for most species, giving 

the stem a furrowed or ribbed appearance more or less 

pronounced depending on the species. In Aylacophora deserticola Cabrera and Nardophyllum chiliotrichioides, the 

ribs are very notorious and continuous over long segments 

of the stem; in these species the ribs are broad and shiny 

with hardly any pubescence. In Katinasia cabrerae, the interrib area is wider than each individual rib, and the whole 

stem is covered by glandular trichomes (biseriate vesicular 

glandular trichomes) and a dense tomentose pubescence. 

In Nardophyllum lanatum (Meyen) Cabrera, the tomentose pubescence is more important and is not overshadowed by the underlying presence of glandular trichomes. 

In Haroldia mendocina and Nardophyllum genistoides, 

ribs are evident and approximately of the same width as 

the interrib areas. In the rest of the species, the presence 

of ribs and/or furrows remains more or less hidden under 

the dense tomentose pubescence. (e.g., Chiliotrichiopsis 

keidelii Cabrera, Chiliotrichiopsis peruviana G. L. Nesom, 

H. Rob., and A. Granda, Nardophyllum bryoides, Nardophyllum patagonicum). In Lepidophyllum cupressiforme

and Chiliophyllum densifolium there are no evident ribs 

along the branches.

Leaves: phyllotaxy. Species belonging to 

the Chiliotrichum Group sensu stricto have alternate phyllotaxy, except for Lepidophyllum cupressiforme, which 

has decussate leaves.

Leaves: shape. Leaf shape presents a broad 

variation among and within the species belonging to the 

Chiliotrichum Group sensu stricto (Figure 2). Leaves 

can range from being elliptic to narrowly elliptic (Figure 

2A,G,K) (e.g., Aylacophora deserticola, Chiliotrichiopsis 

ledifolia, Chiliotrichum diffusum, Chiliotrichum fuegianum, Katinasia cabrerae, Nardophyllum chiliotrichioides, Nardophyllum genistoides, Nardophyllum lanatum); 

oblong to linear (Figure 2B,D,F) (e.g., Cabreraea andina, 

Chiliotrichiopsis keidelii, Chiliotrichiopsis peruviana, 

Nardophyllum bryoides, Nardophyllum chiliotrichioides, 

Chiliotrichum diffusum); obovate (Figure 2C) to narrowly 

obovate (Figure 2I,P,Q) (e.g., Chiliophyllum densifolium, 

Nardophyllum patagonicum); ovate to narrowly elliptic 

(Figure 2E,J) (e.g., Haroldia mendocina) or ovate-triangular (e.g., Lepidophyllum cupressiforme, Figure 2L).

Leaves: size. Leaf size varies both among and 

within species; shortest leaves (2−4 mm long, Figure 2L) 

are found in Lepidophyllum cupressiforme, and also in 

Chiliophyllum densifolium and Nardophyllum bryoides, 

while the longest leaves in the group (30−50 mm long, 
number 92 • 7

FIGURE 1. Stem anatomy of the Chiliotrichum Group sensu stricto representative taxa. A. Aylacophora deserticola. B. Nardophyllum chiliotrichioides. C. Nardophyllum bryoides. D. Chiliotrichum diffusum. E. Chiliophyllum densifolium. F. Chiliotrichiopsis peruviana. G. Cabreraea 

andina. H. Haroldia mendocina. I. Nardophyllum patagonicum. J. Lepidophyllum cupressiforme. (A, Bonifacino et al. 335, LP; B, Bonifacino 

& Donato 791, LP; C, Bonifacino & Donato 699, LP; D, Bonifacino et al. 563, LP; E, Bonifacino et al. 77, LP; F, Flores et al. 2109, LP; G, 

Simón & Bonifacino 765, LP; H, Simón & Bonifacino 782, LP; I, Bonifacino & Donato 691, LP; J, Bonifacino et al. 787, LP).
8 • SMITHSONIAN CONTRIBUTIONS TO BOTANY

FIGURE 2. Leaf morphology of the Chiliotrichum Group sensu stricto species, leaves shown as seen on adaxial side. A. Aylacophora deserticola. 

B. Cabreraea andina. C. Chiliophyllum densifolium. D. Chiliotrichiopsis keidelii. E. Chiliotrichiopsis ledifolia. F. Chiliotrichiopsis peruviana. 

G, H. Chiliotrichum diffusum. I. Chiliotrichum fuegianum. J. Haroldia mendocina. K. Katinasia cabrerae. L. Lepidophyllum cupressiforme. 

M. Nardophyllum bryoides. N. Nardophyllum chiliotrichioides. O. Nardophyllum genistoides. P. Nardophyllum lanatum. Q. Nardophyllum 

patagonicum. (A, Bonifacino et al. 335, LP; B, Simón & Bonifacino 765, LP; C, Bonifacino et al. 77, LP; D. Simón & Bonifacino 520, LP; E. 

Simón & Bonifacino 725, LP; F. Flores et al. 2109, LP; G, Bonifacino et al. 563, LP; H, Bonifacino & Romano 824, LP; I, Bonifacino et al. 595, 

LP; J, Simón & Bonifacino 782, LP; K, Bonifacino & Romano 825, LP; L, Bonifacino et al. 787, LP; M, Bonifacino & Donato 699, LP; N, 

Bonifacino & Donato 791, LP; O, Germain s.n., LP; P, Bonifacino & Romano 811, LP; Q, Bonifacino & Donato 691, LP).
number 92 • 9

Figure 2F) are found in Chiliotrichiopsis ledifolia, Chiliotrichiopsis peruviana, Haroldia mendocina, and Chiliotrichum diffusum.

Leaves: Base. Leaf base can be attenuated 

(Figure 2Q), slightly attenuated (Figure 2E), or truncate 

(Figure 2L).

Leaves: petiole. All the species, with the exception of Lepidophyllum cupressiforme, have a petiole 

broadened towards the stem and constricted above; it has 

a softer texture and is glabrous on the adaxial side. Some 

species (i.e., Aylacophora deserticola, Chiliotrichiopsis 

ledifolia y Haroldia mendocina) have a noticeable tuft of 

aseptate-fl agellate trichomes in the axil of the leaves. As 

mentioned above, Lepidophyllum cupressiforme does not 

have a petiole, and the base is not differentiated from the 

rest of the altogether reduced leaf; the leaves in this species 

are actually attached to the stems throughout all or most 

of their length.

Leaves: apex. Leaf apex varies from acute 

(e.g., Aylacophora deserticola, Katinasia cabrerae, Figure 

2A,K) to obtuse (e.g., Chiliophyllum densifolium, Nardophyllum patagonicum, Figure 2Q).

Leaves: margin. The margin of the leaves of 

the Chiliotrichum Group sensu stricto is always entire, notably varying among the constituent species in the degree 

of in-rolling, with some species showing notably revolute 

margins (e.g., Cabreraea andina, Chiliotrichiopsis keidelii, 

Chiliotrichiopsis peruviana, Chiliotrichum diffusum, Nardophyllum bryoides, Nardophyllum chiliotrichioides, 

Nardophyllum patagonicum, Figure 3B,C,E,J,L), some 

others showing slightly revolute margins (e.g., Aylacophora deserticola, Chiliotrichiopsis ledifolia, Chiliotrichum 

diffusum, Chiliotrichum fuegianum, Haroldia mendocina, 

Figure 3H,K), or some others exhibiting plane, non-revolute margins (e.g., Chiliophyllum densifolium, Katinasia 

cabrerae, Figure 3D,F). It is important to note that for 

some species this character might vary from live specimens 

to herbarium material (e.g., the leaf margin in Katinasia 

cabrerae is totally plane in live specimens, while in herbarium specimens they are slightly revolute).

Leaves: texture. Most species have coriaceous leaves; Chiliotrichiopsis ledifolia, Haroldia mendocina, and Nardophyllum genistoides have chartaceous 

leaves.

Leaves: indumentum. The pubescence type 

and density is extremely variable among and within the 

species of the Chiliotrichum Group sensu stricto. Most 

species have the abaxial side of the leaves densely covered 

by tomentose pubescence, which is responsible for the discolorous leaves so characteristic of the group; this tomentose pubescence is formed by aseptate-fl agellate trichomes 

(Figure 4E); some species also exhibit some tomentose pubescence on the adaxial surface, although much more laxly 

arranged (e.g., Aylacophora deserticola, Chiliotrichiopsis 

ledifolia, Nardophyllum lanatum, Katinasia cabrerae); 

Nardophyllum lanatum exhibits dense tomentose pubescence on both sides. The leaves of all species have glandular trichomes (biseriate vesicular glandular trichomes, 

Figure 4A−C) on both sides with variable density, from 

laxly arranged (e.g., Chiliotrichum diffusum, Nardophyllum spp.) to densely arranged (e.g., Cabreraea andina, 

Chiliotrichiopsis keidelii, Nardophyllum bryoides); some 

species have leaves densely gland-dotted on both surfaces 

(e.g., Chiliophyllum densifolium and Lepidophyllum cupressiforme). In addition to the above-mentioned glandular trichomes, some species (i.e., Chiliotrichiopsis ledifolia, 

Haroldia mendocina, and Nardophyllum genistoides) also 

present noticeable glandular trichomes (biseriate vesicular 

capitate glandular trichomes, Figure 4D).

Leaves: anatomy. The adaxial epidermis 

on the leaves of the Chiliotrichum Group sensu stricto is 

covered in general by a thick cuticle; epidermal cells on 

adaxial surface in general have a prismatic shape with 

more or less thick walls (Figure 39G). Abaxial epidermal cells have thinner walls and a more undulate outline 

than adaxial counterparts (Figure 39E,F). Stomata are 

usually anomocitic (Figure 39E,F), and in the particular 

case of Chiliotrichum, actinocitic (Figure 39H); they are 

in most cases found on the abaxial epidermis only, but in 

Chiliotrichiopsis ledifolia, Haroldia mendocina, Lepidophyllum cupressiforme, and Katinasia cabrerae they are 

on both sides of the leaves; in most species, stomata are 

somewhat elevated (Figure 39C), while in Lepidophyllum 

cupressiforme and Katinasia cabrerae they are at the same 

level as the rest of epidermal cells or even slightly sunken 

(Figure 39D). Mesophyll in most species of the Chiliotrichum Group sensu stricto is dorsiventral with palisade 

parenchyma composed of one to three layers of cells in 

the adaxial side, and diffuse parenchyma on the abaxial 

side (Figure 3B,C,E,G−L). Lepidophyllum cupressiforme 

also has a dorsiventral mesophyll, but inverted, with the 

diffuse parenchyma on the adaxial side and the palisade 

parenchyma on the abaxial side (Figure 3A). In Katinasia cabrerae, the mesophyll is isobilateral, with palisade 

parenchyma on both sides of the leaves and diffuse parenchyma restricted to the center (Figure 3D), while Chiliophyllum densifolium shows a mesophyll entirely composed 

of palisade parenchyma (Figure 3F).

Capitula: arrangement. Capitula are solitary and terminal (Figure 5A1

) for most species in the 
1 0 • SMITHSONIAN CONTRIBUTIONS TO BOTANY

FIGURE 3. Leaf anatomy of the Chiliotrichum Group sensu stricto representative taxa. A. Lepidophyllum cupressiforme. B. Chiliotrichiopsis 

peruviana. C. Nardophyllum chiliotrichioides. D. Katinasia cabrerae. E. Nardophyllum patagonicum. F. Chiliophyllum densifolium. G. Aylacophora deserticola. H, I. Chiliotrichum diffusum. J. Cabreraea andina. K. Haroldia mendocina. L. Chiliotrichiopsis keidelii. (A, Bonifacino et 

al. 787, LP; B, Flores et al. 2109, LP; C, Bonifacino & Donato 791, LP; D, Bonifacino & Romano 825, LP; E, Bonifacino & Donato 691, LP; 

F, Bonifacino et al. 77, LP; G, Bonifacino et al. 335, LP; H, Bonifacino et al. 563, LP; I, Bonifacino & Romano 824, LP; J, Simón & Bonifacino 

765, LP; K, Simón & Bonifacino 782, LP; L, Simón & Bonifacino 520, LP).
number 92 • 1 1

FIGURE 4. Trichomes in the Chiliotrichum Group sensu stricto. A−C. Biseriate vesicular glandular trichomes present on leaves, stems, achenes, 

and disk corolla lobes. D. Biseriate vesicular capitate glandular trichomes present on stems, leaves, and outer phyllaries of Chiliotrichiopsis 

ledifolia, Haroldia mendocina, and Nardophyllum genistoides. E. Aseptate-fl agellate trichomes present on stems, leaves, and outer phyllaries of 

all species. F−I. Twin trichomes present on the achenes (A, B, E, Chiliotrichum diffusum, Bonifacino et al. 563, LP; C–D, Haroldia mendocina, 

Simón & Bonifacino 782, LP; F−I, Nardophyllum bryoides, Bonifacino & Donato 699, LP).
1 2 • SMITHSONIAN CONTRIBUTIONS TO BOTANY

FIGURE 5. Morphology of reproductive features in the Chiliotrichum Group sensu stricto. A. Capitula arrangement: solitary (A1), or in capitulescences (A2). B. Shape of the involucre: cylindrical (B1), cylindrical-campanulate (B2), or globose (B3). C. Type of capitula and sex arrangement of fl orets: radiate capitulum with female fl orets in the periphery and hermaphroditic (functionally male) fl orets in the disk (C1), radiate 

capitulum with female fl orets in the periphery and hermaphroditic fl orets in the disk (C2), and discoid capitulum with only hermaphroditic 

fl orets (C3). D. Type of the receptacle: naked or with 1 or 2 paleae (D1), partially paleate (D2), paleate, and where every fl oret is subtended by 

a palea (D3); p = palea. E. General shape of the paleae: only subtending, not embracing the achene (E1), subtending and embracing the achene 

with acute apex (E2), or subulate apex (E3); cs = cross section. F. Shape of corollas: ray corolla with a well developed limb (F1), ray corolla 

with limb short or not so well developed (F2, F3), tubular corolla broadened more or less abruptly from mid length (F4), and tubular corolla
number 92 • 1 3

Chiliotrichum Group sensu stricto, except for Katinasia 

cabrerae, in which they are in capitulescences of two to fi ve 

(sometimes seven) capitula (Figure 5A2). In Nardophyllum lanatum, capitula can be either solitary or grouped 

in capitulescences of two to three capitula. Capitula can 

be sessile or shortly pedunculate (e.g., Lepidophyllum cupressiforme, Chiliophyllum densifolium, Chiliotrichiopsis 

keidelii, Chiliotrichiopsis peruviana, Nardophyllum bryoides) or sometimes the peduncles can be rather long (e.g., 

Chiliotrichum diffusum).

Capitula: type. Species belonging to the Chiliotrichum Group sensu stricto have either radiate capitula 

(Figure 5C1,C2) formed by marginal fl orets with true ray 

corollas and disk fl orets with tubular corollas (i.e., Cabreraea, Chiliophyllum, Chiliotrichum, Haroldia, and 

Lepidophyllum) or discoid capitula (Figure 5C3

) formed 

entirely by disk fl orets with tubular corollas (i.e., Aylacophora, Katinasia, and Nardophyllum). Chiliotrichiopsis

has radiate capitula except for C. peruviana, which has 

discoid capitula. Disk fl orets in the Chiliotrichum Group 

sensu stricto are bisexual (Figure 5C2,C3), and marginal 

fl orets are pistillate (Figure 5C1,C2), all fertile with the 

only exception being Lepidophyllum, in which disk fl orets 

are functionally male, and only the marginal fl orets produce viable fruits (Figure 5C1

).

Capitula: involucre shape. The involucre in the Chiliotrichum Group sensu stricto can be cylindrical (i.e., Chiliophyllum, Katinasia, Lepidophyllum,

and Nardophyllum, Figure 5B1), campanulate (i.e., Cabreraea, Haroldia, Chiliotrichiopsis peruviana, Figure 5B2

),

or intermediate between campanulate and cylindrical (i.e., 

Chiliotrichiopsis keidelii, Chiliotrichiopsis ledifolia, Chiliotrichum, and Nardophyllum patagonicum); Aylacophora

presents a globose involucre (Figure 5B3). Phyllaries are 

arranged in three to fi ve series, with variable morphology 

among and within genera. In some species, the external 

outer phyllaries show foliaceous apices (i.e., Chiliotrichiopsis keidelii, Chiliotrichiopsis ledifolia, Nardophyllum 

bryoides, Nardophyllum lanatum, Nardophyllum genistoides, and Nardophyllum patagonicum).

Capitula: receptacle. The receptacle in 

most species is either fl at or slightly convex. Nearly all 

species have paleae in the receptacle, these being totally 

paleate (i.e., Aylacophora, Cabreraea, Haroldia, Chiliotrichiopsis, and Nardophyllum patagonicum, Figure 5D3

) 

or partially paleate with paleae mostly placed towards the 

periphery (i.e., Chiliophyllum, Chiliotrichum, Katinasia, 

Lepidophyllum, Nardophyllum bryoides, Nardophyllum chiliotrichioides, and Nardophyllum lanatum, Figure 

5D2). Nardophyllum genistoides is the only species of the 

group that does not have paleae. In Katinasia cabrerae and 

Nardophyllum lanatum, paleae are sometimes absent (Figure 5D1

). Paleae notably embrace the achenes (i.e., Aylacophora, Chiliotrichiopsis, Haroldia, and Nardophyllum

patagonicum, Figure 5E2

,E3) or simply subtend, but not 

enclose them (i.e., Cabreraea, Chiliophyllum, Chiliotrichum, Katinasia, Lepidophyllum, and Nardophyllum, 

Figure 5E1).

Florets: corolla type. Corollas in the 

Chiliotrichum Group sensu stricto could either be true ray 

or tubular. True ray corollas have a usually short basal 

tube that expands into a limb of elliptic to narrowly elliptic shape; the tube can be wide (length to width ratio 

equal or less than four) and then gradually extended into 

a limb (i.e., Cabreraea, Chiliophyllum, Chiliotrichiopsis, 

Haroldia, and Lepidophyllum, Figure 5F2,F3) or it can be 

rather narrow (length to width ratio equal or greater than 

fi ve) and abruptly extended into a limb (i.e., Chiliotrichum, 

Figure 5F1). The apex of the limb in the true ray corollas has three minute lobes; glandular trichomes (biseriate 

vesicular glandular trichomes, Figure 4A−C) can be present or absent; the corollas can be yellow (i.e., Cabreraea, 

Chiliophyllum, Chiliotrichiopsis, Haroldia, and Lepidophyllum) or white (i.e., Chiliotrichum). Tubular corollas 

have a narrow funnel shape that expands either gradually 

towards the apex (i.e., Cabreraea andina, Chiliophyllum 

gradually broadened towards the apex (F5). G. Shape of anthers showing slightly sagittate thecae and antheropodia well developed and swollen (G1), or developed but not swollen (G2). H. Style diversity: branches acute with long and thin papillae (H1), branches obtuse with long 

and thin papillae (H2), branches acute with short papillae (H3), and style shaft base swollen seated on top of a stomata-bearing nectary (H4); 

n = nectary. I. Achene diversity: glandular (I1), glandular and slightly villose (I2), glandular and densely villose (I3), and villose with twintrichomes restricted to the ribs (I4). J. Achenes in cross section: compressed achene (J1), and terete achene (J2); cs = cross section. K. Type of 

pappus elements: scales, (from left to right) divided towards the apex, elliptic short, elliptic large, and linear (K1), fl attened bristles (from left 

to right) scabrid and barbellate towards the apex, and scabrid all the way to the apex (K2), terete bristle, and scabrid (K3); cs = cross section. 

L. Pappus elements arrangement: 1-seriate (L1), 2-seriate (L2), and 3- to 4-seriate (L3).
1 4 • SMITHSONIAN CONTRIBUTIONS TO BOTANY

densifolium, Chiliotrichiopsis keidelii, Chiliotrichiopsis ledifolia, Haroldia mendocina, Nardophyllum bryoides, Nardophyllum chiliotrichioides, and Nardophyllum 

patagonicum, Figure 5F5) or more or less abruptly (i.e., 

Aylacophora deserticola, Chiliotrichiopsis peruviana Chiliotrichum diffusum, Chiliotrichum fuegianum, Katinasia 

cabrerae, Lepidophyllum cupressiforme, Nardophyllum 

genistoides, and Nardophyllum lanatum, Figure 5F4). 

The corolla tube is glabrous or has very few glandular 

trichomes (biseriate vesicular glandular trichome, Figure. 

4A−C). The apex of the tubular corollas has fi ve triangular 

lobes, with glandular trichomes (biseriate vesicular glandular trichome, Figure 4A−C) on the adaxial side, usually 

grouped towards the lobe apices.

Florets: androecium. Size of thecae and 

apical appendages vary greatly among species of the Chiliotrichum Group sensu stricto. The bases of the thecae 

are for most species slightly sagittate (Figure 5G1

,G2); in 

Nardophyllum genistoides they are sagittate and in Haroldia mendocina they are obtuse. Apical appendages vary 

from ovate to triangular. The upper part of the fi lament 

is characterized by the presence of cells with thick walls 

that defi ne an antheropodium zone; this antheropodium 

can have the same width as the rest of the fi lament (Figure 

5G2) or can be noticeably swollen (Figure 5G1). True ray 

corollas have one to four staminodes more or less developed towards the apical part of the tube.

Florets: gynoecium. All species present a 

more or less developed and fl attened nectary disc at the base 

of the style where stomata are present (Figure 5H4). Styles 

vary greatly in shape among species of the Chiliotrichum

Group sensu stricto; style bases are more or less swollen 

in all species except for Lepidophyllum, in which no appreciable swelling is visible. Style branches are totally free, 

with the stigmatic surface reduced to two ventro-marginal 

bands departing from the base and extending 1/2 to 3/4 

of the length of the style branches; the style branches are 

narrowly elliptic to oblong, and the apices vary from acute 

(Figure 5H1,H3) to obtuse (Figure 5H2). Collecting hairs 

are present in all species, usually covering the upper 1/2 to 

3/4, ranging from short papillae (Figure 5H3) to more or 

less developed thin papillae (Figure 5H1,H2).

Achenes. The Chiliotrichum Group sensu 

stricto is characterized by its narrowly obovate achenes, 

more or less terete in transversal section (Figure 5J2), except for Aylacophora, whose achenes are rather fl attened 

(Figure 5J1). All species bear ribs in the achenes, usually 

from two to ten, more or less developed (Figure 5J1,J2). 

Achenes bear biseriate trichomes, glandular and nonglandular (Figure 4A−C,F−I), and they can have either glandular trichomes only (i.e., Lepidophyllum, Figure 5I1

), mostly 

glandular trichomes but also some sparse twin trichomes 

(i.e., Chiliotrichum), or lax (Figure 5I2) to densely (Figure 

5I3) disposed twin trichomes with an important presence 

of glandular trichomes (i.e., Cabreraea, Chiliophyllum, 

Chiliotrichiopsis, Haroldia, Katinasia, Nardophyllum). 

Aylacophora is particularly interesting regarding the pubescence of the achenes since they bear unusually long 

twin trichomes for the group (~3 mm long) totally concealing the pappus, and mostly on the ribs only (Figure 

5I4). Glandular trichomes on the achenes are of the type 

biseriate vesicular glandular, and the twin trichomes are 

type basic fi liform. All species bear a more or less developed carpopodium.

Pappus. The pappus is one of the characters 

that shows more variation among genera and facilitates 

identifi cation of taxa involved. Almost all species within 

the Chiliotrichum Group sensu stricto have pappus elements more or less fl attened except for Cabreraea and 

Katinasia, which instead have terete awn-like bristles 

(Figure 5K3

) than fl attened bristles. Pappus elements can 

either be minute scales somewhat triangular to ovate in 

shape (i.e., Aylacophora, Chiliotrichiopsis, Haroldia, 

Figure 5K1), large narrowly ovate scales (i.e., Nardophyllum patagonicum, Figure 5K1

), narrowly elliptic to linear 

scales (i.e., Lepidophyllum, Figure 5K1), linear scales to 

fl attened bristles (i.e., Chiliophyllum, Chiliotrichum, Nardophyllum bryoides, Nardophyllum chiliotrichioides, and 

Nardophyllum lanatum, Figure 5K1,K2), or as previously 

mentioned, terete awn-like bristles (i.e., Cabreraea, Katinasia, and Nardophyllum genistoides). All of these pappus 

elements, especially the linear scales, fl attened bristles, and 

terete bristles are either scabrid along all their length or 

scabrid and barbellate towards the apex. Pappus elements 

are arranged in one series (i.e., Aylacophora, Chiliophyllum, Chiliotrichiopsis, Haroldia, Figure 5L1

), in two series 

(i.e., Cabreraea, Chiliotrichum, Lepidophyllum, Nardophyllum, Figure 5L2

), or in three to four series (i.e., Katinasia, Figure 5L3

).

CHROMOSOME NUMBERS

Little is known about the chromosome number of the 

genera included in the Chiliotrichum Group sensu stricto. 

Most chromosome counts are by Moore (1981), who 

determined the chromosome numbers of Chiliotrichum 

diffusum 2n = 54, Chiliotrichum fuegianum 2n = ~54, 

Lepidophyllum cupressiforme 2n = 40, and Nardophyllum bryoides 2n = ~50. In a more recent contribution, 
number 92 • 1 5

Hunziker et al. (1989) determined the chromosome number for Chiliotrichiopsis keidelii 2n = 54.

POLLEN

There are no studies describing the pollen grains 

for species belonging to the Chiliotrichum Group sensu 

stricto. Pollen grains in the group are echinate like the 

pollen of most Compositae: Astereae, espheroidal, tricolpate; with colpi tapering towards both ends; the equatorial diameter of the species of the Chiliotrichum Group 

sensu stricto varies between 22 and 34 μm, and the polar 

diameter between 22 and 31 μm. Lepidophyllum has the 

smallest pollen grains and Katinasia, Chiliotrichiopsis, 

and Nardophyllum have the largest pollen grains in the 

group (Figures 40−43).

DISTRIBUTION AND HABITAT

It is useful to look at the distribution of the Chiliotrichum Group sensu stricto, in the context of the larger 

clade, the Chiliotrichum Group sensu lato (see Introduction). The Chiliotrichum Group sensu lato is distributed 

along the Andes from Colombia to Tierra del Fuego, extending to both Argentinean and Chilean extra-Andean 

Patagonia.

Most genera in the Chiliotrichum Group sensu stricto 

are restricted to southern South America, inhabiting areas 

south of 29o S latitude mostly along the Andes and in both 

Argentinean and Chilean Patagonia. Only Chiliotrichiopsis extends northward, reaching southern Peru and southern Bolivia (see Figure 6 and fi gures with distribution maps 

for each species for detailed information on their known 

distribution).

The species belonging to Chiliotrichum Group sensu 

lato grow among an interesting array of habitats ranging 

from sea level to above 4,000 m. Most of the species occur 

in semiarid environments so characteristic of Patagonian, 

Puna, and Monte biogeographic provinces (Cabrera and 

Willink, 1973), but some such as Llerasia occur in the 

wet sub-Paramo of Colombia, Ecuador, and Peru, and in 

the upper part of the Yungas in Bolivia. Several taxa of 

the Chiliotrichum Group sensu stricto (e.g., Aylacophora, 

Chiliophyllum, Chiliotrichum, Katinasia, Lepidophyllum, and Nardophyllum) are important elements of the 

shrubby stratum that characterizes the vegetation in certain areas of Puna, Monte, and Patagonia. They usually 

grow in open areas. However, Chiliotrichum diffusum is 

a rather ubiquitous species also found in subantarctic forests (dominated by Nothofagus spp.), clearings, and even 

forming part of the understory, and Llerasia grows amid 

dwarf forest close to the tree line along central and northwestern Andes.

FIGURE 6. Geographic distribution (shaded area) of the Chiliotrichum group sensu stricto (Compositae: Astereae).
1 6 • SMITHSONIAN CONTRIBUTIONS TO BOTANY

TAXONOMIC TREATMENT

KEY TO GENERA OF THE CHILIOTRICHUM GROUP SENSU LATO (COMPOSITAE: ASTEREAE)

 1a. Capitula discoid; receptacles naked or sometimes with 1 or 2 paleae (Llerasia); disk corollas yellow with glandular 

trichomes along the tube, gradually broadening, with a globose swelling at the base; pappus (2)3- to 5-seriate, pappus 

elements terete bristles . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2

 1b. Capitula discoid or radiate; receptacles paleate or partially paleate; disk corollas yellow (rarely white) with none or 

(rarely) very few glandular trichomes along the tube, gradually or abruptly broadening from middle, but never with a 

globose swelling at the base; pappus 1- or 2-seriate (rarely 3- to 4-seriate, but then corollas white), pappus elements scales 

or more or less fl attened bristles (rarely terete, but then with none of the above characteristics) . . . . . . . . . . . . . . . . . .3

 2a. Erect shrubs; branches rigid and thorny; leaves minute (up to 12 mm long), arranged in macro- and brachyblasts; capitula solitary; pappus 3- to 5(7)-seriate; achenes densely villose. (San Juan to Jujuy in Argentina, northern Chile, and 

southern Bolivia) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Ocyroe

 2b. Scandent shrubs; branches never thorny; leaves large (always more than 40 mm long), arranged in macroblasts only; 

capitula numerous, arranged in profuse corymbiform capitulescences; pappus (2)3-seriate; achenes gland-dotted. (From 

Colombia to southern Bolivia) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Llerasia

 3a. Leaves decussate, triangular, scale-like. (Argentina and Chile, southern Santa Cruz and Magallanes and northern Tierra 

del Fuego) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Lepidophyllum

 3b. Leaves alternate, elliptic to narrowly elliptic or narrowly obovate, oblong, or linear; never triangular and scale-like . . . 

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .4

 4a. Pappus 1-seriate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5

 4b. Pappus 2- to 4-seriate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8

 5a. Branches with long and well-defi ned ribs, noticeably wider than the interrib areas, more or less continuous throughout 

the stem length; leaves sparsely arranged, early deciduous; involucre globose; capitula discoid. (Argentina, restricted to 

Neuquén) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .Aylacophora

 5b. Branches with ribs absent or not well marked, if present of the same width or narrower than the interribs, not continuous 

throughout the length of the stem; leaves more or less densely arranged, persistent; involucres cylindric to campanulate; 

capitula radiate (if discoid with none of the above characteristics) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6

 6a. Leaves obovate, margins entire, non-revolute; pappus elements linear scales, not divided towards the apex, scabrid along 

the margins. (Argentina, restricted to Mendoza) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Chiliophyllum

 6b. Leaves ovate to linear, never obovate, margins revolute; pappus elements narrowly ovate scales, usually divided towards 

the apex, if not, then thicker and with protrusions . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7

 7a. Paleae subulate, rigid towards the apex; corolla veins thick towards the base, protruding on outer surface when dry; pappus elements coarse and undivided towards the apex. (Argentina, known from northern Mendoza) . . . . . . . . Haroldia

 7b. Paleae acute, not subulate, softer towards the apex; corolla veins not thick, not protruding on corolla surface when dry, 

pappus elements thin and fl at, divided towards the apex. (Southern Peru and Bolivia, and northern Argentina: La Rioja, 

Catamarca, Salta, Tucumán, and Jujuy) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Chiliotrichiopsis

 8a. Capitula radiate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9

 8b. Capitula discoid . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10

 9a. Leaves linear; ray corollas yellow; pappus elements terete awn-like bristles, not fl attened, heteromorphic, outer series 

slightly shorter, scabrid throughout their length, inner series barbellate towards the apex. (Argentina, La Rioja and 

Catamarca) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Cabreraea

 9b. Leaves narrowly elliptic to obovate or linear; ray corollas white; pappus elements linear scales, isomorphic, subequal, 

scabrid along the margins. (Chile, from Metropolitana Region southwards to Tierra del Fuego, Argentina, from southern 

Mendoza to Tierra del Fuego) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Chiliotrichum

 10a. Leaf margins non-revolute, entire; corollas white; pappus 3- to 4-seriate, pappus elements terete bristles, outer series 

notably shorter. (Argentina, Mendoza and Neuquén) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Katinasia

 10b. Leaf margins revolute; corollas yellow; pappus 2-seriate, subequal, pappus elements fl attened bristles. (Argentina and 

Chile, widespread) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Nardophyllum
number 92 • 1 7

Aylacophora Cabrera

Taxonomic History

1953. Cabrera describes the genus from material collected 

in central Neuquén (Argentina), and funds Aylacophora deserticola.

1993. Nesom (1993b) transfers Aylacophora to the synonymy of Nardophyllum, making the combination Nardophyllum deserticola (Cabrera) G. L. 

Nesom.

2001. Bonifacino and Sancho reinstate Aylacophora as a 

distinct genus from Nardophyllum.

Aylacophora Cabrera, Bol. Soc. Argent. Bot. 4(4): 266, 

1953. Type: Aylacophora deserticola Cabrera, Bol. Soc. 

Argent. Bot. 4(4): 268, 1953.

Shrubs globose; branches more or less densely disposed, 

ascending, almost leafl ess, 8-ribbed, interrib furrows densely 

tomentose, base of branches with triangular, brownish scales. 

Leaves sparsely arranged, early deciduous, alternate, ascending, gradually reduced towards branch apices, with a tuft of 

woolly trichomes in the axile, narrowly elliptic, base attenuated, petiole broadened towards the stem and constricted 

above, apex acute, margins slightly revolute, tomentose and 

gland-dotted on both surfaces, medium vein slightly marked, 

slightly discolorous, coriaceous. Capitula solitary, terminal, 

pedunculate, homogamous, discoid. Involucres globose; 

phyllaries arranged in 4 to 5 series, gland-dotted, densely 

tomentose on abaxial surface, coriaceous, margins membranaceous; outer phyllaries triangular-ovate; inner phyllaries 

elliptic. Receptacles fl at to slightly convex, paleate; paleae 

narrowly elliptic, boat-shaped, embracing the achenes, apex 

recurved, gland- dotted, margins membranaceous. Florets 25 

to 40, bisexual; corollas yellow, tubular, 5-lobed, broadened 

more or less abruptly from mid-length, lobes triangular; anther thecae base slightly sagittate, apical appendages triangular, antheropodium present, not swollen; style bases notably 

swollen, style branches linear-oblong, apex acute, collecting 

hairs on upper 1/3 to 1/2. Achenes obovate, compressed, 2- 

or 3-ribbed, with long twin trichomes on ribs only, sparse 

glandular trichomes towards the apex of achenes inbetween 

ribs, carpopodium present. Pappus 1-seriate, formed by 

small, narrowly elliptic to triangular scales, scabrid.

Etymology: Name derived from Greek “aylacos” 

(furrow), and “phorus” (to carry), referring to the notable 

furrows that run along the stem.

Aylacophora is a monotypic genus restricted to an 

arid area of western Argentina in Neuquén province. It 

inhabits sand dunes with a very scanty vegetation cover 

amid creosote bushes (Larrea spp.) in the Monte biogeographic province (Cabrera and Willink, 1973).

The genus is characterized by its furrowed and almost 

leafl ess stems, discoid heads, globose involucres, paleate 

receptacles, yellow fl orets, 2- or 3-ribbed achenes compressed with long twin trichomes on the ribs only, and 

1-seriate pappus composed of minute scales.

It differs from similar taxa such as Nardophyllum 

and Katinasia, most notably in the shape of the involucre, 

shape and number of ribs of the achene, and the type and 

size of the pappus. Aylacophora appears to be sister to 

the clade Lepidophyllum + Chiliophyllum + Chiliotrichum 

(Bonifacino and Funk, unpublished data).

Aylacophora deserticola Cabrera, Bol. Soc. Argent. Bot. 

4(4): 268, 1953. Nardophyllum deserticola (Cabrera) 

G. L. Nesom, Phytologia 75(5): 362, 1993. Type: “Argentina, Neuquén, Plaza Huincul, 4-XII-1952, Cabrera 

11053” (holotype: LP 562212-A!; isotypes: GH, LP 

562212-B!, LP 562212-C!, US!).

FIGURES 7, 8, 44

Shrubs 0.5−1 m tall, globose; branches more or less 

densely disposed, ascending, branches 8-ribbed, interrib 

furrows densely tomentose, internodes (0.5−) 1.5−3 cm, 

scales at the branch bases, 0.8−3 × 1−1.5 mm, triangular, 

brownish. Leaves sparsely arranged, early deciduous, alternate, ascending, gradually reduced toward branch apices, 

with a tuft of woolly trichomes in the axils, 8−11 × 1.5−2.5 

mm, narrowly elliptic, base attenuated, petiole broadened 

towards the stem and constricted above, apex acute, margins slightly revolute, tomentose and gland-dotted on both 

surfaces, midvein shallowly marked, coriaceous. Capitula

solitary, terminal, pedunculate, peduncles 1−3 cm long, 

homogamous, discoid. Involucres 6−9 × 6−8 mm, globose; 

phyllaries arranged in 4 to 5 series, gland-dotted, dense tomentose pubescence on abaxial surfaces, coriaceous, margins membranaceous; outer phyllaries 2.5−6.0 × 1−2 mm, 

triangular-ovate, apex acute; inner phyllaries 6−8 × 2 mm, 

elliptic, apex acute-mucronate. Receptacles fl at to slightly 

convex, paleate; paleae 7−9 × 0.8−1.5 mm, narrowly elliptic, boat-shaped, embracing the achenes, apex acute, recurved, gland-dotted, with small tuft of woolly trichomes 

towards the apex, margins membranaceous. Florets 25 to 

40, bisexual; corollas yellow, tubular, 5-lobed, broadened 

more or less abruptly from mid-length, tube 2−3 mm long, 

throat 2.5−3 mm long, lobes 1.5−2 × 0.6−1 mm, triangular, gland-dotted abaxially; anther thecae ~ 2.5 mm long, 

bases slightly sagittate, apical appendages ~ 1 × 0.3 mm, 

triangular, antheropodium present, not swollen; style 
1 8 • SMITHSONIAN CONTRIBUTIONS TO BOTANY

FIGURE 7. Aylacophora deserticola Cabrera. A. Habit. B. Detail of stem. C. Leaves, adaxial view (left), abaxial view (right). D. Involucre. 

E. Outer to inner phyllaries (from left to right). F. Receptacular paleae, lateral view (left), dorsal view (right). G. Detail of pappus (trichomes on 

upper part of achene not drawn). H. Achene with pappus (pappus occluded by notably long twin trichomes). I. Style. J. Corolla (A−J, Bonifacino 

et al. 335, LP).
number 92 • 1 9

shafts 5−6 mm long, base notably swollen, style branches 

3−4 mm long, linear-oblong, apex acute, collecting hairs 

on upper 1/3 to 1/2. Achenes 6−6.5 × 2−2.5 mm, obovate, 

compressed, 2- or 3-ribbed, ribs densely covered with long 

twin trichomes up to ~ 3 mm long, glandular trichomes 

sparsely distributed inbetween nerves towards the apex of 

achenes, carpopodium present. Pappus 0.8−1.6 mm long, 

1-seriate, composed of 13 to 31 narrowly elliptic to triangular scales, scabrid, usually fused towards the base.

Vernacular names: Not known.

Phenology: Flowering specimens have been collected from January to February.

Distribution: Argentina, endemic to Neuquén province (Figure 8).

Habitat: Inhabits arid areas of Monte biogeographic province (Cabrera and Willink, 1973) at 500−650 

m, with annual rainfall less than 150 mm the species inhabits sand dunes with sparse vegetation (~ 40% cover), 

associated to Larrea divaricata Cav., Atriplex lampa Gill. 

ex Moq., Prosopis fl exuosa var. depressa F. A. Roig, 

Senna aphylla subsp. divaricata (Hieron.) L. Bravo, Chuquiraga erinacea D. Don, Gutierrezia solbrigii Cabrera, 

Fabiana patagonica Speg., and Larrea cuneifolia Cav. 

(Cabrera, 1953; Bonifacino and Sancho, 2001: Roig, 

1998; pers. obs.)

Uses: Not known.

Etymology: The specifi c epithet deserticola refers 

to the arid areas where the species ocurrs.

Additional specimens examined: ARGENTINA. PROV. 

NEUQUÉN. Dpto. Añelo. Ruta Prov. 17, 38 km al E de Añelo, 

cerca de la meseta alta, s.date, Valla s.n. (BAA 17790). 

Dpto. Confl uencia. Plaza Huincul, Junto al Cementerio 

38°56'8.2"S, 69°10'50.4"W, 6-I-2002, Bonifacino et al. 

334 (LP); Ruta 10, entre Cutral-Có y Paso de los Indios, 

FIGURE 8. Geographic distribution of Aylacophora deserticola Cabrera.
2 0 • SMITHSONIAN CONTRIBUTIONS TO BOTANY

38°47'23.1"S, 69°16'25.9"W, 6-I-2002, Bonifacino et al. 

335 (LP); al N de Cutralcó, 38°47'23"S, 69°16'26"W, 630 

m, 8-XII-2006, Bonifacino et al. 2907 (US);10 km al W de 

Cutralcó, 8-II-1983, León 3362 (BAA); Plaza Huincul, 27-

XI-1943, Plotnick 47 (LP); 1−2 km de Plaza Huincul, 530 

m, 26-I-1970, Ruiz Leal 27108 (LP); Paso de los Indios a 

Cutral-Co, 5-II-1999, Steibel & Troiani s.n. (ex SRFA 14074 

in LP); Plaza Huincul, junto al Cementerio, s. date, Troiani 

et al. s.n. (ex SRFA 12503 in LP). Dpto. Pehuenches. Paso 

de las Bardas, 2-II-1999, Troiani & Steibel s.n. (ex SRFA 

13920 in LP). Dpto. Zapala. 15 km al SW de Paso de los 

Indios, 5-II-1999, Steibel & Troiani s.n. (ex SRFA 14069 in 

LP); 20 km al SE de Paso de los Indios, 5-II-1999, Steibel & 

Troiani s.n. (ex SRFA 14073 in LP).

Cabreraea Bonif. gen. nov.

Cabreraea Bonif. gen. nov. Type: Chiliophyllum andinum 

Cabrera, Notas Mus. La Plata, Bot. 19(92): 193, 1959 

(≡ Cabreraea andina (Cabrera) Bonif. comb. nov.).

