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The systematic revision of Chaetanthera Ruiz & Pav., and the reinstatement of the genus Oriastrum Poepp. & Endl. (Asteraceae; Mutisieae) [Elektronische Ressource] / vorgelegt von Alison Margarete Robertson Davies

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A systematic revision of Chaetanthera Ruiz & Pav., and the reinstatement of Oriastrum Poepp. & Endl. (Asteraceae: Mutisieae) Alison Margaret Robertson Davies München 2010 A systematic revision of Chaetanthera Ruiz & Pav., and the reinstatement of Oriastrum Poepp. & Endl. (Asteraceae: Mutisieae) Dissertation der Fakultät für Biologie der Ludwig-Maximilians-Universität München vorgelegt von Alison Margaret Robertson Davies München, den 03. November 2009 Erstgutachter: Prof. Dr. Jürke Grau Zweitgutachter: Prof. Dr. Günther Heubl Tag der mündlichen Prüfung: 28. April 2010 For Ric, Tim, Isabel & Nicolas Of all the floures in the meade, Thanne love I most those floures white and rede, Such as men callen daysyes. CHAUCER, ‘Legend of Good Women’, Prol. 43 (c. 1385) “…a traveller should be a botanist, for in all views plants form the chief embellishment.” DARWIN, ‘Darwin’s Journal of a Voyage round the World’, p. 599 (1896) Acknowledgements The successful completion of this work is due in great part to numerous people who have contributed both directly and indirectly. Thank you. Especial thanks goes to my husband Dr.

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A systematic revision of
Chaetanthera Ruiz & Pav., and the
reinstatement of Oriastrum Poepp.
& Endl. (Asteraceae: Mutisieae)


Alison Margaret Robertson Davies



















München 2010








A systematic revision of
Chaetanthera Ruiz & Pav.,
and the reinstatement of
Oriastrum Poepp. & Endl.
(Asteraceae: Mutisieae)













Dissertation der Fakultät für Biologie
der Ludwig-Maximilians-Universität
München




vorgelegt von
Alison Margaret Robertson Davies






München, den 03. November 2009































Erstgutachter: Prof. Dr. Jürke Grau
Zweitgutachter: Prof. Dr. Günther Heubl
Tag der mündlichen Prüfung: 28. April 2010

























For Ric, Tim, Isabel & Nicolas




Of all the floures in the meade,
Thanne love I most those floures white and rede,
Such as men callen daysyes.

CHAUCER, ‘Legend of Good Women’, Prol. 43 (c. 1385)


“…a traveller should be a botanist, for in all views plants form the chief embellishment.”

DARWIN, ‘Darwin’s Journal of a Voyage round the World’, p. 599 (1896)










Acknowledgements

The successful completion of this work is due in great part to numerous people who have
contributed both directly and indirectly. Thank you. Especial thanks goes to my husband Dr.
Ric Davies who has provided unwavering support and encouragement throughout.

I am deeply indebted to my supervisor, Jürke Grau, who made this research possible. Thank
you for your support and guidance, and for your compassionate understanding of wider
issues.

The research for this study was funded by part-time employment on digital archiving projects
coordinated via the Botanische Staatssammlung Munchen (INFOCOMP, 2000 – 2003; API-
Projekt, 2005). Appreciative thanks go to the many friends and colleagues from both the
Botanische Staatssammlung and the Botanical Institute who have provided scientific and
social support over the years. Philomena Bodensteiner, Peter Döbbeler, Tini Ehrhart,
Matthias Erben and Lilo Klingenberg deserve a special mention.

Much of the evidence supporting the final conclusions in the thesis was acquired through
laboratory research and S.E.M. imaging. This would not have been possible without the
patient schooling and friendly assistance of Mrs. E. Vosyka, Dr. E. Facher and Mr E.
Marksteiner.

Travel grants were received from the DAAD (the German Academic Exchange Service) to
visit Chile twice, in 2001 and 2002. My stay in Chile and the field excursions were made far
easier thanks to Sunke Nef & family, Diettrich & Arnulf Becker, and Melica Muñoz-Schick
in Santiago, and Roberto Rodriguez, Señora Gleisner & family, and Marcelo Baeza & family
in Concepción. Special thanks are extended to Mary Kalin Arroyo, an active botanist and
ecologist in Chile, who has an ongoing scientific interest in Chaetanthera.

Permission to use digital images was kindly granted by Michal Belov (www.Chileflora.com),
Mauricio Bonifacino (MVFA, Uruguay), Michael Dillon, María Terese Eyzaguirre
(Fundación R.A. Philippi), Jürke Grau, Michel Grenon, Irene Till-Bottraud (Station Alpin
Joseph Fourier, Grenoble) and Mauricio Zuñiga.

Finally, a huge “Thank you” to my parents Jean & Iain Robertson, and my parents-in-law
Elizabeth & Peter (sadly recently died).
1

Contents
Abstract .................................................................................................................................. 5 
Zusammenfassung .................................................................................................................. 6 
List of Figures ........................................................................................................................ 7 
List of Tables ....................................................................................................................... 12 
I  Introduction ...................................................................................................................... 15 
1  Philosophy of taxonomic research ............................................................................... 15 
2  Goals of taxonomic research ........................................................................................ 16 
3  Application to Chaetanthera & Oriastrum .................................................................. 16 
3.1  Historical background .......................................................................................... 16 
3.2  Novel sources of variation ................................................................................... 17 
3.3  Re-defining taxonomic relationships ................................................................... 18 
II  Materials, methods & measurements 20 
1  Materials ...................................................................................................................... 20 
2  Methods........................................................................................................................ 20 
2.1  Capitula & floral dissection ................................................................................. 20 
2.2  Involucral bract anatomy studies ......................................................................... 20 
2.3  Achene & pappus studies ..................................................................................... 20 
2.4  Pollen studies ....................................................................................................... 21 
2.5  Chromosome studies ............................................................................................ 21 
2.6  Images & illustrations .......................................................................................... 22 
2.7  Geographical data ................................................................................................ 22 
3  Measurements .............................................................................................................. 22 
III  Variation of characters in Chaetanthera & Oriastrum .................................................... 24 
1  General remarks ........................................................................................................... 24 
2  Morphological & anatomical variation ........................................................................ 24 
2.1  Life cycle & habit 24 
2.2  Leaves .................................................................................................................. 29 
2.3  Indumentum ......................................................................................................... 30 
2.4  Involucral bracts................................................................................................... 30 
2.4.1  Arrangement .................................................................................................... 30 
2.4.2  Morphology of the inner involucral bracts ...................................................... 31 
2.4.3  Anatomy of the inner involucral bracts ........................................................... 33 
2.5  Pappus .................................................................................................................. 34 
2.6  Floret variation: ray dimorphism & showiness .................................................... 36 
2.7  Anthers & pollen .................................................................................................. 38 
2.8  Styles & stigmas 41 
2.9  Achenes ................................................................................................................ 43 
2.9.1  Achene shape ................................................................................................... 43 
2.9.2  Carpopodium.................................................................................................... 44 
2.9.3  Achene hairs..................................................................................................... 44 
2.9.4  Achene testa epidermis .................................................................................... 48 
3  Cytological variation .................................................................................................... 50 
4  Genetic variation .......................................................................................................... 54 
IV  Form, function & habitat.................................................................................................. 56 
1  Introduction .................................................................................................................. 56 
2  Homoplasy ................................................................................................................... 56 
3  Habitats & stress 57 
4  Form & function 57 
4.1  Rosettes, succulence & sclerophylly in Chaetanthera ........................................ 57  2