Frutex, ramis teretibus lanuginosus (tomentosis) et 

glandulo-punctatis. Folia anguste elliptica base amplexicaulia margine revoluta. Capitula plurifl ora heterogama 

radiata solitaria, involucris campanulatis, receptaculis 

paleatis. Flosculi lutei dimorphi, fl osculis peripheralibus 

femineis radiatis, fl osculis disci hermaphroditis; corollae 

tubiformes glanduliferae 5-lobatae sensim ampliatae; rami 

stylorum in trientis superioribus papillis collectoribus obsiti. Achenia villosissima. Aristae pappi inaequales scabridae biseriatae.

Shrubs subglobose; branches more or less loosely 

disposed, ascending, not ribbed, densely tomentose and 

gland-dotted. Leaves densely arranged, persistent, alternate, patent, linear-oblong, base truncate, petiole broadened towards the stem and slightly constricted above, 

apex obtuse, slightly uncinate, refl exed, margins notably 

revolute, gland-dotted on both surfaces, densely tomentose on abaxial surface, discolorous, coriaceous. Capitula

solitary, terminal, shortly pedunculate, heterogamous, radiate. Involucres campanulate; phyllaries arranged in 3 to 

5 series, coriaceous; outer phyllaries ovate-elliptic to oblong, apex acute, tomentose on abaxial surface, margins 

membranaceous, the outermost phyllaries with foliaceous 

apices; inner phyllaries oblong to narrowly elliptic, apex 

subacute to obtuse, with scanty tomentose pubescence, 

margins membranaceous, notably broadened towards 

the apex. Receptacles convex, paleate; paleae narrowly 

elliptic, boat-shaped, not embracing the achenes, apex 

acute, villose and gland-dotted, margins membranaceous, 

fi mbriate. Ray fl orets 9 to 13, pistillate; corollas yellow, 

radiate, limb elliptic, 3-lobed; staminodes present; style 

base swollen, style branches narrowly ovate, subequal in 

length. Disk fl orets 20 to 25, bisexual; corollas yellow, 

tubular, 5-lobed, gradually broadened towards the apex, 

lobes triangular; anther thecae bases slightly sagittate, apical appendages narrowly ovate, antheropodium present, 

swollen; style base broadened, style branches narrowly elliptic, apex acute, collecting hairs on upper 1/3. Achenes

narrowly obovate, terete, 3- to 5-ribbed, densely villose, 

with very short glandular trichomes, carpopodium present. Pappus 2-seriate, formed by awn-like terete bristles, 

slightly fl attened towards the base, external series gradually tapered towards the apex, inner series slightly thicker 

and barbellate towards the apex, slightly longer than outer 

series, some fused among each other at the base.

Etymology: I dedicate this genus to the late illustrious synantherologist Angel Lulio Cabrera (1908−1999), 

who made signifi cant contributions to the understanding of 

Compositae systematics, and established the basis of Compositae fl oristics in southern South America.

Cabreraea is a monotypic genus herein described as 

new to include the rare Chiliophyllum andinum. Cabreraea inhabits mountain areas of central-western Argentina 

in the altitudinal belt between 2,000 and 3,000 m.

Morphological, as well as molecular data (Bonifacino 

and Funk, unpublished data) support the segregation of 

Chiliophyllum andinum from Chiliophyllum, and its 

placement in a distinct genus. Cabreraea has a paleate receptacle, acute style branches, and 2-seriate pappus composed of awn-like terete bristles (versus partially paleate 

receptacle, obtuse style branches and a 1-seriate pappus 

composed of fl attened bristles in Chiliophyllum). In addition, there are important differences in the shape and 

anatomy of leaves that set Cabreraea apart from Chiliophyllum (Cabreraea has notably revolute leaves with a white 

tomentose pubescence on the abaxial surface, and has dorsiventral mesophyll (Figure 3J), while Chiliophyllum has 

entire, non-revolute leaf margins, notably glanduliferous 

on both surfaces, not tomentose, and has a homogeneous 

palisade mesophyll (Figure 3F). Cabreraea clearly belongs 

to the Nardophyllum + Chiliotrichiopsis clade, but it is 

unclear to which of those two genera it is more related 

(Bonifacino and Funk, unpublished data).

Cabreraea andina (Cabrera) Bonif. comb. nov. Chiliophyllum andinum Cabrera, Notas Mus. La Plata, Bot. 

19(92): 193, 1959. Type: “[Argentina] San Juan, Quebrada de Gualcamayo, entre Chepical y El Salto, 13-
number 92 • 2 1

XII-1957, Ruiz Leal & F. Roig 18960” (holotype: LP 

18870!).

FIGURES 9, 10, 45

Shrubs up to 2 m tall, subglobose; branches more or 

less loosely disposed, ascending, not ribbed, internodes 

short, up to 3 mm long. Leaves densely disposed, persistent, alternate, patent, 13−22 × 1.5−2 mm, linear-oblong, 

base truncate, petiole broadened towards the stem and 

slightly constricted above, apex obtuse, slightly uncinate, 

refl exed, margins notably revolute, gland-dotted on both 

surfaces, densely tomentose on abaxial side, discolorous, 

coriaceous. Capitula solitary, terminal, shortly pedunculate, peduncles 3−8 mm long, heterogamous, radiate. Involucres 7−8 × 7−8 mm, campanulate; phyllaries arranged 

in 3 to 5 series, coriaceous; outer phyllaries 5−6 × 2 mm, 

ovate-elliptic to oblong, apex acute, tomentose on abaxial 

surface, margins membranaceous, the outermost phyllaries 

with foliaceous apices; inner phyllaries 7−9 × 1.8−2 mm, 

oblong to narrowly elliptic, apex subacute to obtuse, with 

scanty tomentose pubescence, margins membranaceous 

notably broadened towards the apex. Receptacles convex, 

paleate; paleae 7−8 × 1.3−1.8 mm, boat-shaped, not embracing the achenes, narrowly elliptic, apex acute, villose 

and gland-dotted, margins membranaceous, fi mbriate. 

Ray fl orets 9 to 13, pistillate; corollas yellow, radiate, tube 

2.5−4 mm long, limb 5−7.5 × 2.5−3.5 mm, elliptic, 3-lobed; 

staminodes 2 to 4, reduced; style shafts ~ 4 mm long, base 

swollen, style branches 2.5−2.8 mm long, narrowly ovate, 

subequal in length. Disk fl orets 20 to 25, bisexual; corollas 

yellow, tubular, 5-lobed, gradually broadened towards the 

apex, tube 4.5−9 mm long, lobes 1.2−1.5 × 0.5−0.7 mm, 

triangular, without glandular trichomes; anther thecae 

2−3 mm long, bases slightly sagittate, apical appendages 

~ 1 × 0.3 mm, narrowly ovate, antheropodium present, 

swollen; style shafts 4−7 mm long, base swollen, style 

branches 2−2.5 mm long, narrowly elliptic, apex acute, 

collecting hairs short, on upper 1/3. Achenes 4.5−5.5 mm 

long, narrowly obovate, terete, 3- to 5-ribbed, densely villose, with very short glandular trichomes, carpopodium 

present. Pappus 5−6.5 mm long, 2-seriate, composed of 42 

to 53 awn-like terete bristles, slightly fl attened towards the 

base, external series gradually thinner towards the apex, 

inner series slightly with awns thicker and barbellate towards the apex, slightly longer than outer series, some 

fused among each other at the base.

Vernacular names: Not known.

Phenology: Flowering specimens have been collected from November to February.

Distribution: Argentina, mountainous areas of La 

Rioja and San Juan provinces (Figure 10).

Habitat: Scrub dominated areas between 2,000 

and 3,000 m in the transition of biogeographic provinces 

Monte and Puna (Cabrera and Willink, 1973).

Uses: Not known.

Etymology: The specifi c epithet andina refers to 

the Andes mountain range, where the species occurs.

Additional specimens examined: ARGENTINA. PROV. 

LA RIOJA. Dpto. Sarmiento. Entre El Peñón y Jagüé, proximadamente a 9 km del primero, Quebrada del Río El Peñón, 

15-XII-1996, Biurrun & Molina 4574 (CORD); Quebrada 

de Ciénaga Grande, 3,100 m, 8-II-1949, Krapovickas & 

Hunziker 5872 (BAB, LP). PROV. SAN JUAN. Dpto. Angaco. 

Sierra Pie de Palo, subiendo por el Camino a Mogote de 

los Corralitos, en la Quebrada del Molle, ca. 2,500 m, 28-

XI-1980, Hunziker et al. 23776 (CORD); íd., 28-XI-1980, 

Hunziker et al. 23774 (CORD), íd., ca. 2150 m, 28-XI1980, Hunziker et al. 23747 (CORD); íd., ca. 1950−2050 

m, 28-XI-1980, Hunziker et al. 23678 (CORD); íd., 

ca. 1,950−2,050 m, 28-XI-1980, Hunziker et al. 23690 

(CORD); Sierra Pie de Palo, Camino a Mogote de los Corralitos, 1,800−2,100 m, 13-XI-1982, Kiesling & Saenz 

4106 (K, SI); íd., 25-I-2001, Simón & Bonifacino 765 (LP). 

Dpto. Iglesia. Entre Cordón de Punilla y Leoncito, 8-XII1915, Hosseus 1080 (CORD).

Chiliophyllum Phil. nom. cons.

Taxonomic History

1862. Philippi describes a new species from Mendoza (Argentina): Chiliophyllum densifolium.

1900. Hoffmann describes a new species, Chiliophyllum fuegianum, based on material from Tierra 

del Fuego (in the present paper transferred to 

Chiliotrichum).

1935. As a result of changes on the validity of later homonyms after the International Botanical Congress 

at Cambridge in 1930, Rehder et al. present the 

list of later homonyms with recommendations on 

how they should be treated. These authors point 

out that, given the acceptance by most authors of 

Chiliophyllum DC. as a synonym of Zaluzania

Pers., and the fact that Chiliophyllum Phil. has no 

synonyms, the name coined by Philippi must be 

renamed or conserved.

1937. Cabrera renames Chiliophyllum Phil., coining it 

Phyllochilium Cabrera, and creates the combination Phyllochilium densifolium (Phil.) Cabrera.
2 2 • SMITHSONIAN CONTRIBUTIONS TO BOTANY

FIGURE 9. Cabreraea andina (Cabrera) Bonif. A. Habit. B. Leaves, adaxial view (left), abaxial view (right). C. Involucre. D. Outer to inner 

phyllaries (from left to right). E. Receptacular paleae, lateral view (left), dorsal view (right). F. Detail of terete bristle from pappus, inner series 

(left), outer series (right). G. Achene with pappus. H. Style of ray fl oret. I. Style of disk fl oret. J. Corolla of disk fl oret. K. Corolla of ray fl oret 

(A−K, Biurrun & Molina 4574, CORD).
number 92 • 2 3

1940. Green et al. present the list of nomina conservanda, 

offi cially granting Chiliophyllum Phil. the status 

of conserved generic name.

1944. Cabrera recognizes the legitimacy of Chiliophyllum

Phil. as a conserved name, and places Phyllochilium Cabrera in the synonymy of Chiliophyllum

Phil.

1959. Cabrera describes Chiliophyllum andinum from 

material of San Juan, Argentina (in the present 

paper placed in its own genus Cabreraea Bonif. 

gen. nov., herein described as new).

Chiliophyllum Phil. nom. cons., Anales Univ. Chile 21: 

397, 1862 (Non DC., Prodr. 5: 554, 1836).

Phyllochilium Cabrera, Notas Mus. La Plata, Bot. 2(16): 

171, 1937. Type: Chiliophyllum densifolium Phil., 

Anales Univ. Chile 21(2): 397, 1862.

Shrubs subglobose, spreading; branches densely disposed, ascending to erect, not ribbed, notably gland-dotted 

sometimes tomentose. Leaves densely arranged, persistent, 

alternate, patent, obovate, base slightly attenuated, petiole 

FIGURE 10. Geographic distribution of Cabreraea andina (Cabrera) Bonif.
2 4 • SMITHSONIAN CONTRIBUTIONS TO BOTANY

broadened towards the stem and constricted above, apex 

obtuse, margins fl at (non-revolute), notably gland- dotted 

on both surfaces, rarely some tomentose pubescence on 

abaxial side on midvein and margins, midvein elevated on 

abaxial surface, concolorous, coriaceous. Capitula solitary, terminal, shortly pedunculate, heterogamous, radiate. Involucres cylindrical; phyllaries arranged in 3 to 4 

series; outer phyllaries elliptic, apex acute, gland-dotted 

throughout abaxial surface, chartaceous, margins membranaceous; inner phyllaries narrowly elliptic, apex acute, 

gland-dotted towards the apex, chartaceous, margins 

membranaceous. Receptacles slightly convex, partially paleate; paleae linear to narrowly elliptic, boat-shaped, not 

embracing the achenes, with a tuft of woolly trichomes towards the apex, apex acute, margins membranaceous -fi mbriate. Ray fl orets 4 to 8, pistillate; corollas yellow, radiate, 

limb elliptic, 3-lobed; staminodes present; style base swollen, style branches oblong, subequal in length. Disk fl orets 7 to 12, bisexual; corollas yellow, tubular, 5-lobed, 

gradually broadened towards the apex, lobes triangular; 

anther thecae bases slightly sagittate, apical appendages 

triangular -ovate, anteropodium notably swollen; style base 

swollen, style branches oblong, apex obtuse, collecting 

hairs on upper 1/2. Achenes narrowly obovate, terete, 3- to 

6-ribbed, gland-dotted, with short twin trichomes sparsely 

distributed, carpopodium present. Pappus 1-seriate, formed 

by linear scales, minutely scabrid along the margins.

Etymology: Name derived from Greek “chilios” 

(thousand), and “phyllon” (leaf), referring to the many 

leaves that characterize the genus.

Chiliophyllum is a monotypic genus restricted to the 

Andes of central Argentina in Mendoza province. It inhabits stony slopes in an ecotonal area between high Andean 

and Monte biogeographic provinces.

Chiliophyllum is characterized by its densely disposed 

and notoriously gland-dotted leaves with fl at (non- revolute) 

margins, radiate capitula, partially paleate receptacles, yellow ray fl oret corollas, obtuse style branches and 1-seriate 

pappus.

Hitherto, Chiliophyllum was composed of three species, i.e.: C. densifolium, C. fuegianum, and C. andinum. 

However, based on morphological and molecular evidence (Karaman, 2006; Bonifacino and Funk, unpublished 

data), the polyphyletic nature of Chiliophyllum has been 

established. Besides vegetative characters that would set 

the three species apart, reproductive characters including 

the paleate nature of receptacle, type of pappus, color of 

the ray corollas, as well as the shape of style branches support splitting of the genus. As a result, Chiliophyllum is 

rendered monotypic, composed solely of C. densifolium.

Molecular evidence (Bonifacino and Funk, unpublished 

data) place Chiliophyllum in close proximity to Lepidophyllum and Chiliotrichum, as part of the strongly supported 

Chiliotrichum clade. These three genera share the partially 

paleate receptacle, styles branches obtuse, achenes mostly 

gland-dotted with no or very few twin trichomes towards 

the apex, and a pappus composed of linear scales.

Chiliophyllum densifolium Phil., Anales Univ. Chile 

21(2): 397, 1862. Phyllochilium densifolium (Phil.) Cabrera, Notas Mus. La Plata, Bot. 2(16): 172, 1937. Type: 

“[Argentina] Portillo, lado de Mendoza, 1861−1862, 

W. Díaz [s.n.]” (holotype: SGO!; isotype: LP!).

(FIGURES 11, 12, 46)

Shrubs up to 1.7 m tall, subglobose, spreading; 

branches densely disposed, ascending to erect, glanduliferous to rarely tomentose; internodes shorter than 1 mm 

long. Leaves densely arranged, persistent, alternate, patent, 5−6 × 2−2.5 mm, obovate, base slightly attenuated, 

petiole broadened towards the stem and constricted 

above, apex obtuse, margins fl at (non-revolute), notably gland-dotted on both surfaces, sometimes aseptatefl agellate trichomes on midveins and margins, midvein 

prominent abaxially, concolorous, coriaceous. Capitula

solitary, terminal, shortly pedunculate, peduncles 1−1.5 

mm long, heterogamous, radiate. Involucres 7−8 × 4−5 

mm, cylindrical; phyllaries arranged in 3 to 4 series; outer 

phyllaries 5 × 1−1.5 mm, elliptic, apex acute, gland-dotted throughout abaxial surface, chartaceous, margins 

membranaceous; inner phyllaries 5−6 × 1 mm, narrowly 

elliptic, apex acute, gland-dotted towards the apex, 

chartaceous, margin membranaceous. Receptacles slightly 

convex, partially paleate; paleae 3 to 10, 6−7 × 1 mm, 

linear to narrowly elliptic, boat-shaped, not embracing the 

achenes, with a tuft of woolly trichomes towards the apex, 

apex acute, margins membranaceous-fi mbriate. Ray fl orets 4 to 8, pistillate; corollas yellow, radiate, tube ~ 2 mm 

long, limb 4.5−6 × 2−3 mm, elliptic, 3-lobed; staminodes 

3 to 5, reduced; style shafts 2−3 mm long, base swollen, 

style branches 2−3 mm long, oblong, subequal in length. 

Disk fl orets 7 to 12, bisexual; corollas yellow, tubular, 

5-lobed, gradually broadened towards the apex, tube 

5−6 mm long, lobes 1.8−2.2 × 0.5−0.8 mm, triangular, 

gland-dotted abaxially; anther thecae 1.8−2 mm long, bases 

slightly sagittate, apical appendages 0.3−0.6 × 0.2−0.3 

mm, triangular-ovate, antheropodium notably swollen; 

style shafts 3.5−5 mm long, base swollen, style branches 

1−1.2 mm long, oblong, obtuse, collecting hairs on upper 
number 92 • 2 5

FIGURE 11. Chiliophyllum densifolium Phil. A. Habit. B. Leaves, adaxial view (left), abaxial view (right). C. Involucre. D. Outer to inner phyllaries (from left to right). E. Receptacular paleae, lateral view (left), dorsal view (right). F. Detail of pappus scale. G. Achene with pappus. H. 

Style of ray fl oret. I. Style of disk fl oret. J. Corolla of disk fl oret. K. Corolla of ray fl oret (A−K, Bonifacino et al. 77, LP).
2 6 • SMITHSONIAN CONTRIBUTIONS TO BOTANY

1/2. Achenes 4−6 mm long, narrowly obovate, terete, 3- 

to 6-ribbed, gland-dotted, with short twin trichomes very 

sparsely distributed, carpopodium present, conspicuous. 

Pappus 3−4 mm long, 1-seriate, composed of 50 to 60 linear scales, minutely scabrid along the margins.

Vernacular names: “Romero del piche” (Boelcke 

10137, LP); “Romero Pichi” (Ruiz Leal 7210, LP), 

“Romero chico, Piche” (Roig, 2001). In mapuche tongue, 

“Piche” means small, and it is a name that is also applied to 

Fabiana imbricata Ruiz and Pav. (Roig, 2001).

Phenology: Flowering specimens have been collected from January to March.

Distribution: Argentina, endemic to the Andes of 

Central Mendoza (Figure 12).

Habitat: Rocky slopes of the ecotonal area between 

Monte and High Andes biogeographic provinces (Cabrera 

and Willink, 1973) at 2,000−2,600 m range. It occurs together with Nassauvia axillaris D. Don, Berberis buxifolia 

Lam., Ephedra spp., Adesmia spp. among others.

Uses: Diuretic, used with “mate”, infusion of Ilex 

paraguariensis A. St.-Hil. – (Boelcke 10137, LP); according 

to Roig (2001) it is used for “sandstone in kidneys, liver, 

etc., in infusions taken several times a day”.

Etymology: The epithet densifolium refers to the 

leaves densely arranged along the branches.

Additional specimens examined: ARGENTINA. PROV. 

MENDOZA. Dpto. San Carlos. Camino a Laguna Diamante, 

Refugio militar Gral. Alvarado, 19-I-1963, Boelcke et al. 

FIGURE 12. Geographic distribution of Chiliophyllum densifolium Phil.
number 92 • 2 7

10137 (BAB, LP); íd., 27-I-1950, Cuezzo & Barkley 459 

(NY); Arroyo de la Quebrada “Casa de Piedra”, 19-I-1952, 

Serra 70 (F, LP); Quebrada del Paso de la Cruz de Piedra, 

2,000 m, 20-I-1941, Ruiz Leal 7279 (LP); íd., 18-I-1965, 

Ruiz Leal 23530 (LP); Rincón de los leones (Estancia La 

Plancha), 18-I-1941, Ruiz Leal 7210 (LP). Dpto. Tunuyán. 

S.loc, I-1950, s.leg. s.n. (BA 30237); al W de La Consulta, 

I-1950, s.leg. s.n. (BA 30240); Puesto Gendarmería Alférez 

Portinari, Arroyo Cascada de la Vieja, 20-I-1963, Boelcke 

et al. 10094 (BAB, LP); Arroyo Grande, Puesto Gendarmería Nacional El Portillo, 12-I-2000, Bonifacino et al. 77 

(LP); Bella Vista, 2,400 m, 4-XII-1946, Covas 18043 (SI); 

Ruta Prov. 94, Río Grande, Refugio Lemos, 2,250 m, 22-I1985, Hunziker et al. 11263 (SI); Rincón Colorado, 2,500 

m, 8-III-1933, Ruiz Leal 1281 (LP); Cordillera de Tunuyán, 

Vallecito, I-1917, Sanzin 1313 (BA); Portillo de Tunuyán, 

33o 37'S, 69o 36'W, 2,600 m, 27-I-2001, Simón & Bonifacino 826 (LP); íd,, 33o 37'S, 69o 35'W, 27-I-2001, Simón 

& Bonifacino 817 (US).

Chiliotrichiopsis Cabrera

Taxonomic History

1937. Cabrera describes Chiliotrichiopsis to include Chiliotrichiopsis keidelii, a new species based on material from Jujuy (Argentina).

1944. Cabrera transfers Gutierrezia ledifolia Griseb. 

to Chiliotrichiopsis, making the combination 

C. ledifolia (Griseb.) Cabrera.

1954. Cabrera (1954b) describes Chiliotrichiopsis mendocina (Haroldia mendocina in this paper) 

from material collected in northern Mendoza 

( Argentina).

2001. Nesom et al. describe Chiliotrichiopsis peruviana, a 

new species from southern Peru, based on material collected in Ayacucho.

Chiliotrichiopsis Cabrera, Notas Mus. La Plata, Bot. 

2(16): 172, 1937. Type: Chiliotrichiopsis keidelii (i)

Cabrera, Notas Mus. La Plata, Bot. 2(16): 172, 1937.

Shrubs more or less globose; branches densely disposed, 

ascending, 6- to 8-ribbed, branches gland-dotted and tomentose. Leaves dense to laxly arranged, persistent, alternate, 

oblong to linear or narrowly elliptic, base slightly attenuated, petiole broadened towards the stem and constricted 

above, apex obtuse to acute, margins notably revolute, 

densely tomentose on abaxial surface, discolorous, coriaceous or chartaceous. Capitula solitary (rarely arranged in 

3 or 4 capitula capitulescences), terminal, sessile to shortly 

pedunculate, homogamous, radiate or discoid. Involucres

cylindrical-campanulate, phyllaries arranged in 3 to 5 series; 

outer phyllaries with foliaceous apices, ovate to narrowly 

ovate, apex acute, tomentose and gland-dotted on abaxial 

surface, coriaceous; inner phyllaries oblong to narrowly elliptic, apex acute to subacute, tomentose and gland- dotted 

towards the apex, papery. Receptacles convex, paleate; 

paleae narrowly elliptic to narrowly ovate, embracing the 

achenes, with or without a small tuft of woolly trichomes 

towards the apex, apex acute to subacute, margins membranaceous, fi mbriated or not. Ray fl orets (when present) 5 

to 9 (−13), pistillate; corollas yellow, radiate, limb elliptic 

to oblong, 3-lobed; staminodes present; style base swollen, 

style branches linear-triangular, subequal in length. Disk

fl orets (11−) 13 to 35, bisexual; corollas yellow, tubular, 

5-lobed, broadened gradually or more or less abruptly from 

mid-length, lobes triangular; anther thecae base slightly 

sagittate, apical appendages ovate, antheropodium present, 

not swollen; style base swollen, style branches oblong to 

narrowly elliptic, apex acute, collecting hairs very short (papillae) on upper 1/2 to 4/5. Achenes obovate, terete, 3- to 

5-ribbed, dense to laxly villose, gland-dotted, carpopodium 

present. Pappus 1-seriate, composed of scales, divided towards the apex, margins fi mbriate, somewhat fused among 

each other at the base.

Etymology: Derived from Chiliotrichum and Greek 

“opsis” (similar to), denoting the resemblance of these 

plants with those belonging to Chiliotrichum.

Chiliotrichiopsis is distributed along the Andes from 

southern Peru and Bolivia to La Rioja in Argentina. The 

genus comprises three species that live in high Andean 

and Puna biogeographic provinces (Cabrera and Willink, 

1973) from 2,200 to 4,200 m.

Chiliotrichiopsis is characterized by the presence of 

external phyllaries with foliaceous apices, a paleate receptacle, gradually or more or less abruptly broadened disk 

corollas, and a 1-seriate pappus, made of fl at pale-like 

elements, divided and fi mbriate towards the apex. Chiliotrichiopsis can either have radiate or discoid heads (Chiliotrichiopsis peruviana).

Chiliotrichiopsis forms together with Nardophyllum 

and Cabreraea, a strongly supported polytomy that is diffi cult to characterize morphologically. The three genera 

are defi ned by rather contrasting morphological characters, and I have been unable to fi nd a major morphological character uniting them. Future morphological research 

focused on microcharacters might yield some interesting 

results on this respect.
2 8 • SMITHSONIAN CONTRIBUTIONS TO BOTANY

Chiliotrichiopsis mendocina, a rather enigmatic species, only known from the type and a few recent collections made near the type locality, is here segregated from 

Chiliotrichiopsis based on molecular evidence (Bonifacino 

and Funk, unpublished data) but also on morphological 

grounds. It is placed in monotypic Haroldia (see discussion under Haroldia).

Chiliotrichiopsis keidelii (i) Cabrera, Notas Mus. La Plata, 

Bot. 2(16): 172, 1937. Type: “[Argentina, Prov. Salta] 

Quebrada de la Quesera, parte inferior, 3550 m, 12-II1932, J. Keidel [s.n.]” (holotype: LP 2149!, isotype: LP 

60261!).

(FIGURES 13, 14, 47)

Shrubs 0.5−1.5 m tall, more or less globose, densely 

branched; branches erect, 6- to 7-ribbed, tomentose and 

gland-dotted, internodes 1−3 mm long. Leaves densely disposed, persistent, alternate, patent, without a tuft of woolly 

trichomes on axils, 5−15(−25) × 1−2 (−4) mm, linear to 

narrowly elliptic or oblong, base attenuated, petiole broadened towards the stem and constricted above, apex obtuse, 

sometimes acute, margins revolute, densely tomentose, and 

gland-dotted on abaxial side, densely gland-dotted on adaxial side, discolorous, coriaceous. Capitula solitary, terminal, sessile, heterogamous, radiate. Involucres 6−8 × 4−6 

mm, cylindrical-campanulate; phyllaries arranged in 3 to 4 

series; outer phyllaries 3−4 × 2−2.5 mm, ovate, apex acute, 

outermost ones with foliaceous apices, tomentose on abaxial surface, gland-dotted towards the apex, coriaceous, 

margins membranaceous; inner phyllaries 5.5−7 × 1.5−2 

mm, oblong to narrowly elliptic, apex subacute, glabrous 

to slightly gland-dotted, papery, margins membranaceous. 

Receptacles convex, paleate; paleae 7.5−9 × 1−1.5 mm, narrowly obovate, embracing the achenes, apex acute, tomentose and gland-dotted towards the apex, with a tuft of woolly 

trichomes towards the apex, margins membranaceous. Ray 

fl orets 5 to 9 (−13), pistillate; corollas yellow, radiate, tube 

2−2.5 mm long, limb 6−7 × 1.5−3 mm, narrowly elliptic to 

elliptic, 3-lobed; staminodes (2−) 3 or 4; style shafts 3−4 

mm long, base not swollen, style branches ~ 2 mm long, 

triangular, subequal in length. Disk fl orets (11−)13 to 18 

(−24), bisexual; corollas yellow, tubular, 5-lobed, gradually broadened towards the apex, tube 4−7 mm long, lobes 

1.5−2 × 0.7 mm, triangular, gland-dotted abaxially; anther 

thecae 2.5−3.5 mm long, bases slightly sagittate, apical appendages 0.5−0.7 × 0.2−0.3 mm, ovate, antheropodium 

present, not swollen; style shafts 3−5 mm long, base slightly 

swollen, style branches 2−3 mm long, narrowly elliptic, apex 

acute, collecting hairs very short (papillae) on upper 3/4. 

Achenes 4−6 mm long, obovate, terete, densely villose and 

gland-dotted, twin trichomes ~ 1 mm long, carpopodium 

present; ray fl orets achenes 3- to 4-ribbed, mainly villose 

along ribs; disk fl orets achenes 2 or 3(−4)-ribbed, uniformly 

villose throughout the achene surface. Pappus 2.5−3.5 (−5) 

mm long, 1-seriate, composed of (11−) 17−22 (−29) scales, 

divided towards the apex, margins laciniate towards the 

apex, fused to each other at the base.

Vernacular names: “Trompo” (Venturi 3012b, LP); 

“Tola” (Venturi 8724, SI); “Tola’e vizcacha”, “Tola tuni” 

(Arenas and Martinez s.n., SI); “Tola viscacha” (Claren 

11495, CORD); “legia ploma” (Massy 396, US).

Phenology: Flowering specimens have been collected 

from December to April.

Distribution: Argentina, common in Jujuy, Salta, 

and Tucuman, also found in Catamarca. In Bolivia in department of Potosí (Figure 14).

Habitat: Puna (Cabrera and Willink, 1973) from 

2,200 to 4,300 m, grows on rocky mountain slopes with a 

poor vegetation cover or associated to tussok grasses formations (Festuca spp., Stipa spp., etc.). It is usually associated to 

other Puna shrubs like Parastrephia spp., not forming dense 

consociations.

KEY TO THE SPECIES OF CHILIOTRICHIOPSIS CABRERA

 1a. Capitula discoid; achenes laxly villose throughout the whole surface. (Endemic to Peru: Ayacucho) . . . . . C. peruviana

 1b. Capitula radiate; achenes densely villose throughout the whole surface or villose pubescence mostly confi ned to upper 

part . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2

 2a. Branches erect; leaves linear no narrowly elliptic or oblong, coriaceous; tuft of woolly trichomes absent from leaf axils;

achenes densely villose throughout the whole surface, most twin trichomes ~ 1 mm long. (Northern Argentina and southern Bolivia). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. keidelii

 2b. Branches defl exed; leaves narrowly elliptic, chartaceous; tuft of woolly trichomes present in leaf axils; achenes villose 

pubescence mostly confi ned to the upper part, most twin trichomes not longer than 0.5 mm. (Restricted to Argentina: 

Catamarca and La Rioja) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. ledifolia
number 92 • 2 9

FIGURE 13. Chiliotrichiopsis keidelii Cabrera. A. Habit. B. Leaves, adaxial view (left), abaxial view (right). C. Involucre. D. Outer to inner 

phyllaries (from left to right). E. Receptacular paleae, lateral view (left), dorsal view (right). F. Detail of pappus scales. G. Achene with pappus. 

H. Style of ray fl oret. I. Style of disk fl oret. J. Corolla of disk fl oret. K. Corolla of ray fl oret (A−K, Simón & Bonifacino 520, LP).
3 0 • SMITHSONIAN CONTRIBUTIONS TO BOTANY

FIGURE 14. Geographic distribution of Chiliotrichiopsis keidelii Cabrera. Note the distribution of the intermediate specimens (triangles), occupying an area more or less placed inbetween the known distributions for Chiliotrichiopsis keidelii and Chiliotrichiopsis ledifolia.
number 92 • 3 1

Uses: Used as fi rewood (Burkart and Troncoso 

11898, LP).

Etymology: The specifi c epithet honors geologist 

Juan Keidel, collector of the type specimen.

Note 1: There are a few specimens that have characteristics that set them apart from typical Chiliotrichiopsis keidelii. In these specimens, most characteristics are 

like Chiliotrichiopsis keidelii, but the shape of the leaves 

and the pubescence of the achenes are similar to those of 

Chiliotrichiopsis ledifolia. These structurally intermediate 

specimens have been collected in an area also somewhat 

intermediate between the geographic distributions of the 

two species (Figure 14). I consider that given the lack of 

a complete series of morphologically intermediate populations between both species, they are clearly separate and are 

maintained as distinct specifi c entities here.

Additional specimens examined: ARGENTINA. PROV. 

CATAMARCA. Not defi ned Dpto. Region Andina, 1893, s. leg.

(LP). Dpto. Andalgalá. Entre cerro de Las Capillitas y Muschaca, XI-1874, Schickendantz 39 (CORD). PROV. JUJUY.

Dpto. Capital. Entre León y Nevado de Chañi, Las Cuevas, 

3,000 m, III-1963, Fabris et al. 4079 (LP, CORD). Dpto. 

Cochinoca. Peñas de Ascalte, a 10 km aprox. al WSW de 

Cochinoca, 22o 46'S, 65o 57'W, 21-XII-1980, Arcipnete s.n. 

(NY, SI); Cochinoca, Cerro de la Candelaria, 22o 43'S, 65o 

54'W, s. date, Arenas & Martinez s.n (SI); Río Despensa, 

4,200 m, 19-I-1971, Boelcke et al. s.n (BAA 7146); Cochinoca, III-1975, Cabezas 21 (SI); íd., III-1975, Cabezas 12

(SI); Abra Pampa, 8 km al S, 3,700 m, 30-I-1971, Ellenberg 4282 (US).Dpto. Humahuaca. Tres Cruces, III-1929, 

Budin 10 (BA); El Aguilar, 4,300 m, 14-I-1948, Cabrera 

9245 (BAB, LP); Mina Aguilar, 4,000 m, 18-II-1970, Cabrera & Frangi 20662 (LP); Mina Aguilar, 4,000 m, 11-II1968, Cabrera et al. 18955 (LP); íd., 4,200 m, 23-II-1963, 

Cabrera et al. 15463 (LP, NY); Tres Cruces, Puente del 

Diablo, 4,000 m, 21-II-1959, Fabris & Marchionni 1728

(LP); íd., 4,000 m, 21-II-1959, Fabris & Marchionni 1748

(LP); Mina Aguilar, Arroyo Padrioc, 3,800 m, XII-1964, 

Fernandez s.n. (BAA 4760); Sierra del Mal Paso, entre el 

pie de la sierra y el cerro Tocante, I-1988, Fernandez s.n. 

(BA 76676); íd., I-1988, Fernandez s.n. (BA 76677); Mina 

Aguilar, vega a 4 km al N del Molino, 4,000 m, III-1983, 

Hunziker 10549 et al. (SI); 19 km al E de Humahuaca, 

ruta prov 73, 5-V-1999, Krapovickas & Seijo 47710

(NY); Mina Aguilar, Río Despensas, 4,100 m, 19-I-1971, 

Ruthasatz s.n. (BAA 9987); Mina Aguilar, 4,000−4,300 m, 

7-I-1950, Schwabe 441 (BAB); Tres Cruces, 3,820 m, 11-I1950 Schwabe 482 (BAB); Mina Aguilar, arriba del molino, 

4,300−4,400 m, 16-I-1953, Sleumer 3397 (LIL); Tres Cruces, 2-II-1944, Soriano 652 (SI); Sierra de Aguilar, 4,000 m, 

29-III-1929, Venturi 8724 (SI, US). Dpto. Rinconada. Mina 

Pirquitas, 4,200 m, 20-I-1948, Cabrera 9344 (BAB, LP); 

camino a Abra-Pampa, 4,160 m, 24-I-1971, Boelcke et al. 

s.n. (BAA 7229). Dpto. Santa Catalina. El Alto, W de Santa 

Catalina, 4,000 m, 21-I-1966, Cabrera et al. 17677 (LP, 

BAA); Mina Perdida, ca. 4,100 m, 25-26-I-1901, Claren 

11495 (CORD). Dpto. Susques. Al S de Coranzuli, 3937 m, 

23o1'41"S, 66o24'11"W, 6-I-2005, Bonifacino & Muttoni 

1448 (US). Dpto. Tilcara. San Gregorio, above Tilcara, 

12,000 ft, 11-II-1939, Balls 5996 (K, US); subiendo la Quebrada de Huichaira-Pocollio, 3,210 m, 24-I-1999, Beck 

26507 (US); Huacalera, Quebrada de Abra de La Cruz, 

3,900 m, 2-III-1955, Cabrera 12145 (LP); Alto de la Laguna 

Negra, 4,100 m, 3-III-1961, Cabrera & Hernández 13989

(LP); Pampa Corral, 3,800 m, 18-I-1966, Fabris et al. 6380

(LP, BAA); Sisilera, 10-I-1974, Kiesling et al. 376 (LP); Cimarrones (cerca de Co. Mudana), 16-I-1974, Kiesling et al. 

390 (LP); Cerro al E de Huacalera, 3,900 m, 20-III-1967, 

Löerner 395 (LP); Cerro Alto de Lima, W de Huacalera, 

4,200 m, 27-III-1967, Löerner 636 (LP); Tilcara, arriba de 

San Gregorio, 3,850 m, 29-XII-1952, Sleumer 3127 (SI); 

28 km W of junction routes 9 and 52, 3,350 m, 20-II-1993, 

Stuessy & Morales 12980 (LP). Dpto. Tumbaya. camino al 

Abra de Lipan, 3,900 m, III-1979, Cabrera et al. 30553 (SI); 

Subida al Abra de Lipan, 3,800 m, 15-II-1980, Cabrera et 

al. 31657 (SI); Purmamarca a Abra de Pibes, 17-II-1937, 

Castellanos s.n. (LP, BA 20202); Tumbaya, camino de 

Purmamarca a Abra de Lipán, 3,900 m, 14-II-1985, Kiesling et al. 5230 (SI); Peñarco, Volcan, II-1931, Pastore s.n.

(BA 31/84); desde Purmamarca a intersección con ruta 40, 

3,450 m, 5-I-2001, Simon & Bonifacino 519 (LP); camino a 

Lipan, a aprox. 30 km de Purmamarca, 23o42'S, 65o38'W, 

3950 m, 5-I-2001, Simon & Bonifacino 520 (LP); San Antonio de los Pibes, 3,850 m, 5-I-1953, Sleumer 3222 (SI); 

28 km W of Purmamarca on gravel road, Cuesta de Lipán, 

3,600 m, 21-II-1993, Stuessy & Morales 12986 (LP). Dpto. 

Valle Grande. Caspalá, cerros a 3,200 m, 1-III-1940, Burkart & Troncoso 11898 (SI, LP); Caspalá, 3,000 m, 25-

XII-1962, Fabris 3613 (LP); Abra Colorada, 20 km W 

Caspalá, 4,200 m, 26-XII-1962, Fabris 3645 (LP); Ciénaga 

Grande, entre Abra Colorada y Caspalá, 11-III-1967, Fabris & Crisci 6884 (LP); Caspalá, 12-III-1967, Fabris & 

Crisci 6995 (LP). Dpto. Yavi. Cuesta de Toquero, camino 

a Santa Catalina, 4,000 m, 31-I-1943, Cabrera 7797 (BAB, 

LP); Cerrillos a Escaya, 6-II-1937, Castellanos s.n. (LP, 

BA 20212). PROV. SALTA. Dpto. Cachi. Cerro de Cachi, 

I-1897, Spegazzini s.n. (ex LPS 17184 in LP); Cerro de Cachi, 

I-1897, Spegazzini s.n. (ex LPS 17179 in LP). Dpto. Cafayate. Sierra de los Quilmes, 29-I-1943, Castellanos s.n.

(BA 47093). Dpto. Caldera. Alrededores del Nevado 
3 2 • SMITHSONIAN CONTRIBUTIONS TO BOTANY

del Castillo, 19-23-III-1873, Lorentz & Hieronymus 92

(CORD); Subida al Nevado del Castillo, Tres Lagunas-Las 

Cuevas, 4,100−4,200 m, 16-III-1952, Sleumer & Vervoorst 

3011 (LP). Dpto. Iruya. Quebrada de Iruya, 3,750 m, 27-

III-1972, Suti & Montaña s.n. (LP). Dpto. Los Andes. Quebrada de Polvorillas, 4,000 m, 2-II-1944, Cabrera 8393 (LP); 

íd., 4,300 m, 11-II-1945, Cabrera 8635 (LP, US); Región de 

La Puna, camino a Mina Concordia, 23-I-1949, Cabrera & 

Schwabe 55 (BAB, LP). Dpto. Poma. Ruta 51, punto más 

alto de Abra de Muñano, 4,050 m, 23-I-1989, Arriaga et al. 

504 (BA); Abra de Muñano, 4,200 m, 9-II-1946, Cabrera 

8986 (LP). PROV. TUCUMAN. Dpto. Chicligasta. Ea Santa 

Rosa, 4,000 m, 18-I-1927, Venturi 4757 (LP, SI). Dpto. 

Tafí del Valle. Abra de Lara, 4,000 m, II-1903, Baer 136

(BAB); Cuesta de Los Cardones, 3,500 m, 30-I-1933, Burkart 5225 (LP, SI); Quebrada del Baron, Los Planchones, 

3,300 m, 7-II-1958, Fabris 1380 (LP); Cerro Muñoz, ladera 

oriental, 14-II-1958, Fabris 1548 (LP); Cerro El Negrito 

(ladera SE) +/- 3,800 m, 26o37'S, 65o44'W, 19-I-1964, Giusti et al. s.n., (BAA 3861); Cerro Muñoz, 3,300−4,000 m, 

22-II-1905, Lillo 4201 (US); Cañada del Alegau, 3,800 m, 

5-IV-1926, Schreiter 4252 (NY); El Infi ernillo, Cerro El 

Negrito, 26o43'S, 65o46'W, 3,200 m, 5-I-2001, Simon & 

Bonifacino 443 (LP); Quebrada del Chorro, 3,100 m, 23-

IV-1926, Venturi 4105 (LP, SI); Sierra del Cajón, 3,400 m, 

27-IV-1926, Venturi 6229 (US); Cumbres Calchaquíes, km 

93, entre Zanja de los Cardones y Alto del Tío, 2,800 m, 

15-XII-1969, Vervoorst 7635 (LIL); Sierras Calchaquíes, 

Peñas Azules, 19-III-1974, Zardini 330 (LP). Dpto. Tafi 

Viejo. La Cienaguita, Cumbres Calchaquíes, 2,200 m, 

4-I-1913, Rodriguez 1240 (LP). Dpto. Trancas. Trancas, 

Cuesta del Arca, XII-1896, Spegazzini s.n. (ex LPS 17186 

in LP). BOLIVIA. NOT DEFINED DPTO. Buen Retiro, 4,000 m, 

XII-1901, Cárdenas 46 (LP). DPTO. POTOSÍ. Tupiza, 3,700 

m, II-1904, Fiebrig 3003 (BM); Salitre pr. Yavi in Argentina, 6-I-1902, Fries 1043 (CORD); 8 km al E de Chuqui, 

4,000 m, 30-VII-1999, Massy 396 (US).

Additional specimens examined with intermediate 

characteristics between Chiliotrichiopsis keidelii and Chiliotrichiopsis ledifolia: ARGENTINA. PROV. CATAMARCA.

Not defi ned Dpto. Vayas altas, alpen von Catamarca, 

9,000–11,000 ft, I-1872, Lorentz 609 (CORD). Dpto. 

Ambato. Sierra de Ambato, Falda E, subiendo desde El 

Rodeo hacia el Cerro Manchado, 2,700−2,900 m, 23-

25-II-1967, Hunziker 19162 (CORD); Sierra de Ambato 

(falda E, frente a El Rodeo), Los Cajones, ~3,100 m, 15-

17-XII-1971, Hunziker 21667 (CORD); Sierra de Ambato, Falda E, subiendo desde El Rodeo hacia el Cerro 

Manchado, 2,600−2,700 m, 23−25-II-1967, Hunziker 

19125 (CORD); íd., 2,700−2,900 m, 23−25-II-1967, 

Hunziker 19179 (CORD); íd., 2,900−3,100 m, 23−25-

II-1967, Hunziker 19286 (CORD); íd., rumbo a Casa de 

Cubas, 3,100−3,200 m, 28−30-I-1968, Hunziker & Di 

Fulvio 19737 (CORD, F); íd., rumbo a Casa de Cubas, 

3,000 m, 28−30-I-1968, Hunziker & Di Fulvio 19821

(CORD). Dpto. Andalgalá. Candado, 6-XII-1896, Bruch 

s.n. (US); El Candado, 2,700 m, II-1915, Jörgensen 1272

(BA, BAB, SI, US). PROV. SALTA. Dpto. Guachipas. Pampa 

Grande, XII-1897, Spegazzini s.n. (ex LPS 17187 in LP). 

PROV. TUCUMÁN. Dpto. Chicligasta. Estancia Las Pavas, 

en faldas de cerro, 2,650 m, 15-III-1924, Venturi 3012b 

(BAB, LP); Estancia Las Pavas, Puesto La Cascada, 2,600 

m, 10-III-1924, Venturi 3012 (NY, SI); Estancia Santa 

Rosa, 3,000 m, 2-XII-1926, Venturi 4639 (BA, BAB, F, 

LP, SI); Estancia Santa Rosa, 4,000 m, 18-I-1927, Venturi 4757 (BA, BAB). Dpto. Tafi del Valle. Abra de Lara, 

4,000 m, II-1902, Baer s.n. (BA 24906); Sierras Calchaquíes, Peñas Azules, 3,400 m, s. date, Burkart 5210

(LP, SI); Cumbres Calchaquíes, Quebrada del Barón, 

3,400 m, 26o45'S, 65o46'W, 16-III-1984, Gómez-Soza & 

Múlgura 244 (SI); Lara, 3,200 m, 25-I-1912, Rodríguez 

275 (BA, SI). Dpto. Tafi Viejo. Alto de Anfamá, I-1943, 

Descole 1618 (BAB, BM).

Chiliotrichiopsis ledifolia (Griseb.) Cabrera, Notas Mus. 

La Plata, Bot. 9(46): 244, 1944. Gutierrezia ledifolia 

Griseb., Pl. Lorentz.: 125, 1874. Type: “[Argentina, 

Prov. Catamarca] Vayas altas, alpen von Catamarca, 

9,000−11,000 ft, mitte Jan 1872, Lorentz 610” (lectotype: CORD!, isolectotype: CORD!, designated by 

Cabrera, 1944).

(FIGURES 15, 16, 48)

Shrubs up to 1.5 m tall, more or less globose, spreading, densely branched; branches defl exed, 6- to 7-ribbed, 

gland-dotted, slightly tomentose, internodes 3−7 mm 

long. Leaves laxly disposed, persistent, alternate, ascending with a tuft of woolly trichomes on leaves axils, 

17−30 × 2−4 mm, narrowly elliptic, base attenuated, petiole broadened towards the stem and constricted above, 

apex acute, margins revolute, tomentose on abaxial side, 

and laxly tomentose on adaxial side, gland-dotted on 

both surfaces with prominent vesicular capitate glandular trichomes, discolorous, chartaceous. Capitula solitary, 

terminal, pedunculate, peduncles 5−20 mm long, heterogamous, radiate. Involucres 7−8 × 4−5 mm, cylindricalcampanulate; phyllaries arranged in 3 to 4 series; outer 

phyllaries 5−6 × 1.5−1.7 mm, narrowly ovate, apex acute, 

outermost ones with foliaceous apices, tomentose and 
number 92 • 3 3

FIGURE 15. Chiliotrichiopsis ledifolia (Griseb.) Cabrera. A. Habit. B. Leaves, adaxial view (left), abaxial view (right). C. Involucre. D. Outer 

to inner phyllaries (from left to right). E. Receptacular paleae, lateral view (left), dorsal view (right). F. Detail of pappus scales. G. Achene with 

pappus. H. Style of ray fl oret. I. Style of disk fl oret. J. Corolla of disk fl oret. K. Corolla of ray fl oret (A−K, Simón & Bonifacino 725, LP).
3 4 • SMITHSONIAN CONTRIBUTIONS TO BOTANY

gland-dotted on abaxial surface, coriaceous, margins membranaceous, fi mbriate; inner phyllaries 6−7 × 1.7−2 mm, 

oblong to narrowly elliptic, apex acute, with a small tuft 

of woolly trichomes towards the apex, tomentose and 

gland-dotted towards the apex, papery, margins membranaceous, fi mbriate. Receptacles slightly convex, paleate; paleae 7−8 × 1.5 mm, narrowly obovate, embracing 

the achenes, apex acute, tomentose and gland-dotted 

towards the apex, margins membranaceous, fi mbriate 

towards the apex. Ray fl orets 7 to 8, pistillate; corollas 

yellow, radiate, tube 1−2 mm long, limb 6−9 × 3−4 mm, 

narrowly elliptic, 3-lobed; staminodes 2 to 3; style shafts 

4−5 mm long, base swollen, style branches ~ 2 mm long, 

triangular, subequal in length. Disk fl orets 13 to 19, bisexual; corollas yellow, tubular, 5-lobed, gradually 

broadened towards the apex, tube 4.5−7 mm long, lobes 

1.5−2 × 0.6−0.8 mm, triangular, gland-dotted abaxially; 

anther thecae 2.3−3.0 mm long, bases slightly sagittate, 

apical appendages 0.5−0.8 × 0.2−0.3 mm, ovate-triangular, antheropodium present, swollen; style shafts 4−6 mm 

long, base slightly swollen, style branches 2−2.3 mm long, 

triangular ovate, apex acute, collecting hairs short on 

upper 1/2 to 3/4. Achenes similar in disk and ray fl orets , 

3.5−5 mm long, obovate, terete, 3- to 5-ribbed, laxly vilFIGURE 16. Geographic distribution of Chiliotrichiopsis ledifolia (Griseb.) Cabrera.
number 92 • 3 5

lose and gland-dotted, twin trichomes denser towards the 

apex, twin trichomes shorter than 0.5 mm long. Pappus

1.5−2.5 mm long, 1-seriate, composed of 19 to 25 scales, 

triangular ovate, divided towards the apex, margins laciniate towards the apex, somewhat fused to each other 

at the base.

Vernacular names: Not known.

Phenology: Flowering specimens have been collected from January to February.

Distribution: Argentina, restricted to central mountains in Catamarca and La Rioja (Figure 16).

Habitat: Puna and Puna-like vegetation (Cabrera 

and Willink, 1973) from 2,200 to 3,300 m, grows associated to tussok grasses formations (Festuca spp., Stipa spp., 

etc.), isolated or forming dense consociations.

Uses: Not known.

Etymology: The specifi c epithet is derived from 

Greek “ledon”, shrub belonging to genus Cistus from Cyprus, from which a particular kind of resin is obtained. Apparently, the name would refer to the similarity between the 

shape and smell of the leaves of Chiliotrichiopsis ledifolia 

with those of Cistus.

Note 1: At CORD there are two specimens collected 

by Lorentz (Lorentz 609 and Lorentz 610) whose labels perfectly match the type locality mentioned in the protologue 

of Gutierrezia ledifolia and that were identifi ed as such by 

Grisebach (1874). When transferring the species to Chiliotrichiopsis, Cabrera (1944) noticed that only one of those 

specimens (Lorentz 610) corresponded with Chiliotrichiopsis ledifolia and was selected as lectotype. The other specimen, Lorentz 609, clearly belongs to C. keidelii, although it 

presents some characteristics intermediate between this species and C. ledifolia (see Note 1 under C. keidelii).

Additional specimens examined: ARGENTINA. PROV. 

CATAMARCA. Dpto. Belén. Las Bayas, en cerros al W de 

Granadillas, 2,800 m, 26-I-1952, Sleumer & Vervoorst 

2566 (LP, US). PROV. LA RIOJA. Dpto. Chilecito. Río del Oro, 

I-1912, Bodenbender 16267 (CORD); proximidades de 

Mina El Oro, 29o 6' 49.9"S, 67o 40'21.7", 2222 m, 3-XII2006, Bonifacino et al. 2847 (US); W Sierra de Famatina, 

Mina El Oro, 3,200 m, 7-II-1956, Calderón 1141 (BAA, 

CORD, SI); Camino a Mina El Oro, 10-I-1959, Dawson & 

Guarrera 3132 (LP, BA); Chilecito, Camino a Mina El Oro, 

29o 06'S, 67o 42'W, 2,300−2,500 m, 23-I-2001, Simon & 

Bonifacino 725 (LP); íd., 23-I-2001, Simon & Bonifacino 

740 (LP). Dpto. Famatina. Sierra Famatina, entre la Mina 

Teresa y la altura del Espíritu Santo, 25-I-1879, Hieronymus & Niederlein 787 (CORD); Sierra Famatina, en las 

cercanías del Pie de la Cuesta, más arriba del Vallecito, 16-

20-I-1879, Hieronymus & Niederlein 765 (CORD); Sierra 

de Famatina, arriba del Vallecito: La Hoyada, ~2,500 m, 

10-16-I-1908, Jiménez 14948 (CORD); íd., 10-16-I-1908, 

Jiménez 14966 (CORD); Sierra de Famatina, La Vega de 

la Hoyada, 22-I-1908, Jiménez 15132 (CORD); Sierra de 

Famatina, arriba de Vallecito, La Hoyada, ca. 2,500 m, 31-

I-1908, Kurtz 15001 (CORD).

Chiliotrichiopsis peruviana G. L. Nesom, H. Rob. and 

A. Granda, Brittonia 53: 430, 2001. Type: “Peru. Ayacucho: prov. Puquio, Puente Torre Muestre at 76 km 

on rd. Nazca-Puquio, sep quebrada, N facing, 3,600 m, 

2-XII-1997, M. Weigend & H. Förther 97/676” (holotype: US!, isotypes: USM, MSB-n.v.; isoparatype: “Perú, 

Dpto. Ayacucho, Prov. Lucanas, carretera NazcaPuquios, 7-VIII-1997, Flores et al. 2109”: LP!).

(FIGURES 17, 18, 49)

Shrubs up to 1.5 m tall, more or less globose, lax to 

densely branched; branches erect, 6- to 8-ribbed, ribs discontinuous more or less evident, tomentose and gland-dotted, 

internodes 1−5 mm long. Leaves laxly disposed, persistent, 

alternate, patent, without a tuft of woolly trichomes on axils, 

15−45 (−50) × 1.5−2.5 mm, linear, base attenuated, petiole 

broadened towards the stem and constricted above, apex 

obtuse, recurved and slightly mucronate, margins revolute, 

densely tomentose and gland-dotted on abaxial side, densely 

gland-dotted on adaxial side, discolorous, coriaceous. Capitula solitary or more often arranged in capitulescences of 2 

to 3 capitula, terminal, pedunculate, peduncles up to ~ 8 mm 

long, homogamous, discoid. Involucres 10−12 × 8−10 mm, 

campanulate; phyllaries arranged in 4 to 5 series; outer 

phyllaries 6 × 2−3 mm, narrowly ovate, apex acute, coriaceous, tomentose and gland-dotted on abaxial surface, margins membranaceous; inner phyllaries 7−10 × 2.5−3 mm, 

narrowly elliptic, apex acute, gland-dotted towards the 

apex, papery, margins membranaceous, broad. Receptacles

convex, paleate; paleae 10−11 × 1−2 mm, narrowly elliptic, embracing the achenes, apex acute-subulate, glanddotted towards the apex, margins membranaceous. Florets

30 to 35, bisexual; corollas yellow, tubular, 5-lobed, more 

or less abruptly broadened from mid-length, tube ~ 3 mm 

long, throat 2−3 mm long, lobes 1.5−2.2 × 0.8−1 mm, 

triangular, gland-dotted abaxially; anther thecae 2.3−

2.5 mm long, bases slightly sagittate, apical appendages 

0.7−1 × 0.5 mm, ovate, antheropodium present, not swollen; style shafts 4−6 mm long, base notably swollen, style 

branches 2.7−3.2 mm long, oblong, apex acute, collecting 

hairs very short (papillae) on upper 3/4 to 4/5. Achenes

6−8 mm long, narrowly obovate, terete, 3- to 4-ribbed, laxly 
3 6 • SMITHSONIAN CONTRIBUTIONS TO BOTANY

FIGURE 17. Chiliotrichiopsis peruviana G. L. Nesom, H. Rob., and A. Granda. A. Habit. B. Leaves, adaxial view (left), abaxial view (right). 

C. Involucre. D. Receptacular paleae, lateral view (left), dorsal view (right). E. Outer to inner phyllaries (from left to right). F. Detail of pappus 

scales. G. Achene with pappus. H. Style. I. Corolla (A−I, Flores 2109, LP).
number 92 • 3 7

villose and gland-dotted throughout the whole surface, twin 

trichomes shorter than 0.5 mm long, carpopodium present. 

Pappus ~ 2.5 mm long, 1-seriate, composed of 11−16 scales, 

triangular to narrowly elliptic, margins laciniate towards 

the apex, somewhat fused among each other at the base.

Vernacular names: Not known.

Phenology: Flowering specimens have been collected from August to December.

Distribution: Peru, restricted to department of Ayacucho (Figure 18).

Habitat: Puna (Cabrera and Willink, 1973) from 

3,500 to 3,600 m, along steep mountain slopes dominated 

by shrubs amid Polylepis spp. (Nesom et al., 2001).

Uses: Not known.

Etymology: The specifi c epithet refers to the country to which this species is an endemic.

Chiliotrichum Cass.

Taxonomic History

1788. Forster describes the species Amellus diffusus from 

material collected by himself in Tierra del Fuego.

1817. Cassini establishes Chiliotrichum and creates the 

species Chiliotrichum amelloideum. Cassini based 

his description on Forster species but did not employ the epithet used by Forster creating in consequence an illegitimate name.

1826. Sprengel transfers Amellus diffusus to Aster L. (Aster 

magellanicus Spreng.) without taking Forster’s 

epithet and then creating a superfl uous name.

1829. Tausch considers that the differences between the 

species described by Forster (1787) as Amellus

FIGURE 18. Geographic distribution of Chiliotrichiopsis peruviana G. L. Nesom, H. Rob., and A. Granda.
3 8 • SMITHSONIAN CONTRIBUTIONS TO BOTANY

diffusus and the rest of the species belonging to 

the genus Amellus are enough to create a new 

genus, and apparently unaware of the genus created by Cassini in 1817 (Chiliotrichum), describes 

Tropidolepis Tausch, making the combination 

Tropidolepis diffusa (G. Forst.) Tausch.

1831. Lessing, based on material collected by Poeppig in 

Chile, describes Chiliotrichum rosmarinifolium 

Less., pointing out that the main difference of his 

new species from the previous one (Chiliotrichum 

amelloideum) is in the shape of the leaf.

1833. Nees lowers the rank of the two species described 

to the date, Chiliotrichum amelloideum y C. rosmarinifolium, to varieties of Chiliotrichum amelloideum, C. amelloideum var rosmarinifolium, C.

amelloideum var. diffusum and adds a third variety, C. amelloideum var. lanceolatum.

1836. Candolle (1836a), in agreement with Nees, accepts 

only one species, Chiliotrichum amelloideum, 

but only two of Nees’ varieties, C. amelloideum

var. rosmarinifolium and C. amelloideum var. 

lanceolatum.

1848. Hombron and Jacquinot name and present analytical illustrations for two new species, Chiliotrichum ovatifolium and C. feliciae, and one variety 

C. ovatifolium var. retortum.

1858. Mueller places the two names created by Hombron 

and Jacquinot, and C. rosmarinifolium in the synonymy of Chiliotrichum amelloideum.

1873. Philippi describes a new species, Chiliotrichum 

virgatum.

1894. Philippi describes three new species, Chiliotrichum 

angustifolium, C. longifolium, and C. tenue and a 

variety of Chiliotrichum rosmarinifolium, C. rosmarinifolium var. glabrescens.

1898. Kuntze gives priority to the name coined by Forster 

(1788) and makes the combination Chiliotrichum 

diffusum.

1899. Spegazzini describes three forms in Chiliotrichum diffusum, C. diffusum f. typica, C. diffusum f. media,

and C. diffusum f. angustifolium (“angustifolia”).

1901. Reiche accepts only two species in Chiliotrichum, 

C. diffusum and C. rosmarinifolium.

1971. Cabrera also accepts only two species, Chiliotrichum diffusum and C. rosmarinifolium, pointing 

out that the main difference between them is the 

leaf width.

Chiliotrichum Cass., Bull. Sci. Soc. Philom. Paris 4(2): 69, 

1817. Tropidolepis Tausch, Flora 12 (5): 68, 1829. Chiliotrichum sect. Euchiliotrichum Hook. f., Bot. Antarct. 

Voy., Vol. 1, Fl. Antarct. 2: 304, 1846 [1847]. Type: 

Amellus diffusus G. Forst., Fasc. Plant. Magellan.: 39, 

1788 (≡ Chiliotrichum diffusum (G. Forst.) Kuntze, 

Rev. Gen. Pl. 3(2): 141, 1898).

Shrubs hemispheric to globose; branches densely disposed, ascending, not ribbed young branches gland-dotted 

to tomentose and gland-dotted. Leaves densely arranged, 

persistent, alternate, patent to ascending, linear to elliptic, narrowly elliptic or obovate, base attenuated, petiole 

broadened towards the stem and constricted above, apex 

obtuse to acute, margins revolute to slightly revolute, 

densely tomentose on abaxial surface, and sparsely tomentose on adaxial surface or with no aseptate- fl agellate 

trichomes on this surface, gland-dotted on both sides, 

midvein sunken on adaxial side, discolorous, coriaceous. 

Capitula solitary, terminal, pedunculate, heterogamous, 

radiate. Involucres cylindrical-campanulate, phyllaries 

arranged in 3 to 5 series; outer phyllaries triangular to 

ovate, apex acute, tomentose towards the apex on abaxial 

side, coriaceous; inner phyllaries narrowly elliptic, apex 

acute, glabrous with a tuft of woolly trichomes towards 

the apex, membranaceous. Receptacles convex to slightly 

convex, partially paleate; paleae linear to narrowly elliptic, not embracing the achenes, with a tuft of woolly trichomes towards the apex, apex acute, margins narrow, 

membranose-fi mbriate. Ray fl orets 3 to 14, pistillate; corollas white, radiate, limb narrowly elliptic, 3-lobed; staminodes present; style base slightly swollen, style branches 

narrowly ovate, subequal in length. Disk fl orets 7 to 27, 

bisexual; corollas yellow, tubular, 5-lobed, more or less 

abruptly broadened from about mid-length, lobes triangular; anther thecae base slightly sagittate, apical appendages ovate, antheropodia present, not swollen; style base 

swollen, style branches oblong, apex subacute to obtuse, 

collecting hairs on upper 2/5 to 1/2. Achenes narrowly obovate, terete, 4- to 6-ribbed, gland-dotted, sometimes with 

few twin trichomes sparsely distributed towards the apex, 

carpopodium present, conspicuous. Pappus 2- to 3-seriate, 

composed of linear scales, scabrid along the margins.

Etymology: Name derived from Greek “chilios” 

(thousand) and “trich” (hair). Cabrera (1971) explained 

that Cassini was probably referring to the tomentose indument on the abaxial surface of the leaves. On the original 

description, Cassini was rather emphatic on the “très nombreouses” elements of the pappus, so it is also likely that the 

French synantherologist might have been referring to this 

trait instead of the leaf pubescence when coining the name 

for his new genus.
number 92 • 3 9

Chiliotrichum is composed of two species distributed 

from central Argentina and Chile, to Tierra del Fuego 

along the Andes and in Patagonia. This includes the vast 

archipelago west of Chile and Islas Malvinas. The genus is 

the most evenly and continuously distributed of the Chiliotrichum group, and it occurs in very different habitats 

that range from Andean meadows above the tree line to 

the windswept barren grasslands of northern Tierra del 

Fuego.

Chiliotrichum is morphologically well defi ned, with 

radiate heads, partially paleate receptacles, white ray corollas, and 2- to 3-seriate pappus formed by linear scales.

Traditionally, Chiliotrichum was considered to be 

composed of two species, C. diffusum and C. rosmarinifolium. Nees (1833), Candolle (1836a), and more recently 

Skottsberg (1916) suggested that these two names should 

be regarded as heterotypic synonyms instead. However, 

later authors (Mueller, 1858; Reiche, 1901; Cabrera, 

1971) considered the species as distinct entities. The study 

of many herbarium specimens, and fi eld observations carried out throughout the geographic range of Chiliotrichum

has led me to confi rm that the two traditionally recognized species of Chiliotrichum, namely C. diffusum and 

C. rosmarinifolium, are one vegetatively variable species 

that shows a morphological cline that follows a latitudinal 

gradient.

In addition to this, I have observed that one of the species of Chiliophyllum, Chiliophyllum fuegianum, could be 

better treated as a species belonging to the genus Chiliotrichum. This hypothesis is supported by several major morphological traits such as the color of ray corollas (white), 

the partially paleate receptacles, the style branches obtuse, 

the pappus composed of linear scales arranged in 2 to 3 

series, and also molecular data (Bonifacino and Funk, unpublished data). On the basis of these molecular data as 

well as on morphological grounds (i.e.: paleate nature of 

the receptacle, radiate capitula, and pappus composed of 

linear scales), the closest allies to Chiliotrichum among 

the core Chiliotrichum Group are Chiliophyllum and 

Lepidophyllum.

In this study, Chiliotrichum is considered to have two 

species: Chiliotrichum diffusum, a widely distributed species in southern South America, and Chiliotrichum fuegianum restricted to northern Tierra del Fuego.

Chiliotrichum diffusum (G. Forst.) Kuntze, Rev. Gen. Pl. 

3(2): 141, 1898. Amellus diffusus G. Forst., Fasc. Plant. 

Magellan.: 39, 1788. Chiliotrichum amelloideum Cass.

nom. illeg., in Cuvier, Dict. Sci. Nat. 8: 577, 1817. Aster 

magellanicus Spreng. nom. illeg., Syst. Veg. 3: 526, 1826. 

Tropidolepis diffusa (G. Forst.) Tausch, Flora 12 (5): 68, 

1829. Chiliotrichum amelloideum var. diffusum (G. Forst.) 

Nees, Gen. Spec. Aster.: 240, 1833. Chiliotrichum diffusum f. typica Speg. nom. illeg., Anales Soc. Ci. Argent. 

48: 186, 1899. Type: “Terra del Fuego, J. R. & G. Forster

[s.n]” (Lectotype: BM 528784!, designated here).

Chiliotrichum rosmarinifolium Less., Linnaea 6: 109, 1831. Amellus rosmarinifolius Poepp. ex Less. pro syn., Linnaea 6: 109, 1831. Chiliotrichum 

amelloideum var. rosmarinifolium (Less.) Nees, Gen. Spec. Aster.: 240, 1833. 

Type: “Flos chilens in rapibus Pico de Pilque ad pagum Antuco summis, frigides 

[7100'] Jan. 1829, Poeppig 870” (holotype: W; isotypes: BM!, G, NY!, W; 

scanned image of holotype at W: LP!; photo of isotype at G: LP!).

Chiliotrichum amelloideum var. lanceolatum Nees, Gen. Spec. Aster.: 240, 

1833. Ind. Loc. Type: “in Rigni Chilensis montibus, Poeppig [s.n.]” Type: not 

found.

Chiliotrichum feliciae Hombr. and Jacquin., in d’Urv., Voy. Pôle Sud, Atlas: 28, 1848. Type: “Silvarum ad oras, D’Etroit de Magellan, Port Famine, 

1838−1840, Hombron [s.n.]” (holotype: P 222878, scanned image of holotype 

at P: LP!)

Chiliotrichum ovatifolium Hombr. and Jacquin., in d’Urv., Voy. Pôle Sud, Atlas: 28, 1848. Type: “In locis apertis arenosisque, D’ Etroit de Magellan, Havre 

Pecket, 1838−1840, Hombron [s.n.]” (holotype: P 222877, scanned image of 

holotype at P: LP!).

Chiliotrichum ovatifolium var. retortum Hombr. and Jacquin., in d’Urv., Voy. 

Pôle Sud, Atlas: 28, 1848. Type: “in abruptis littoribus Porteis Gallant et in 

locis a vento soud [defansix ?] Porteis famis, H. Hombron [s.n.]” (holotype: P, 

scanned image of holotype at P: US!).

KEY TO THE SPECIES OF CHILIOTRICHUM CASS.

1a. Leaves patent, margins revolute, generally 10−40 mm long; capitula notably pedunculate, peduncles (5−) 25−40 mm 

long. (Widespread along the Andes and Patagonia, from Mendoza in Argentina and Libertador O’Higgins in Chile to 

Tierra del Fuego archipelago and Islas Malvinas). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. diffusum

 1b. Leaves ascendent, margins fl at to slightly revolute, generally 4−6 mm long; capitula sessile to shortly pedunculate, peduncles up to 1.5 mm long. (Restricted to northern Tierra del Fuego in Argentina and Chile) . . . . . . . . . . . . . C. fuegianum
4 0 • SMITHSONIAN CONTRIBUTIONS TO BOTANY

Chiliotrichum virgatum Phil., Anales Univ. Chile 43: 485, 1873. Type: 

“[Chile] Magallanes, aestate 1864−1865 [s. leg. s.n.]” (holotype: SGO!; isotypes: B destroyed, NY!, SGO!; photo of the isotype at B: F!).

Chiliotrichum angustifolium Phil., Anales Univ. Chile 87: 401, 1894. Type: 

“[Chile] Andes Prov. Curicó, aestate 1889 [s. leg. s.n.]” (holotype: SGO!).

Chiliotrichum longifolium Phil., Anales Univ. Chile 87: 401, 1894. Type: 

“[Chile] Trancura, Feb. 1891, O. Philippi [s.n.]” (lectotype: SGO 44760!, 

designated here; syntype: “Trapatrapa, Febr. 89, Rahmer [s.n.]”, SGO 

44761!, SGO 65012!).

Chiliotrichum rosmarinifolium var. glabrescens Phil., Anales Univ. Chile 

87: 401, 1894. Type: “[Chile] Araucania, C. Sage [s.n.]” (holotype: SGO 

65011!).

Chiliotrichum tenue Phil., Anales Univ. Chile 87: 402, 1894. Type: “[Chile] 

Andibus prov. Curicó, 1892, Manuel Vidal [s.n.]” (holotype: SGO 65013!).

Chiliotrichum diffusum f. angustifolium (“angustifolia”) Speg., Anales Soc. 

Ci. Argent. 48: 186, 1899. Type: “[Argentina] Nahuel Huapi [Spegazzini 

s.n.]” (holotype: LPS 17189 in LP!).

Chiliotrichum diffusum f. media Speg., Anales Soc. Ci. Argent. 48: 186, 

1899. Ind. Loc. Type: “[Argentina] Hab. In Chubut centrali, Nov. 1898, 

J. Koslowsky s.n.” (holotype: not found).

(FIGURES 19, 20, 50, 51)

Shrubs (0.1−) 0.3−1.5 (−2) m tall, globose; branches 

densely disposed, ascending, not ribbed, tomentose and 

gland-dotted, internodes 1−3 (−10) mm long. Leaves

densely arranged, persistent, alternate, patent, (6−)10−40 × 

(0.8−)1−5 (−8.5) mm, linear to narrowly elliptic or narrowly obovate, base attenuated, petiole broadened towards the stem and constricted above, apex acute to 

obtuse, margins revolute, densely tomentose on abaxial 

side, glabrous to arachnoid on adaxial side, gland-dotted 

in both sides, midvein impressed on adaxial side, discolorous, coriaceous. Capitula solitary, terminal, notably 

pedunculate, peduncles (5−) 25−40 mm long, heterogamous, radiate. Involucres 6−10 × 4−5 mm, cylindricalcampanulate; phyllaries arranged in 4 to 5 series; outer 

phyllaries 4−5 × 1−2.3 mm, triangular, apex acute, tomentose abaxially, coriaceous, margins membranaceous, 

fi mbriate; inner phyllaries 5.5−8.5 × 1.5−2 mm, narrowly 

elliptic, apex acute, with a tuft of woolly trichomes towards the apex, membranaceous, margins membranaceous, fi mbriate. Receptacles convex, partially paleate; 

paleae 9 to 16, 8.5−9 × 0.5−1 mm, linear to narrowly 

elliptic, not embracing the achenes, apex acute, with 

a tuft of woolly trichomes towards the apex, margins 

membranaceous, fi mbriate. Ray fl orets 8 to 14, pistillate; corollas white, radiate, tube 3.5−5 mm long, limb 

(5−) 7−14 × 2.8−5 (−6) mm, elliptic, 3-lobed; staminodes 

3 or 4; style shafts 4.5−5 mm long, base slightly swollen, 

style branches 1−2 mm long, narrowly ovate, subequal in 

length. Disk fl orets 12 to 27, bisexual; corollas yellow, tubular, 5-lobed, broadened more or less abruptly from midlength, tube 2.5−3 mm long, throat 2.5−3 mm long, lobes 

0.7−2 × 0.4−0.5 mm, triangular, gland-dotted abaxially; 

anther thecae 1.5−1.8 mm long, bases slightly sagittate, 

apical appendages 0.5−0.7 mm long, ovate, antheropodium present, not swollen; style shafts 3.5−6 mm long, 

base swollen, style branches 1.8−3.2 mm long, oblong, 

apex acute, collecting hairs on upper 2/5 to 1/2. Achenes

3.5−6.5 mm long, narrowly obovate, terete, 4- to 6-ribbed, 

gland-dotted, sometimes with few twin trichomes sparsely 

distributed towards the apex, carpopodium present, conspicuous. Pappus (3.5−) 4−7 mm long, (1) 2 to 3-seriate, 

composed of 50 to 70 linear scales, outer series slightly 

shorter, scabrid along the margins.

Vernacular names: “Mata negra,” “Incienso” 

(Rothkogel s.n., SI 8466), “Fachine” (in Islas Malvinas; 

Moore, 1968), “Margarita” (Kurtz 5942, CORD), “Mata 

azul” (Cozzo s.n., BA 53064), “Mata negra o Romerillo” 

(Villagrán et al. 8133, CONC); “Kó’or” or “Kóor” (lengua Ona; Haloua and Padín, 1999); “Gorhj” (Borla and 

Vereda, 2001).

Phenology: The species shows a wide blooming 

season, and fl owering specimens have been collected from 

October to April.

Distribution: Chiliotrichum diffusum occupies a 

wide latitudinal range extending from southern Mendoza 

Province in Argentina and Libertador O’Higgins in Chile 

to Tierra del Fuego, along the Andes including the immense archipelago west of Chile, Isla de los Estados and 

Islas Malvinas. It is an important element in the Patagonian steppe of southern Santa Cruz and northern Tierra del 

Fuego (Figure 20).

Habitat: Chiliotrichum diffusum inhabits high Andean, Insular, Patagonian, and subantarctic biogeographic 

provinces (Cabrera and Willink, 1973). It is a quite ubiquitous species that grows at full sun on dry sandy soils in 

northern Mendoza and Neuquén or in moister areas such 

as the NW coast of Tierra del Fuego. Chiliotrichum diffusum can occur in rather scattered populations or forming 

denser associations, constituting the dominant species (NW 

and S Tierra del Fuego). In Tierra del Fuego, a considerable part of the Patagonian steppe is covered by associations 
number 92 • 4 1

FIGURE 19. Chiliotrichum diffusum (G. Forst.) Kuntze. A. Habit. B. Leaves, for each pair, adaxial view (left), abaxial view (right). C. Involucre. D. Outer to inner phyllaries (from left to right). E. Receptacular paleae, lateral view (left), dorsal view (right). F. Detail of pappus scales. 

G. Achene with pappus. H. Style of ray fl oret. I. Style of disk fl oret. J. Corolla of disk fl oret. K. Corolla of ray fl oret (A, B1, C−K, Bonifacino 

et al. 563, LP; B2, Bonifacino et al. 336, LP; B3, Maldonado 650, LP).
4 2 • SMITHSONIAN CONTRIBUTIONS TO BOTANY

FIGURE 20. Geographic distribution of Chiliotrichum diffusum (G. Forst.) Kuntze. Note leaf shape and size variation along the latitudinal 

gradient.
number 92 • 4 3

of Chiliotrichum diffusum and Festuca gracillima Hook. f. 

(Tuhkanen et al., 1989, 1990; pers. obs.). It can also grow 

in the shade in the understory of the Nothofagus spp. forest 

and also in very swampy areas at the ecotonal area between 

subantarctic forest and the high Andean vegetation (Moore, 

1983; Roig, 1998; pers. obs.). It can grow scattered or form 

dense associations intermingled with other shrubs typical of 

subantarctic forest such as Berberis buxifolia Lam., Ribes 

magellanicum Poir., and Pernettya mucronata Gaudich.ex 

G. Don. (Pisano, 1977; León et al. 1998; pers. obs.). Chiliotrichum diffusum grows from sea level in Tierra del Fuego 

to 2,700 m in Mendoza (Argentina) and neighboring areas 

in Chile. According to Davis and McAdam (1989), Chiliotrichum diffusum used to be much more common in Islas 

Malvinas than it is today, having diminished because of intense grazing; however, it still remains an important element 

of the native vegetation of this archipelago, being among the 

few plant species that grow to almost 2 m tall. Moore (1968) 

points out that Chiliotrichum diffusum is common in Islas 

Malvinas, on wet soils beside streams together with Cortaderia pilosa (d’Urv.) Hack., growing from sea level to 300 

m, but it can also grow on sandy soils near the coast and on 

occasion forms dense thickets away from streams. Woods 

(2000) reports it as a species used in certain areas as song 

perches by “Falkland Grass Wren” Cistothorus platensis.

Uses: The Onas (aboriginal tribes from Tierra del 

Fuego) used branches to make a particular kind of tatoo 

known as “lóiste” (Haloua and Padín, 1999). They also 

used Chiliotrichum diffusum to make arrows, and the fl owers were used to remediate vision problems by means of 

rubbing the fl orets in the eyes (Borla and Vereda, 2001). It 

is cultivated and commercialized in the United Kingdom for 

ornamental purposes (Bean, 1981). Brander (1977) selected 

and named one cultivar of Chiliotrichum diffusum, “Chiliotrichum diffusum SISKA”. This cultivar is widely offered 

online in European nurseries because of its impressive and 

attractive blossoms.

Etymology: The epithet diffusum, derived from 

Latin, refers to the irregular and widely spreading branching pattern that characterizes the species.

Note 1: The branches of Chiliotrichum diffusum

exhibit a clearly marked seasonal growth, as evidenced by 

irregular intervals of modifi ed leaves with a scale-like appearance followed by elliptic to narrowly obovate leaves.

Note 2: Forster (1788), in the original diagnosis of 

Amellus diffusus, stated “Habitat in Terra Ignis, Insulisque 

Noui Anni, in scopulis mari imminentibus” as the type locality. The Resolution sailed by Tierra del Fuego between 

December 21, 1774 and January 3, 1775, and anchored 

apparently at two locations: west of Navarino in Christmas 

Sound (Chile) and in New Year’s Island north of Isla de los 

Estados (Argentina). By that time, Chiliotrichum diffusum

should have been in full blooming season, and being such 

an attractive and fairly common species it is not unlikely 

that the Forsters might have collected it profusely on both 

land excursions. There is a wealth of type material related 

to this species distributed in several herbaria (Nicolson and 

Fosberg, 2004), however BM holds the richest collection 

of the Forsters’ materials. I have studied two specimens deposited at BM (BM 528784 and BM6082) that perfectly 

match the original diagnosis, and that have been identifi ed 

by BM staff, and by Nicolson and Fosberg (2004) as type 

material. Therefore, one of this specimens (BM 528784), 

which holds more information on the scanty label “Terra 

del Fuego, J. R. & G. Forster [s.n]” is selected here as the 

lectotype.

Note 3: When transferring Amellus diffusus to 

Aster, Sprengel (1826) did not consider the original epithet 

and instead coined a new and superfl uous (i.e.: illegitimate) 

name.

Note 4: Cassini (1817) gave a detailed explanation about the reason why he created Chiliotrichum in 

order to transfer the species Amellus diffusus to this new 

genus. However, Cassini did not take the original epithet 

and coined a new and illegitimate name. Based on this observation, it should be noted that the names of varieties of 

Chiliotrichum amelloideum by Nees (1833) would have to 

be corrected using the correct name of the species. However, given the fact that none of the varieties are recognized 

here, there is no need to further clutter the synonymy of 

this taxon.

Note 5: In the original diagnosis of Chiliotrichum 

longifolium, Philippi (1894) mentions two specimens: 

“Habitat in Andibus Araucaniæ ad Trapatrapa, (C. Rahmer) et Valdiviæ l. D. Trancura (O. Philippi)”. I’ve seen 

both specimens at SGO, and they perfectly match the original diagnosis. I choose as lectotype the specimen collected 

by Otto Philippi (grandson of R. A. Philippi).

Note 6: According to Stafl eu and Cowan (1976), 

the publication date of the Fasciculus plantarum magellanicarum, when Amellus diffusus G. Forst. was described, 

was after September 18, 1787. The year 1787 is the date 

that most commonly appears in the literature relevant to 

this species (e.g.: Cabrera, 1971; Moore, 1983; Sancho and 

Ariza Espinar, 2003), while Moore (1968) gives 1789 as the 

original publication date. However, according to Nicolson 

and Fosberg (2004), the date of the publication would have 

been between March and April 1788, time when paged 

preprints that appeared between January 16 and 17, 1788 

were distributed.
4 4 • SMITHSONIAN CONTRIBUTIONS TO BOTANY

Additional specimens examined: ARGENTINA. PROV. 

CHUBUT. Dpto. Cushamen. Cerro Hielo Azul, 28-XII-2000, 

Angelino 14 (BAA); Ruta 15, 2 km al W de intersección con 

Ruta 40, 42°23'37"S, 71°9'54"W, 11-I-2002, Bonifacino 

et al. 404 (US); Valle de la Laguna Blanca, 42°52'S, 

71°15'W, s. date, Koslowsky 258 (SI). Dpto. Futaleufu.

Región Río Corcovado, s. date, Illin 65 (SI, CORD); Valle 

de la Laguna Blanca, 42°52'S, 71°15'W, 10-I-1901, Koslowsky 129 (BAF, SI, CORD); Cerro Cuche, 28-I-1947, 

Soriano 2509 (LP, BAA); Lago Futalaufquen, 26 -II-1950, 

Soriano 4297 (LP); Parque Nacional Los Alerces, Lago Futalaufquen, 9-I-1948, Soriano 2907 K (BAA); Lago Futalaufquen. Parque Nacional Los Alerces, 18-II-1950, Soriano 

4177 (BAA). Dpto. Languiñeo. Carrenleufu, 1-III-1900, 

Illin s.n. (LP); Ruta 13, 21 km al SW de Paso del Sapo, 12-

IV-1988, León 4018 (BAA); Río Tecka, Estancia Pampa 

Chica, 25-I-1947, Soriano 2482 (LP). Dpto. Río Senguerr.

Lago La Plata, 26-I-1969, Guerrero 56 (SI); márgenes y alrededores del Arroyo Blanco, costa S del Lago Fontana, 20-

22-II-1999, Gutiérrez 108 (US); Río Unión (Lago La Plata), 

10-I-1969, Iglesias & Scotbeer 10 (LP); Río Union, Lago La 

Plata, 13-I-1969, Iglesias & Scotbeer 23 (LP); Estancia La 

Pepita, 11-II-1945, Soriano 1510 (SI); Lago Fontana, 14-II1947, Soriano 2601 (BAA); Lago Fontana, Estancia Pepita, 

29-I-1960, Soriano 5656 (BAA); Pampa de Chalía, Estancia 

La Media Luna, 3-XII-1981, Villamil et al. 2205 (SI). Dpto. 

Tehuelches. Lago Gral. Paz, 4-III-1903, Gerling s.n. (SI 

8461); Río Pico, campo del Sr. Torres, 23-I-1960, Soriano 

5568 (BAA). PROV. MENDOZA. Not defi ned Dpto. Entre Piedra Parada y Valle Rahuí, 14-II-1888, Kurtz 6046 (CORD); 

Entre Río Grande y Arroyo Calquegue, 30-I-1888, Kurtz 

5942 (CORD); Planchón de los Ciegos, I-1897, Stuckert 

2851 (CORD). Dpto. Malargue. Valle de Calmuco, 14-II1942, Burkart et al. 14359 (SI, LP); Calmuco, 14-II-1942, 

Covas 225 (LP); Calmuco, 14-II-1942, Covas 478 (LP). 