4.2  Cushions, stem buds & scleroid bracts in Oriastrum .......................................... 58 
4.3  Fragile pappus & indehiscent achenes: dispersal brakes ..................................... 59 
4.4  Xeromorphic adaptations in pollen, pericarp & testa epidermis .......................... 60 
4.5  Competitive ability............................................................................................... 60 
4.6  Reproductive strategy .......................................................................................... 61 
V  The biogeography of Chaetanthera & Oriastrum ........................................................... 63 
1  The distribution of ................................................................................. 63 
1.1  Species distributions ............................................................................................ 63 
1.1.1  Species endemic to the Altiplano ..................................................................... 63 
1.1.2  Species endemic to Chile ................................................................................. 63 
1.1.3  Species from Chile & Argentina ...................................................................... 64 
1.2  Species hotspots ................................................................................................... 64 
1.2.1  The diversity hotspot “Coquimbo” (29° - 30°S) .............................................. 66 
1.2.2  The diversity hotspot “Santiago” (33° - 34°S) ................................................. 67 
1.3  Trichome diversity: evidence of species radiation ............................................... 67 
1.4  Lamina morphology & genetic affinity: evidence of species radiations ............. 70 
1.4.1  Linear-leaved group ......................................................................................... 70 
1.4.2  Dentate-ciliate leaved group. ........................................................................... 73 
2  The distribution of Oriastrum ...................................................................................... 75 
2.1  Species distributions ............................................................................................ 75 
2.1.1  Species endemic to either Argentina or Chile or Peru ..................................... 76 
2.1.2  Species from Argentina & Chile ...................................................................... 77 
2.1.3  Species from Argentina, Chile, Bolivia & Peru ............................................... 77 
2.2  Species hotspots ................................................................................................... 77 
2.3  Species radiations................................................................................................. 78 
2.3.1  Santiago............................................................................................................ 78 
2.3.2  Altiplano group ................................................................................................ 78 
2.3.3  Altoandino group ............................................................................................. 81 
2.3.4  Andino group 81 
VI  “The variety of all things forms a pleasure” .................................................................... 84 
1  Introduction .................................................................................................................. 84 
2  C. glabrata: polymorphism & the El Niño effect ........................................................ 85 
2.1  Introduction .......................................................................................................... 85 
2.2  Variation & distribution ....................................................................................... 85 
2.3  El Niño Southern Oscillation (ENSO) ................................................................. 88 
2.4  Results: observed phenotypic plasticity ............................................................... 88 
2.5  Conclusion ........................................................................................................... 91 
3  C. linearis & C. albiflora: active hybridisation ........................................................... 92 
3.1 92 
3.2  Variation & distribution 92 
3.3  Results: hybrid zones ........................................................................................... 93 
3.4  Conclusions .......................................................................................................... 96 
4  Perennial, scapose Chaetanthera: polymorphism or hybridisation ............................. 96 
4.1  Introduction .......................................................................................................... 96 
4.2  Variation & distribution ....................................................................................... 97 
4.3  Analysing variation: are the traditional characters truly diagnostic? ................... 97 
4.3.1  Results .............................................................................................................. 98 
4.3.2  Summary .......................................................................................................... 99 
4.4  Analysing variation: principle components analysis ......................................... 100 
4.4.1  Methods.......................................................................................................... 100  3

4.4.2  Results ............................................................................................................ 102 
4.5  Analysing variation: HYWIN ............................................................................ 104 
4.5.1  Results of the HYWIN analysis ..................................................................... 105 
4.5.2  Discussion ...................................................................................................... 108 
4.5.3  Summary ........................................................................................................ 108 
4.6  Extinction or botanist’s pride? ........................................................................... 109 
VII  Discussion & conclusions of systematic analysis ...................................................... 111 
1  Uniting characters - grouping species ........................................................................ 111 
2  Evolutionary adaptation to environmental stress ....................................................... 115 
3  Current dynamic change in Chaetanthera ................................................................. 116 
4  Conclusions ................................................................................................................ 119 
VIII  Chaetanthera & Oriastrum: an historic review ......................................................... 123 
1  Introduction ................................................................................................................ 123 
2  Chronological changes in Chaetanthera & Oriastrum .............................................. 123 
3  Nomenclatural notes for & ................................................ 126 
3.1  Chaetanthera ciliata: lectotype of Chaetanthera Ruiz & Pav. .......................... 126 
3.2  Oriastrum pusillum: generitype of Oriastrum Poepp. & Endl. ......................... 127 
3.3  David Don: nomenclatural types & nomenclatural obscurity ............................ 127 
3.3.1  Don’s nome............................................................................ 127 
3.3.2  Nomenclatural obscurity of Don names ........................................................ 128 
3.4  Rudolfo Philippi: lectotypes & nomina nuda ..................................................... 128 
IX  Chaetanthera Ruíz & Pav. ............................................................................................. 130 
1  Diagnosis.................................................................................................................... 130 
2  Subgeneric division of Chaetanthera ........................................................................ 131 
2.1  Chaetanthera Ruíz & Pav. subgenus Chaetanthera .......................................... 131 
2.2  subgenus Tylloma (D.Don) Less. ................................................ 132 
3  Key to the species ...................................................................................................... 133 
4  Species descriptions ................................................................................................... 136 
4.1  ChaetantheraChaetanthera136 
1.  albiflora (Phil.) A.M.R. Davies .......................................... 136 
2.  Chaetanthera chilensis (Willd.) DC .......................................................... 140 
3.  Chaetanthera ciliata Ruiz & Pav.. ............................................................. 146 
4.  Chaetanthera depauperata (Hook. & Arn.) A.M.R. Davies ..................... 149 
5.  Chaetanthera elegans s.l. Phil ................................................................... 152 
6.  Chaetanthera glandulosa J. Rémy var. glandulosa. .................................. 156 
7.  Chaetanthera glandulosa var. gracilis A.M.R. Davies ............................. 159 
8.  Chaetanthera incana Poepp. ex Less......................................................... 161 
9.  Chaetanthera linearis Poepp. ex Less. ...................................................... 164 
10.  Chaetanthera linearis x albiflora .......................................................... 167 
11.  Chaetanthera microphylla (Cass.) Hook. & Arn. .................................. 168 
12.  Chaetanthera moenchioides Less. ......................................................... 171 
13.  Chaetanthera multicaulis DC. ............................................................... 176 
14.  Chaetanthera perpusilla (Wedd.) Anderb. & S. E. Freire. .................... 179 
15.  Chaetanthera peruviana A. Gray .......................................................... 181 
16.  Chaetanthera ramosissima D. Don ex Taylor & Phillips ...................... 184 
17.  Chaetanthera x serrata Ruíz & Pav. ..................................................... 187 
18.  Chaetanthera taltalensis (Cabrera) A.M.R. Davies............................... 190 
4.2  Chaetanthera subgenus Tylloma (D.Don) Less. ................................................ 193 
19.  Chaetanthera euphrasioides (DC.) F. Meigen ...................................... 193 
20.  Chaetanthera flabellata D. Don ex Taylor & Phillips. .......................... 196  4