PROV. NEUQUEN. Not defi ned Dpto. Mallín Hue Culle, 

Parque Nacional Lanín, 1-III-1985, Cusato 3973 (BAA); a 

2 km de Bajada de Sainuco, 14-I-1968, Ruiz Leal 25813 

(LP). Dpto. Aluminé. Entre Sainuco y Lago Aluminé, 7-XII1981, Cabrera et al. 32893 (SI, LP); Valle Escondido, 

Ñorquinco, 12-XII-1925, Comber 348 (E);Parque Nacional Lanín, subida a Quillen a Ruca Choroi, 28-II-1985, Cusato 3909 (BAA); Laguna Ruca Choroi, 1-II-1968, Eskuche 

& Klein 109 (SI). Dpto. Catan Lil. Estancia Yao Yao, 

8-XII-1946, Dawson 1178 (BAA); Sierra del Chachil, 29-I1965, Rúgolo & Agrasar 407 (BAA). Dpto. Chos Malal.

Riscos Bayos, Valle Arroyo Olletas, 36°51'S, 70°20'W, 25-

I-1964, Boelcke & Correa 11183 (SI, BAA). Dpto. Huiliches. 

Parque Nacional Lanín, Lago Huechulaufquen, subida a 

Cerro Los Angeles, 7-II-1948, Dawson & Schwabe 2603 

(BAA); Parque Nacional Lanín, Lago Huechulafquen, 

subida al Cerro de los Angeles, 7-II-1948, Dawson & 

Schwabe 2611 (BAA); Potrero del Tromen, Parque Nacional Lanín, 8-II-1983, Tarak s.n. (BAA). Dpto. Lacar. San 

Martín de los Andes, I-1941, Bridarolli 2069 (LP); San 

Martín de los Andes, Cerro Chapelco, 16-II-1968, Cabrera 

& Crisci 19157 (LP); Cerro Chapelco, laderas encima de 

refugio, expuestas al N, 23-XI-1974, Correa 5893 (LP); 

Parque Nacional Lanín, Cerro Chapelco, 12-II-1961, León 

& Calderon 869 (BAA); Parque Nacional Lanín, Lago 

Lacar, Cerro Malo, mallíPueston en ladera SW, 22-II-1961, 

León & Calderon 1332 (BAA); Cordón Chapelco, entre 

Cerro Chapelco y Cerro Negro, Parque Nacional Lanín, 

20-III-1983, Mermoz s.n. (BAA). Dpto. Loncopué. Cajón 

Chenque Pehuen. Valle a la entrada del Cajón, 14-I-1965, 

Rúgolo & Agrasar 142 (BAA); Chenque Pehuen, 8-I-1967, 

Schajovskoy s.n. (LP). Dpto. Los Lagos. Puesto Manzano, 

Cerro Cortinario, 19-I-1965, s. leg. s.n. (BAA 11273); 

Parque Nacional Nahuel Huapi, Laguna Ortiz Basualdo, 

7-I-1952, Bolecke & Correa 5365 (BAA, SI); Parque Nacional Nahuel Huapi, Refugio Cerro Colorado, 14-II-1953, 

Boelcke & Correa 6899 (BAA); Paso Cardenal A. Samoré, 

40°42'48"S, 71°56'37"W, 24-I-2003, Bonifacino & Romano 824 (LP); Cerro Belvedere en Correntoso, 5-II-1934, 

Burkart 6401 (LP); Laguna del Toro, Rincón Grande, I1942, De Jones 67 (LP); Parque Nacional Nahuel Huapi, 

cerca del refugio Cerro Colorado, 5-II-1951, Diem 1824 

(SI); s. loc., 10-I-1993, González Arzac 977 (LP); Correntoso, 6-II-1934, Parodi 11694 (BAA); Parque Nacional Nahuel Huapi, Punta Inferior Mallín Grande, Río Manso 

Superior, XII-1974, Richencq 204 (SI); Lago Nahuel Huapi, 

cuenca del Río Machete, 1-II-1914, Rothkugel 43 (SI); Lago 

Nahuel Huapi, Cerro Belvedere, 21-III-1934, Spegazzini s.n 

(LP 61233). Dpto. Minas. 18 km de las Ovejas camino a 

Laguna Epu-Lauquen, Mallin de la Culebra, 36°54'S, 

70°55'W, 14-I-1964, Boelcke et al. 10780 (SI); Paso del 

Macho, bajada a laguna proxima al paso, 36°26'S, 70°46'W, 

26-I-1970, Boelcke et al. 13951 (SI); Confl uencia de los 

Ríos Pichi-Neuquén y Neuquén, Cerro de Las Yeguas, 

36°35'S, 70°45'W, 23-I-1970, Boelcke et al. 13767 (BAA); 

Paso del Macho, bajada a la laguna, 36°27'S, 70°40'W, 26-

I-1970, Boelcke et al. 13957 (BAA); Sierra de Cochicó, 

Cajón de la Botica, 36°21'22"S, 70°34'36"W, 29-I-1970, 

Boelcke et al. 14117 (SI); Extremo N Laguna Varvarco 

Campos, Arroyo Benitez, curso medio, 36°17'S, 70°39'W, 

30-I-1970, Boelcke et al. 14264 (BAA, LP, SI). Dpto. 

Ñorquin. Termas de Copahue, 17-II-1940, Cabrera 6156 

(LP); along the road from Baños de Copahue, ~5 km of the 

town, 12-I-1985, King 9399 (SI); Trolope, 26-XII-1944,

O’Donell 2134 (BM); Parque Nacional Copahue, Estancia 
number 92 • 4 5

Trolope, 9-I-1973, Rúgolo de Agrasar et al. s. n (BAA 

12272). Dpto. Picunches. Paso Pino Hachado, cerca de la 

frontera con Chile, 38o 39'S, 70o 53'W, 16-I-2000, Bonifacino et al. 191 (LP); Pino Hachado, 6-III-1939, Burkart 

9696 (SI); Pino Hachado, 24-I-1948, Dawson & Schwabe 

2232 (BAA); Paso Pino Hachado, 30-I-1974, Gentili 148 

(LP); Portezuelo Sanguileo, SW de Paso Pino Hachado, 

20-II-1983, León 3410 (BAA); Pino Hachado, 5-II-1920, 

Parodi 2236 (BAA). PROV. RÍO NEGRO. Dpto. 25 de Mayo.

Pendiente Norte del Cerro Anecón Grande, 22-XII-1938, 

Ferruglio 29 (LP); Cerro Anecón, pendiente norte, 11-I1939, Ferruglio 43 (LP). Dpto. Bariloche. Bariloche, Cerro 

Ñireco, II-1946, Bernasconi s.n. (SI 15936); Cerro Otto, 

Parque Nacional Nahuel Huapi, 7-I-1946, Boelcke 1746

(LP, BAA); Parque Nacional Nahuel Huapi, Cumbre Cerro 

Rigi, 11-I-1952, Boelcke & Correa 5476 (BAA); Parque 

Nacional Nahuel Huapi, Cerro Tronador, entre Mallin 

Chileno y Refugio, 17-I-1952, Boelcke & Correa 5705 (SI); 

Parque Nacional Nahuel Huapi, Cerro Tronador, Mallin 

Chileno, 18-I-1952, Boelcke & Correa 5785 (SI); Parque 

Nacional Nahuel Huapi, subida al Granito, 22-I-1952, 

Boelcke & Correa 5898 (BAA, SI); Parque Nacional Nahuel Huapi, faldeos Cerro Santa Elena, 31-I-1952, Boelcke 

& Correa 6208 (SI); Cerro Catedral, 1584 m, 41°10'40.6"S, 

71°27'36.6W, 9-XII-2006, Bonifacino et al. 2922 (US); 

Parque Nacional Nahuel Huapi, Tronador, Picada a Laguna Los Césares, 26-I-1983, Cusato 2453 (BAA); Cerro 

Catedral, II-1954, Cabrera 11501 (LP); Parque Nacional 

Nahuel Huapi, 13-II-1978, Corte 160 (LP); Cerro Lopez, 

I-1960, Fabris 2175 (LP); Cerro Catedral, I-1960, Fabris 

2236 (LP, BAA); Lago Nahuel Huapi, s. date, Grüner 140 

(LP); Sierra de Lopez, 14-II-1914, Hosseus 301 (CORD); 

Lago Nahuel Huapi, Cerro Utne, 20-II-1914, Hosseus 487 

(CORD); Lago Nahuel Huapi, Sierra Lopez, II-1933, 

Ljungner 259 (BM); Parque Nacional Nahuel Huapi, Cerro 

Riggi, Limite Arg-Chile, II-1958, Marchionni s.n. (LP); 

Cerro Catedral, 17-I-1946, Montiel 211 (LP); Nahuel 

Huapi, 1915, Rothkogel s.n. (SI 8466). Dpto. Pilcaniyeu.

Cerro Colorado, 2-II-1914, Hosseus 1356A (CORD). 

SANTA CRUZ. Dpto. Guer Aike. Estancia Las Viscachas, 

Cerro Sin Nombre, Faldeo NE, 50°46'S, 72°8'W, 29-I1977, Arroyo & Boelcke TBPA 2631 (SI); La Balsa, sobre 

Río Gallegos, I-1967, Boelcke 12418 (BAA); Estancia Sofía, 

3 km W del Casco, ladera exp. SW, 51°44'S, 71°20'W, 8-II1978, Boelcke & Correa TBPA 2977 (SI); Estancia Stag 

River, 10 km N casco sobre río Venados, 51°38'S, 71°57'W, 

13-II-1978, Boelcke & Correa TBPA 3172 (SI); Estancia 

Stag River, afl . W del río Venados, SE Meseta la Torre, 

51°34'S, 71°57'W, 17-II-1978, Boelcke & Correa TBPA 

3305 (SI); Ruta 293, Punta Alta, 14-I-1967, Boelcke et al. 

12418 (SI, BAA); Río Gallegos, al frente del cerro Buitres, 

15 leguas de la costa, II-1899, Hauthal s.n. (CORD); Estancia Glencross, Río Turbio, 100 m, 6-XII-1950, Sleumer 

1004 (US); s. loc., 19-XII-1967, Zeballos s.n. (BAA 10650); 

Río Gallegos, 19-XII-1967, Zeballos s.n. (BAA 10680). 

Dpto. Lago Argentino. Lago Argentino, Brazo Norte, Ventisquero Upsala, Bahia Cipres, 50°0'S, 73°20'W, 19-I-1967, 

Boelcke et al. 12577 (SI, BAA); Parque Nacional Los Glaciares, 291 m, 50°28'24.1"S, 72°59'41.2"W, 12-XII-2006, 

Bonifacino et al. 3000 (US); Lago Argentino, canal de los 

Témpanos, Glaciar Ameghino, 10-XII-1963, Correa 3098 

(BAA); Ventisquero Perito Moreno, Parque Nacional los 

Glaciares, 17-XII-1981, Cusato s.n. (BAA 17883); Lago 

Roca, 16-XII-1969, Guerrero 58 (SI); Lago Argentino, 17-

XII-1968, Guerrero 63 (SI); Lago Argentino, XII-1958/III1959, James 144 (SI); Cerro Buenos Aires y Ventisquero 

Moreno, 4-I-1959, James 666 (BM); Lago Frío, southern 

arm of Lago Argentino, 13-I-1959, James 818 (BM); Lago 

Argentino, near Ventisquero Uppsala, Estancia Master’s, 

21-XII-1958, James 1441 (BM); s. loc., 1904-1905, Koslowsky 75 (CORD); Lago San Martin, 8-XII-1914, Pemberton 2 (SI); Parque Nacional Los Glaciares, 29-II-1953, 

Perez Moreau s.n. HERB B. RIV. 70004 (CORD); Parque 

Nacional Los Glaciares, zona Lago Argentino, 29-III-1953, 

Perez Moreau s.n. (BAA); Lago Argentino, II-1982, Suarez 

s.n. (SI 432); Fitz Roy, entre Río Blanco y Lago de Los Tres, 

7-II-1957, Suti 3680 (CORD); Glaciar Perito Moreno, 26-

XI-1984, Valla s.n. (BAA 19251). Dpto. Lago Buenos Aires.

Ing. Pollavicini, X-1973, Suarez 823 (SI); Río Jeinemeni, 

5-II-1903, Von Platen & Greiner 42 (SI). Dpto. Río Chico.

Lago Belgrano, 1-III-1903, Von Platen & Greiner 44 (SI). 

PROV. TIERRA DEL FUEGO. Not defi ned Dpto. Arroyo Pico, 

28-XII-1918, Parodi 2931 (BAA). Dpto. Río Grande. Ruta 

3, km 2775, 53°29'53"S, 68°6'14"W, 28-I-2002, Bonifacino et al. 589 (US). Dpto. Ushuaia. Isla de los Estados, 

Puerto Cook, 20-I-1912, s. leg. s.n. (SI); Ushuaia, 5-II-1896, 

Alboff s.n. (CORD); Ushuaia, 6-II-1896, Alboff s.n. (SI 

8460); Bahia Buen Suceso, 22-XI-1998, Biganzoli 465 (SI); 

Parque Nacional Tierra del Fuego, Cerro Guanaco, 

54°47'48"S, 68°33'29"W, 24-I-2002, Bonifacino et al. 538

(US); íd., 54°47'48"S, 68°33'29"W, 24-I-2002, Bonifacino 

et al. 549 (US); Ruta costera al E de Ushuaia, a 7 km del 

Puerto, 54°49'3"S, 68°11'19"W, 25-I-2002, Bonifacino et 

al. 562 (US); Paso Garibaldi, 54°41'22"S, 67°50'29"W, 

26-I-2002, Bonifacino et al. 563 (US); Ruta J, al E de Estancia Harberton, 81 m, 54°54'11.5"S, 67°6'48.2"W, 14-XII2006, Bonifacino et al. 3026 (US); Parque Nacional Tierra 

del Fuego, Bahía La Pataia, 54°51'20"S, 68°34'24"W, 16-

XII-2006, Bonifacino et al. 3049 (US); La Pataia, 12-II1962, Crespo s.n. (SI 21548); Península Mitre, Bahía Buen 
4 6 • SMITHSONIAN CONTRIBUTIONS TO BOTANY

Suceso, N facing slopes at S end of the Bahia, 13-X-1971, 

Dudley & Goodall 149 (SI); Peninsula Mitre, Bahía Buen 

Suceso, 54°48'S, 65°15'W, 14-X-1971, Dudley & Goodall 

172 (SI); Península Mitre, Bahía Valentín near Punta Blanca, 

54°53'S, 65°25'W, 15-X-1971, Dudley & Goodall 413 

(SI); Puerto Cook, 54°45'S, 64°3'W, 24-X-1971, Dudley & 

Goodall 829 (SI); Rabbit island, Estancia Harberton, 14-

XII-1967, Goodall 1056 (SI); Top Mountain, 15-XII-1967, 

Goodall 1066 (SI); Bahia Thetis, N of the settlement towards Cabo San Vicente, 54°40'S, 65°0'W, 20-XI-1969, 

Goodall 2307 (SI); Gable Island, near Estancia Harberton, 

12-III-1970, Goodall 2477 (SI); Pampa de los Indios, Canal 

del Beagle, 14-I-1960, Grondona s.n. (BAA 12974); alrededores del Aeropuerto de Ushuaia, 10-I-1955, Grondona 

4185 (BAA); cerca del río Pipo, orillas del Canal Beagle, 

13-I-1959, Grondona 7094 (BAA); Bahia Thetis, Estancia 

El Arroyo, 10 km camino a Lago Fagnano, 16-I-1959, 

Grondona 7119 (BAA); camino a La Pataia, 13-XII-1985, 

Hermann s.n. (SI); Ushuaia, 3-I-1904, Hicken 11 (SI); Ushuaia, alrededores del Matadero-Frigorifi co, 21-XII-1949, 

Luti Herbera 1417 (CORD); Isla de los Estados, Puerto 

Cook, ladera E, 54o45'S, 64o05'W, 25-XI-1967, Nicora & 

Crespo 7200 (BAA, SI); Isla de los Estados, Bahía Porto 

Roca, 12-I-1974, Ristorti 7449 (SI); Ushuaia, 12-II-1953, 

Ruiz Leal & Roig 1810 (CORD); Hills 1 km NW of Ushuaia, 18-III-1979, Solomon 4751 (SI); Estancia Sara, Bahia 

La Pataia, 2-III-1962, Vallerini 39 (BAA). Islas Malvinas.

Puerto Stanley, 12-II-1979, Dimitri & Orfi la 318 (SI); Isla 

Soledad, Puerto Stanley, 27-XII-1903, Hicken 23 (SI); near 

Port Stanley, s. date, Lellman s.n. (US); East Falkland, Sparrow Cove, N of Port Stanley, 5-I-1950, Sladen s.n. (BM 

608566); Isla Soledad, Puerto Stanley, 12-II-1979, Ulibarri 

& Dimitri 1214 (SI). CHILE. REG. DE AISÉN. Prov. Coihaique. 

Balmaceda, camino a Coihaique. Puente El Ministro, 7-I1989, Bartoli & Rua s.n. (BAA 21589); Río Coihaique, 22-

I-1939, Von Rentzell 6210 (SI). Prov. General Carrera. 

Región del Lago Bs As, fi n del Valle León, 13-II-1939, Von 

Rentzell 6262 (SI). REG. DEL BÍO BÍO. Prov. Ñuble. Termas 

de Chillán, Cerro Pirigallo, 6-II-1936, Cabrera 3641 (LP); 

Chillan, s. date, s.leg. 933 (US); Baños de Chillan, Nieblas, 

III-1927, Werdermann 1326 (SI, US). REG. DE LA ARAUCANÍA. Prov. Malleco. Cordillera Las Raíces, Lonquimay, 

2-III-1939, Burkart 9537 (SI); Malleco, Angol, Parque Nacional Nahuelbuta, 5-XII-1971, Mahu 8112 (LP). REG. DE

LOS LAGOS. Prov. Llanquihue. Puerto Montt, II-1973, Lagiglia 3851 (SI). REG. DE MAGALLANES. Prov. Antártica Chilena. Cape Horn, Hermite island, 1839-1843, Hooker s.n. 

(BM 608555); Canal Beagle, Puerto Villarino, 28-II-1903, 

Pennington 211 (SI, CORD). Prov. Magallanes. Punta Arenas, 25-XI-1950, Brooke 7051 (BM); 70 km NW of Punta 

Arenas, 4-I-1939, Eyerdam & Beetle 24098 (SI); Punta 

Arenas, II-1927, Guiñazú 274 (BAA); Punta Arenas, 

I-1911, Hauman s.n. (BAA 8156); Punta Arenas, 5-XII1910, Hicken 349 (SI); Punta Arenas, 10-I-1904, Hicken 1

(SI); Punta Arenas, 14-II-1863, Isern 8509 (SI); Punta Arenas, 14-II-1863, Isern 8511 (SI); Puerto del Hambre, 21-II1863, Isern 8520 (SI); Strait of Magellan, Elizabeth Island, 

19-I-1888, Lee s.n. (US); Río Pescado, 40 km al N de Punta 

Arenas, 2-XI-1985, León 3716 (BAA); Punta Arenas, 16-II1906, Sargent s.n. (US). Prov. Tierra del Fuego. Tierra del 

Fuego, I-1769, Banks & Solander s.n. (BM 608544); Tierra 

del Fuego, Estrecho de Magallanes, III-1930, Canessa s.n. 

(BAA); Porvenir, 16-I-1904, Hicken 206 (SI). Prov. Ultima 

Esperanza. Estancia Eberhard, a aprox. 10 km camino a 

Puerto Natales, 24-XI-1955, Boelcke 7477 (BAA); Estancia 

Cerro Castillo, Lago Maravilla, NE shore, 51°19'S, 

72°43'W, 21-XII-1975, Dollenz & Moore TBPA 827 (SI); 

Seno Unión, N side, Ancon Sin Salida, 52°9'S, 73°21'W, 

10-I-1976, Dollenz & Moore TBPA 875 (SI); Seno Unión, 

N side, Ancon Sin Salida, 52°9'S, 73°21'W, 10-I-1976, 

Dollenz & Moore TBPA 909 (SI); Isla Piazzi, Caleta 

Ocación, Abra Leackey’s retreat, 51°44'S, 74°1'W, 16-I1976, Dollenz & Moore TBPA 1028 (SI); Isla Rennel N, 

Canal Smith, 51°54'S, 74°12'W, 23-I-1976, Dollenz & 

Moore TBPA 1136 (SI); Isla Vidal Gormaz, Seno Nantuel, 

Bahia María Angélica, 51°53'S, 74°41'W, 1-II-1976, Dollenz & Moore TBPA 1254 (SI); Río Verde, 150 km al N del 

estrecho, XII-1926, Guiñazú 129 (BAA); Puerto Prat, 4-II1904, Hicken 150 (SI); Seno Ultima Esperanza, Puerto 

Toro, Río Serrano, 2 km from mouth, 51°24'S, 73°5'W, 

17-I-1977, Moore & Pisano TBPA 1777 (SI); Seno Ultima 

Esperanza, Puerto Toro, W of río Serrano, N of ventisquero 

Balmaceda, 51°25'S, 73°5'W, 18-I-1977, Moore & Pisano 

TBPA 1810 (SI); Seno Ultima Esperanza, Puerto Toro, 

costa E río Serrano, 51°25'S, 73°4'W, 22-I-1977, Moore & 

Pisano TBPA 1866 (SI); Seno Ultima Esperanza, Península 

Roca, Seno Resi, 51°51'S, 73°2'W, 21-I-1978, Pisano s.n. 

TBPA 2802 (SI); Seno Ultima Esperanza, Península Roca, 

Seno Resi, 51°51'S, 73°2'W, 24-I-1978, Pisano s.n. TBPA 

2875 (SI); Seno Ultima Esperanza, Puerto Bella Vista, 

51°31'S, 73°16'W, 11-I-1977, Pisano 74 (SI). REG. DEL

MAULE. Prov. Curicó. Cord. Volcán Peteroa, I-1925, Werdermann 607 (SI, BM). Prov. Talca. Cordillera Talca-Picazo, 

26-I-1939, Barros 1858 (SI); Laguna del Maule, 30-I-1963, 

Boelcke & Bacigalupo 10336 (BAA, LP, SI); Laguna del 

Maule, 36o01'S, 70o33'W, 24-I-1990, Gardner & Knees

4528 (E); Cordillera del Maule, 1855, Germain s.n. (BM); 

Laguna del Maule, 24-I-1981, Grau 2828 (BM). REG. LIBERTADOR O’HIGGINS. Prov. Conchagua. El Flaco, 26-I-1948, 

Barros 7366 (LP). Specimens from cultivation: England, 
number 92 • 4 7

Hampshire, Ampfi eld, Hillier’s Arboretum, 14-V-1974, 

s. leg. 853 (BM); Royal Botanic Garden Edinburgh, V-1956, 

s. leg C1258 (BM).

Chiliotrichum fuegianum (O. Hoffm.) Bonif. comb. 

nov. Chiliophyllum fuegianum O. Hoffm., in Dusén, Wissench. Erg. Schwed. Exped. Magellansländer 

1895−1897, 3 (5): 96, 1900. Type: “[Argentina] Tierra 

del Fuego, Río Cullen, 1891, B. Ansorge 429” (holotype: UPS; photo of holotype at UPS: LP!).

(FIGURES 21, 22, 52)

Shrubs up to 1 m tall, globose, spreading; branches 

densely disposed, ascending, not ribbed, tomentose, internodes short, up to 1 (−2) mm long. Leaves densely disposed, 

persistent, alternate, ascending, 4−6 (−8) × 1−1.5 mm, 

elliptic to obovate, base attenuated, petiole broadened 

towards the stem and constricted above, apex obtuse, 

margins fl at to slightly revolute, gland-dotted in both 

surfaces, laxly to densely tomentose on abaxial surface, 

midvein notorious on abaxial side, forming furrow on 

adaxial side, discolorous, coriaceous. Capitula solitary, 

terminal, peduncles short, up to 1.5 mm long, heterogamous, radiate. Involucres 7−8 × 3.5−4.5 mm, cylindricalcampanulate; phyllaries arranged in 3 to 4 series; outer 

phyllaries 3.0−5.0 × 1.5−2.5 mm, triangular-ovate, apex 

acute, gland-dotted to slightly tomentose abaxially, coriaceous, margins membranaceous, fi mbriate; inner phyllaries 5−8 × 1.2−2 mm, narrowly elliptic, apex acute, 

gland-dotted abaxially with a tuft of woolly trichomes 

towards the apex, membranaceous, margins slightly fi mbriate. Receptacles slightly convex, partially paleate; paleae 

4 to 7, 7−9 × 0.5−1 mm, linear to narrowly elliptic, not 

embracing the achenes, apex acute, gland-dotted towards 

the apex, margins membranaceous, fi mbriate. Ray fl orets

3 to 6 (−8), pistillate; corollas white, radiate, tube 3−4.2 mm 

long, limb 5−8.3 × 1−2.6 mm, narrowly elliptic, 2- to 

3-lobed; staminodes 1 to 2, very reduced; style shafts 

4−4.2 mm long, base slightly swollen, style branches 

2−3 mm long, narrowly ovate, subequal in length. 

Disk fl orets 7 to 15, bisexual; corollas yellow, tubular, 

5-lobed, broadened more or less abruptly from midlength, tube 2−4 mm long, throat 1.5−2 mm long, lobes 

1.5−2.1 × 0.5−0.7 mm, triangular, gland-dotted abaxially; 

anther thecae 2−2.3 mm long, bases slightly sagittate, apical appendages 0.4−0.7 mm long, triangular-ovate, antheropodium present, not swollen; style shafts 3.7−6 mm 

long, base swollen, style branches 1.8−2.5 mm long, narrowly obovate, apex acute, collecting hairs on upper 1/2 to 

2/3. Achenes 4.5−5.5 mm long, narrowly obovate, terete, 

4- to 6-ribbed, gland-dotted, sometimes with few twin trichomes sparsely distributed towards the apex, carpopodium present, conspicuous. Pappus 4−5 (−6) mm long, 

2- to 3-seriate, composed of 65 to 70 linear scales, outer 

series slightly shorter, scabrid along the margins.

Vernacular names: Not known.

Phenology: Flowering specimens have been collected from December to February. Specimens with mature 

fruits have been collected in May and July.

Distribution: Argentina and Chile, endemic to 

northern Tierra del Fuego (Figure 22)

Habitat: Sand dunes and clay-sandy soil depressions in Patagonian steppe (Cabrera and Willink, 1973). It 

grows in coastal grasslands together with Lepidophyllum 

cupressiforme and Chiliotrichum diffusum.

Uses: Not known.

Etymology: The specifi c epithet fuegianum refers 

to the geographic area where the species occurs.

Note 1: Chiliotrichum fuegianum (O. Hoffm.) 

Bonif., was originally described in Chiliophyllum DC. 

(= Zaluzania Pers.). However, Hoffmann (1900) was unequivocally referring to Chiliophyllum Phil. since in the 

same publication of the new species he mentioned the differences between the new species and Chiliophyllum densifolium Phil. It is important to highlight that Hoffmann 

himself was aware of the differences between Chiliophyllum fuegianum and Chiliophyllum densifolium, especially 

those concerning the pappus (2- to 3-seriate in C. fuegianum versus 1-seriate in C. densifolium). Reiche (1901) also 

noticed that the characteristics described for this species 

were not totally congruent with those of the genus in which 

it was placed.

Note 2: As mentioned before, morphological characters such as the type of pappus, pubescence of achenes, 

color of marginal fl orets corolla as well as molecular characters (Bonifacino and Funk, unpublished data) put this 

species in close vicinity to Chiliotrichum diffusum.

Additional specimens examined: ARGENTINA. PROV. 

TIERRA DEL FUEGO. Dpto. Río Grande. Ruta 3, Bahía San 

Sebastián, 53°14'1"S, 68°34'6"W, 28-I-2002, Boni facino 

et al. 595 (LP); al N de Bahía San Sebastián, próximo 

a planta “TOTAL”, 52°53'19"S, 68°21'38"W, 13-XII2006, Bonifacino et al. 3017 (US); Estancia Cullen, Puesto 

Beta, 7-I-1971, Goodall 3249 (BAB, SI, herb. Goodall); 

Estancia Cullen, Arroyo Alfa, 20-V-1972, Goodall 4301 

(BAB, herb. Goodall); Estancia Cullen, Arroyo Alfa, 

20-V-1972, Goodall 4302 (BAB, herb. Goodall); Estancia Cullen, beach near outlet of Río Cullen, 19-V-1972, 

Goodall 4296 (BAB, herb. Goodall); Estancia Cullen, 
4 8 • SMITHSONIAN CONTRIBUTIONS TO BOTANY

FIGURE 21. Chiliotrichum fuegianum (O. Hoffm.) Bonif. A. Habit. B. Leaves, adaxial view (left), abaxial view (right). C. Involucre. D. Outer 

to inner phyllaries (from left to right). E. Receptacular paleae, lateral view (left), dorsal view (right). F. Detail of pappus scales. G. Achene with 

pappus. H. Style of ray fl oret. I. Style of disk fl oret. J. Corolla of disk fl oret. K. Corolla of ray fl oret (A−K, Bonifacino et al. 595, LP).
number 92 • 4 9

Arroyo Beta, 20-XII-1975, Goodall 5033 (herb. Goodall); Estancia Cullen, 1-VI-1977, Goodall 5167 (herb. 

Goodall); Estancia San Martín, Golfo San Sebastián, 6-

II-1955, Grondona 4477 (BAB); Bahía San Sebastián, Estancia San Martin, 6-II-1955, Hunziker 6827 (BAB, LP); 

Estancia Cullen, Arroyo Beta, 52o44'S, 68o 35'W, I-1972, 

Moore & Goodall 338 (LP, BAB, herb. Goodall); Estancia Cullen, N side of Río Cullen, 52o51'S, 68o24'W, 5-I1972, Moore & Goodall 356 (BAB, K, herb. Goodall). 

CHILE. REG. DE MAGALLANES. Prov. Tierra del Fuego. Bahía 

Inútil, fi rst line of hills to the S of Río Marazzi, 25-VII1977, Goodall 5168 (herb. Goodall); NE end of Bahia 

Inútil, road to Porvenir, 7 km west of Puerto Nuevo, 26-

VII-1977, Goodall 5171 (herb. Goodall); Bahía Lomas, 

S side of bay, just E of Estancia Cuarto Chorrillo, 29-

V-1978, Goodall 5207 (herb. Goodall); Estancia Pepita, 

NW part of Bahía Lomas, 29-V-1978, Goodall 5210 

(herb. Goodall); Estrecho de Magallanes, Primera Angostura, Punta Espora, 29-V-1978, Goodall 5211 (herb. 

Goodall); Bahía Lomas, SE part of the bay, 28-V-1978, 

Goodall 5294 (herb. Goodall); Puerto Sara, I-1954, Magens s.n. (F, SGO).

FIGURE 22. Geographic distribution of Chiliotrichum fuegianum (O. Hoffm.) Bonif.
5 0 • SMITHSONIAN CONTRIBUTIONS TO BOTANY

Haroldia Bonif. gen. nov.

Haroldia Bonif. gen. nov. Type: Chiliotrichiopsis mendocina

Cabrera, Notas Mus. La Plata, Bot. 17(84): 73, 1954 

(≡ Haroldia mendocina (Cabrera) Bonif. comb. nov.).

Frutex, ramis teretibus glandiferis, folia anguste elliptico-ovata base amplexicaulia margine revoluta, insigniter 

glandulifera. Capitula plurifl ora hetergama radiata solitaria, involucris campanulatis, receptaculis paleatis, paleis ad corollam disci aequalibus vel longioribus apicibus 

subulatis. Flosculi lutei dimorphi, fl osculis peripheralibus 

femineis radiatis, fl osculis disci hermaphroditis. Corollae 

tubifl ormes glanduliferes 5-lobatae abrupte ampliatae in 

venis basim versus incrassatis; thecae antherarum base non 

breviter sagittatis; rami stylorum in quadrantibus superioribus papillis collectoribus obtecti. Achenia ad extremum 

constricta, base stipitatea, villosa; squamae pappi crassae 

scabridae 1-seriatae.

Shrubs more or less globose; branches densely disposed, ascending, 6- to 8-ribbed, with notable glandular 

vesicular capitate trichomes. Leaves laxly disposed, persistent, alternate, ascending with a tuft of woolly trichomes 

on leaves axils, gradually reduced towards the apex, narrowly elliptic to narrowly ovate, base attenuated, petiole broadened towards the stem and constricted above, 

apex acute, margins revolute, notable glandular vesicular 

capitate trichomes on both sides, laxly tomentose pubescence on abaxial side, discolorous, chartaceous. Capitula

solitary, terminal, pedunculate, heterogamous, radiate. 

Involucres campanulate, phyllaries arranged in 2 to 3 

series; outer phyllaries linear-triangular to oblong, apex 

acute, notable glandular vesicular capitate trichomes on 

abaxial surface, chartaceous, margins membranaceous; 

inner phyllaries, narrowly elliptic, apex subulate, notable 

glandular vesicular capitate trichomes on abaxial surface, 

chartaceous, margins broadly membranaceous. Receptacles slightly convex, paleate; paleae linear to narrowly 

elliptic, embracing the achenes, apex subulate, rigid, when 

dry equaling or surpassing the corolla, notable glandular 

vesicular capitate trichomes towards the apex, margins 

membranaceous. Ray fl orets ~9, pistillate; corollas yellow, 

radiate, limb elliptic, 3-lobed; staminodes present; style 

base swollen, style branches oblong, subequal in lenght. 

Disk fl orets 21 to 30, bisexual; corollas yellow, tubular, 

gradually broadened towards the apex, 5-lobed, lobes triangular, corolla veins thick towards the base, protruding 

on outer surface when dry; anther thecae bases obtuse, not 

sagittate, apical appendages triangular, antheropodium 

present, not swollen; style base swollen, style branches 

ovate-oblong, apex acute, collecting hairs on upper 3/4. 

Achenes narrowly obovate, terete, (3)4- to 6-ribbed, with 

a noticeable constriction towards the apex, laxly and uniformly villose and gland-dotted. Pappus 1-seriate, composed of scales, triangular ovate, seldom divided towards 

the apex, notably scabrid.

Etymology: I dedicate this genus to Harold Robinson, renowned synanterologist who has made notable 

contributions to the taxonomy and systematics of the Compositae.

Haroldia is a new monotypic genus described herein, 

only known from northern Mendoza (Argentina), although probably extending over neighboring San Juan 

province. It inhabits dry rocky slopes in the ecotonal range 

between the high Andean and Monte biogeographic provinces ( Cabrera and Willink, 1973).

The genus is characterized by its minute pappus made 

of coarse scales, radiate heads, campanulate involucres, 

paleate receptacles, and yellow corollas with noticeable 

veins towards the base.

Although similar to Chiliotrichiopsis, Haroldia differs 

in the above mentioned characteristics, and also in the subulate and stiff nature of the receptacular paleae apices 

(versus acute and soft textured in Chiliotrichiopsis), the 

usually apically constricted achenes, and the stouter appearance of the twin trichomes on the achenes, very shortly 

divided towards the apex (versus twin trichomes, thinner 

and delicate and usually deeply divided towards the apex 

in Chiliotrichiopsis).

The systematic position of Haroldia is somewhat 

unresolved according to current molecular data (Bonifacino and Funk, unpublished data). Although forming 

part of the Chiliotrichiopsis + Nardophyllum + Cabreraea + Katinasia clade, it is clear that it does not belong to 

Chiliotrichiopsis.

Haroldia mendocina (Cabrera) Bonif. comb. nov. Chiliotrichiopsis mendocina Cabrera, Notas Mus. La Plata, 

Bot. 17(84): 73, 1954. Type: “[Argentina, Mendoza, 

Quebrada Chacay, Cordillera del Tigre, 2200 m, 9-XII1927, D. O. King 333” (holotype: LP 561922!).

(FIGURES 23, 24, 53)

Shrubs up to 1 m tall, more or less globose; branches 

densely disposed, ascending, 6- to 8-ribbed, with notable glandular vesicular capitate trichomes, internodes 

3−6 mm long. Leaves laxly disposed, persistent, alternate, 

ascending with a tuft of woolly trichomes on leaves axils, 
number 92 • 5 1

FIGURE 23. Haroldia mendocina (Cabrera) Bonif. A. Habit. B. Leaves, adaxial view (left), abaxial view (right). C. Involucre. D. Outer to inner 

phyllaries (from left to right). E. Receptacular paleae, lateral view (left), dorsal view (right). F. Detail of pappus scales. G. Achene with pappus. 

H. Style of ray fl oret. I. Style of disk fl oret. J. Corolla of disk fl oret. K. Corolla of ray fl oret (A−K, Simón & Bonifacino 782, LP).
5 2 • SMITHSONIAN CONTRIBUTIONS TO BOTANY

10−20 (−27) × 3−5 (−6.5) mm, gradually reduced towards 

the apex, narrowly elliptic to narrowly ovate, base attenuated, petiole broadened towards the stem and constricted 

above, apex acute, margins revolute, with notable glandular vesicular capitate trichomes on both sides, laxly tomentose pubescence on abaxial side, chartaceous. Capitula

solitary, terminal, pedunculate, peduncles (5−) 10−15 mm 

long, heterogamous, radiate. Involucres 5−6 × 4−5 mm, 

campanulate; phyllaries arranged in 2 to 3 series; outer 

phyllaries 3−4 × 0.5−1 mm, linear-triangular to oblong, 

apex acute, with notable glandular vesicular capitate trichomes on abaxial surface, coriaceous, margins membranaceous; inner phyllaries 4−6.5 × 1−1.5 mm, narrowly 

elliptic, apex subulate, with notable glandular vesicular 

capitate trichomes on abaxial surface, margins broadly 

membranaceous. Receptacles slightly convex, paleate; paleae 6−7 × 1−1.5 mm, linear to narrowly elliptic, embracing the achenes, apex subulate, rigid, when dry equaling 

or surpassing the corolla, with notable glandular vesicular 

capitate trichomes towards the apex, margins membranaceous. Ray fl orets ~ 9, pistillate; corollas yellow, radiate, tube 1−1.5 mm long, limb 4−7 × 2.5−4 mm, elliptic, 

3-lobed; staminodes 2 or 3; style shafts 1.8−3 mm long, 

base swollen, style branches 1.5−4.5 mm long, oblong, 

FIGURE 24. Geographic distribution of Haroldia mendocina (Cabrera) Bonif.
number 92 • 5 3

subequal in length. Disk fl orets 21 to 30, bisexual; corollas 

yellow, tubular, 5-lobed, gradually broadened towards the 

apex, tube 4.5−5 mm long, lobes 1.5−1.7 × 0.7−0.8 mm, 

triangular, gland-dotted abaxially, corolla veins thick towards the base, protruding on outer surface when dry; 

anther thecae 2−2.5 mm long, bases obtuse, not sagittate, 

apical appendages 0.5−0.6 × 0.3 mm, triangular, antheropodium present, not swollen; style shafts 3.5−4.5 mm 

long, base slightly swollen, style branches 1.5−2 mm long, 

ovate-oblong, apex acute, collecting hairs on upper 3/4. 

Achenes 3.5−5 mm long, narrowly obovate, terete, (3)4- 

to 6-ribbed, with a noticeable constriction towards the 

apex, laxly and uniformly villose and gland-dotted. Pappus 1−1.5 mm long, 1-seriate, composed of 18 to 25 fl attened bristles, triangular ovate, seldom divided towards 

the apex, notably scabrid.

Vernacular names: Not known.

Phenology: Flowering specimens have been collected from December to January.

Distribution: Argentina, endemic to Mendoza province, probably extending into southern San Juan (Figure 24).

Habitat: Ecotonal area bewteen Monte and high 

Andean vegetation (Cabrera and Willink, 1973), from 

2,200 to 2,600 m, along rocky mountain slopes, not forming big consociations

Uses: Not known.

Etymology: The specifi c epithet refers to the geographic area where the type specimen was collected.

Additional specimens examined: ARGENTINA. PROV. 

MENDOZA. Dpto. Las Heras. Cordillera del Tigre, Estancia 

San Alberto, 32o27'49"S, 69o25'47"W, 2,500 m, 21-I-2005, 

Bonifacino & Muttoni 1646 (US); Cordillera del Tigre, 

Estancia San Alberto, 32o24'37"S, 69o28'21"W, 2,970 m, 

22-I-2005, Bonifacino & Muttoni 1656 (US); tres km al 

W de Uspallata, camino al NW, 32o31.7'S, 69o30.5'W, 

2,600 m, 26-I-2001, 26-I-2001, Simon & Bonifacino 775

(LP); íd., Simon & Bonifacino 782 (LP); íd., 26-I-2001, 

Simon & Bonifacino 791 (LP).

Katinasia Bonif. gen. nov.

Katinasia Bonif. gen. nov. Type: Nardophyllum cabrerae

Bonif., Taxon 54(3): 688−689 (≡ Katinasia cabrerae

(Bonif.) Bonif. comb. nov.).

Frutex, ramis teretibus lanuginosis (tomentosis) et 

glanduliferis. Folia base amplexicaulia, marginibus integris non revolutis. Capitula plurifl ora homogama discoidea solitaria vel 2−5 (−7) in corymbis terminalibus laxe 

disposita, involucris cylindricis, receptaculis paleatis vel 

nudis. Flosculi hermaphroditi, corollis tubiformibus albis 

glanduliferis 5-lobatis abrupte ampliatis; rami stylorum in 

dimidiis superioribus papillis collectoribus obsiti. Achenia 

glandulo-punctata etiam in pilis geminatis brevibus leviter pubescentia; aristae pappi inaequales scabridae 3- vel 

4-seriatae.

Shrubs hemispheric; branches more or less densely disposed, ascending, 8-ribbed, tomentose and gland-dotted. 

Leaves more or less laxly arranged, persistent, alternate, 

ascending, narrowly elliptic, base slightly attenuated, base 

attenuated, petiole broadened towards the stem and slightly 

constricted above, apex acute, margins entire, non-revolute, 

tomentose and gland-dotted on both sides, secondary veins 

evident (one at each side of midvein), concolorous, coriaceous. Capitula arranged in corymbiform capitulescenses 

composed by 2 to 5 (7) heads, sometimes solitary, terminal, 

sessile or pedunculate, homogamous, discoid. Involucres

cylindrical; phyllaries arranged in 5 to 6 series; outer phyllaries ovate-elliptic, apex acute-acuminate, gland-dotted 

to slightly tomentose on abaxial surface, coriaceous; inner 

phyllaries narrowly-elliptic to oblong, apex acute-acuminate, gland-dotted towards the apex, papery. Receptacles

fl at, naked, occasionally with 1 or 2 paleae; paleae linear 

to narrowly elliptic, not embracing the achenes, apex acute, 

margins membranaceous. Florets 4 to 6, bisexual; corollas 

white, tubular, 5-lobed, broadened more or less abruptly 

from mid-length, lobes triangular; anther thecae bases 

slightly sagittate, apical appendages triangular, antheropodium present, not swollen; style bases slightly swollen, style 

branches linear to oblong, apex acute, collecting hairs on 

upper 1/3. Achenes obconic, terete, 5- to 8-ribbed, sparsely 

villose and gland-dotted, villose pubescence mostly on the 

ribs, carpopodium present. Pappus 3- to 4-seriate, composed of terete bristles, scabrid, the inner series shortly barbellate towards the apex, outer series notably shorter.

Etymology: I dedicate this genus to Liliana Katinas, 

leading Argentinean synantherologist, who has worked extensively on southern South American Compositae.

Katinasia is a new monotypic genus described herein, 

that it is restricted to an arid area of western Argentina 

in both southern Mendoza and Neuquén provinces. It inhabits the ecotonal area between Patagonian and Monte 

biogeographic provinces (Cabrera and Willink, 1973).

The genus is characterized by its discoid heads with 

4 to 6 fl orets, white corollas and a 3- to 4-seriate pappus, 

composed of scabrid terete bristles, the outer series noticeably shorter, and the leaves having entire (non-revolute) 

margins.

Specimens belonging to this genus had been traditionally identifi ed as Nardophyllum lanatum, a species endemic 
5 4 • SMITHSONIAN CONTRIBUTIONS TO BOTANY

to Chile, mainly because of the resemblance in the shape of 

the involucre (Cabrera, 1954a, 1971; Bonifacino and Sancho, 2001; Sancho and Ariza Espinar, 2003). After a detailed 

study of all available specimens and detailed fi eld observations by Bonifacino (2005), it was concluded that the Argentinean populations ascribed to Nardophyllum lanatum

actually belonged to a distinct undescribed species, which 

was described as Nardophyllum cabrerae Bonifacino. Nardophyllum cabrerae differed from the rest of Nardophyllum species, in several noticeable characteristics, the most 

notable being the white color of the corollas and the nature 

of the pappus. Notwithstanding those differences, from the 

narrow phylogenetic information available in 2005, segregation from Nardophyllum did not seem needed. Recent 

molecular research on the whole Chiliotrichum Group 

(Bonifacino and Funk, unpublished data) has yielded interesting results concerning the position of this species, clearly 

showing its distinction from Nardophyllum, and placing it 

as the sister group to the Nardophyllum + Chiliotrichiopsis

clade. The characteristics that marked this species as new, 

when looked at in a broader phylogenetic context (i.e.: the 

whole Chiliotrichum Group), proved suffi cient to support 

a generic segregation on morphological grounds. Further, 

Katinasia is the only genus inside Chiliotrichum Group 

with iso- bilateral mesophyll in the leaves.

Katinasia cabrerae (Bonif.) Bonif. comb. nov. Nardophyllum cabrerae Bonif., Taxon 54(3): 688−689. Type: 

“Argentina, Prov. Neuquén, Dpto. Catan Lil, al W de 

Ruta 40, cerca de Arroyo China Muerta, 39°31'26.9"S, 

70°18'49.6"W, 29-I-2003, Bonifacino & Romano 825”

(holotype: LP!; isotypes: MVFA!, SI!).

(FIGURES 25, 26, 54)

Shrubs 40–100 cm tall, hemispheric; branches more 

or less densely disposed, ascending, branches 8-ribbed, tomentose and gland-dotted, internodes 1–6 mm long. Leaves

more or less laxly arranged, persistent, alternate, ascending, 

7−17 × 0.6−2.5 mm, gradually reduced towards the branch 

apex, narrowly elliptic, base slightly attenuated, petiole 

broadened towards the stem and slightly constricted above, 

apex acute, margins entire, non- revolute, slightly tomentose 

and gland-dotted on both sides, secondary veins evident (one 

at each side of midvein), concolorous, coriaceous. Capitula

arranged in corymbiform capitulescenses composed by 2 to 

5 (7) heads, sometimes solitary, terminal, sessile or with peduncles up to 15 mm long, homogamous, discoid. Involucres 10−12 × 2−4 mm, cylindrical; phyllaries arranged in 5 

to 6 series; outer phyllaries 4−5 × 0.5−2 mm, ovate-elliptic, 

apex acute-acuminate, gland-dotted to slightly tomentose 

on abaxial surface, coriaceous; inner phyllaries 6−10 × 1−2 

mm, narrowly-elliptic to oblong, apex acute-acuminate, 

gland-dotted towards the apex, papery. Receptacles fl at, 

naked, occasionally partially paleate; paleae 1 or 2, ~ 10 × 1 

mm, linear to narrowly elliptic, not embracing the achenes, 

apex acute, margins membranaceous. Florets 4 to 6, bisexual; corollas white, tubular, 5-lobed, broadened more 

or less abruptly from mid-length, tube 2.5−3.5 mm long, 

throat 3.5−5 mm long, lobes 1−1.6 × 0.5 mm, triangular, 

gland-dotted abaxially; anther thecae 2−2.5 mm long, bases 

slightly sagittate, apical appendages 0.8−1.1 × 0.2−0.3 mm, 

triangular, antheropodium present, not swollen; style shafts 

8−12 mm long, base slightly swollen, style branches 2.5−3.5 

mm long, linear to oblong, acute, collecting hairs on upper 

1/3. Achenes 4.5−6 mm long, obconic, terete, 5- to 8-ribbed, 

sparsely villose and gland-dotted, villose pubescence mostly 

on the ribs, carpopodium present. Pappus 6−8 mm long, 3- 

to 4-seriate, composed of 110 to 140 terete bristles, scabrid, 

the inner series shortly barbellate towards the apex, outer 

series shorter, 2−3 mm long.

Vernacular names: Not known.

Phenology: Flowering specimens have been collected from December to February.

Distribution: Argentina, endemic to southern Mendoza and Neuquén (Figure 26).

Habitat: Semiarid steppes between 700 and 1,580 m 

in the Patagonia-Monte ecotonal range (Cabrera and 

Willink, 1973). Considering the relatively few previous 

collections, Katinasia cabrerae seems to be a rare species. However, it is locally abundant at the type locality. 

It grows associated with Colliguaja integerrima Gill. and 

Hook., Larrea nitida Cav., Prosopis denudans Benth., 

Nassauvia glomerulosa D. Don, Tetraglochin ameghinoii 

Speg., and Schinus spp.

Uses: Not known.

Etymology: The specifi c epithet refers to late great 

synantherologist Angel Lulio Cabrera (1908−1999).

Additional specimens examined: ARGENTINA. PROV. 

MENDOZA. Dpto. Malargue. Ruta 40, 20 km S de Calmuco, 

1-II-1963, Boelcke et al. 10420 (BAA, LP, SI); Laguna de 

Coipo Lauquen, 36°35'19"S, 69°49'40.6"W, 1555 m, 

8-XII-2006, Bonifacino et al. 2906 (US); Laguna de Coipo 

Lauquen, al Sur de Calmuco, 15-I-1942, Burkart et al. 14425

(LP, SI); Laguna de Coipo Lauquen, 11-I-1973, Cabrera et 

al. 22843 (LP); entre Calmuco y Río Barrancas, 15-II-1942, 

Covas 280 (LP); entre Ranquil Norte y Río Barrancas, 5-I1966, Ruiz Leal 24371 (LP); El Manzano, 1580 m, 27-I-1941, 

Ruiz Leal 7430 (LP); Laguna de Coipo Lauquen, 36°34.5'S, 

69°49.5'W, 1,500 m, 30-I-2001, Simón & Bonifacino 847

(LP). PROV. NEUQUÉN. Dpto. Pehuenches. Entre Río Barrancas y Buta Ranquil, 8-II-1950, Boelcke 4235 (BAA, LP).
number 92 • 5 5

FIGURE 25. Katinasia cabrerae (Bonif.) Bonif. A. Habit. B. Leaves, adaxial view (left), abaxial view (right). C. Involucre. D. Outer to inner phyllaries (from left to right). E. Palea. F. Detail of pappus terete bristle. G. Achene with pappus. H. Style. I. Corolla (A−I, Bonifacino & Romano 

825, LP).
5 6 • SMITHSONIAN CONTRIBUTIONS TO BOTANY

Lepidophyllum Cass.

Taxonomic History

1816. Cassini creates Lepidophyllum to include a species 

originally described by Lamark from the Straits of 

Magellan in 1786 as Conyza cupressiformis.

1823. Cassini makes the combination Lepidophyllum 

cupressiforme.

1839. Meisner does not accept Cassini’s Lepidophyllum

and, without explanation, coins Brachyridium

to replace Cassini’s name; presents also the combination Brachyridium cupressiforme (Lam.) 

Meisner.

FIGURE 26. Geographic distribution of Katinasia cabrerae (Bonif.) Bonif.
number 92 • 5 7

1855. Schultz Bipontinus (1855a) creates the combination 

Gutierrezia cupressiformis (Lam.) Sch. Bip.

1862−1931. Several authors (see “Excluded Species”) coin 

species names or made combinations in Lepidophyllum; however, all those taxa are now referred 

to the genus Parastrephia Nutt. emend. Cabrera.

1945. Cabrera presents a synopsis for Lepidophyllum

(which includes Parastrephia Nutt. emend. Cabrera) and creates two sections, Eulepidophyllum

and Tola to place current Lepidophyllum and 

Parastrephia respectively.

1954. Cabrera (1954a), in his revision of the genus Nardophyllum, and after detailed studies of related South 

American Compositae revises the circumscription 

of Lepidophyllum, excluding the species from the 

Puna region and transferring them to Parastrephia, 

thus rendering Lepidophyllum monotipic, with the 

single species, L. cupressiforme.

Lepidophyllum Cass., Bull. Sci. Soc. Philom. Paris 1: 199, 

1816. Brachyridium Meisn. nom. illeg., Pl. Vasc. Gen., 

Diagn., 187, 1839. Lepidophyllum sect. Eulepidophyllum Cabrera nom. illeg., Bol. Soc. Argent. Bot. 1: 49, 

1945. Type: Conyza cupressiformis Lam., Encycl. 2: 91, 

1786 (≡ Lepidophyllum cupressiforme (Lam.) Cass., in 

Cuvier, Dict. Sci. Nat., ed. 2, 26: 37, 1823).

Shrubs globose; branches densely disposed, ascending, 

not ribbed, gland-dotted. Leaves very densely arranged, 

persistent, opposite, decussate, sessile, triangular-ovate, 

base truncate, apex obtuse, margins slightly involute, 

mostly glabrous, with a few aseptate-fl agellate trichomes 

along margins in younger leaves, coriaceous. Capitula

solitary, terminal, sessile, heterogamous, radiate. Involucres cylindrical-ovoid, phyllaries arranged in 3 to 4 series; 

outer phyllaries ovate-elliptic, boat-shaped, apex obtuse, 

with a few glandular trichomes towards the apex, coriaceous, margins membranaceous; inner phyllaries narrowly 

obovate, apex obtuse, with a few glandular trichomes towards the apex, papery, margins membranaceous. Receptacles fl at, partially paleate; paleae 1 (2), linear to narrowly 

obovate, not embracing the achenes, sometimes reduced 

to a single awn, apex obtuse, sometimes acute, with a few 

glandular trichomes towards the apex, margins membranaceous. Ray fl orets (1) 3 to 6, pistillate; corollas yellow, 

radiate, limb elliptic to oblong, 3-lobed; staminodes present; style base not swollen, style branches linear-triangular, subequal in length. Disk fl orets (3) 4 to 7, bisexual, 

functionally male; corollas yellow, tubular, 5-lobed, tube 

broadened more or less abruptly from mid-length, lobes 

triangular; anther thecae bases slightly sagittate, apical appendages ovate, anteropodium present, swollen; style base 

slightly swollen, style branches oblong, obtuse, collecting 

hairs on upper 1/2. Achenes narrowly obovate, terete, 

7- to 8-ribbed, gland-dotted, carpopodium present. Pappus

2- to 3-seriate, composed of linear scales, scabrid along 

the margins, slightly shorter on ray fl orets.

Etymology: Name derived from Latin lepido (scale) 

and phyllum (leaves).

Lepidophyllum is monotypic and is distributed in 

Argentina from northern Santa Cruz to Tierra del Fuego, 

and in Chile in the provinces of Magallanes and Tierra del 

Fuego of Region de Magallanes. It inhabits both coastal 

and interior saline plains in the Patagonian biogeographic 

province (Cabrera and Willink, 1973). It has also been 

found on rocky slopes in northern Santa Cruz.

Lepidophyllum is characterized by the reduced, opposite and decussate leaves, radiate capitula, partially paleate 

receptacle, yellow ray corollas and gland-dotted achenes. 

According to molecular evidence (Bonifacino and Funk, 

unpublished data), the closest relatives to Lepidophyllum

are Chiliophyllum and Chiliotrichum. It differs from Chiliotrichum and Chiliophyllum (most notably) in the opposite phylotaxy, a unique feature in the Chiliotrichum

Group. Lepidophyllum Cass. is monotypic, and composed 

of the species Lepidophyllum cupressiforme (Lam.) Cass.

Lepidophyllum cupressiforme (Lam.) Cass., in Cuvier, 

Dict. Sci. Nat., ed. 2, 26: 37, 1823. Conyza cupressiformis Lam., Encycl. 2: 91, 1786. Baccharis cupressiformis (Lam.) Pers., Syn. Pl. 2: 425, 1807. Athanasia 

cupressiformis (Lam.) Commerson ex Cass., in Cuvier 

pro syn., Dict. Sci. Nat., ed. 2, 26: 37, 1817. Brachyridium cupressiforme (Lam.) Meisn. nom illeg., Pl. Vasc. 

Gen., Comment., 127, 1839. Gutierrezia cupressiformis 

(Lam.) Sch. Bip., Flora 8: 114, 1855. Grindelia cupressiformis (Lam.) Sch. Bip. ex Reiche pro syn., Anales Univ. 

Chile 109: 25, 1901. Type: “[Chile] Magellan, au cotè 

du patagons, Baye Boucault, M. Commerson s.n.” (holotype: P 279523; scanned image of holotype at P: LP!; 

isotype: P 222885; photos of isotype: F!, SI!, LP!).

(FIGURES 27, 28, 55)

Shrubs up to 1.5 m tall, globose; branches densely 

disposed, ascending, not ribbed, gland-dotted, internodes short, shorter than 0.5 mm. Leaves very densely 

arranged, persistent, opposite, decussate, sessile, 1.5−3 × 

1.3−2.2 mm, triangular-ovate, base truncate, apex obtuse, 
5 8 • SMITHSONIAN CONTRIBUTIONS TO BOTANY

margins slightly involute, mostly glabrous, with a few 

aseptate-fl agellate trichomes along margins in younger 

leaves, coriaceous. Capitula solitary, terminal, sessile, 

heterogamous, radiate. Involucres 5−7 × 3−4 mm, cylindrical-ovoid, phyllaries arranged in 3 to 4 series; outer 

phyllaries 3−5.5 × 2−2.5, ovate-elliptic, boat-shaped, apex 

obtuse, glabrous, with a few glandular trichomes towards 

the apex, coriaceous, margins membranaceous; inner 

phyllaries 5−7 × 1.2−1.6 mm, narrowly obovate, apex obtuse, with a few glandular trichomes towards the apex, papery, margins membranaceous. Receptacles fl at, partially 

paleate; paleae 1 (−2), 5.5−7 × 1 mm, linear to narrowly 

obovate, not embracing the achenes, sometimes reduced 

to a single awn, apex obtuse, sometimes acute, margins 

membranaceous, with a few glandular trichomes towards 

the apex. Ray fl orets (1−) 3 to 6, pistillate; corollas yellow, 

radiate, tube 2−3.6 mm long, limb 2−4 × 1−1.2 mm, elliptic to oblong, 3-lobed; staminodes 1, very reduced; 

style base not swollen, style branches ~ 2 mm long, linear-triangular, subequal in length. Disk fl orets (3−) 4 to 

7, bisexual, functionally male; corollas yellow, tubular, 

5-lobed, broadened more or less abruptly from midlength, tube 3−3.5 mm long, throat 1−1.5 mm long, lobes 

1.5 × 0.6−0.7 mm, triangular, gland-dotted abaxially; anther thecae ~2 mm long, anther bases slightly sagittate, 

apical appendages 0.5 × 0.2−0.3 mm, ovate, antheropodium present, notably swollen; style shafts 5−6 mm long, 

base slightly swollen, style branches 1−1.3 mm long, oblong, apex obtuse, collecting hairs on upper 1/2. Achenes

4−6 mm long, narrowly obovate, terete, 7- to 8-ribbed, 

gland-dotted towards the apex, carpopodium present. 

Pappus 3.5−5 mm long, 2- to 3-seriate, composed of linear scales, scabrid along the margins, slightly shorter on 

ray fl orets.

Vernacular names: “Mata verde” (Morrison s.n., 

SI); “Pichi” (Philippi s.n., SGO); “Mata negra” (Haloua 

and Padín, 1999); “Pine scented daisy” (Borla and Vereda, 

2001).

Phenology: Flowering specimens have been collected from October to April.

Distribution: Argentina, provinces of Santa Cruz 

and Tierra del Fuego, and Chile in the Región of Magallanes in the provinces of Magallanes and Tierra del 

Fuego (Figure 28).

Habitat: Coastal and interior saline plains, however it has also been found on rocky slopes of Bosques 

Petrifi cados National Monument in Santa Cruz province 

(Argentina), always in the Patagonian biogeographic province (Cabrera and Willink, 1973). The commonest habitat 

is along estuary banks where it can form dense consociations. It can also grow associated to Berberis buxifolia 

Lam., Senecio patagonicus Hook. and Arn., Anarthrophyllum desideratum (DC.) Benth., Acaena spp. Festuca gracillima Hook. f., Chiliotrichum diffusum (G. Forst.) Kuntze, 

Chiliotrichum fuegianum (O. Hoffm.) Bonif. (Roig, 1998; 

Pisano, 1977; pers. obs.). On the coastal saline plains where 

grasses are nonexistent or very sparse Lepidophyllum cupressiforme grows associated to Sarcocornia perennis 

(Mill.) A.J. Scott.

Uses: This species is highly promissory as a resin 

producer, given the impressive amounts of this substance 

in the leaves. Used as ornamental (Goodall, pers. comm.; 

Haloua and Padín, 1999). Branches used as fuel (Haloua 

and Padín, 1999).

Etymology: The specifi c epithet refers to the resemblance of the leaves to those of Cupressus L.

Note 1: In the original description of Conyza cupressiformis, Lamarck (1786) indicated: “Cette plante a 

été découverte dans les terres de Magellan par M. Commerson”. I have observed two scanned images of specimens 

at P that, corresponding entirely with the diagnosis, bear 

the following labels: “Magellan, au cotè du patagons, Baye 

Boucault, M. Commerson s.n, 1767” (P 279532), and “Magellan, Coste des Patagons. Baye Boucault, sur les collines, 

M. Commerson, 1767” (P 222895). Both specimens have a 

handwritten description by Commerson where the species 

is named Athanasia cupressiformis. Given the total conformity of these specimens with the species characteristics, 

as well as the locality data, they are identifi ed as holotype 

(P279532) and isotype (P222895). The image distributed 

by F as the type of the species corresponds with the holotype specimen (P279532) at P.

Note 2: The publication date for Lepidophyllum 

cupressiforme has traditionally been associated to the date 

in which Cassini published Lepidophyllum (1816) in “Bulletin des Sciences, par la Societe Philomatique”. However, 

Cassini made the combination Lepidophyllum cupressiforme (Lam.) Cass. in Cuvier in 1823 in “Dictionnaire des 

Sciences Naturelles, ed. 2”.

Note 3: C. Meisner (1839) did not accept Cassini’s 

Lepidophyllum and without explanation he coined Brachyridium. According to article 52.1 of the present code (Viena 

Code, Mcneill et al, 2006) Brachyridium represents a superfl uous renaming and therfore consitutues an illegitimate 

name. In the same publication, Meisner also created the 

combination Brachyridium cupresiforme (Lam.) Meisner 

which also bears the status of illegitimate. It is interesting 

that Meisner’s description of Brachyridium is a word-byword re-shuffl ed copy of Candolle’s (1836b) description of 

Lepidophyllum, even including an erroneous description 
number 92 • 5 9

FIGURE 27. Lepidophyllum cupressiforme (Lam.) Cass. A. Habit. B. Leaves, adaxial view (left), abaxial view (right). C. Detail of a branch 

showing opposite phyllotaxy. D. Involucre. E. Receptacular paleae, dorsal view (left), lateral view (right). F. Outer to inner phyllaries (from left 

to right). G. Detail of pappus scales. H. Achene with pappus. I. Style of ray fl oret. J. Style of disk fl oret. K. Corolla of disk fl oret. L. Corolla of 

ray fl oret (A−L, Bonifacino et al. 787 L; FIGURE 27A by Alice Tangerini).
6 0 • SMITHSONIAN CONTRIBUTIONS TO BOTANY

of the pappus by the Swiss botanist. The description of the 

pappus presented by Cassini perfectly matches the specimen. In the description that Candolle presents in his prodromus, he comments on the similarities of Lepidophyllum

with Brachyris, and it is probably from this assertion that 

Meisner took the idea for the new name.

Note 4: In his synopsis for Lepidophyllum (which 

included Parastrephia Nutt. emend. Cabrera) Cabrera 

(1945) created two sections, Eulepidophyllum and Tola,

to contain current Lepidophyllum and Parastrephia respectively. However, he did not provide a Latin diagnosis and so 

the names he created are rendered illegitimate under article 

36.1 of the current Vienna Code (McNeill et al., 2006).

Note 5: When capitula have a reduced number of fl orets (2 or 3 ray fl orets, and ~3 disk fl orets) palea are absent.

Additional specimens examined: ARGENTINA. PROV. 

SANTA CRUZ. Not defi ned Dpto. Patagonia, II-1891, Beaufi ls 603 (NY); Río Santa Cruz, 24-XII-1896, Hatcher 119 

FIGURE 28. Geographic distribution of Lepidophyllum cupressiforme (Lam.) Cass.
number 92 • 6 1

(NY); Patagonia, 1911, Marelli s.n. (SI); íd., Moreno & 

Tonini 161 (NY); íd., 1904, Tessleff s.n. (BAB). Dpto. 

Corpen Aike. Santa Cruz, s. date, Burmeister s.n. (SI); íd., 

4-XII-1932, Castellanos s.n. (BA 7928); Puerto Santa Cruz, 

12-XII-1975, Correa et al. 6577 (BAB); Santa Cruz, I-1914, 

Hicken & Haumamn 823 (SI); Puerto Santa Cruz, 7-XII1945, O’ Donell 3943 (BAA, NY); Meseta del Río Santa 

Cruz y Chico, 12-XII-1902, Spegazzini s.n. (BAB). Dpto. 

Deseado. Puerto Deseado, 25-I-1896, Alboff s.n. (LP); 

Monumento Bosques Petrifi cados, cañadón al NE de la estación del Parque, 47°39'17"S, 67°58'37"W, 3-XII-2002, 

Bonifacino & Donato 772 (LP); Ruta prov 47, 48°9'23"S, 

67°17'32"W, 3-XII-2002, Bonifacino & Donato 785 (LP); 

Ruta prov 47, 47°45'37"S, 66°10'59"W, 3-XII-2002, 

Bonifacino & Donato 787 (LP); Puerto Deseado, camino 

Cabo Blanco a Tellier, 13-XI-1965, Correa & Nicora 3358 

(BAB); a 12 km de Puerto Deseado camino a Cabo Blanco, 

19-XI-1963, Correa et al. 2630 (BAB); Puerto Deseado, 

13-I-1914, Hicken & Haumamn 140 (LP); Puerto Deseado, 13-I-1914, Hicken & Haumamn 143 (SI); Deseado, 

1899−1900, Spegazzini s.n. (BA 5116); Antonio de Biedma, 

25-XI-1969, Vidal 29 (LP). Dpto. Güer Aike. margen derecha Río Coyle, junto a puente sobre Ruta 3, 13-I-1970, s. 

leg. (LP); Márgenes del Río Gallegos, 26-XI-1976, Ambrosetti 26618 (BAB); Cabo Virgenes, 26-IV-1968, Anliot 

6051 (SGO); Río Gallegos, Estancia Los Pozos, 51°33'S, 

69°20'W, 5-XII-1975, Arroyo et al. (BAB); Estación exp. 

Río Chico, s. date, Boelcke et al. 12325 (BAA, SI); Río 

Gallegos, Boelcke et al. 15297 (BAB); Río Gallegos, 22-III1917, Bonarelli s.n. (BA 5115); Ruta prov 57, Cañadón Boliche 51°0'37"S, 69°21'25"W, 29-I-2002, Bonifacino et al. 

600 (LP); Río Gallegos, s. date, Brown 85 (NY); íd., 5-XII1932, Castellanos 7937 (LP); íd., 5-XII-1932, Castellanos 

s.n. (BA 7937); íd., 6-I-1960, Correa & Perez Moreau 1789 

(BAB, LP); camino Lemarchand a Puerto Coig, 24-XI-1963, 

Correa et al. 2791 (BAB); Estancia Cabo Buen Tiempo, 

2-XII-1975, Correa et al 6559 (BAB); Río Gallegos, 10 m, 

20-XI-1975, Gentili 324 (BAB, LP, SI); Río Gallegos, 5-X1971, Goodall 3897 (BAB); Río Gallegos, Ruta 3 sobre el 

río, 17-XII-1985, Hermann s.n. (SI); Río Gallegos, 27-III1914, Hicken & Hauman s.n. (SI 9386); Estancia Montedinero, Cabo Vírgenes, XI-1974, Molina 6 (LP); Estancia 

Las Vegas, valle del Río Coyle, I-1914, Morrison s.n. (SI); 

S. loc., 15-XII-1945, O’Donell 4079 (US); Río Gallegos, 

12-XII-1945, O’Donell 4086 (LP); íd., 28-II-1959, Perrone 

s.n. (BA 58435); Río Gallegos, 28-II-1959, Perrone s.n.

(BA 70460); a 70 km de Gallegos rumbo a Esperanza, 8-III1936, Scott 231 (LP); entre Gallegos y Coyle, Monte Tigre, 

12-III-1936, Scott & Biraben 239 (LP); entre Gallegos y 

Monte Aymond, 22-I-1949, Soriano 3280 (BAB); Río Gallegos, 5-I-1941, Spegazzini s.n. (BAB). Dpto. Lago Buenos 

Aires. Ruta provincial 502, a 28 km al N de Dos Manantiales, 48o00'S, 69o43'W, 8-II-975, Boelcke et al. 16227 

(BAB, LP). Dpto. Magallanes. A 52 km de El Salado, sobre 

Ruta 3, 170 m, 13-I-1970, s. leg. (LP); Puerto San Julián, 

XII-1933, Blake 208 (LP); San Julián, 22-II-1944, Blake 

437 (MO, SI); Puerto San Julián, s. date, Carette s.n. (LP); 

San Julián, 20 km al S de San Julián, Ruta 3, 22-XI-1963, 

Correa et al. 2686 (BAB); 35 km N of San Julián, road to 

puerto Deseado, 20 m, s. date, Eyerdam et al. 23975 (F, 

SI); San Julián, a 3 km camino a Piedra Buena, 18-I-1949, 

Grondona 2008 (BAB). PROV. TIERRA DEL FUEGO. Dpto. Río 

Grande. Caleta Josefi na, 8-IV-1921, s. leg. 53 (BA 5113); 

Ruta 3, Bahía San Sebastián 53°14'01"S, 68°34'6"W, 28-

I-2002, Bonifacino et al. 590 (LP); Extremo N de Península 

Páramo, 52°59'14"S, 68°16'12"W, 28-I-2002, Bonifacino 

et al. 594 (LP); Ruta 3 Bahía San Sebastián, 53°14'1"S, 

68°34'6"W, 28-I-2002, Bonifacino et al. 598 (LP); al N de 

Bahía San Sebastián, 52°53'19"S, 68°21'38"W, proximo a 

planta “TOTAL, 13-XII-2006, Bonifacino et al. 3016 (US); 

al N de San Sebastián, 16-I-1933, Castellanos s.n. (LP, BA 

7946); desembocadura del Arroyo Alfa, 20-II-1942, Castellanos s.n. (BA 45793); borde de playa sobre Bahía San Sebastián, 53°17'S, 68°26'W, 29-I-1995, Fortunato & Elechosa 

4892 (NY, US); Bahía San Sebastián, Estancia San Martín, 

6-II-1955, Hunziker 6828 (BAB, LP); Estancia Cullen, 7-

IV-1968, Goodall 1718 (NY); Estancia Cullen, NW edge of 

San Sebastian Bay, north end of salt plateau, 14-XI-1970, 

Goodall 2572 (SI); Bahía San Sebastián, Estancia San Martín, near Cullen, 19-V-1972, Goodall 4292 (BAB); Bahía 

San Sebastián, Estancia San Martín, 6-II-1955, Grondona 

4475 (BAB); Estancia Cullen, 52o54'S, 68o25'W, 13-I-1968, 

Moore 1467 (LP); Estancia Cullen, Arroyo Beta, 52o44'S, 

68o33'W, 5-I-1972, Moore & Goodall 339 (BAB). Dpto. 

Ushuaia. Isla de los Estados y Canal de Beagle, 7-VIII-1913, 

Hogberg 180 (BAB); Isla de los Estados y Canal de Beagle, 

verano-otoño-1902, Rodriguez 31 (BAB). CHILE. REG. DE

MAGALLANES. Prov. Magallanes. Estrecho de Magallanes, 

próximo Primera Angostura, 52o27'28"S, 69o32'56.3"W, 

2 m, 13-XII-2006, Bonifacino et al. 3014 (US); San Gregorio, 26-X-1968, Cekalovic s.n. (BAB, LP); Angostura, 300 

m, 14-I-1930, Donat 213 (F, NY); Ruta 255, 120 km E of 

Punta Arenas, 2 km E of Bahía San Gregorio, 52°32'–34'S, 

52°32'–34'W, 9-X-1971, Dudley et al. 82 (MO, SGO); 

Cabo Fairweather, s. date, King 2 (BM); Gregory Bay, 

Strait of Magellan, 19-I-1888, Lee s.n. (US); Estancia Punta 

Delgada, ~180 km NE of Punta Arenas, coast of Bahía Posesión, 18-III-1964, Moore 1058 (K, SGO); próximo a Primera Angostura, 52°20'S, 69°40'W, 22-XII-1976, Seibert et 

al. (BAB). Prov. Tierra del Fuego. Bahía Inútil, 1-III-1917, 

Bonarelli s.n. (SI 9385); Bahía Inútil, extremo N, 53o20'S, 

69o35'W, 14-III-1968, Moore 2253 (LP); SW de Punta 
6 2 • SMITHSONIAN CONTRIBUTIONS TO BOTANY

Catalina, Cuarto Chorillo, 52o39'S, 68o59'W, 7-XI-1971, 

Moore 2355 (LP); Fuegia ad litt. Maris, II-1879, Philippi 

s.n. (SGO); 84.7 km from Porvenir along road to San Sebastián, 31-III-1985, Wallace & Forlonge 039/85 (SGO).

Nardophyllum (Hook. and Arn.) 

Hook. and Arn.

Taxonomic History

1786. Lamarck describes Conyza bryoides based on material collected by Commerson in the Straits of 

Magellan.

1807. Persoon includes Lamarck’s species in Baccharis, 

coining Baccharis bryoides (Lam.) Pers.

1834. Meyen describes Barnadesia lanata based on material from “Río del Volcán”, in the Andes of Region Metropolitana, Chile.

1835. Hooker and Arnott describe Gochnatia revoluta. 

They were uncertain about the placement of this 

species in Gochnatia so they described Gochnatia

subgenus Nardophyllum? (with a question mark) 

to accomodate the species.

1836. Hooker and Arnott elevate Gochnatia subg. Nardophyllum to generic status, and provide a more 

detailed description of the taxon. They also publish the combination Nardophyllum revolutum

(Hook. and Arn.) Hook. and Arn., and describe a 

new species, Nardophyllum obtusifolium.

1838. Candolle describes Dolichogyne, apparently unaware that it corresponds with the same taxonomic entity described earlier by Hooker and 

Arnott (1836) as Nardophyllum, along with 

which he also describes two new species, Dolichogyne staehelinoides and D. gnaphalioides.

1846. Hooker describes Chiliotrichum sect. Anactinia, and 

describes three new species: Chiliotrichum humile, 

C. kingii, and C. darwinii. He commented on the 

similarities between the typical section of Chiliotrichum and the newly described section pointing 

out that the difference was mainly in the absence 

of ray fl orets in Chiliotrichum sect. Anactinia.

1849. Rémy elevates Chiliotrichum sect. Anactinia to genus 

rank creating Anactinia (Hook. f.) J. Rémy, and 

proposes Anactinia hookeri for Chiliotrichum 

humile, creating an illegitimate name. Rémy also 

describes a new species, Dolichogyne chiliotrichioides, from material collected in Sotaqui. In the same 

publication, Rémy also considers Dolichogyne staehelinoides and D. gnaphalioides to be heterotypic 

synonyms, proposing a new and illegitimate name 

to refer to these taxa: Dolichogyne candollei.

1855. Schultz Bipontinus (1855b) creates the combination 

Microchaete humilis (Hook. f.) Sch. Bip.

1856. Philippi describes Dolichogyne genistoides based 

on material collected by Germain in the Andes of 

Santiago (Chile).

1856. Weddell describes Dolichogyne armata (currently 

placed in Ocyroe) based on material from Bolivia.

1862. Gray makes an important contribution to the understanding of Nardophyllum, identifying synonyms 

and making several combinations. He identifi es 

Dolichogyne staehelinoides, D. gnaphalioides,

and D. candollei as heterotypic synonyms of Nardophyllum revolutum. Gray also recognizes the 

identity between Nardophyllum and Chiliotrichum sect. Anactinia, forming the combinations 

Nardophyllum kingii, N. humile, and N. darwinii. Finally, he notices that Dolichogyne chiliotrichioides also belongs to Nardophyllum, and 

accordingly makes the new combination.

1873. Gray notes similarities between Dolichogyne genistoides and the rest of Nardophyllum species and 

makes the combination Nardophyllum genistoides.

1894. Philippi describes three new species of Nardophyllum: Nardophyllum scoparium, N. parvifolium, 

and N. paniculatum.

1898. Kuntze places Nardophyllum in the synonymy of 

Aster, publishes the new species Aster nardophyllum, and makes the combination Aster staehelinoides (DC.) Kuntze.

1901. Reiche includes Dolichogyne armata in Nardophyllum, and makes the combination Nardophyllum 

armatum (currently placed in Ocyroe). Reiche also 

places Ocyroe spinosa Phil. in the synonymy of 

N. armatum, and Microchaete humilis (Hook. f.) 

Sch. Bip in the synonymy of Nardophyllum humile (Hook. f.) A. Gray.

1905. Macloskie makes the combination Nardophyllum 

staehelinoides (DC.) Macloskie. However, the 

specimen which he refers to this new combination was collected in Santa Cruz (Argentina) and 

does not belong to the taxon coined by Candolle 

(1838), Dolichogyne staehelinoides.

1918. Hauman describes Nardophyllum bracteolatum 

based on material from Mendoza (Argentina).

1954. Cabrera (1954a) writes the most complete generic 

revision of Nardophyllum. In this important 

contribution, he establishes several synonymies, 

makes new combinations and excludes species, 
number 92 • 6 3

drastically reducing the number of valid names in 

Nardophyllum from ~15 to only 7: Nardophyllum armatum (Wedd.) Reiche, N. bracteolatum 

Hauman, N. bryoides (Lam.) Cabrera, N. chiliotrichioides (J. Rémy) A. Gray, N. genistoides 

(Phil.) A. Gray, N. lanatum (Meyen) Cabrera, and 

N. obtusifolium Hook. and Arn.

1993. Nesom (1993b) places the monotypic genera Aylacophora and Paleaepappus in the synonymy of 

Nardophyllum, coining the combinations Nardophyllum deserticola and Nardophyllum patagonicum. Nesom also reinstates Philippi’s species 

Nardophyllum scoparium after its exclusion from 

the genus by Cabrera (1954a).

2001. Bonifacino and Sancho reinstate monotypic Aylacophora and Paleaepappus, returning to Cabrera’s 

defi nition of Nardophyllum.

2004. Bonifacino and Sancho exclude Nardophyllum scoparium from Nardophyllum, placing this latter 

species in monotypic Guynesomia Bonif. and Sancho, coining the name Guynesomia scoparia (Phil.) 

Bonif. and Sancho.

2005. Bonifacino points out differences between Argentinean and Chilean populations assigned to 

Nardophyllum lanatum, describing new species 

Nardophyllum cabrerae to refer to the populations on the eastern side of the Andes. In the 

same publication, he places Nardophyllum obtusifolium and Nardophyllum bracteolatum in the 

synonymy of Nardophyllum bryoides and N. chiliotrichioides respectively.

Finally, in this treatment, and as a result of recent molecular evidence (Bonifacino and Funk, unpublished 

data) but also based on morphological grounds, 

the composition of Nardophyllum is reconsidered. 

Nardophyllum is composed of fi ve species: Nardophyllum chiliotrichioides, N. lanatum, N. bryoides, 

N. genistoides, and N. patagonicum (see discussion under N. patagonicum and N. genistoides for 

evidence supporting their inclusion, and discussion 

under Katinasia cabrereae (≡ Nardophyllum cabrerae) for evidence supporting its exclusion).

Nardophyllum (Hook. and Arn.) Hook. and Arn. in 

Hooker, Comp. Bot. Mag. 2: 44, 1836. Gochnatia subg. 

Nardophyllum Hook. and Arn. in Hooker, Comp. Bot. 

Mag. 1: 109, 1835. Type: Gochnatia revoluta D. Don 

ex Hook. and Arn. in Hooker, Comp. Bot. Mag. 1:109, 

1835 (= Barnadesia lanata Meyen, Reise Erde I: 347, 

1834 ≡ Nardophyllum lanatum (Meyen) Cabrera, Notas Mus. La Plata, Bot. 17(83): 63, 1954).

Dolichogyne DC., Prodr. (DC.) 7(1): 256, 1838. Type: Dolichogyne staehelinoides DC., Prodr. 7(1): 256, 1838 (= Barnadesia lanata Meyen, Reise Erde 

I: 347, 1834 ≡ Nardophyllum lanatum (Meyen) Cabrera, loc. cit.).

Chiliotrichum sect. Anactinia Hook. f., Bot. Antarct. Voy., Vol. 1, Fl. Antarct. 2: 304, 1846 [1847]. Anactinia (Hook. f.) J. Rémy in Gay, Fl. Chil. 4: 

8, 1849. Type: Chiliotrichum humile Hook. f., Bot. Antarct. Voy., Vol. 1, 

Fl. Antarct. 2: 304, 1846 [1847] (= Conyza bryoides Lam., Encycl. 2: 91, 

1786 ≡ Nardophyllum bryoides (Lam.) Cabrera, Notas Mus. La Plata, Bot. 

17(83): 61, 1954).

Paleaepappus Cabrera, Bol. Soc. Argent. Bot. 11(4): 273, 1969. Type: Paleaepappus patagonicus Cabrera, Bol. Soc. Argent. Bot. 11(4): 273, 1969 

(≡ Nardophyllum patagonicum (Cabrera) G. L. Nesom, Phytologia 75(5): 

358−365, 1993).

Shrubs cushion shaped to globose; branches dense to 

sparsely disposed, gland-dotted to densely tomentose and 

gland-dotted, 5- to 8-ribbed, sharp-ended or not. Leaves

dense to laxly arranged, persistent, alternate, narrowly obovate, elliptic or oblong, base slightly attenuated, petiole 

broadened towards the stem and slightly constricted above, 

apex obtuse, lax to densely tomentose on abaxial surface, 

gland-dotted on both surfaces, margins notably revolute, 

concolorous or discolorous, coriaceous or chartaceous. 

Capitula solitary, terminal, sessile to shortly pedunculate, 

homogamous, discoid. Involucres cylindrical, phyllaries 

arranged in 3 to 6 series; outer phyllaries with foliaceous 

apices, ovate to narrowly triangular, apex subacute to 

subulate, laxly tomentose and gland-dotted on abaxial 

surface, rarely with notable glandular vesicular capitate 

trichomes (in N. genistoides), coriaceous; inner phyllaries 

narrowly elliptic to narrowly ovate, apex acute to subulate, laxly tomentose and gland-dotted towards the apex 

on abaxial surface, papery. Receptacles fl at to convex, 

paleate to partially paleate, rarely naked (N. genistoides); 

paleae narrowly elliptic to narrowly ovate, not embracing 

the achenes (except in N. patagonicum), gland-dotted towards the apex, margins membranaceous. Florets 5 to 22, 

bisexual; corollas yellow, tubular, 5-lobed, tube gradually broadened or more or less abruptly broadened from 

mid-length, lobes triangular; anther thecae bases slightly 

sagittate, apical appendages ovate, antheropodia present 

not swollen; style bases swollen, style branches oblong to 

narrowly elliptic, apex acute to subacute, collecting hairs 

very short on upper 1/2 to 3/4. Achenes narrowly obovate, 

terete, 5- to 10-ribbed, laxly villose and gland-dotted, to 
6 4 • SMITHSONIAN CONTRIBUTIONS TO BOTANY

densely villose, carpopodium present. Pappus 2 (3)-seriate, 

composed of fl attened bristles (scales narrowly ovate in 

N. patagonicum), scabrid, sometimes barbellate towards 

the apex.

Etymology: From “Nardus” (genus belonging to 

Poaceae) and the Latin “phyllum” (leaf), denoting the resemblance of this genus with Nardus L.