21.  Chaetanthera flabellifolia Cabrera ........................................................ 198 
22.  Chaetanthera frayjorgensis A.M.R. Davies .......................................... 200 
23.  Chaetanthera glabrata (DC) F. Meigen ................................................ 203 
24.  Chaetanthera kalinae A.M.R.Davies..................................................... 207 
25.  Chaetanthera limbata (D. Don) Less. ................................................... 209 
26.  Chaetanthera philippii B. L. Rob. 211 
27.  Chaetanthera pubescens A.M.R. Davies. .............................................. 214 
28.  Chaetanthera renifolia (J. Rémy) Cabrera. ........................................... 217 
29.  Chaetanthera schroederi G. F. Grandjot & K. Grandjot ....................... 219 
30.  Chaetanthera spathulifolia Cabrera. ...................................................... 222 
31.  Chaetanthera splendens (J. Rémy) B. L. Rob. ...................................... 225 
32.  Chaetanthera villosa D. Don ex Taylor & Phillips. .............................. 227 
X  Oriastrum Poepp. & Endl. ............................................................................................. 230 
1  Diagnosis.................................................................................................................... 230 
2  Subgeneric division of Oriastrum .............................................................................. 231 
2.1  Oriastrum Poepp. & Endl. subgenus Oriastrum ................................................ 231 
2.2 Egania (J.Rémy) .................................... 232 
3  Key to the species ...................................................................................................... 233 
4  Species descriptions ................................................................................................... 235 
4.1  Oriastrum subgenus Oriastrum ......................................................................... 235 
1.  Oriastrum chilense (J. Rémy) Wedd. ........................................................ 235 
2.  Oriastrum gnaphalioides (J. Rémy) Wedd.. .............................................. 238 
3.  Oriastrum lycopodioidesy) Wedd. ............................................... 242 
4.  Oriastrum pusillum Poepp. & Endl.. ......................................................... 245 
5.  Oriastrum werdermannii A.M.R.Davies ................................................... 248 
4.2  Oriastrum subgenus Egania .............................................................................. 250 
6.  Oriastrum abbreviatum (Cabrera) A.M.R. Davies .................................... 250 
7.  Oriastrum acerosum (J. Rémy) Phil. 253 
8.  Oriastrum achenohirsutum (Tombesi) A.M.R. Davies ............................. 256 
9. piculatum (J. Rémy) A.M.R. Davies ...................................... 258 
10.  Oriastrum cochlearifolium A. Gray. ...................................................... 261 
11.  Oriastrum dioicum (J. Rémy) Phil.. ....................................................... 263 
12.  Oriastrum famatinae A.M.R. Davies ..................................................... 266 
13.  Oriastrum polymallum Phil.................................................................... 268 
14. ulvinatum Phil. ................................................................... 272 
15.  Oriastrum revolutum (Phil.) A.M.R. Davies ......................................... 275 
16.  Oriastrum stuebelii (Hieron.) A.M.R. Davies var. stuebelii .................. 279 
17.  var. cryptum A.M.R.Davies. .................................. 282 
18.  Oriastrum tarapacensis A.M.R. Davies ................................................ 285 
19.  Oriastrum tontalensis................................................... 287 
XI  Summary of taxonomic revision .................................................................................... 290 
1  Changes in the nomenclature of Chaetanthera & Oriastrum .................................... 290 
2  Updates in the descriptive taxonomy of Chaetanthera & Oriastrum species ........... 290 
3  Additional information concerning & Oriastrum ............................... 291 
XII  References .................................................................................................................. 292 
XIII  Appendix 1 ................................................................................................................. 300