Nardophyllum is the largest genus in the Chiliotrichum Group sensu stricto, with fi ve species distributed 

from southern San Juan in Argentina and neighboring 

areas in Chile to Tierra del Fuego throughout the southern 

Andes and Patagonia. Its habitats diverge greatly, ranging 

from sea level up to ~3,800 m occurring in high Andean, 

Patagonian, Monte, and subantarctic biogeographic provinces (Cabrera and Willink, 1973).

The genus is characterized by discoid heads, cylindrical involucres, paleate to partially paleate receptacles, 

and yellow corollas. Molecular evidence (Bonifacino and 

Funk, unpublished data) suggests that its closest relatives 

are Chiliotrichiopsis and Cabreraea, from which it differs 

mainly in the absence of ray fl orets (i.e: discoid heads) and 

the nature of the pappus.

Phylogenetic analysis based on both nuclear and chloroplast data (Bonifacino and Funk, unpublished data) 

fi rmly indicate the position of monotypic Paleaepappus deeply nested inside Nardophyllum. Although being 

a highly derived taxon, Paleaepappus is actually part of 

Nardophyllum, an idea earlier proposed by Nesom (1993a 

and 1993b), who actually coined the combination Nardophyllum patagonicum (Cabrera) Nesom. Nardophyllum 

armatum, a very interesting species from the Puna region 

in northern Argentina extending to neighboring areas in 

Chile and Bolivia is here excluded from Nardophyllum

and also from Chiliotrichum Group, an idea that is supported on both morphological and molecular grounds 

( Bonifacino and Funk, unpublished data; Bonifacino, 

2008). Nardophyllum genistoides, a rare species from 

central Chile, is maintained in Nardophyllum until more 

specimens are available to study the morphology of this 

species that it is only known from two collections made in 

the mid-1800s.

In light of molecular evidence, and after careful reevaluation of its diagnostic characteristics, Nardophyllum 

cabrerae, a recently described species from northern Patagonia (Bonifacino, 2005), is here excluded from Nardophyllum and placed in its own genus (see discussion under 

Katinasia Bonif.).

Nardophyllum bryoides (Lam.) Cabrera, Notas Mus. La 

Plata, Bot. 17(83): 61, 1954. Conyza bryoides Lam., Encycl. 2: 91, 1786. Baccharis bryoides (Lam.) Pers., Syn. 

Pl. 2: 425, 1807. Type: “[Chile] Magellan, côte des Patagons, baye Boucault, XII-1767, M. Commerson [s.n.]”

(holotype: P 226751; scanned image of holotype at P: 

LP!; isotype: P 222887; scanned image of isotype at 

P: LP!).

KEY TO THE SPECIES OF NARDOPHYLLUM (HOOK. AND ARN.) HOOK. AND ARN.

 1a. Leaves more or less laxly arranged, both macro and brachyblasts present; pappus elements fl at scales, narrowly ovate 

0.4−1.2 mm wide, smooth. (Restricted to Chubut, Argentina) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . N. patagonicum

 1b. Leaves more or less densely arranged, brachyblasts absent; pappus elements fl attened bristles, up to 0.1 mm wide, 

scabrid . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2

 2a. Leaves chartaceous, with evident glandular trichomes (biseriate vesicular capitate glandular trichomes); involucres 

~ 8 mm wide; receptacles naked. (Endemic to the Andes near Santiago, Chile) . . . . . . . . . . . . . . . . . . . . N. genistoides

 2b. Leaves coriaceous, with no such evident glandular trichomes (only biseriate vesicular glandular trichomes); involucres up 

to 5 mm wide; receptacles partially paleate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3

 3a. Branches tapering towards the end, sometimes ending in sharp tips, with notable ribs separated by narrow furrows; 

leaves laxly arranged. (Widespread along W and S Argentina and central Chile) . . . . . . . . . . . . . . N. chiliotrichioides

 3b. Branches not evidently tapering towards the end, never ending in sharp tips, with very narrow ribs and wide furrows, 

sometimes not visible due to abundant tomentose pubescence; leaves densely to more laxly disposed . . . . . . . . . . . . . 4

 4a. Ribs on stems not evident, usually hidden under dense tomentose pubescence; leaves densely disposed, notably discolorous, with a dense tomentose pubescence on abaxial surface only, sometimes with lax tomentose pubescence on adaxial 

side; involucres 9−13 × 4−5 mm; phyllary apices subacute to obtuse; fl orets 8 to 15 (−20). (Widespread in Patagonia, 

southern Chile and western and southern Argentina) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . N. bryoides

 4b. Ribs on stems narrow and noticeable; leaves laxly disposed, more or less concolorous, with tomentose pubescence uniformly covering the branches and both surfaces of the leaves; involucres 10−15 (−17) × 2.5−4.5 mm; phyllary apices 

acute to subulate; fl orets 4 to 6. (Restricted to central Chile) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . N. lanatum
number 92 • 6 5

Nardophyllum obtusifolium Hook. and Arn., in Hooker, Comp. Bot. Mag. 

2: 44, 1836. Type: “[Argentina] Port Desire, C. Darwin 325” (holotype: 

CGE; isotype: K!).

Chiliotrichum humile Hook. f., Bot. Antarct. Voy., Vol. 1, Fl. Antarct. 2: 

304, 1846 [1847]. Nardophyllum humile (Hook. f.) A. Gray, Proc. Amer. 

Acad. Arts, 5: 123, 1862. Anactinia hookeri J. Rémy nom. illeg., in Gay, Fl. 

Chil. 4: 8, 1849. Microchaete humilis (Hook. f.) Sch. Bip., Flora 38(8): 121, 

1855. Type: “Port Gregory, Capt. King’s exp. 13” (lectotype: BM 608512!, 

designated here; isolectotypes: BM!, K!; paratype: “Cape Fairwarther 9”, 

BM 608514!; isoparatypes: BM!, K!).

Chiliotrichum kingii Hook. f., Bot. Antarct. Voy., Vol. 1, Fl. Antarct. 2: 

305, 1846 [1847]. Nardophyllum kingii (Hook. f.) A. Gray, Proc. Amer. 

Acad. Arts 5: 123, 1862. Type: “Patagonia, Capt. Middleton [s.n.]” 

(lecto type: K!, designated here; isolectotype: K!; syntype: “port St. Elena, 

Capt. King 2”, K!).

Chiliotrichum darwinii Hook. f., Bot. Antarct. Voy., Vol. 1, Fl. Antarct. 2: 

305, 1846 [1847]. Nardophyllum darwinii (Hook. and Arn.) A. Gray, Proc. 

Amer. Acad. Arts 5: 123, 1862. Type: “Port Desire, C. Darwin [s.n.]” (holotype: K!).

Nardophyllum parvifolium Phil., Anales Univ. Chile 87: 434, 1894. Dolichogyne parvifolia Phil. ex Kuntze pro. syn., Revis. Gen. Pl. 3(3): 130, 1898. 

Type: “[Chile] Pucaullu, Feb, 1887, Otto Ph[ilippi] [s.n.]” (holotype: SGO 

62035!; isotypes: 43809 SGO!, LP!, B, destroyed; photos of isotype at B: 

F 14799!, SGO!, US!).

Aster nardophyllum Kuntze, Revis. Gen. Pl. 3(3): 130, 1898. Nardophyllum nardophyllum (Kuntze) Macloskie nom. illeg., Rep. Princeton 

Univ. Exp. Patagonia, Botany, 8: 784, 1905. Type: “Aster Nardophyllum, Patagon. 50/3°, 1882/4, F. P. Moreno & Tonini [s.n.]” (lectotype: 

NY 162130!, designated here; isolectotype: US!; syntype: “Aster nardophyllum OK Patagonien, Sa. Cruz, 1891/2, legit Beaufi ls 604”, NY 

162129!).

(FIGURES 29, 30, 56, 57)

Shrubs 5−100 cm tall, cushion shaped to globose;

branches densely to sparsely disposed, ascending, (in dense 

cushion shaped individuals the branches creeping along 

the soil, ascending distally), gland-dotted to densely tomentose and gland-dotted, 5-ribbed (sometimes ribs concealed by dense tomentum), internodes very short up to 

2 (−4) mm long. Leaves densely arranged, persistent, alternate, patent to slightly ascendant, (1−) 2−9 × 1−1.5 mm, 

narrowly elliptic to narrowly obovate, base slightly attenuated, petiole broadened towards the stem and slightly 

constricted above, apex obtuse, margins notably revolute, 

gland-dotted to tomentose and gland-dotted on adaxial 

surface, densely tomentose and gland-dotted on abaxial 

surface, discolorous, coriaceous. Capitula solitary, terminal, sessile to shortly pedunculate, peduncles up to 4 mm 

long, homogamous, discoid. Involucres 9−13 × 4−5 mm, 

cylindrical, phyllaries arranged in 4 or 5 series; outer phyllaries with foliaceous apices, 2.5−6 × 1.5−2.5 mm, ovate 

to elliptic, with a notorious longitudinal mid-ridge towards the base, apex acute to subacute, laxly tomentose 

and gland-dotted on abaxial surface, coriaceous, margins 

membranaceous, fi mbriate towards the apex; inner phyllaries 6−12 × 1−2 mm, narrowly elliptic, apex subacute, 

gland-dotted towards the apex on abaxial surface, papery, 

margins membranaceous, fi mbriate towards the apex. Receptacles fl at to slightly convex, partially paleate; paleae 1 

to 6, 8−13 × 0.5−1 mm, linear to oblong, boat-shaped, not 

embracing the achenes, apex acute, gland-dotted towards 

the apex, margins membranaceous, fi mbriate. Florets 8 

to 15 (−20), bisexual; corollas yellow, tubular, 5-lobed, 

gradually broadened towards the apex, tube 4.5−9 mm 

long, lobes 1.2−1.7 × 0.5−0.8 mm, triangular, gland-dotted abaxially; anther thecae 2.5−3 mm long, bases slightly 

sagittate, apical appendages 0.5−0.7 × 0.2−0.3 mm, triangular-ovate, antheropodium present, not swollen; 

style shafts 3.5−7 mm long, base swollen, style branches 

3.5−5 mm long, narrowly elliptic to oblong, subacute, 

collecting hairs very short on upper 3/5 to 2/3. Achenes 

4−6.5 mm long, narrowly obovate, terete, 5- to 7-ribbed, 

gland-dotted, sparsely villose, carpopodium present. Pappus (4.5−) 6−8 mm long, 2-seriate, composed of 40 to 70 

fl attened bristles, up to 0.1 mm wide, some fused among 

each other at the base, outer series slightly shorter, scabrid, inner series barbellate towards the apex.

Vernacular names: “Leña de pobre” (Hicken s.n., 

SI 1862); “Mata torcida” (León et al., 1998).

Phenology: Flowering specimens have been collected from October to February.

Distribution: Argentina, broadly distributed 

through Río Negro, Nequen, Chubut, Santa Cruz and 

Tierra del Fuego. In Chile, it is common in Región de Aisén 

and Región de Magallanes (Figure 30).

Habitat: It grows from sea level up to 1,200 m. 

Nardophyllum bryoides is a common species in both Andean and extra-Andean Patagonia provinces (Cabrera and 

Willink, 1973), constituting in certain areas the dominant 

species of the shrub stratum, like in SW Santa Cruz, Argentina, where it occurs associated with Festuca pallescens 

(St.-Yves) Parodi (León et al. 1998; Bonifacino, 2005). Further, the arid steppe that covers most of the extra-Andean 

Patagonia is characterized by the abundance of Nassauvia glomerulosa D. Don and Nassauvia ulicina Macloskie, 

which depending on the particular area are associated 
6 6 • SMITHSONIAN CONTRIBUTIONS TO BOTANY

FIGURE 29. Nardophyllum bryoides (Lam.) Cabrera. A. Detail of a branch. B. Habit. C. Leaves, adaxial view (left), abaxial view (right). 

D. Involucre. E. Outer to inner phyllaries (from left to right). F. Receptacular paleae, dorsal view (left), lateral view (right). G. Achene with 

pappus. H. Detail of pappus bristles (left, from inner series; right, from outer series). I. Style. J. Corolla (A−J, Bonifacino & Donato 699, LP; 

Figure 29B by Alice Tangerini).
number 92 • 6 7

with variable amounts of Nardophyllum bryoides (Lam.) 

Cabrera, Ephedra frustillata Miers, Chuquiraga aurea 

Skottsb., Brachyclados caespitosus Speg., Tetraglochin 

caespitosum Phil., etc. (Roig, 1998; Bonifacino, 2005). 

Bonifacino (2005) comments on the variable habitat of this 

species, and hypothesizes about the apparent association 

of the dense cushion shaped habit with areas with notable 

water and temperature stresses (high altitude and coastal 

areas), and the more spreading branching pattern whit less 

stressful conditions (interior of Patagonian steppe).

FIGURE 30. Geographic distribution of Nardophyllum bryoides (Lam.) Cabrera.
6 8 • SMITHSONIAN CONTRIBUTIONS TO BOTANY

Uses: Cultivated in the United Kingdom as a species suitable for rock gardens.

Etymology: The specifi c epithet refers to the resemblance of the plant (especially in pulvinate populations) 

with Bryophytes.

Note 1: In the protologue of Chiliotrichum humile,

Hooker (1846) mentions “Strait of Magalhaens [sic], Cape 

Gregory; Capt. King”. Additionally but apart from the diagnosis, Hooker also points out that the species is also present 

at “Cape Fairweather on the east cost of Patagonia”. I’ve 

studied six specimens from BM mounted on the same sheet 

with the headline “Strait of Magellan, Anderson in King’s 

voyage”, with the following individual labels 1) “Cape 

Gregory, 13”, 2) “Port Gregory, Capt. King’s exped., 

13. Herb. John Miers, bequeathed 1879”, 3) “Cape Fairweather, 9”, 4) “Cape Fairweather, Capt. King’s exped., 

13. Herb. John Miers, bequeathed 1879”, 5) “Nardophyllum humile, Magallanes, 1800, Temperate South America, 

comm. F. Philippi 1904”, and 6) “Port. St. Elena, 34. Collected by James Anderson on Capt. P. P. King’s Voyage 

to South America in H. M. S. Adventure” 1826−1830”. 

Further, I’ve studied another specimen at K with no label 

but with the following annotation on the sheet: “Voyage 

of the Beagle 1826−1830, Cape Fairweather, E Patagonia 

and Port Gregory, Capt. King”. All these materials match 

the original diagnosis. I select specimen 1(BM 608512) as 

lectotype since it is the one that is more in agreement with 

the diagnosis. Specimen 2 (also holding BM 608512) apparently was formerly in a different herbarium and it is 

considered a duplicate of the lectotype designated here. 

Specimens 3 and 4 (BM 608514) are considered paratypes 

since they were mentioned apart from the diagnosis in the 

original publication. Specimens 5 (BM 608513) and 6 (BM 

608515) correspond with material not directly related to 

the original description. As regards the material deposited 

at K, I consider it to be a set of mixed isolectotype and isoparatype mounted on the same sheet.

Note 2: Regarding the type of Chiliotrichum kingii Hook. f., I’ve studied four specimens deposited at K, all 

of which match the original diagnosis and the labels have 

the same localities that appear on the protologue. I select 

the specimen with label “Patagonia, Capt. Middleton s.n.” 

as lectotype since it is the more complete and with more 

abundant reproductive material. Additionally, two sheets 

deposited at K with material assigned to King and collected 

in St. Elena, one with “2” and the other with “34” written 

on the labels, are identifi ed as syntypes.

Note 3: In the protologue of Chiliotrichum darwinii Hook. f., J. D. Hooker cites a specimen from “Patagonia; 

Port Desire, C. Darwin”. I’ve studied a specimen originally 

belonging to Hooker’s personal herbarium, deposited at K 

with no label but with annotations made by K staff “Port 

Desire, Darwin”. The specimen match the original diagnosis and is considered here as the holotype.

Note 4: Kuntze (1898) mentions at least two specimens in the protologue of Aster nardophyllum: “490; 604 

Moreno and Tonini, Beaufi ls”. I’ve studied two specimens 

deposited at NY, one with the label “Aster Nardophyllum, 

Patagon. 50/3°, 1882/4, F. P. Moreno and Tonini [s.n.]”,

and the other “Aster nardophyllum OK Patagonien, Sa. 

Cruz, 1891/2, legit Beaufi ls 604”. Both specimens match 

the original diagnosis, however the numeration seems to 

have been somehow misplaced in the publication, since 

number “604” appears in the Beaufi ls specimen, while in 

Moreno and Tonini specimen there is no collection number on the label. The most complete specimen is the one 

collected by Moreno and Tonini, and it is here selected as 

the lectotype.

Note 5: Bonifacino (2005), when transferring 

Nardophyllum obtusifolium to the synonymy of Nardophyllum bryoides commented: “In his revision of Nardophyllum, Cabrera (1954a) stated that the only differences 

between N. bryoides and N. obtusifolium were: (1) habit, 

plants dwarf, cushion-shaped (N. bryoides), versus plants 

taller, non-cushion-shaped but frequently forming hemispheric shrubs (N. obtusifolium), and (2) receptacle paleate (N. bryoides) versus epaleate (N. obtusifolium). Moore 

(1983) pointed out that “Fuegian plants vary from dense 

cushions to rather lax mats, apparently dependent upon soil 

moisture and exposure, and perhaps the intensity of grazing, while receptacular bracts may be present or absent in 

adjacent plants, clearly showing no certainty of the need to 

keep such species as two distinct taxa”. My observations 

have led me to consider Nardophyllum obtusifolium and 

Nardophyllum bryoides as morphological extremes of a 

polymorphic species distributed throughout a large area 

characterized by very diverse habitats. The habit varies 

from hemispheric shrubs with branches laxly arranged to 

dwarf shrubs with branches densely disposed and even appressed to the soil. This trend towards plants with a denser 

branching pattern is particularly marked along the altitude 

gradient. Plants at high altitude exhibit denser branching, 

although as explained above, dwarf-looking plants also 

occur in open areas of the southernmost tip of the Patagonian scrub. The stem and leaf indument is extremely 

variable, from entirely covered with aseptate-fl agellate and 

biseriate vesicular glandular trichomes, which render a 

whitish tomentose and gland-dotted nature to both organs, 

or entirely glanduliferous (only biseriate vesicular glandular 

trichomes), even in individuals growing in close proximity. 
number 92 • 6 9

The receptacles are partially paleate, both in dwarf-cushion 

shaped individuals and hemispheric shrubs.”

Note 6: In dried herbarium specimens, the involucre usually adopts an obconic shape.

Additional specimens examined: ARGENTINA. NOT

DEFINED PROV. Laguna Amarga, 300 m, 14-I-1931, Donat 

366A (SI); Cañadon León, 30-XII-1955, Fischer s.n. (CONC 

19082). PROV. CHUBUT. Not defi ned Dpto. Meseta de Espinosa, 1929, Feruglio 100 (BA). Dpto. Ameghino. Estancia 

Lochiel, 30 km al W de Camarones, 25-X-1946, Soriano 

1995 (LP, SI). Dpto. Cushamen. Ruta 15, 2 km al oeste de 

intersección de Ruta 15 y Ruta 40, 42°23'37.1"S, 

71°09'54.3"W, 11-I-2002, Bonifacino et al. 403 (LP); íd., 

11-I-2002, Bonifacino et al. 405 (LP); al S de Leleque, 12-

XII-1981, Cabrera et al. 33076 (SI); Leleque, 13-I-1947, 

Soriano 2341 (LP). Dpto. Escalante. Ruta 27, entre Estancia 

Las Mellizas y Estancia La Guillermina, 45°10'34.4"S, 

67°57'51.4"W, 14-I-2002, Bonifacino et al. 433 (LP); Manantiales Behr, 26-I-1932, Castellanos s.n. (BA 6272); Pampa 

del Castillo, 15-XII-1929, Feruglio 48 (BA); Comodoro 

Rivadavia, Alta Pampa del Castillo, 17-XII-1952, Ruiz Leal 

14686 (LP). Dpto. Futaleufú. Esquel, XI-1940, Kühne mann 

s.n. (BA 37419); íd., 5-XII-1941, Kühnemann 592 (BA); íd., 

2-IV-1946, Scolnik 306 (LP); íd., 30-I-1945, Soriano 1409

(LP, SI). Dpto. Languiñeo. 10 km al E de Tecka, 12-XII1981, Cabrera et al. 33115 (SI); Región Río Corcovado, 

43°S, 71°W, 5-I-1901, Illín 203 (CORD, SI); Quinta experimental Río Corcovado, s. date, Illin 98 (CORD). Dpto. 

Mártires. Valle de las Plumas, s.date, Hosseus 96 (CORD). 

Dpto. Paso de Los Indios. R 24 a 50 km S de Paso de Los 

Indios, desvío a 23 km al SW, Estancia Laguna Blanca, 

44°28'S, 69°33'W, 30-XI-1976, Arroyo et al. 160 (SI); Ruta 

24, al S de Paso de Los Indios, 760 m, 44°21'32"S, 

69°22'56.7"W, 10-XII-2006, Bonifacino et al. 2966 (US); 

Ruta Prov 53, 44°05'48.2"S, 67°58'15.2"W, 26-XI-2002,

Bonifacino & Donato 699 (LP). Dpto. Río Senguerr. Valle 

del Lago Blanco, 1901, Koslowsky s.n. (BA 5138); Valle de 

la Laguna Blanca, 45°52'S, 71°15'W, 15-II-1901, Koslowsky s.n. (BA 5132); íd., 15-XII-1901, Koslowsky 174

(SI); 60 km al S de Alto Río Senguerr, Estancia Pepita, 13-II1947, Soriano 2585 (LP); 60 km al S de Alto Río Senguerr, 

21-I-1948, Soriano 3175 (LP, SI). Dpto. Sarmiento. Ruta 

24, 91 km al norte de Sarmiento, laderas al oesta de Laguna 

Seca, 44°53'17.3"S, 69°12'51.6"W, 15-I-2002, Bonifacino 

et al. 443 (LP); Ruta Prov 24, Laguna Palacios, cerros al N, 

44°44'28.4"S, 69°7'13.0"W, 27-XI-2002, Bonifacino & 

Donato 709 (LP); Ruta Prov 24, Cerros al W de Laguna 

Seca, 44°53'10.9"S, 69°13'25.6"W, 27-XI-2002, Bonifacino & Donato 718 (LP); Bajo del Avestruz, 45°15'29.2"S, 

68°58'1.6"W, 28-XI-2002, Bonifacino & Donato 726 (LP); 

Ruta Prov 23, 45°10'39.5"S, 69°15'52.4"W, 28-XI-2002, 

Bonifacino & Donato 742 (LP); Margen SW Lago Musters, 

15-XII-1981, Cabrera et al. 33205 (LP, SI); Sierra de San 

Bernardo, 15-XII-1981, Cabrera et al. 33221 (SI); NNW 

del Lago Musters, alrededores de la Laguna Colorada, 28-

XII-1939, Feruglio s.n. (BA 34784); NNW del Lago Musters, alrededores del puesto B. Crespo, 15-I-1940, Feruglio 

s.n. (BA 34783). PROV. NEUQUÉN. Not defi ned Dpto. Charahuilla, Arroyo Lapa, 13-II-1939, Chicchi 123 (LP). Dpto. 

Aluminé. Rahue, 17-II-1942, Perez Moreau s.n. (BA 47267). 

Dpto. Catan Lil. Casa de Lata, I-1972, Cabrera 21886 

(CONC, LP); Cumbre de la Bajada de Rahue, 20-I-1948, 

Dawson & Schwabe 2099 (BAA, CONC); Espinazo del 

Zorro, 17-II-1941, Perez Moreau s.n. (BA 47266); Cuesta 

de Rahue, II-1942, Perez Moreau s.n. (BA 48796); camino 

Aluminé-Las Coloradas, más adelante del puente sobre el 

Río Picún Leufú, 1485 m, 6-I-1970, Ruiz Leal 26764 (LP). 

Dpto. Collon Curá. Piedra Pintada, II-1940, Castagnet 148 

(LP). Dpto. Huiliches. Ruta 23, entrada a Reserva Indígena 

Aucapan, 39°41'38.4"S, 70°59'51.3"W, 7-I-2002, Bonifacino et al. 355 (LP). Dpto. Lacar. Parque Nacional Nahuel 

Huapí, estancia Chacabuco, 800 m, 26-I-1946, Boelcke 

1939 (BAA, LP). Dpto. Los Lagos. Rincón Grande, 1,000 

m, 22-XII-1940, Neumeyer 366 (LP). Dpto. Minas. Cordillera del Viento, Arroyo Piedra Blanca, 15-III-1912, Hicken 

38 (SI). Dpto. Zapala. Parque Nacional Laguna Blanca, 33 

km al SW de Zapala, II-1958, Perrone s.n. (BA 58874); Laguna Blanca, 9-I-1966, Ruiz Leal 24451 (LP). PROV. RÍO

NEGRO. Dpto. Ñorquinco. Cerca de Río Chico, hacia 

Ñorquinco, 11-XII-1981, Cabrera et al. 33063 (SI). Dpto. 

Pilcaniyeu. Pilcaniyeu, Estancia Rayhman, 26-I-1951, 

Boelcke 4473 (BAA, LP); Cerro Leones, al E de San Carlos 

de Bariloche, 41°05'S, 71°08'W, 21-I-2000, Bonifacino et 

al. 265 (LP, MVFA); Cerro Leones, al E de San Carlos de 

Bariloche, 1-II-1934, Burkart 6225 (BA); Comallo, 8-XII1938; Cabrera 4848 (F, LP); Camino a Ñirihuau, 16-I-1935, 

Cabrera & Job 353 (LP); Río Pichi Leufú, 5 km más abajo 

del camino de Bariloche a Pilcaniyeu, IV-1914, Hosseus 

1217 (CORD); 21 km de Pilcaniyeu, 27-I-1944, Nicora 

3661 (SI); 6 km de Pilcaniyeu camino a Bariloche, 28-I1944, Nicora 3681 (SI). PROV. SANTA CRUZ. Not defi ned 

Dpto. Río Santa Cruz, XII-1896, Hatcher s.n. (NY). Dpto. 

Corpen Aike. Ruta 3, 20 km al N de Piedra Buena, 3-XII1971, Boelcke et al. 15339 (BAA, LP); al S de Puerto San 

Julián, 159 m, 49°40'27"S, 68°14.5'55.8"W, 16-XII-2006, 

Bonifacino et al. 3037 (US); ~24 km S of Comandante Piedrabuena, road to Río Gallegos, 60 m, 1-I-1939, Eyerdam 

et al. 24014 (K, SI). Dpto. Deseado. Colonia Gral. Las 

Heras, 19-I-1914, s. leg. s.n. (SI); Puerto Deseado, 24-I1896, Alboff s.n. (CORD); Deseado y Golfo San Jorge, 
7 0 • SMITHSONIAN CONTRIBUTIONS TO BOTANY

1899–1900, Ameghino 145 (BA); entre Tellier y San Julián 

(km 108/109), a aprox. 39 km al S de Río Deseado, 18-XII1979, Bernardello & Figueroa 336 (CORD); Ruta Prov. 

281, km 14, alrededores de la gruta de Lourdes, 8-I-1967, 

Boelcke 12143 (BAA, SI); Puerto Deseado, 26-III-1917, 

Bonarelli s.n. (BA 5136); Ruta prov 12, 46°51'54.8"S, 

68°3'11.2"W, 2-XII-2002, Bonifacino & Donato 763 (LP); 

Ruta prov 12, 46°53'41.5"S, 68°11'7.1"W, 2-XII-2002, 

Bonifacino & Donato 776 (LP); Jaramillo a Cerro Blanco, 

Ruta 281, 8-XII-1970, Crepo & Troncoso 1685 (SI); Tehuelches, 300 m, 15-XII-1928, Donat 126 (F, NY); Puerto 

Deseado, 13-I-1914, Hicken & Hauman 144 (SI); Puerto 

Deseado, 21-XI-1945, O’ Donell 3600 (F); Holdich a Colonia Las Heras, 11-II-1936, Scott & Birabén 6 (LP). Dpto. 

Güer Aike. Estancia Stag River, 51°34'S, 72°00'W, 670 m, 

2-II-1978, Ambrosetti & Mendez s.n. (BAB); Estancia Las 

Viscachas, Laguna Viscachas, ladera SSE de la laguna, 

50°43'S, 71°58'W, 1,200 m, 28-I-1977, Arroyo et al. 526A

(SI); Estancia Cabo Buen Tiempo, 51°35'S, 69°06'W, 3-

XII-1975, Arroyo et al. s.n. (SI); Río Gallegos, Estancia La 

Angélica, 51°27'S, 69°07'W, 4-XII-1975, Arroyo et al. s.n.

(BAB); Estancia La Verdadera Argentina, S del Co León, 

50°53'S, 72°13'W, 16-I-1977, Arroyo et al. s.n. (SI); Estancia La Verdadera Argentina, Cerro de la Virgen, laderas y 

fi lo N, 50°50'S, 72°14'W, 525 m, 17-I-1977, Arroyo et al. 

s.n. (SI); Estancia La Verdadera Argentina, Cerro de la Virgen, 50°49'S, 72°14'W, 900 m, 18-I-1977, Arroyo et al. s.n.

(SI); Estancia Las Viscachas, Cerro Viscachas, 50°46'S, 

72°01'W, 850 m, 25-I-1977, Arroyo et al. s.n. (SI); Estancia 

Las Viscachas, Cerro Viscachas, en el fi lo, 50°46'S, 

72°01"W, 850 m, 25-I-1977, Arroyo et al. s.n. (SI); Estancia Las Viscachas, Cerro Viscachas, lomada a 2 km de Laguna Viscachas, 50°44'S, 71°58'W, 23-I-1977, Arroyo et 

al. s.n. (SI); Estancia Las Viscachas, ladera SSE de la Laguna, 50°43'S, 71°58'W, 1,200 m, 28-I-1977, Arroyo et 

al. s.n. (SI); Estancia Las Viscachas, lomada a 2 km de 

Laguna Viscachas, 50°44'S, 71°58'W, 23-I-1977, Arroyo 

et al. s.n. (BAB); Estancia Stag River, ladera superior S y 

fi lo de meseta Latorre, 51°34'S, 72°01'W, 750 m, 15-II1978, Boelcke et al. s.n. (BAB); Ruta 40, a 25 km de El 

Cerrito, 50°45'S, 71°27'W, 17-I-1967, Boelcke 12499

(BAA, BAB, SI); Próximo a intersección Rutas 3 y 40, 43 

m, 51°36'59"S, 69°37'36.9"W, 16-XII-2006, Bonifacino 

et al. 3031 (US); Río Gallegos, 5-XII-1932, Castellanos s.

n. (BA 7938, LP); Estancia Cabo Buen Tiempo, 2-XII1975, Correa et al. 6563 (BAB); Estancia Las Vegas, Río 

Coyle, 18-I-1917, Dauber 150 (BAA, CORD); 26 km SW 

of Río Gallegos, road to Punta Arenas, 30 m, 4-I-1939, 

Eyerdam et al. 24083 (SI); Estancia Flora, ruta prov. 7, a 

2.5 km al N del cruce con ruta prov. 2, 50°59'S, 70°47'W, 

80 m, 2-II-1995, Fortunato & Elechosa 4928 (NY); Cancha Carrera, 240 m, 26-I-1976, Gentili 397 (LP); Río Gallegos airport, 26-X-1969, Goodall 2187 (herb. Goodall); 

Estancia Cabo Buen Tiempo, Punta El Gancho, 7-XII1975, Goodall 4903 (herb. Goodall); Estancia Cabo Buen 

Tiempo, Río Gallegos in front of Isla Deseada, 4-XII1975, Goodall 4964 (herb. Goodall); Estancia Los Pozos, 

hills N of cliffs on N side of Río Gallegos, 5-XII-1975, 

Goodall 4981 (herb. Goodall); Sud de Río Gallegos, 22-I1949, Grondona 2067 (BAB); 10 km camino Río Gallegos 

a Río Turbio, 23-I-1949, Grondona 2159 (LP); Estancia 

Monte dinero, Cabo Vírgenes, XI-1974, Molina 1 (LP); 

Sección San Antonio, 51°24'S, 71°34'W, 19-I-1978, Roig 

et al. s.n. (SI); Estancia Stag River, 25-I-1958, Tweedie 

298 (K); íd., 25-I-1958, Tweedie 297 (K); íd., 14-I-1958,

Tweedie 252 (LP). Dpto. Lago Argentino. Alrededores de 

Co Fitz Roy, II-1932, Agostini B-46 (LP); Lago San Martín, Estancia La Federica, 48°54'S, 72°18'W, 12-II-1975,

Boelcke et al. 16363 (LP); Lago Argentino, Península Magallanes, Estancia Co Buenos Aires, 900−1,000 m, 17-II1975, Boelcke et al. 16498 (BAB, LP); Lago San Martin, 

Brazo de la Lancha, 48°59'26.9"S, 72°13'26.0"W, 19-I2002, Bonifacino et al. 514 (LP); íd., 19-I-2002, Bonifacino et al. 515 (LP); Parque Nacional Los Glaciares, Fitz 

Roy, 14-II-1975, Cabrera et al. 25864 (LP, SI); entrada a 

Parque Nacional Los Glaciares, 120 m, 3-II-1995, Fortunato & Elechosa 4935 (BAB); South shore of Lago Argentino, from 30−50 km westward from Santa Cruz river, 

XI-1907, Furlong 153 (NY); Lago Argentino, 1904−1905, 

Koslowsky 26 (CORD); íd., 1904−1905, Koslowsky 29

(CORD); ca. 90 km da Monte León, lungo la strada per 

Calafate, 12-I-1974, Pichi Sermolli & Bizzarri 7429 (K); 

Burmeister Penninsula, Lake Argentino, I-1900, Prichard 

s.n. (BM, NY). Dpto. Lago Buenos Aires. Ruta nacional 

520, a 21 km al W de Perito Moreno, 46°47'S, 71°09'W, 

4-II-1975, Boelcke et al. 16090 (BAA, LP); al S de Perito 

Moreno, 597 m, 47°3'19.9"S, 70°46'27.3"W, 11-XII2006, Bonifacino et al. 2984 (US); Los Antiguos camino a 

Perito Moreno, 24-XI-1965, Correa & Nicora 3638 (BAA, 

LP); Lago Buenos Aires, III-1928, Guiñazú s.n. (BA 

31/1725); Lago San Martín, 8-XII-1914, Hicken s.n. (SI 

1862); entre Arroyo Telca y Perito Moreno, 640 m, 12-

XII-1965, Ruiz Leal 24219 (LP); Patagonia andina, Lago 

Buenos Aires, 12-XII-1908, Skottsberg s.n. (SGO 58893). 

Dpto. Magallanes. San Julián, 15-XII-1926, Blake 71 (K); 

íd., II-1933, Blake 219 (K); íd., 10-XII-1944, Blake 378

(SI); íd., 10-XII-1944, Blake 382 (SI); íd., 3-XII-1944; 

Blake 438 (SI); San Julián, 25-III-1917, Bonarelli s.n. (BA 

5133); íd., XII-1917, Bonarelli s.n. (BA 5139); Puerto San 

Julián, 1915, Carette s.n. (LP); 50 km N San Julián, road 

to Puerto Deseado, 30-XII-1938, Eyerdam et al. 23896

(K, SI); Puerto San Julián, 19-I-1949, Grondona 2026
number 92 • 7 1

(LP); San Julián, 24-XI-1945, O’ Donell 3709 (BM, F, K).

Dpto. Río Chico. Lago Pueyrredón, cerros al norte de Río 

Oro, 47°24'25.4"S, 71°59'30.4"W, 17-I-2002, Bonifacino et al. 464 (LP); Parque Nacional F. P. Moreno, Estancia La Oriental, Lago Belgrano, 29-I-1971, Capurro 1587

(BA); entre Estancia Tres Cerritos y Lago Cardiel, 9-XII1965, Ruiz Leal 24115 (LP); entre Las Horuqetas y Río 

Olnie, 1,000 m, 11-XII-1965, Ruiz Leal 24245 (LP); 

Parque Nacional F. P. Moreno, Estancia La Oriental, 12-

II-1973, Rumball s.n. (BA 69248); íd., 8-II-1973, Rumball 

s.n. (BA 69244); íd., 15-II-1973, Rumball s.n. (BA 69188); 

Lago Belgrano, Península de los Ciervos, 46°10'–48°15'S, 

71°–72°20'W, 800 m, 1-III-1903, Von Platen & Greiner 

53 (SI). PROV. TIERRA DEL FUEGO. Dpto. Río Grande. Estancia Cullen, tall steep hill halfway between Argentine-Chilean limit and the Arroyo Alfa, 6-I-1971, Goodall 3110 

(BAB, herb. Goodall); Estancia Cullen, Puesto Beta along 

Arroyo Beta, near its mouth, 7-I-1971, Goodall 3244

(BAB, herb. Goodall); Estancia Cullen, beyond and N of 

Arroyo Alfa, 20-V-1972, Goodall 4299 (BAB, herb. 

Goodall); Estancia Cullen, Arroyo Beta, 52°44'S, 68°33'W, 

5-I-1972, Moore & Goodall 336 (herb. Goodall, K, LP); 

Estancia Cullen, hills to W of Río Alfa, 52°43'S, 68°36'W, 

5-I-1972, Moore & Goodall 354 (herb. Goodall, SI). 

Dpto. Ushuaia. Isla de los Estados, Canal de Beagle, verano-1902, Rodríguez 25 (BAB). CHILE. REG. DE AISÉN. 

Prov. Gral. Carrera. Camino de Cerro Castillo a Puerto 

Ibañez, 46°15'S, 72°00'W, 700 m, 15-III-1977, Schlegel 

6997 (CONC); Chile Chico, 46°33'S, 71°44'W, 20 m, 12-

III-1967, Seki 403 (CONC). REG. DE MAGALLANES. Prov. 

Magallanes. San Gregorio, 26-X-1968, Cekalovic s.n.

(LP); íd., 25-X-1968, Cekalovic s.n. (CONC 33903); íd., 

II-1954, Díaz s.n. (CONC 93942); Ruta 255, 120 km E of 

Punta Arenas, 2 km E of Bahia San Gregorio, 9-X-1971, 

Dudley et al. 84 (CONC, SGO); NW face of Mt. Aymond 

on Chile-Argentina Border, ca. 2 km on right side of Route 

255-N heading to Río Gallegos, 52°09-10'S, 69°25–29'W, 

9-X-1971, Dudley et al. 107 (herb. Goodall); 67 km E of 

Punta Arenas-Puerto Natales Road, just past entrance to 

Est. San Gregorio, 52°30'S, 70°00'W, 10 m, 26-XII-1994, 

Landrum & Landrum 8427 (CONC); San Gregorio, I1953, Magens s.n. (SGO); San Gregorio, Estrecho de Magallanes, I-1954, Magens s.n. (CONC 14817); Estancia 

Punta Delgada, ca. 180 km NE of Punta Arenas, Cañadón 

Grande, 30 km ENE of settlement, 18-III-1964, Moore 

1053 (K). Prov. Tierra del Fuego. 30 km al S de Manantiales, 2-XI-1973, Dollenz 78 (CONC); Bahia Lee, 52°52'S, 

70°16'W, 6-XI-1971, Moore 2339 (LP); Bahia Felipe, 23-

I-1952, Pfi ster s.n. (CONC 12264); Bahia Lee, 6-XI-1971, 

Pisano 3157 (CONC); Punta Espora, XII-1975, Pisano & 

Dollenz 4502 (CONC); Estrecho de Magallanes, I-1966, 

Tsujii 679 (CONC). Prov. Ultima Esperanza. Estancia 

Cerro Castillo, 30 km N Rincón Negro, 51°00'S, 72°16'W, 

400 m, 16-XII-1975, s. leg. s.n. (CONC 50915); Sierra del 

Toro, 51°10'S, 72°50'W, 700−750 m, 9-II-1992, Arroyo et 

al 92-39 (CONC); Sierra de los Baguales, Cerro Santa 

Lucía, 50°44'S, 72°20'W, 750 m, 6-I-1985, Arroyo 850092

(CONC); Cerro Donoso, sector Río de las Chinas, 50°44'S, 

72°31'W, 900 m, 9-II-1987, Arroyo et al. 870322 (CONC); 

Estancia Cerro Castillo, 51°11'S, 72°22'W, 700 m, 14-XII1975, Boelcke et al. s.n. (SI); Estancia Cerro Castillo, 

51°00'S, 72°16'W, 400 m, 16-XII-1975, Boelcke et al. s.n. 

(SI); Sierra de los Baguales, Estancia La Cumbre, Cerro Sin 

Nombre, 50°42'S, 72°22'W, 800 m, 18-XII-1975, Boelcke 

et al. s.n. (BAB, CONC, SI); Parque Nacional Torres del 

Paine, 1 km E of E end of Lago Nordenskjöld, 51°00'S, 

72°51'W, 100 m, 24-XI-1996, Elvebakk 96:667 (CONC); 

Sierra de los Baguales, Estancia La Cumbre, Campo de la 

Tropilla, 50°44'S, 72°51'W, 800 m, 15-XII-1986, Landero 

686 (CONC); Near Guardaparque’s house, Paine, 25-III1972, Leontsinis 23 (herb. Goodall); Torres del Paine, 

50°58'S, 73°02'W, 8-XII-1962, Matte s.n. (CONC 40418); 

Salto Grande del Paine, 22-XII-1969, Pisano 2330 (CONC, 

herb. Goodall, LP); Estancia La Cumbre, Sierra de los Baguales, 600 m, 18-III-1972, Pisano 3639 (CONC); Puesto 

Weber, Estancia Paine, 20-II-1974, Pisano 4300 (CONC); 

Estancia Guido, Cerro Guido, 700−900 m, 16-I-1952, 

Pfi ster & Ricardi s.n. (CONC 12155); Las Cumbres, Baguales, 550−850 m, 6-II-1962, Ricardi & Matthei 399 

(CONC); Angostura, cerca de Lago Sarmiento, 8-II-1962, 

Ricardi & Matthei 466 (CONC).

Nardophyllum chiliotrichioides (J. Rémy) A. Gray, Proc. 

Amer. Acad. Arts 5: 123, 1862. Dolichogyne chiliotrichioides J. Rémy, in Gay, Fl. Chile, 4: 104, 1849. Type: 

“Cordillere de Sotaqui, Chili, M. Cl. Gay [s.n.]” (holotype: P 222883, scanned image of holotype at P: LP!; 

isotypes: P 222884, F!, NY!; scanned image of isotype at 

P: LP!; photos of isotype at P: F 37660!, LP!, SI!).

Nardophyllum bracteolatum Hauman, Anales Soc. Ci. Argent. 86: 324, 

1918. Type: “[Argentina, Prov. Mendoza] Pedernal, Cordillera de San Carlos, 3,700 m, 26-III-1918, R. Sanzin 1810” (holotype: BRLU?; isotype: BA!, 

LP!; photo of holotype at BRLU: BA!).

(FIGURES 31, 32, 58, 59)

Shrubs 40−100 cm tall, globose; branches densely 

disposed, ascending, tapering towards the end, 8-ribbed, 

interrib areas densely tomentose, internodes (2−) 4−9 mm 

long. Leaves laxly arranged, persistent, alternate, 

(3−) 8−16 × (0.5−) 1−2 mm, narrowly elliptic to narrowly 
7 2 • SMITHSONIAN CONTRIBUTIONS TO BOTANY

FIGURE 31. Nardophyllum chiliotrichioides (J. Rémy) A. Gray. A. Habit. B. Leaves, adaxial view (left), abaxial view (right). C. Involucre. 

D. Outer to inner phyllaries (from left to right). E. Receptacular paleae, dorsal view (left), lateral view (right). F. Detail of pappus bristle. 

G. Achene with pappus. H. Style. I. Corolla (A−I, Bonifacino et al. 407, LP; Figure 31A by Alice Tangerini).
number 92 • 7 3

obovate, base slightly attenuated, petiole broadened towards the stem and constricted above, apex acute, recurved, margins notably revolute, densely tomentose and 

gland-dotted on abaxial side, gland-dotted on adaxial 

side, discolorous, coriaceous. Capitula solitary, terminal, 

pedunculate, peduncles 3−15 mm long, homogamous, discoid. Involucres 7−8 × 4−5 mm, cylindrical; phyllaries arranged in 3 to 4 series; outer phyllaries 3−6 × 0.7−1.8 mm, 

triangular-ovate, mid-ridge clearly marked, apex acute-subulate, gland-dotted on abaxial surface, coriaceous, margins membranaceous; inner phyllaries 6−7 × 1.3−2 mm, 

narrowly elliptic to narrowly ovate, apex acute-subulate, 

gland-dotted towards the apex on abaxial surface, papery, 

margins membranaceous. Receptacles fl at to slightly convex, partially paleate; paleae 8 to 12, 6−8.5 × 1.5 mm, 

narrowly ovate to ovate-triangular, boat-shaped, not 

embracing the achenes, apex acute, membranaceous, 

gland-dotted on adaxial surface, margins membranaceous. Florets 10 to 22, bisexual; corollas yellow, tubular, 

5-lobed, broadened more or less abruptly from midlength, tube 2.5−5 mm long, throat 3−5 mm long, lobes 

0.8−2 × 0.5−0.7 mm, triangular, gland-dotted abaxially; 

anther thecae 1.8−2.5 mm long, bases slightly sagittate, 

apical appendages 0.8−1.2 × 0.3 mm, triangular, antheropodium present, not swollen; style shafts 4−6 mm long, 

base swollen, style branches 4−5 mm long, narrowly elliptic to oblong, subacute, collecting hairs short on upper 

1/2. Achenes 3−6.5 mm long, narrowly obovate, terete, 

6- to 8-ribbed, sparse to densely villose and gland-dotted, 

carpopodium present. Pappus 6−8.5 (−10) mm long, 2-

seriate, composed of 55 to 70 fl attened bristles, up to 0.1 

mm wide, scabrid, outer bristles somewhat shorter.

Vernacular names: Not known.

Phenology: Flowering specimens have been collected from October to February. There seems to be a correlation between the time of blooming and the geographic 

distribution, with populations towards the north fl owering 

before the southern populations, and populations close to the 

Atlantic ocean fl owering before those close to the Andes.

Distribution: Argentina: very common in Chubut, 

but also present in Mendoza, Neuquén, San Juan, and 

Santa Cruz. Chile: present in Coquimbo and Metropolitana 

regions (Figure 32). The scattered distribution shown in 

Figure 32 probably refl ects an incomplete sampling rather 

than the actual distribution.

Habitat: Nardophyllum chiliotrichioides occurs 

from sea level to 3,800 m in San Juan and Mendoza provinces (Argentina) in the high Andean and Patagonian 

biogeographic provinces (Cabrera and Willink, 1973). In 

Chile it grows above 2,500 m. It is a common species in 

the patagonian steppe of Chubut where it occurs associated 

with Nassauvia axillaris D. Don, Nassauvia glomerulosa 

D. Don, Nassauvia ulicina Macloskie, and Junellia spp. In 

many areas it is the dominant species in the shrub stratum.

Uses: Not known.

Etymology: The specifi c epithet refers to the similarity of this species with Chiliotrichum, especially on the 

leaves and the habit.

Note 1: In the original publication of Dolichogyne 

chiliotrichioides, J. Rémy (1849) stated that the species 

was from “Cordilleras de Sotaqui” in Chile. I’ve studied 

images of two specimens deposited at P (P 222883 and P 

222884). Both specimens match perfectly the original diagnosis, however their labels are slightly different. One of the 

labels (specimen P 222883) reads “Cordilleras de Sotaqui”, 

while the other (specimen P 222884) reads “Chili, Prov. 

Coquimbo”. While both specimens are from Coquimbo, 

the locality on specimen P 222883 is exactly the same that 

appears in the original description and it is therefore identifi ed as the holotype, and specimen P 222884 as isotype. The 

photograph distributed by F as phototype of Dolichogyne 

chiliotrichioides corresponds with the isotype at P.

Note 2: According to Stafl eu and Cowan (1979) 

and the staff at BA, Hauman deposited most of his types 

at BRLU. I have studied a photograph at BA that apparently would correspond with a photo of the holotype of 

Nardophyllum bracteolatum Hauman, but since I have not 

received confi rmation of its presence at BRLU, I am leaving 

the location of the holotype at BRLU as probable. I have 

studied two other specimens, one at BA and the other at LP. 

Both specimens match the original diagnosis and the data 

that appear on the labels match those of the protologue, 

therefore, I identify them as isotypes. However, the label of 

the specimen deposited at LP has some inconsistencies in 

the collection data: the date and elevation are different (25-

III-1916, 3600 m). It is my opinion that these differences are 

due to errors when transcribing the label since both specimens have the same collector and collection number as well 

as the locality that appeared in the original description.

Note 3: Bonifacino (2005), when transferring Nardophyllum bracteolatum to the synonymy of Nardophyllum 

chiliotrichioides commented: “Nardophyllum chiliotrichioides is a highly polymorphic species occurring on both 

sides of the Andes Mountain Range in Chile (Coquimbo 

and Metropolitana regions) and in Argentina (Santa Cruz, 

Chubut, Neuquén, Mendoza, and San Juan provinces). 

Hauman (1918) described Nardophyllum bracteolatum

based on herbarium material from Mendoza (Argentina) 

and made explicit his doubts concerning the distinctness of his new species when compared to Nardophyllum 
7 4 • SMITHSONIAN CONTRIBUTIONS TO BOTANY

c hiliotrichioides [“Il existe dans l ́Herbier [BA] un fragment 

(Ameghino 205), ne presentant qu’un seúl capitule en mauvais état, mais qui me semble appartenir à une espèce extrêmement voisine, si pas identique, à la mienne”].” I have 

studied the specimen Ameghino 205 at BA and it perfectly 

corresponds with typical Nardophyllum chiliotrichioides

and matches the characteristics of southern populations. 

Bonifacino (2005) added: “Cabrera (1954a) pointed out 

the great similarity between Nardophyllum bracteolatum

and Nardophyllum chiliotrichioides, noting that the only 

FIGURE 32. Geographic distribution of Nardophyllum chiliotrichioides (J. Rémy) A. Gray.
number 92 • 7 5

difference between those species was the number of fl orets 

per capitulum. My observations, based on a large number of 

specimens collected throughout the whole geographic range 

of the species, indicate that the number of fl orets per capitulum in Nardophyllum chiliotrichioides and Nardophyllum bracteolatum populations overlaps, although northern 

populations (Nardophyllum bracteolatum) tend to exhibit 

larger numbers of fl orets per capitulum (15–22) than southern populations (Nardophyllum chiliotrichioides, 10–18 

fl orets per capitulum). Nardophyllum chiliotrichioides is a 

fairly common species in the Patagonian scrub of Chubut 

(Argentina) and is the dominant species in the shrub stratum of certain areas. In Mendoza and San Juan, the species 

has only been collected above 2,200 m; these populations 

are more similar to Chilean populations and exhibit leaves 

smaller than the populations from Chubut and Santa Cruz. 

The only difference between northern (central Chile, San 

Juan, and Mendoza in Argentina) and southern populations 

(Chubut and Santa Cruz in Argentina) lie in the branching 

pattern (branches form sharper angles in northern populations than in southern ones). According to my observations, 

there are no clear boundaries between the northern and 

southern populations, and therefore there is no evidence to 

support Nardophyllum bracteolatum and N. chiliotrichioides as distinct taxa. The latter name has priority prescribing the treatment of Nardophyllum bracteolatum as 

its heterotypic synonym”. Molecular evidence (Bonifacino 

and Funk, unpublished data) seems to suggest that material 

originally ascribed to N. bracteolatum would slightly differ 

from N. chiliotrichioides, but at the light of morphological 

evidence mentioned above, until more specimens are obtained from the Andes of Mendoza, I reaffi rm my earlier 

hypothesis (Bonifacino, 2005) of adopting a broad concept 

of this taxon, keeping N. bracteolatum in the synonymy of 

N. chiliotrichioides.

Additional specimens examined: ARGENTINA. PROV. 

CHUBUT. Not defi ned Dpto. Río Chico, aut-1900, Ameghino 

s.n. (LP); Travesía del Chubut, XII-1903, Illín s.n. (BAB). 

Dpto. Ameghino. Ameghino, 380 m, 24-XII-1967, Ruiz 

Leal 25672 (LP). Dpto. Cushamen. Ruta Prov. 12, entre 

Ruta Nac. 40 y Gualjaina, 42°43'50.8"S, 70°47'16.8"W, 

12-I-2002, Bonifacino et al. 407 (LP); 20 km al E de 

Cushamen, 31-XII-1947, Soriano 2786 (LP). Dpto. Escalante. 100 km N of Comodoro Rivadavia, 200 m, 23-

XII-1938, Eyerdam et al. 23776 (K, SI). Dpto. Florentino 

Ameghino. Próximo a Garayalde, 417 m, 44°38'29.1'S, 

66°34'32.1"W, 17-XII-2006, Bonifacino et al. 3046 (US); 

Ruta Prov. 29, entre Garayalde y Estancia La Chiquita, 

44°36'33.6"S, 66°52'21.3"W, 4-XII-2002, Bonifacino & 

Donato 788 (LP); Ruta Prov 31, entre Ruta Nac. 3 y Dique 

Ameghino, 44°03'21.2"S, 66°10'10.2"W, 4-XII-2002, 

Bonifacino & Donato 789 (LP); Ruta Prov 31, entre Ruta 

Nac. 3 y Dique Ameghino, 44°01'13.4"S, 66°09'36.0"W, 

4-XII-2002, Bonifacino & Donato 790 (LP); Ruta 3 a 

129 km al NW de Camarones, 15-I-1978, Romanzuk et 

al. 895 (BAB). Dpto. Gaiman. Ruta Prov 31, entre Ruta 

Nac. 3 y Dique Ameghino, 43°46'16.2"S, 66°17'3.8"W, 

4-XII-2002, Bonifacino & Donato 791 (LP); Ruta Nac. 

25, 43°35'45.2"S, 66°29'17.1"W, 5-XII-2002, Bonifacino 

& Donato 799 (LP). Dpto. Languiñeo. Río Corcovado, 

I-1904, Illín s.n. (BA 26041). Dpto. Martires. Ruta Prov 53, 

~ 1 km al SW de Intersección con Ruta 25, 43°46"28.1"S, 

67°29'26.2"W, 26-XI-2002, Bonifacino & Donato 679

(LP); Ruta Prov 53, ~1 km al SW de Intersección con 

Ruta 25, 43°46'28.1"S, 67°29'26.2"W, 26-XI-2002, Bonifacino & Donato 683 (LP); Ruta Prov 53, ca. Sierra Negra, 

43°59'24.0"S, 67°35'32.2"W, 26-XI-2002, Bonifacino & 

Donato 692 (LP); Ruta Prov. 48, 30 km al sur de intersección con Ruta Prov. 25, 43°58'17.0"S, 67°16'24.6"W, 13-

I-2002, Bonifacino et al. 425 (LP); Ruta Nac. 3, km 1661, 

44°44'30.1"S, 66°40'30.4"W, 30-I-2002, Bonifacino et 

al. 602 (LP); alta planicie cerca de la Estación Las Plumas, 

4-XI-1925, Hosseus 152 (CORD). Dpto. Paso de los Indios. Ruta Prov. 12, entre Paso Berwin y Ruta Prov. 25, 

43°42'30.8"S, 68°56'12.1"W, 12-I-2002, Bonifacino et al. 

419 (LP); Ruta Prov. 27, 3 km al sur de intersección con 

Ruta Prov. 29, 44°37'50.2"S, 67°50'00.0"W, 14-I-2002, 

Bonifacino et al. 430 (LP); Valle de los Altares, 19-XII1981, Cabrera et al. 33291 (LP, SI); El Sombrero, 6-I-1948, 

Krapovickas 3893 (BAB); a 5 km de Paso de los Indios, en 

el cruce con Yacimiento Condor, 19-XII-1993, Molina et 

al. 4670 (NY); entre Estancia Cañadón Grande 900 m y 

Paso de los Indios 520 m, 28-XII-1967, Ruiz Leal 25696

(LP); 23 km al E de El Sombrero, 6-I-1948, Soriano 2840

(LP). Dpto. Rawson. 10 km al S de Trelew, 18-XII-1981, 

Cabrera et al. 33268 (LP, SI); Trelew, 12 km al S, 7-XII1980, Castroviejo & Lopez 2313 (SI). Dpto. Sarmiento. 

Ruta Prov. 24, 91 km al norte de Sarmiento, laderas al oeste 

de Laguna Seca, 44°53'17.3"S, 69°12'51.6"W, 15-I-2002, 

Bonifacino et al. 447 (LP). PROV. MENDOZA. Dpto. San Carlos. Arroyo de la Quebrada Casa de Piedra, 17-I-1952, Serra 

77 (LP); Rincón de los Leones, 18-I-1941, Ruiz Leal 7212

(LP). Dpto. Tunuyán. Portillo, de Tunuyán, 33°37'17"S, 

69°35'57"W, 2,800 m, 27-I-2001, Simón & Bonifacino 

819 (LP); Portillo, Cuesta de los Afl igidos, 29-I-1934, Ruiz 

Leal 2052 (LP). PROV. NEUQUÉN. Dpto. Zapala. Alrededores 

de Cerro Lotena, 20-XI-1983, Correa et al. 8837 (BAB); 

Zapala, II-1920, Hauman s.n. (BA 5144). PROV. SANTA

CRUZ. Dpto. Deseado. Deseado, 1899-1900, Ameghino 

205 (BA). PROV. SAN JUAN. Dpto. Calingasta. Quebrada Los 
7 6 • SMITHSONIAN CONTRIBUTIONS TO BOTANY

Avestruces, W de Cerro Castaño, II-1960, Fabris & Marchionni 2354 (LP); Río Manrique a portezuelo de Potrerillos, 31°00'S, 69°50'W, ~3,000 m, 23-I-1991, Kiesling et al. 

7689 (SI); Río Mondaca (o Río Blanco o Patillos), Alojo 

de Mondaca, 30°50'S, 70°10'W, 2,900 m, 19-II-1990, 

Kiesling et al. 7539 (MO, SI). CHILE. REG. DE COQUIMBO. 

Prov. Limari. Cordillera San Miguel, 2,800 m, 12-I-1959, 

Jiles 3599 (CONC); Los Molles, 3,000 m, 17-I-1972, Zöllner 5567 (CONC). REG. METROPOLITANA. Not defi ned Prov.

Cordillera de las Arañas, I-1861, Lambert s.n. (SGO 43816). 

Prov. Area Metropolitana. Entre Farellones y Valle Verde, 

33°22'12.1"S, 70°16'44.2"W, 16-I-2003, Bonifacino & Romano 809 (LP); Fierro Carrera, 2,800−3,800 m, I-1930, 

Montero 1035 (CONC); Las Condes, XII-1948, Morales 

s.n. (CONC 93945); entre Farellones y La Parva, hasta 

2,700 m, 18-I-1980, Muñoz 1582 (SGO); La Parva, faldeo 

norte hacia el Cerro Colorado, 7-I-1979, Muñoz & Meza 

1399 (SGO); Río San Francisco, Fierro Carrera, ~3,000 m, 

I-1925, Weerderman 621 (F, K, LP, NY, SI). Prov. Cordillera. Parque Nacional El Morado, 33o49'S, 70o05'W, 14-

I-1991, Teillier et al. 2410 (MO).

Nardophyllum genistoides (Phil.) A. Gray, Proc. Amer. 

Acad. Arts 8: 636, 1873. Dolichogyne genistoides Phil., 

Linnaea 28: 738, 1856. Type: “[Chile] Dolichogyne 

genistoides Phil., Cordillera de Santiago, Febr. 1857, 

Germ. [s.n.]” (holotype: SGO 62028!; isotypes: BM 

608509!, LP!).

(FIGURES 33, 34, 60)

Shrubs 10−30 cm tall, branches more or less densely 

disposed, ascending, 8-ribbed, tomentose when young, 

later covered with notable glandular vesicular capitate 

trichomes, internodes short, up to 3 mm long. Leaves

densely arranged, persistent, ascending, alternate, 11−15 × 

2−3 mm, narrowly elliptic to obovate, base slightly attenuated, petiole broadened towards the stem and constricted 

above, apex acute, margins notably revolute, densely covered by notable glandular vesicular capitate trichomes on 

both surfaces, concolorous, chartaceous. Capitula solitary, 

terminal, shortly pedunculate, peduncles 3−12 mm long, 

homogamous, discoid. Involucres ~ 18 × 8 mm, cylindrical-campanulate; phyllaries arranged in 3 to 4 series; outer 

phyllaries with foliaceous apices, 8−10 × 3 mm, or narrowly triangular-ovate to narrowly elliptic, apex acute, 

gland-dotted on abaxial surface with notable glandular 

vesicular capitate trichomes, coriaceous; inner phyllaries 

15−17 × 2−3 mm, narrowly ovate-elliptic, apex acutesubulate, gland-dotted towards the apex on abaxial surface, papery, margins membranaceous. Receptacles fl at, 

naked. Florets ~ 20, bisexual; corollas yellow, tubular, 

5-lobed, gradually broadened towards the apex, tube 10−14 

mm long, lobes 1.7−2 × 0.7−0.8 mm, triangular, glanddotted abaxially; anther thecae ~ 3 mm long, bases slightly 

sagittate, apical appendages 1−1.5 × 0.5 mm, triangular, 

antheropodium present, not swollen; style shafts ~ 9 mm 

long, base swollen, style branches 4−5 mm long, narrowly 

obovate, apex acute, collecting hairs very short on upper 

2/3. Achenes 6−8 mm long, obconic, terete, ~ 6-ribbed, 

gland-dotted, sparsely villose (long twin trichomes of up 

to 1.5 mm long), carpopodium present. Pappus 12−13 mm 

long, 2-seriate, composed of 33 to 53 fl attened bristles, up 

to 0.1 mm wide, scabrid, barbellate towards the apex.

Vernacular names: Not known.

Phenology: Flowering specimens have been collected from January to February.

Distribution: Chile, the only three collections are 

from Región Metropolitana (Figure 34).

Habitat: Little is known about this species whose 

only three collections are from the mid–19th century. Probably it occurs at mid elevations in the ecotonal zone between 

Central Chile and high Andean biogeographical provinces 

(Cabrera and Willink, 1973).

Uses: Not known.

Etymology: The specifi c epithet probably refers to 

the similarity of this species with those of the genus Genista 

L. (Leguminosae), which have yellow fl owers and small elliptic leaves.

Note 1: The photograph distributed by F as the 

phototype of Dolichogyne genistoides (negative number 

37661) taken from material deposited at P (P 279524), 

doesn’t match the description of this species, nor the characteristics of the holotype at SGO. This specimen at P notably resembles genus Nassauvia Comm. ex Juss., and it 

probably corresponds with species Nassauvia planifolia 

Wedd.

Additional specimens examined: CHILE. REG. METROPOLITANA. Not defi ned Dpto. Andes de Santiago, s. date, 

s. leg. s.n. (K); Cordillera de las Arañas, I-1861, Lambert 

s.n., (SGO 71787).

Nardophyllum lanatum (Meyen) Cabrera, Notas Mus. La 

Plata, Bot. 17(83): 63, 1954. Barnadesia lanata Meyen, 

Reise Erde I: 347, 1834. Type: B, destroyed (Neotype, 

designated here: “CHILE, Región Metropolitana, Prov. 

Area Metropolitana, entre Farellones y Corral Quemado, 33°21'21.2"S, 70°19'38.8"W, 16-I-2003, Bonifacino & Romano 811” (neotype: SGO 154610!; isoneotypes: LP!, SI!).
number 92 • 7 7

FIGURE 33. Nardophyllum genistoides (Phil.) A. Gray. A. Habit. B. Leaves, adaxial view (left), abaxial view (right). C. Involucre. D. Outer 

to inner phyllaries (from left to right). E. Detail of pappus bristle. F. Achene with pappus. G. Style. H. Corolla (A, Germain s.n., BM; B−H, 

Germain s.n., LP).
7 8 • SMITHSONIAN CONTRIBUTIONS TO BOTANY

Gochnatia revoluta D. Don ex Hook. and Arn., in Hooker, Comp. Bot. 

Mag. 1: 109, 1835. Nardophyllum revolutum (D. Don ex Hook. and Arn.) 

Hook. and Arn., in Hooker, Comp. Bot. Mag. 2: 44, 1836. Pentaphorus 

rosmarinifolius Gill. ex Hook. and Arn. pro syn., Comp. Bot. Mag. 1: 109, 

1835. Type: “Gochnatia revoluta Don, but it differs from Gochnatia by 

the branches of style pret long and slightly acute and pappus deciduous 

plumose, ca. on bud, Compos. Cordillera, Cuming 185” (lectotype: K!, 

designated here; isolectotype: K!; syntype: “Vernonia, Gochnatia revoluta 

Don., Travesia Provincia San Luis, Dr. Gillies 153”, K!; isosyntype: BM!).

Dolichogyne staehelinoides DC., Prodr. 7(1): 256, 1838. Dolichogyne 

candollei J. Rémy nom. illeg., in Gay, Fl. Chile 4: 103, tab. 45, 1849. 

Nardophyllum candollei (J. Rémy) F. Meigen nom. illeg., Bot. Jahrb. 

Syst. 17: 281, 1893. Aster staehelinoides (DC.) Kuntze, Revis. Gen. 

Pl. 3(3): 131, 1898. Nardophyllum staehelinoides (DC.) Macloskie, 

Rep. Princeton Univ. Exp. Patagonia, Botany, 8: 784, 1905. Type: “De 

Peña Blanca al Cerro en las Pampas, Nee [s.n.], herb. Thibaud.” (holotype: G; scanned image of holotype at G: LP!; photos of holotype at 

G: LP!, SI!).

FIGURE 34. Geographic distribution (shaded area) of Nardophyllum genistoides (Phil.) A. Gray.
number 92 • 7 9

Dolichogyne gnaphalioides DC., Prodr. 7(1): 256, 1838. Dolichogyne 

candollei J. Rémy nom. illeg., in Gay, Fl. Chile 4: 103, tab. 45, 1849. 

Nardophyllum candollei (J. Rémy) F. Meigen nom. illeg., Bot. Jahrb. 

Syst. 17: 281, 1893. Type: “[Chile] Hab. Cordiliera de Talcaragné, D. C. 

Gay de Draguignan, 1833” (holotype: G; scanned image of holotype at 

G: LP!; isotypes: F!, US!).

(FIGURES 35, 36, 61)

Shrubs 30−100 cm tall, globose, usually spreading; 

branches densely disposed, ascending, 8-ribbed, internodes 

up to 3 mm long. Leaves densely arranged, persistent, alternate, ascending, 9−18 × 1.3−2 mm, narrowly obovate, base 

slightly attenuated, petiole broadened towards the stem and 

constricted above, apex acute to subacute, margins notably revolute, densely tomentose and gland-dotted on both 

surfaces, more or less concolorous, coriaceous. Capitula 

solitary, more rarely arranged in cimose syn fl orescences 

composed of 2 to 3 capitula, terminal, sessile to shortly 

pedunculate, peduncles up to 10 mm long, homogamous, 

discoid. Involucres 10−15(−17) × 2.5−4.5 mm, cylindrical; 

phyllaries arranged in 4 to 6 series; outer phyllaries with foliaceous apices, 4.5−7 × 1.5−2.5 mm, triangular-ovate to elliptic, with a notorious longitudinal mid-ridge towards the 

base, apex acute to subulate, tomentose and gland-dotted on 

abaxial surface, coriaceous, margins membranaceous; inner 

phyllaries 9−16 × 2−3 mm, narrowly elliptic, apex acute 

to subulate, tomentose gland-dotted towards the apex on 

abaxial surface, papery, margins membranaceous. Receptacles fl at, naked or rarely partially paleate; when present, 

paleae 1, ~ 10−15 × 1 mm, narrowly elliptic, not embracing the achenes, apex acute, gland-dotted towards the apex 

on abaxial surface, margins membranaceous. Florets 4 to 

6, bisexual; corollas yellow, tubular, 5-lobed, broadened 

more or less abruptly from mid-length, tube 5−6 mm long, 

throat 4−5 mm long, lobes 1.4−3 × 0.6−0.9 mm, triangular, 

gland-dotted abaxially; anther thecae 3−4 mm long, bases 

slightly sagittate, apical appendages 1−1.5 × 0.3−0.4 mm, 

triangular, antheropodium present, not swollen; style shafts 

8−10 mm long, base swollen, style branches 4−6.5 mm 

long, narrowly elliptic to oblong, subacute, collecting hairs 

short on upper 1/2. Achenes 6.5−9 mm long, obconic, terete, ~ 6-ribbed, gland-dotted, sparsely villose, carpopodium 

present. Pappus 9−13 mm long, 2 (3)-seriate, composed of 

50 to 110 fl attened bristles, up to 0.1 mm wide, scabrid, 

barbellate towards the apex.

Vernacular names: Not known.

Phenology: Flowering specimens have been collected from October to April.

Distribution: Restricted to Chile in Regions of 

Atacama, Coquimbo, Valparaíso, Metropolitana, and General O’Higgins (Figure 36).

Habitat: Nardophyllum lanatum occurs in rocky 

slopes between 1,200 and 3,200 m in the ecotonal zone 

between Central Chile and high Andean biogeographical 

provinces (Cabrera and Willink, 1973). I have seen it growing amid sparse vegetation in close proximity to chaparral 

vegetation characterized by Lithraea caustica Hook. and 

Arn. and Quillaja saponaria Molina.

Uses: Not known.

Etymology: The specifi c epithet refers to the dense 

and profuse tomentose pubescence that covers leaves and 

stems.

Note 1: The type of Barnadesia lanata Meyen, was 

collected near Queseria and Río del Volcán in the Andes 

of the Metropolitan Region in Chile. According to Stafl eu 

and Cowan (1981) Meyen’s herbarium was acquired by B. 

However, according to B staff (pers. comm.) this specimen 

is not present at B, and it was probably destroyed during 

the fi res caused by the bombing of Berlin-Dahlem in March 

1943. I’ve been unable to locate photos of this material in 

the F collection of negatives of material originally deposited 

at B. I therefore think designation of a neotype is in order. 

I’ve selected a specimen collected near the area where the 

holotype was collected. The material here proposed as neotype has abundant reproductive material and has duplicates 

distributed in SGO, SI, and LP.

Note 2: Hooker and Arnott (1835) in the protologue of Gochnatia revoluta mentioned two specimens “La 

travesia and La Punta, Province of San Luis, Dr. Gillies” 

and “Cordillera of Chili, Cuming (185)”. I have studied 

both specimens deposited at K and they perfectly match the 

original diagnosis. I select Cuming 185 as Lectotype since 

is the material utilized by Hooker and Arnott in the original descripition since Gillies’ specimen was not available to 

them. Cuming’s material is immature but it doesn’t pose 

any doubts regarding his taxonomic identity, and is more 

abundant than Gillie’s material.

Note 3: Regarding the syntype of Gochnatia 

revoluta D. Don ex Hook. and Arn., the label of Gillie’s 

material at K says: “Vernonia, 153, Gochnatia revoluta

Don [an illegible word between parenthesis], Travesia 

Province San Luis, Gillies”. The information on the labels 

match to a great extent what appears on the original publication by Hooker and Arnott (1835); however, a special 

comment should be made on the locality of this specimen. 

Province of San Luis in Argentina is mentioned as the locality for this specimen, while all the rest of ancient and 
8 0 • SMITHSONIAN CONTRIBUTIONS TO BOTANY

FIGURE 35. Nardophyllum lanatum (Meyen) Cabrera. A. Habit. B. Leaves, adaxial view (left), abaxial view (right). C. Involucre. D. Outer to 

inner phyllaries (from left to right). E. Palea (dorsal view). F. Detail of pappus bristle. G. Achene with pappus. H. Style. I. Corolla (A−I, Bonifacino & Romano 811, LP).
number 92 • 8 1

modern records for the species on the herbaria to which I 

have access are restricted to the western side of the Andes. 

According to Britten and Boulger (1893), Gillies resided 

in Mendoza and traveled widely between Argentina and 

Chile, as well as in the provinces of Mendoza, San Luis, 

and Cordoba, so it is probable that the locality for this 

specimen might have been somehow incorrectly noted 

on the label. There is a specimen deposited at BM (BM 

608510) with a handwritten label “No 153, Vernonia”, 

and with a printed label “Mendoza Dr. Gillies”. This 

specimen at BM, although holding a different locality, has 

the same ID and number of specimen attributed to Gillies, 

however the locality doesn’t match the locality mentioned 

in the protologue. It is very likely that this printed label 

was added at a later time when the material was distributed and incorrectly assigned to the specimen. Since the 

FIGURE 36. Geographic distribution of Nardophyllum lanatum (Meyen) Cabrera.
8 2 • SMITHSONIAN CONTRIBUTIONS TO BOTANY

specimen matched the original diagnosis and the handwritten label matched the one in the specimen identifi ed 

as syntype at K, the specimen at BM (BM 608510) is here 

identifi ed as isosyntype.

Note 4: J. Rémy (1849) considered Dolichogyne 

gnaphalioides DC. and Dolichogyne staehelinoides DC. 

heterotypic synonyms and instead of considering only one 

and transferring the other to the synonymy, he decided to 

coin a new and illegitimate superfl uous name Dolichogyne 

candollei J. Rémy. Because Rémy’s name applies to Dolichogyne gnaphalioides DC. and Dolichogyne staehelinoides DC. the superfl uous name appears under the synonymy 

of both names.

Additional specimens examined: CHILE. NOT DEFINED REG. s. loc., s. date, Gay s.n. (F). REG. DE ATACAMA. 

Prov. Huasco. Km 28, Río del Estrecho, 29°12'S, 70°09'W, 

3,150 m, 19-I-1994, Arancia et al. 94180 (CONC). REG. 

DE COQUIMBO. Prov. Elqui. Estación Junta de Chingoles, 

I-1944, Pizarro de Urizar s.n. (SGO). Prov. Limari. Camara 

de Carga, Los Molles, 12-I-1975, Bassano s.n. (SGO); Tulahuen, s. date, Geisse s.n. (SGO 43814); La Hualtata, 

16-I-1949, Jiles 1225 (CONC, LP); Río Molles, 2450 m, 

7-III-1951, Jiles 1963 (CONC, LP); Gordito, 3,000 m, 

1-II-1954, Jiles 2569 (CONC); La Hualtata, 2,400 m, 

30-III-1956, Jiles 2982 (CONC); Río Illapel, 2,800 m, 

16-II-1962, Jiles 4171 (CONC); Cordillera de Ovalle, Río 

Mostazal, 30°50'S, 70°47'W, 2,600 m, 25-IV-1962, Jiles 

4280-a (CONC); Potrero Grande, 2,700 m, 19-II-1965, 

Jiles 4562 (CONC). REG. DE VALPARAISO. Prov. Los Andes.

Entre Caracoles de Portillo y Juncal, 32°50'S, 70°12'W, 

2,300−2,800 m, 12-I-1981, Arroyo 81336 (CONC); Uspallata Pass der Chilenishen Hochcordillere (33°S breite) 

Juncal, auf den Bergen 2,100 m, 8-II-1903, Buchtien s.n.

(SI 8565); Juncal, 8,000−9,000 ft, I-1930, Elliott 635 (K); 

Paso Aconcagua, Llano de Juncalillo, 2,600−2,700 m, 

9-III-1954, Ricardi 2918 (CONC); Paso Aconcagua, Juncal, Quebrada Chépica, 2,400−2,500 m, 10-III-1954, Ricardi 2963 (CONC); Camino a la Laguna Negra, 2,130 m, 

15-I-1990, Teillier et al. 1979 (CONC); Río Colorado, 

2,500 m, 24-II-1967, Zöllner 1403 (LP). Maitencillo, Río 

Blanco, 3,000 m, 18-II-1972, Zöllner 5522 (CONC). Prov. 

Petorca. Río Sobrante above Chincolco, 2,500 m, 14-II1939, Morrison 17321 (K). Prov. Valparaíso. Colliguay, 

1928, Jaffuel s.n. (CONC 51957); Co Roble, Cordillera de 

la costa, 6-III-1966, Zöllner 1402 (LP). REG. METROPOLITANA. Not defi ned Prov. Alhué, Monte Cantillana, 2-I-1939, 

Barros 2587 (LP); Cordillera de Santiago, 1856−1857, 

Germain s.n. (F); Mountains near Santiago, Morro El 

Buey, ~8,000 ft, 21-IX-1958, Godley 28a (BM); Cordillera, 

Paso Cruz, 34oS, 1,500 m, s. date, Kuntze 192 (NY). Prov. 

Area Metropolitana. Entre Farellones y Paso de Jorquera, 

2,000−2,200 m, 9-II-1957, Garaventa 5347 (CONC); 

Cerro San Ramón, 2,250 m, II-1933, Grandjot s.n. (CONC 

1078, SGO); Colorado, Tal bei, I-1939, Grandjot s.n. (SI); 

Potrero Grande, I-1936, Grandjot s.n. (CONC 20773); 

Cerro Abanico, 2,200 m, II-1950, Gunckel 22071 (CONC); 

Cordillera del Abanico, 2,220 m, 30-XII-1928, Looser 687

(CONC); Refugio de Farellones, Cordillera de Las Condes, 

2,300 m, 9-II-1957, Looser 5754 (LP); Farellones, en 

faldeos, 2,100−2,300 m, 3-III-1942, Pisano 1395 (CONC); 

Cerro Noviciado, cumbre, 33°23'S, 70°57'W, 2,000 m, 

4-II-1956, Schlegel 1004 (CONC). Prov. Chacabuco.

Tiltil, I-1920, Jaffuel s.n. (CONC 46297); Cerros de Tiltil, 

XII-1950, Moreno s.n. (CONC 93936); road to Laguna 

Chicauma, 10 km from the lake, 30 km SW of Tiltil, 

1,960 m, 21-I-1939, Morrison & Wagenknecht 17123

(K, SI). Prov. Cordillera. Cajón del Maipo, 11-I-1924, Jaffuel s.n. (CONC 51954); Cajón del Maipo, Tramo Estero 

Coyanco -El Manzano, Estero El Sauce, 1,380 m, 23-III1995, Muñoz & Moreira 3745 (SGO); Bajando hacia 

el Embalse del Yeso, zona árida cerca de Campamento 

Minero, 20-I-1995, Muñoz et al. 3645 (SGO); Bajando 

hacia el Embalse del Yeso, zona árida cerca de Campamento Minero, 20-I-1995, Muñoz et al. 3656 (SGO); Cajón 

del Yeso, Las Romazas, 2,100 m, 22-I-1995, Muñoz et al. 

3668 (SGO); Lagunillas, Cajón del Volcán San José, 2,200 

m, 19-II-1966, Peña 10 (CONC); Lo Valdes, Cajón Morales, 2,200 m, 6-III-1953, Ricardi 2435 (CONC); Entre 

Lo Valdes y la Yesera, 2,450 m, 11-II-1963, Ricardi et al. 

864 (CONC); Aguas Negras, El Volcán, 2,300 m, II-1967,

Richter s.n. (CONC 93937); Parque Nacional El Morado, 

2,200−2,800 m, 14-II-1978, Romero & Flores 24 (SGO); 

Parque Nacional El Morado, 33°49'S, 70°05'W, 2,150 m, 

14-I-1991, Teillier et al. 2409 (SGO); Cajón del Maipo, 

Trayecto Campamento Cruz de Piedra hasta 2,100 m, 

2,100−2,400 m, 19-II-1995, Villagrán et al. 8567 (SGO). 

REG. GRAL O’HIGGINS. Prov. Colchagua. Vegas del Flaco al 

E de la Quebrada de los Ríos, 1,800 m, 7-II-1974, Mahu 

9858 (LP); San Fernando, Termas del Flaco, lado sur, 

1,700 m, 15-II-1983, Montero 12486 (CONC); Termas 

del Flaco, 34°56'S, 70°25'W, 1,800 m, 3-II-1989, Niemeyer & Fernández 8907 (CONC); íd., ribera norte del río 

Tinguiririca, II-1986, Núñez & Labra s.n. (SGO 105284); 

San Fernando, Vegas del Flaco, Cerro del Arroyo, 2,300 m, 

9-II-1955, Ricardi 3209 (CONC); junto al Río Tinguiririca 

y Río Azufre, 34°49'S, 70°34'W, 1,220 m, 9-III-1979, Villagrán & Arroyo s.n. (CONC 55922); Termas del Flaco, 

19-II-1966, Zöllner 833 (LP); íd., 2,500 m, 19-II-1966, 
number 92 • 8 3

Zöllner 891 (CONC); near of the baths El Flaco, 2,500 m, 

9-IV-1977, Zöllner 10168 (MO).

Nardophyllum patagonicum (Cabrera) G. L. Nesom, 

Phytologia 75(5): 358−365, 1993. Paleaepappus patagonicus Cabrera, Bol. Soc. Argent. Bot. 11(4): 273, 

1969. Type: “Chubut, Río Chico, Aut. 1900, Ameghino 

[s.n.]” (holotype: LP!).

(FIGURES 37, 38, 62)

Shrubs 10−40 cm tall, irregular to globose, with thick 

and notably twisted stems; branches densely disposed and 

intricate, ascending, tapering towards the end, 8-ribbed 

(ribs occluded by dense tomentum), densely tomentose. 

Leaves arranged in macroblasts and brachyblasts, when in 

macroblasts more or less laxly arranged, internodes up to 

5 mm long, persistent, alternate, ascending, (3.5−) 8−13 × 

1−2.5 mm, narrowly obovate, base slightly attenuated, petiole broadened towards the stem and constricted above apex, 

obtuse, margins notably revolute, tomentose on abaxial surface, glabrous to sparsely tomentose on adaxial surface, discolorous, coriaceous. Capitula solitary, terminal, sessile to 

shortly pedunculate, peduncles up to 10 mm long, homogamous, discoid. Involucres 9−10 × 6−8 mm, cylindrical-campanulate; phyllaries arranged in 3 to 4 series; outer phyllaries 

with foliaceous apices, 7−8 × 1.5−2 mm, narrowly ovate, 

mid-ridge clearly marked, apex acute-subulate, tomentose 

and gland-dotted on abaxial surface, coriaceous; inner 

phyllaries 9−10 × 2−2.5 mm, narrowly ovate, apex acutesubulate, laxly tomentose and gland-dotted towards the 

apex on abaxial surface, papery, margins membranaceous . 

Receptacles convex, paleate; paleae 9−10 × 2−2.5 mm, narrowly ovate and boat-shaped, embracing the achenes, apex 

acute-subulate, recurved, tomentose and gland-dotted on 

upper adaxial surface, margins membranaceous. Florets 25 

to 40, bisexual; corollas yellow, tubular, 5-lobed, gradually 

broadened towards the apex, tube 5−6.5 mm long, lobes 

1.5−2 × 0.8−1 mm, triangular, gland-dotted abaxially; anther thecae 1.8−2 mm long, bases slightly sagittate, apical 

appendages 1−1.2 × 0.3−0.4 mm, triangular-ovate, antheropodium present, swollen; style shafts 4.5−5 mm long, base 

swollen, style branches 3−5 mm long, narrowly obovate to 

oblong, apex acute, collecting hairs very short on upper 2/3 

to 3/4. Achenes 3−7 mm long, narrowly obovate, terete, 

8- to 10-ribbed, gland-dotted and densely villose, carpopodium present. Pappus 4−7 mm long, 2-seriate, composed 

of 9 to 11 narrowly ovate scale 0.4−1.2 mm wide, scabrid 

along the margins.

Vernacular names: Not known.

Phenology: Flowering specimens have been collected from November to December.

Distribution: Argentina, endemic to central Chubut 

(Figure 38).

Habitat: Nardophyllum patagonicum occurs in the 

Patagonian biogeographic province (Cabrera and Willink, 

1973), in an area of reduced annual rainfall, close to 

200 mm (Conti, 1998). It grows in rocky areas with very 

scarce vegetation cover, and associated with Chuquiraga

spp., Nassauvia spp., Lycium sp., Fabiana patagonica Speg., 

Junellia spp., Nardophyllum chiliotrichioides (J. Rémy) 

A. Gray, etc.

Uses: Not known.

Etymology: The specifi c epithet refers to the geographic area where the species occur.

Note 1: Nesom (1993b) proposed the inclusion 

of monotypic Paleaepappus in Nardophyllum. Later, 

Bonifacino and Sancho (2001) reinstated Paleaepappus

as a distinct genus, an idea also shared by Nesom et al. 

(2003). Additional morphological evidence enriched with 

specifi c fi eld observations, together with molecular evidence (Bonifacino and Funk, unpublished data) strongly 

suggest that, in spite of distinctive features, this taxon belongs to Nardophyllum. In the light of the phylogenetic 

tree generated via molecular data, it has been possible to 

reexamine certain characters and discover morphological 

traits that fi t the new Nardophyllum circumscription here 

presented.

Additional specimens examined: ARGENTINA. PROV. 

CHUBUT. Dpto. Sarmiento. Bajo del Avestruz 45°15'29"S, 

68°58'2"W, 15-I-2002, Bonifacino et al. 451 (LP); Ruta 

Prov. 24, al S de Paso de Los Indios, 346 m, 44°44'25.8"S, 

69°11'50.4"W, 10-XII-2006, Bonifacino et al. 2967 (US);

Ruta Prov 24, Lado W de Laguna Palacios, 44°44'26"S, 

69°12'46"W, 27-XI-2002, Bonifacino & Donato 713

(LP); Ruta Prov. 24, Lado W de Laguna Palacios, 

44°46'42"S, 69°15'23"W, 27-XI-2002, Bonifacino & 

Donato 716 (LP); Ruta Prov. 24, Cerros al W de Laguna 

Seca, 44°53'11"S, 69°13'26"W, 27-XI-2002, Bonifacino 

& Donato 717 (LP); Bajo del Avestruz, 45°15'29.2"S, 

68°58'1.6"W, 28-XI-2002, Bonifacino & Donato 721

(LP); Ruta Prov. 24, 45°16'0.5"S, 69°01'50"W, 28-XI2002, Bonifacino & Donato 727 (LP); Colinas arcillosas 

al W de Lag. Seca, 94 km al N de Sarmiento, 12-XII-1987, 

León 3846 (BAA). Dpto. Mártires. Ruta Prov. 53, ~Sierra 

Negra, 43°59'24"S, 67°35'32"W, 26-XI-2002, Bonifacino 

& Donato 691 (LP).
8 4 • SMITHSONIAN CONTRIBUTIONS TO BOTANY

FIGURE 37. Nardophyllum patagonicum (Cabrera) G. L. Nesom. A. Habit. B. Leaves, adaxial view (left), abaxial view (right). C. Involucre. D. Outer to inner phyllaries (from left to right). E. Receptacular paleae, lateral view (left), dorsal view (right). F. Detail of pappus scale. 

G. Achene with pappus. H. Style. I. Corolla (A−I, Bonifacino & Donato 691, LP).
number 92 • 8 5

SPECIES EXCLUDED FROM 

CHILIOTRICHUM GROUP SENSU STRICTO

Dolichogyne acaulis Wedd. ex Benth. and Hook. f., Gen. 

Pl. 2(1): 258, 1873. Lepidophyllum acaule (Wedd. ex 

Benth. and Hook. f.) Benth. and Hook. f. ex Jackson, 

Index Kew. 2(3): 58, 1894. Type: “[Bolivia] Prov. Larecaja, Viciniis Sorata, via ad Lacatia ad frontes rivi 

Aparasiri, in graminosis, Alt. Reg. Alpina, 3,900 m, 

8bre 1858-Mai 1859, Mandon 211”, lectotype: BM, 

isolectotype: K, P, US! (≡ Novenia acaulis (Wedd. ex 

Benth. and Hook. f.) S. E. Freire and F. Hellwig, Taxon, 

39 (1): 125, 1990).

Dolichogyne glabra Phil., Anal. Mus. Nac. Chile, Sec. 2, 

Bot. 8: 39, 1891. Type: “[Chile] Lorohuasi, Jan 1885, 

F. Ph[ilippi s.n.]”, holotype: SGO! (= Parastrephia 

lucida (Meyen) Cabrera, Notas Mus. La Plata, Bot. 

17(83): 57, 1954).

Dolichogyne lepidophylla Wedd., Chlor. And. I: 182, 1856. 

Ind. loc. type: “Perou: departement de Tacna, sur le 

plateau de la Cordillere de Tacora, h. 4,000 a 5,000 metres, Pentland, Wedd. [s.n.]; deapartement d’Arequipa, 

Meyen. Bolivie: Cordilleres du departement de La Paz, 

Pentland, Wedd. [s.n.]; punas entre Oruro et Potosi, 

d’Orbigny [s.n.]”, syntypes: P, scanned images of syntypes at P: LP! (= Parastrephia quadrangularis (Meyen) 

Cabrera, Notas Mus. La Plata, Bot. 17(83): 57, 1954).

Dolichogyne rigida Wedd., Chlor. And. I: 182−183, 1856. 

Ind. loc. type: “Perou Austral. (Cordillera de Tacora), 

Wedd. [s.n.]”, holotype: P, scanned image of holotype at 

P: LP!, isotypes: US!, F!, P (= Parastrephia lucida (Meyen) 

Cabrera, Notas Mus. La Plata, Bot. 17(83): 57, 1954).

Dolichogyne rupestris Wedd., Chlor. And. I: 183, 1856. 

Type: “Bolivie, Potosi, d’Orbigny 1382”, holotype: P, 

scanned image of holopype at P: LP!, isotypes: F!, P 

FIGURE 38. Geographic distribution of Nardophyllum patagonicum (Cabrera) G. L. Nesom.
8 6 • SMITHSONIAN CONTRIBUTIONS TO BOTANY

(= Parastrephia lucida (Meyen) Cabrera, Notas Mus. 

La Plata, Bot. 17(83): 57, 1954.

Lepidophyllum abietinum (Phil.) Reiche, Anal. Univ. Chile 

109: 26, 1901. Polyclados abietinus Phil., Anal. Univ. 

Chile 43: 492, 1873. Type: “[Chile] Salitreras de Antofagasta, 1872, W. Doell [s.n.]”, holotype: SGO! (= Parastrephia lucida (Meyen) Cabrera, Notas Mus. La Plata, 

Bot. 17(83): 57, 1954).

Lepidophyllum cupressinum (Phil.) Kuntze, Rev. Gen. Pl. 

3(2): 162, 1898. Polyclados cupressinus Phil., Viage 

Atacama: 208, 1860. Type: “[Chile] Alto de Puquios, in 

deserto Atacamensi, Februario 1854, Ph[ilippi] [s.n.]”, 

holotype: SGO!, isotype: SGO! (= Parastrephia quadrangularis (Meyen) Cabrera, Notas Mus. La Plata, Bot. 

17(83): 57, 1954).

Lepidophyllum lucidum (Meyen) Cabrera, Bol. Soc. Argent. Bot 1(1): 51, 1945. Baccharis lucida Meyen, Reise 

um die Erde 1: 460, 1834. Type: “Peru, circa Tacora, 

14,000−17,000, 31-IV-1833, Meyen”, holotype: B, destroyed; photo of the holotype at B: F! (≡ Parastrephia 

lucida (Meyen) Cabrera, Notas Mus. La Plata, Bot. 

17(83): 57, 1954).

Lepidophyllum meyenii A. Gray nom. illeg., Proceed. 

Amer. Acad. Arts and Sci. 5: 122, 1862. (= Parastrephia 

quadrangularis (Meyen) Cabrera, Notas Mus. La Plata, 

Bot. 17(83): 57, 1954).

Lepidophyllum phylicaeforme (Meyen) Hieron. ex R. E. 

Fr., Nov. Acta Reg. Soc. Scient. Upsaliensis, Ser. 4, 1 (1): 

77, 1905. Baccharis phylicaeformis Meyen, Reise um die 

Erde 2: 31, 1835. Type: “Peru: Arequipa, 14,000−16,000, 

31-IV-1833, Meyen”, holotype: B, destroyed; photos of 

the holotype at B: US!, F! (= Parastrephia lucida (Meyen) 

Cabrera, Notas Mus. La Plata, Bot. 17(83): 57, 1954).

Lepidophyllum phylicaeforme var. resinosum (Walp.) Blake, 

Washington Acad. Scien. Journ. 21(14): 326, 1931. Vernoniaphylicaeformis var. resinosa Walp., Nov. Act. Acad. 

Caes. Leop. Carol., 19(suppl. 1): 253, 1843. Ind. loc. 

type: “Peruvia: in planitie circa Tocoram, alt. 14−17,000 

ped. [s. leg., s.n.]”, holotype: not seen, unknown location

(= Parastrephia lucida (Meyen) Cabrera, Notas Mus. La 

Plata, Bot. 17(83): 57, 1954).

Lepidophyllum quadrangulare (Meyen) Benth. and Hook. 

f., Genera Plantarum 2(1): 258, 1873. Baccharis quadrangularis Meyen, Reise um die Erde 1: 460, 1834. Ind. 

loc. type: “[Peru] Tacora”, holotype: B, destroyed; 

photos of the holotype at B: F!, US! (≡ Parastrephia 

quadrangularis (Meyen) Cabrera, Notas Mus. La Plata, 

Bot. 17(83): 57, 1954).

Lepidophyllum rigidum (Wedd.) Benth. and Hook. f., 

Genera Plantarum 2(1): 258, 1873. Dolichogyne rigida

Wedd., Chlor. And. I: 182-183, 1856. Type: See entry 

under Dolichogyne rigida Wedd. (= Parastrephia lucida

(Meyen) Cabrera, Notas Mus. La Plata, Bot. 17(83): 57, 

1954).

Lepidophyllum teretiusculum Kuntze, Rev. Gen. Pl. 3(2): 

162, 1898. Type: “[Chile] Conchi-Ascotan, 3,000−3,900, 

7-III-1892, Otto Kuntze [s.n.]”, holotype: NY! (≡ Parastrephia teretiuscula (Kuntze) Cabrera, Notas Mus. La 

Plata, Bot. 17(83): 57, 1954).

Lepidophyllum tola Cabrera, Bol. Soc. Argent. Bot 1(1): 

51, 1945. Dolichogyne lepidophylla Wedd., Chlor. And. 

I: 182, 1856. Type: See entry under Dolichogyne lepidophylla Wedd. (= Parastrephia quadrangularis (Meyen) 

Cabrera, Notas Mus. La Plata, Bot. 17(83): 57, 1954).

Nardophyllum armatum (Wedd.) Reiche, Anales Univ. 

Chile, 109: 29, 1901. Dolichogyne armata Wedd., Chlor. 

And. I: 181, 1856. Type: “Dolichogyne armata Wedd. 

Chloris, frutex graveolens 1−2 metres fl or. lut. Potosi 

(Bolivie) M. Weddell # 4084”, lectotype: P, designated 

by Bonifacino, 2008; scanned image of lectotype: LP; 

isolectotype: P, F! (≡ “Ocyroe armata” (Wedd.) Bonif., 

Brittonia 60(3): 207, 2008; = Ocyroe spinosa Phil., Anales Mus. Nac. Chile, Bot. 8: 33, tab. I, fi g. 5, 1891).

Nardophyllum paniculatum Phil., Anal. Univ. Chil. 87: 

435, 1894. Type: “[Chile] Mansel, April 1892, Julius 

Philippi [s.n.]”, holotype: SGO!, isotype: SGO! (= Baccharis philippii Heering, Jahrb. Hamb. Wiss. Anst. 

1913, 21 Beih. 80, 1914).

Nardophyllum scoparium Phil., Anal. Univ. Chil. 87: 

434, 1894. Hinterhubera scoparia (Phil.) Cabrera, 

Notas Mus. La Plata 17: 65. 1954. Type: “[Chile] in 

parte superiore vallis de Huanta dicti provinciæ Coquimbo, F. Philippi s./n.”, holotype: SGO!, isotype: LP! (≡

Guynesomia scoparia (Phil.) Bonif. and Sancho, Taxon 

53(3): 677, 2004).
number 92 • 8 7

FIGURES 39–62

FIGURE 39. Stem and leaf anatomy of the Chiliotrichum Group sensu stricto representative taxa. A. Close-up of a stem cross section of Nardophyllum chiliotrichioides, note the abundant schlrenchyma along the ribs. B. Stem cross section of Nardophyllum chiliotrichioides. C. Leaf 

cross section showing an elevated stoma on abaxial epidermis, in Nardophyllum bryoides. D. Leaf cross section showing a sunken stoma on 

adaxial epidermis, in Katinasia cabrerae. E. Adaxial epidermis of Haroldia mendocina. F. Abaxial epidermis of Haroldia mendocina. G. Adaxial 

epidermis of Chiliotrichum diffusum. H. Abaxial epidermis of Chiliotrichum diffusum (A, B, Simón & Bonifacino 782, LP; C, D, Bonifacino et 

al. 563, LP; E, Bonifacino & Donato 699, LP; F, Bonifacino & Romano 825, LP; G, H, Bonifacino & Donato 791, LP).
8 8 • SMITHSONIAN CONTRIBUTIONS TO BOTANY

FIGURE 40. Pollen of the Chiliotrichum Group sensu stricto. A. Chiliotrichum diffusum, polar view. B. Chiliotrichum diffusum, equatorial 

view. C. Lepidophyllum cupressiforme, polar view. D. Lepidophyllum cupressiforme, equatorial view. E. Chiliophyllum densifolium, polar 

view. F. Chiliophyllum densifolium, equatorial view (A, B, Bonifacino et al. 563, LP; C, D, Bonifacino et al. 77, LP; E, F, Gentili 324, LP).
number 92 • 8 9

FIGURE 41. Pollen of the Chiliotrichum Group sensu stricto (cont.). A. Nardophyllum patagonicum, polar view. B. Nardophyllum patagonicum, equatorial view. C. Nardophyllum bryoides, polar view. D. Nardophyllum bryoides, equatorial view. E. Chiliotrichiopsis peruviana, polar 

view. F. Chiliotrichiopsis peruviana, equatorial view (A, B, Leon 3846, BAA; C, D, Bonifacino et al. 464, LP; E, F, Flores et al. 2109, LP).
9 0 • SMITHSONIAN CONTRIBUTIONS TO BOTANY

FIGURE 42. Pollen of the Chiliotrichum Group sensu stricto (cont.). A. Chiliotrichiopsis keidelii, polar view. B. Chiliotrichiopsis keidelii, equatorial view. C. Aylacophora deserticola, polar view. D. Aylacophora deserticola, equatorial view. E. Cabreraea andina, polar view. F. Cabreraea 

andina, equatorial view (A, B, Simón & Bonifacino 520, LP; C, D, Bonifacino et al. 335, LP; E, F, Biurrun & Molina 4574, CORD).
number 92 • 9 1

FIGURE 43. Pollen of the Chiliotrichum Group sensu stricto (cont.). A. Katinasia cabrerae, polar view. B. Katinasia cabrerae, equatorial view. 

C. Haroldia mendocina, polar view. D. Haroldia mendocina, equatorial view (A, B, Simón & Bonifacino 847, US; C, D, Simón & Bonifacino 

775, US).
9 2 • SMITHSONIAN CONTRIBUTIONS TO BOTANY

FIGURE 44. Aylacophora deserticola Cabrera. A. Habit and habitat, N of Cutral-Co, Neuquén, Argentina; arrow points to one of several 

individuals of Aylacophora deserticola. B. Close-up of capitulum. C. Close-up of stem, notice wide ribs and narrow furrows.
number 92 • 9 3

FIGURE 45. Cabreraea andina (Cabrera) Bonif. A. Habit and habitat (Mogote de los Corralitos, San Juan, Argentina). B. Close-up of 

branches.
9 4 • SMITHSONIAN CONTRIBUTIONS TO BOTANY

FIGURE 46. Chiliophyllum densifolium Phil. A. Branching; notice shiny gland-dotted leaves, and cylindrical shape of involucre. B. Habit and 

habitat (Portillo de Tunuyán, Mendoza, Argentina). C. Close-up of capitula.
number 92 • 9 5

FIGURE 47. Chiliotrichiopsis keidelii Cabrera. A. Close-up of a capitulum. B. Lateral view of involucre. C. Habitat and habit, S of Say, Jujuy, 

Argentina). D. Close-up of branches.
9 6 • SMITHSONIAN CONTRIBUTIONS TO BOTANY

FIGURE 48. Chiliotrichiopsis ledifolia (Griseb.) Cabrera. A. Habitat (road to Mina El Oro, near Chilecito, La Rioja, Argentina). B. Lateral view 

of involucre. C. Close-up of branches. D. Close-up of leaves. E. Detail of the capitulum.
number 92 • 9 7

FIGURE 49. Chiliotrichiopsis peruviana G. L. Nesom, H. Rob. and A. Granda. Image of Holotype at US.
9 8 • SMITHSONIAN CONTRIBUTIONS TO BOTANY

FIGURE 50. Chiliotrichum diffusum (G. Forst.) Kuntze. A. Habit. B. Close-up of involucre. C. Close-up of capitulum. D. Close-up of leaves.
number 92 • 9 9

FIGURE 51. Chiliotrichum diffusum (G. Forst.) Kuntze (cont.). A. Habitat in N Fuegia, close to Bahía San Sebastián, Argentina; notice dense 

consociation and almost exclusive presence on the shrub stratum. B. Habitat in S Fuegia amid Nothofagus forest clearings. C. Chiliotrichum 

diffusum can also be found as part of the understory of Nothofagus forest such as the one shown here. D. Dense consociations of Chiliotrichum 

diffusum in forest clearings in SW Chubut, Argentina.
100 • SMITHSONIAN CONTRIBUTIONS TO BOTANY

FIGURE 52. Chiliotrichum fuegianum (O. Hoffm.) Bonif. A. Close-up of capitula. B. Detail of leaves. C. Habit and Habitat (Bahía San Sebastián, Tierra del Fuego, Argentina). D. Close-up of branches.
number 92 • 101

FIGURE 53. Haroldia mendocina (Cabrera) Bonif. A. Habit. B. Lateral view of capitulum. C. Upper view of capitulum. D. Habitat (Cordillera 

del Tigre, Mendoza, Argentina). E. Close-up of the leaves.
102 • SMITHSONIAN CONTRIBUTIONS TO BOTANY

FIGURE 54. Katinasia cabrerae (Bonif.) Bonif. A. Close-up of branches. B. Lateral view of capitula; notice distinctive cylindrical shape of involucre and white corollas. C. Habit; notice intricate branching pattern. D. Cluster of capitula. E. Habitat at type locality (Arroyo China Muerta, 

Neuquén, Argentina), here Katinasia is among the dominant species in the shrub stratum. F. Close-up of the leaves; notice entire (non-revolute) 

margin of leaves and the slightly noticeable ribs along the stem.
number 92 • 103

FIGURE 55. Lepidophyllum cupressiforme (Lam.) Cass. A. Close-up of the capitulum. B. Lateral view of capitulum. C. Close-up of branches; 

notice dense branching pattern. D. Habit. E. Close-up of a branch; notice opposite phyllotaxy. F. Habitat in N Fuegia (W of Bahía Inútil, 

Chile); notice absolute dominance of Lepidophyllum cupressiforme in the shrub stratum, a common situation in saline fl at and low areas of N 

Fuegia.
104 • SMITHSONIAN CONTRIBUTIONS TO BOTANY

FIGURE 56. Nardophyllum bryoides (Lam.) Cabrera. A. Close-up of capitula in lateral view. B. Close-up of branching pattern. C. Closeup of leaves. D. Close-up of leaves; notice white tomentose pubescence causing a more whitish appearance on leaves. E. Habit of dwarf 

individual appressed to the ground. F. Close-up of dwarf individual showing very dense branching pattern (SE Santa Cruz, close to Río 

Gallegos, Argentina).
number 92 • 105

FIGURE 57. Nardophyllum bryoides (Lam.) Cabrera (cont.). A. Habit in W Santa Cruz, close to Lake San Martín (Argentina); notice compact 

shape of the shrubs and dominance in the vegetation. B. Towards the north end of the distribution, individual plants tend to show a more lax 

branching pattern, as shown here in central Chubut (Argentina). C. Some individual plants can attain a notable size, amassing an important 

amount of soil between their branches; also notice the different pubescence shown on the leaves of these two contiguous individuals.
106 • SMITHSONIAN CONTRIBUTIONS TO BOTANY

FIGURE 58. Nardophyllum chiliotrichioides (J. Rémy) A. Gray. A. Close-up of the capitulum, from a population in Chubut, 300 m elevation 

(Argentina). B. Close-up of the capitulum from a population in Mendoza, 2,800 m elevation (Argentina). C. Close-up of the capitulum from a 

population near Santiago, 2,400 m elevation (Chile). D. Close-up of branches in an individual from central Chubut (Argentina). E. Close-up of 

stem and leaf; notice revolute margin on leaf and ribs along the stem.
number 92 • 107

FIGURE 59. Nardophyllum chiliotrichioides (J. Rémy) A. Gray (cont.). A. Habit in central Chubut (Argentina). B. Habit in Mendoza (Argentina), notice the more dense branching and compact shape. C. Habitat in central Chubut (Argentina).
108 • SMITHSONIAN CONTRIBUTIONS TO BOTANY

FIGURE 60. Nardophyllum genistoides (Phil.) A. Gray. A. Image of holotype at SGO. B. Detail of holotype label. C. Image of Isotype at BM.
number 92 • 109

FIGURE 61. Nardophyllum lanatum (Meyen) Cabrera. A. Close-up of a capitulum; notice the distinctive cylindrical shape of the involucre. 

B. Close-up of a branch and leaves; notice dense whitish tomentose pubescence all over the stem and leaves, and the notably revolute margin, 

also notice the rather minute ribs, notably contrasting with the ribs exhibited by Nardophyllum chiliotrichioides, also growing on the same area 

of the central Andes in Chile. C. Habit and habitat (Farellones, Región Metropolitana, Chile), notice the compact shape and dense branching 

pattern.
110 • SMITHSONIAN CONTRIBUTIONS TO BOTANY

FIGURE 62. Nardophyllum patagonicum (Cabrera) G. L. Nesom. A. Habit; notice dwarf habit and intricate branching pattern. B. Close-up 

of branches; notice dense tomentose pubescence covering the stems and also the leaves arranged in brachyblasts. C. Close-up of a capitulum. 

D. Habitat in central Chubut, near Laguna Palacios (Argentina).
Acknowledgments

I hereby express my most profound gratitude to Jorge Crisci, Vicki Funk, 

and Liliana Katinas for their guidance and encouragement during the development of this work and for their ongoing help during several years of 

daisy-hunting ventures. I am also very grateful to Gisela Sancho, who offered valuable insights on the systematics of Compositae during several stages 

of the present study; and to Diego Gutierrez, Daniel Giuliano, Laura Iharlegui, 

and Silvia Torres, fellows from LP, who have enriched my understanding of the 

Andean Compositae during countless discussions. I devoted a large amount of 

time to fi eldwork (although not as much as I would have liked!), and during that 

time I received help from numerous individuals and institutions that made my 

fi eld trips a success. I especially thank Adrian Azpiroz, Mariano Donato, Laura 

Iharlegui, German Muttoni, Gisela Sancho, and Pablo Simon; they not only 

helped out with the practical aspects of plant collecting but proved to be cheerful companions and great fi eld comrades. I give special thanks to my home university, Universidad de La República Oriental del Uruguay, and particularly to 

the Facultad de Agronomía, which, through the Botany Department, provided 

the vehicle for some of the trips I conducted in Argentina and Chile. I am greatly 

indebted to my fellows at the Botany department in the Facultad de Agronomía 

at Montevideo, especially to Eduardo Marchesi and Gabriela Speroni for their 

continued support during these years. I give special thanks to Dan Nicolson 

for his help with several nomenclatural problems; and to Harold Robinson for 

his help with the Latin diagnoses. I also thank Scott Whittaker (SEM LAB at 

NMNH, Smithsonian Institution) for his assistance and help while securing pollen SEM photographs; and Ann Elizabeth Wolf (Herbier National de Paris) for 

her assistance in locating type material.

I am also very grateful to the following individuals and institutions who 

contributed in several ways to the completion of the present study: Ariza Espinar, Adriana Bartoli, Norma Degginani, Enrique Estramil, Enrique Gamez, Natalie Goodall, Arturo Granda, Alvaro Idarraga, Roberto Kiesling, Rolando León, 

Sirli Leython, Clodomiro Marticorena, Corporación Nacional Forestal (Chile), 

Fabián Michelangeli, INTA Río Gallegos (Argentina), Mélica Muñoz, Robbin 
112 • SMITHSONIAN CONTRIBUTIONS TO BOTANY

Moran, Guy Nesom, Parques Nacionales de Argentina, 

Parques Nacionales de Venezuela, Alejandra Plaza, Pedro 

Steibel, Ivan Tatanov, Sebastián Teillier, Roberto Tortosa, 

Estrella Urtubey, Benjamín Van Ee, Richard Worthington, 

Yacimientos TOTAL (Argentina), and Fernando Zuloaga. 

I thank Alice Tangerini for preparing some of the line illustrations and Paul Peterson and Larry Dorr for editing and 

seeing the manuscript through the review process. I am 

thankful to Fundación Myndel Pedersen, IAPT, the Facultad de Agronomía, the Smithsonian Institution, and the 

National Geographic Society (grant no. 7646-04) for their 

fi nancial support towards the completion of this study. I 

also acknowledge the Smithsonian Institution for its support through a postdoctoral fellowship, thus granting me 

the opportunity to broaden my understanding of South 

American Compositae and providing me with access to 

its vast collections and facilities. I am very thankful to the 

curators at the institutions BAA, BAB, BAF, BM, CONC, 

CORD, E, F, G, K, LIL, LP, MO, MVFA, NY, P, SI, SGO, 

and US for facilitating my visits and loans of specimens.

Finally, on a personal note, I am greatly indebted to 

my family, especially to my wife Valeria—dauntless companion during several of my plant collecting journeys 

along the Andes—and to my two sons, Joaquín and Juan 

Manuel, who endured my absences; I thank the three of 

them for their patience.
References

Anonymous. 1962. Systematics Association Committee for Descriptive Biological Terminology. Terminology of simple symmetrical plane shapes (Chart 1). Taxon 11: 

145−156, 245−247.

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Index of Scientifi c Names

Main entries for taxa are indicated by bold page numbers.

Acaena spp., 58

Adesmia spp., 26

Amellus diffusus G. Forst., 4, 37–39, 43

Amellus rosmarinifolius Poepp. ex Less. pro syn., 39

Anactinia (Hook. f.) J. Rémy in Gay, 62, 63

Anactinia hookeri J. Rémy nom. illeg., 62, 65

Anarthrophyllum desideratum (DC.) Benth., 58

Aster L., 37

Aster magellanicus Spreng. nom. illeg., 37, 39

Aster nardophyllum Kuntze, 62, 65, 68

Aster staehelinoides (DC.) Kuntze, 62, 78

Astereae Cass., 1, 4, 5, 15, 16

Asterinae (Cass.) Dumort., 5

Athanasia cupressiformis (Lam.) Commerson ex Cass., 57, 58

Atriplex lampa Gill. ex Moq., 19

Aylacophora Cabrera, 1, 2 (Tbl. 1), 3, 5, 6, 13–15, 16, 17

Aylacophora deserticola Cabrera, 6, 7 (Fig. 1A), 8 (Fig. 2A), 9, 14, 17–20, 90 (Fig. 

42C,D), 92 (Fig. 44)

Baccharis bryoides (Lam.) Pers., 62, 64

Baccharis cupressiformis (Lam.) Pers., 57

Baccharis L., 2 (Tbl. 1), 4, 62

Baccharis lucida Meyen, 86

Baccharis philippii Heering, 86

Baccharis phylicaeformis Meyen, 86

Baccharis quadrangularis Meyen, 86

Barnadesia lanata Meyen, 62, 63, 76, 79

Berberis buxifolia Lam., 26, 43, 58

Brachyclados caespitosus Speg., 67

Brachyridium cupressiforme (Lam.) Meisn. Nom. illeg., 56–58

Brachyridium Meisn. nom. illeg., 56–58

Cabreraea andina (Cabrera) Bonif., 5, 6, 8 (Fig. 2B), 9, 10 (Fig. 3J), 13, 20–23, 90 

(Fig. 42E,F), 93 (Fig. 45)

Cabreraea Bonif., 2 (Tbl. 1), 3, 5, 6, 13, 14, 16, 20, 23, 27, 64

Celmisia Cass., 3
118 • SMITHSONIAN CONTRIBUTIONS TO BOTANY

Chiliophyllum andinum Cabrera, 20, 23, 24

Chiliophyllum DC., 21, 47

Chiliophyllum densifolium Phil., 2 (Tbl. 1), 6, 7 (Fig. 1E), 8 

(Fig. 2C), 9, 10 (Fig. 3F), 13, 21, 23, 24–27, 47, 88 (Fig. 

40E,F), 94 (Fig. 46)

Chiliophyllum fuegianum O. Hoffm., 21, 24, 39, 47

Chiliophyllum Phil., 1, 2 (Tbl. 1), 3, 5, 6, 13–15, 16, 17, 20, 

21, 23, 24, 39, 47, 57

Chiliotrichiopsis Cabrera, 1, 2 (Tbl. 1), 3, 5, 6, 13–15, 16, 20, 

27, 28, 50, 54, 64

Chiliotrichiopsis keidelii Cabrera, 6, 8 (Fig. 2D), 9, 10 (Fig. 

3L), 13–15, 27, 28–32, 35, 90 (Fig. 42A,B), 95 (Fig. 47)

Chiliotrichiopsis ledifolia (Griseb.) Cabrera, 6, 8 (Fig. 2E), 

9, 11 (Fig. 4D), 13, 14, 27, 28, 30 (Fig. 14), 31, 32–35, 96 

(Fig. 48)

Chiliotrichiopsis mendocina Cabrera, 2 (Tbl. 1), 28, 50

Chiliotrichiopsis peruviana G. L. Nesom, H. Rob. and A. 

Granda, 6, 7 (Fig. 1F), 8 (Fig. 2F), 9, 10 (Fig. 3B), 13, 14, 27, 

28, 35–37, 89 (Fig. 41E,F), 97 (Fig. 49)

Chiliotrichum amelloideum Cass. nom. illeg., 4, 37–39, 43

Chiliotrichum amelloideum var. diffusum (G. Forst.) Nees, 

38, 39

Chiliotrichum amelloideum var. lanceolatum Nees, 39

Chiliotrichum amelloideum var. rosmarinifolium (Less.) Nees, 

38, 39

Chiliotrichum angustifolium Phil., 38, 40

Chiliotrichum Cass., 1, 2 (Tbl. 1), 3–6, 16, 17, 21, 24, 27, 

37–39, 43, 62

Chiliotrichum darwinii Hook. f., 62, 65

Chiliotrichum diffusum (G. Forst.) Kuntze, 1, 6, 7 (Fig. 1D), 8 

(Fig. 2G,H), 9, 10 (Fig. 3H,I), 11 (Fig. 4A,B,E), 13–15, 38, 

39–47, 58, 87 (Fig. 39G,H), 88 (Fig. 40A,B), 98 (Fig. 50), 99 

(Fig. 51)

Chiliotrichum diffusum f. angustifolium (“angustifolia”) Speg., 

38, 40

Chiliotrichum diffusum f. media Speg., 38, 40

Chiliotrichum diffusum f. typica Speg. nom. illeg., 38, 39

Chiliotrichum feliciae Hombr. and Jacquin., 38, 39

Chiliotrichum fuegianum (O. Hoffm.) Bonif., 5, 8 (Fig. 2I), 9, 

14, 21, 24, 39, 47–49, 58, 100 (Fig. 52)

Chiliotrichum humile Hook. f., 62, 63, 65, 68

Chiliotrichum kingii Hook. f., 62, 65

Chiliotrichum longifolium Phil., 38, 40, 43

Chiliotrichum ovatifolium Hombr. and Jacquin., 38, 39

Chiliotrichum ovatifolium var. retortum Hombr. and Jacquin., 

38, 39

Chiliotrichum rosmarinifolium Less., 5, 38, 39

Chiliotrichum rosmarinifolium var. glabrescens Phil., 38, 40

Chiliotrichum sect. Anactinia Hook. f., 4, 62, 63

Chiliotrichum sect. Euchiliotrichum Hook. f., 4, 38

Chiliotrichum tenue Phil., 38, 40

Chiliotrichum virgatum Phil., 38, 40

Chrysothamnus Nutt., 4

Chuquiraga aurea Skottsb., 67

Chuquiraga erinacea D. Don, 19

Chuquiraga spp., 83

Colliguaja integerrima Gill. and Hook., 54

Conyza bryoides Lam., 4, 62–64

Conyza cupressiformis Lam., 4, 56, 57

Cortaderia pilosa (d’Urv.) Hack., 43

Cupressus L., 58

Diplostephium Kunth., 2 (Tbl. 1), 3–5

Dolichogyne acaulis Wedd. ex Benth. and Hook. f., 85

Dolichogyne armata Wedd., 62, 86

Dolichogyne candollei J. Rémy nom. illeg., 62, 78, 79, 82

Dolichogyne chiliotrichioides J. Rémy, 62, 71, 73

Dolichogyne DC., 4, 63

Dolichogyne genistoides Phil., 62, 76

Dolichogyne glabra Phil., 85

Dolichogyne gnaphalioides DC., 4, 62, 79, 82

Dolichogyne lepidophylla Wedd., 85, 86

Dolichogyne parvifolia Phil. ex Kuntze pro. syn., 65

Dolichogyne rigida Wedd., 85, 86

Dolichogyne rupestris Wedd., 85

Dolichogyne section Dolichogyne Wedd., 4

Dolichogyne section Tola Wedd., 4

Dolichogyne staehelinoides DC., 4, 62, 63, 78, 82

Ephedra frustillata Miers, 67

Ephedra spp., 26

Ericameria Nutt., 2 (Tbl. 1), 3–5

Fabiana imbricata Ruiz and Pav., 26

Fabiana patagonica Speg., 19, 83

Festuca gracillima Hook. f., 43, 58

Festuca pallescens (St.-Yves) Parodi, 65

Festuca spp., 28, 35

Genista L., 76

Gochnatia revoluta D. Don ex Hook. and Arn., 62, 63, 78, 79

Gochnatia subg. Nardophyllum Hook. and Arn., 62, 63 

Grindelia cupressiformis (Lam.) Sch. Bip. ex Reiche pro syn., 57

Gutierrezia cupressiformis (Lam.) Sch. Bip., 57

Gutierrezia ledifolia Griseb., 27, 32, 35

Gutierrezia solbrigii Cabrera, 19

Guynesomia Bonif. and Sancho, 63

Guynesomia scoparia (Phil.) Bonif. and Sancho, 63, 86

Haroldia Bonif., 2 (Tbl. 1), 3, 5, 6, 13, 14, 16, 28, 50

Haroldia mendocina (Cabrera) Bonif., 5, 6, 7 (Fig. 1H), 8 (Fig. 

2J), 9, 10 (Fig. 3K), 11 (Fig. 4D), 14, 27, 50–53, 87 (Fig. 

39E,F), 91 (Fig. 43C,D), 101 (Fig. 53)

Hinterhubera Sch. Bip. ex Wedd., 2 (Tbl. 1), 3, 5

Hinterhubera scoparia (Phil.) Cabrera, 86

Hinterhuberinae Cuatrec., 3, 5

Ilex paraguariensis A. St.-Hil., 26

Junellia spp., 73, 83

Katinasia Bonif., 2 (Tbl. 1), 3, 5, 6, 13–15, 16, 17, 50, 53, 54, 

64, 102 (Fig. 54E)

Katinasia cabrerae (Bonif.) Bonif., 6, 8 (Fig. 2K), 9, 10 (Fig. 3D), 

12–14, 53, 54–56, 87 (Fig. 39D), 91 (Fig. 43A,B), 102 (Fig. 54)

Larrea cuneifolia Cav., 19

Larrea divaricata Cav., 19
number 92 • 119

Larrea nitida Cav., 54

Larrea spp., 17

Lepidophyllum abietinum (Phil.) Reiche, 86

Lepidophyllum acaule (Wedd. ex Benth. and Hook. f.) Benth. 

and Hook. f. ex Jackson, 85

Lepidophyllum Cass., 1, 2 (Tbl. 1), 3–6, 13–15, 16, 17, 39, 56, 

57, 58, 60

Lepidophyllum cupressiforme (Lam.) Cass., 1, 6, 7 (Fig. 1J), 

8 (Fig. 2L), 9, 10 (Fig. 3A), 13, 14, 47, 56, 57–62, 88 (Fig. 

40C,D), 103 (Fig. 55)

Lepidophyllum cupressinum (Phil.) Kuntze, 86

Lepidophyllum lucidum (Meyen) Cabrera, 86

Lepidophyllum meyenii A. Gray nom. illeg., 86

Lepidophyllum phylicaeforme (Meyen) Hieron. ex R. E. Fr., 86

Lepidophyllum phylicaeforme var. resinosum (Walp.) Blake, 86

Lepidophyllum quadrangulare (Meyen) Benth. and Hook. f., 86 

Lepidophyllum rigidum (Wedd.) Benth. and Hook. f., 86

Lepidophyllum sect. Eulepidophyllum Cabrera nom. illeg., 

57, 60

Lepidophyllum teretiusculum Kuntze, 86

Lepidophyllum tola Cabrera, 86

Lithraea caustica Hook. and Arn., 79

Llerasia Triana, 1, 2 (Tbl. 1), 3, 5, 6, 15, 16

Lycium sp., 83

Microchaete humilis (Hook. f.) Sch. Bip., 62, 65

Nardophyllum (Hook. and Arn.) Hook. and Arn., 1, 2 (Tbl. 1), 

3–6, 13–15, 16, 17, 20, 27, 50, 54, 62–64, 83

Nardophyllum armatum (Wedd.) Reiche, 2 (Tbl. 1), 3, 62–

64, 86

Nardophyllum bracteolatum Hauman, 62, 63, 71, 73–75

Nardophyllum bryoides (Lam.) Cabrera, 1, 6, 7 (Fig. 1C), 8 

(Fig. 2M), 9, 11 (Fig. 4F–I), 13, 14, 63, 64–71, 87 (Fig. 39C), 

89 (Fig. 41C,D), 104 (Fig. 56), 105 (Fig. 57)

Nardophyllum cabrerae Bonif., 2 (Tbl. 1), 53, 54, 63, 64

Nardophyllum candollei (J. Rémy) F. Meigen nom. illeg., 78, 79

Nardophyllum chiliotrichioides (J. Rémy) A. Gray, 6, 7 (Fig. 

1B), 8 (Fig. 2N), 9, 10 (Fig. 3C), 13, 14, 63, 64, 71–76, 83, 

87 (Fig. 39A,B), 106 (Fig. 58), 107 (Fig. 59), 109

Nardophyllum darwinii (Hook. and Arn.) A. Gray, 65

Nardophyllum deserticola (Cabrera) G. L. Nesom, 17, 63

Nardophyllum genistoides (Phil.) A. Gray, 6, 8 (Fig. 2O), 9, 11 

(Fig. 4D), 13, 14, 62, 63, 64, 76, 77 (Fig. 33), 78 (Fig. 34), 

108 (Fig. 60)

Nardophyllum humile (Hook. f.) A. Gray, 62, 65, 68

Nardophyllum kingii (Hook. f.) A. Gray, 62, 65

Nardophyllum lanatum (Meyen) Cabrera, 6, 8 (Fig. 2P), 9, 13, 

14, 53, 54, 63, 64, 76, 78–83, 109 (Fig. 61)

Nardophyllum nardophyllum (Kuntze) Macloskie nom. illeg., 65

Nardophyllum obtusifolium Hook. and Arn., 62, 63, 65, 68 

Nardophyllum paniculatum Phil., 62, 86

Nardophyllum parvifolium Phil., 62, 65

Nardophyllum patagonicum (Cabrera) G. L. Nesom, 6, 7 (Fig. 

1I), 8 (Fig. 2Q), 9, 10 (Fig. 3E), 13, 14, 63, 64, 83–85, 89 

(Fig. 41A,B), 110 (Fig. 62)

Nardophyllum revolutum (D. Don ex Hook. and Arn.) Hook. 

and Arn., 62, 78

Nardophyllum scoparium Phil., 62, 63, 86

Nardophyllum staehelinoides (DC.) Macloskie, 62, 78

Nardus L., 64

Nassauvia Comm. ex Juss., 76

Nassauvia axillaris D. Don, 26, 73

Nassauvia glomerulosa D. Don, 54, 65, 73

Nassauvia planifolia Wedd., 76

Nassauvia ulicina Macloskie, 65, 73

Nothofagus spp., 1, 15, 43, 99

Novenia acaulis (Wedd. ex Benth. and Hook. f.) S. E. Freire 

and F. Hellwig, 85

Ocyroe Phil., 2 (Tbl. 1), 3–6, 16, 62

Ocyroe armata (Wedd.) Bonif., 86

Ocyroe spinosa Phil., 62, 86

Olearia Moench, 3, 4

Oritrophium (Kunth.) Cuatrec., 2 (Tbl. 1), 3, 5

Paleaepappus Cabrera, 2 (Tbl. 1), 3, 5, 63, 64, 83

Paleaepappus patagonicus Cabrera, 63, 83

Parastrephia lucida (Meyen) Cabrera, 85, 86

Parastrephia Nutt. emend Cabrera, 2 (Tbl. 1), 3–5, 28, 57, 60

Parastrephia quadrangularis (Meyen) Cabrera, 85, 86

Parastrephia teretiuscula (Kuntze) Cabrera, 86

Pentaphorus rosmarinifolius Gill. ex Hook. and Arn. pro. 

syn., 78

Pernettya mucronata Gaudich.ex G. Don., 43

Phyllochilium Cabrera, 21, 23

Phyllochilium densifolium (Phil.) Cabrera, 21, 24

Polyclados abietinus Phil., 86

Polyclados cupressinus Phil., 86

Polylepis spp., 37

Prosopis denudans Benth., 54

Prosopis fl exuosa var. depressa F. A. Roig, 19

Pteronia L., 2 (Tbl. 1), 3–5

Quillaja saponaria Molina, 79

Ribes magellanicum Poir., 43

Sarcocornia perennis (Mill.) A. J. Scott., 58

Schinus spp., 54

Senecio patagonicus Hook. and Arn., 58

Senna aphylla subsp. divaricata (Hieron.) L. Bravo, 19

Solidagininae O. Hoffm., 5

Stipa spp., 28, 35

Tetraglochin ameghinoii Speg., 54

Tetraglochin caespitosum Phil., 67

Tropidolepis diffusa (G. Forst.) Tausch, 38, 39

Tropidolepis Tausch, 38

Vernonia phylicaeformis var. resinosa Walp., 86

Zaluzania Pers., 21, 47
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