CLASSIFICATION AND PHYLOGENY OF SOUTH AMERICAN GNAPHALIEAE (ASTERACEAE) FROM MORPHOLOGY AND MOLECULAR DATA

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1 CLASSIFICATION AND PHYLOGENY OF SOUTH AMERICAN GNAPHALIEAE (ASTERACEAE) FROM MORPHOLOGY AND MOLECULAR DATA Susana E. Freire Instituto Darwinion, San Isidro, Argentina.

2 V Southamerican Gnaphalieae Achyrocline (Less.) DC., 32 spp. Anaphalis DC., c. 110 spp. Antennaria J. Gaertn., c. 40 spp. Belloa J. Rémy, 1 or 12 spp Berroa Beauverd, 1 sp Chevreulia Cass, 5 spp Chionolaena DC.,17 spp (incl. Gnaphaliothamnus Kirp., Leucopholis Gardn.) Cuatrecasasiella H. Rob. 2 spp.(= Luciliopsis Wedd.) Facelis Cass., 3 spp. Gamochaeta Wedd., c. 60 spp Gamochaetopsis Anderb. & S.E. Freire, 1 sp Jalcophila Dillon & Sagástegui, Loricaria Wedd., 19 spp Lucilia Cass. (incl. Oligandra Less.), 8 or 12 spp Luciliocline Anderb. & S.E. Freire, 5 or spp Raouliopsis S.F. Blake, 2 spp Micropsis DC., c. 5 spp Mniodes (A. Gray) Benth., 5 spp. Pseudognaphalium Kirp., c. 90 spp. Psilocarphus Nutt., 8 spp. Stenophalium Anderb. 3 spp Stuckertiella Beauverd, 2 spp. 22 genera 15 exclusively from South America

3 USING MOLECULAR AND MORPHOLOGICAL EVIDENCE: PHYLOGENY OF LUCILIA GROUP To test the monophyly of Lucilia Group To test the monophyly of each genus of Lucilia Group PHYLOGENY OF GAMOCHAETA To test the monophyly of Gamochaeta and to establish the relationships among its species

4 PHYLOGENY OF LUCILIA GROUP (GNAPHALIEAE, ASTERACEAE) INFERRED FROM MORPHOLOGY AND MOLECULAR DATA M. A. Chemisquy1, S. E. Freire1, S. G. Beck 2 y E. Urtubey1 1 Instituto Darwinion, Argentina. 2 Universidad Mayor de San Andrés, Bolivia

5 V Lucilia Group Merxmüller, Leins & Roessler Belloa J. Rémy, 12 especies Berroa Beauverd, 1 sp Chevreulia Cass, 5 spp Chionolaena DC.,17 spp Facelis Cass., 3 spp Gnaphaliothamnus Kirp., 1 sp Leucopholis Gardn., 4 spp Loricaria Wedd. 19 spp Lucilia Cass., 8 spp. Luciliopsis Wedd., 3 spp Mniodes (A. Gray) Benth., 5 spp Oligandra Less., 1 spp Raouliopsis S.F. Blake, 2 spp Jalcophila Dillon & Sagástegui, 2 spp Novenia S.E. Freire, 1 sp 13 géneros

6 SUBTRIBES OF GNAPHALIEAE Anderberg, 1991 LORICARIINAE RELHANIINAE CASSINIINAE ANGIANTHIINAE GNAPHALIINAE

7 Lucilia Group sensu Anderberg, 1991 Belloa J. Rémy, 12 spp Berroa Beauverd, 1 sp Chevreulia Cass., 5 spp Cuatrecasasiella H. Rob., 2 spp Facelis Cass., 3 spp Lucilia Cass., 11 spp. 7 géneros Anderberg, 1991 Jalcophila Dillon & Sagástegui, 2 spp

8 V Lucilia Group Merxmüller et al. 1977/Anderberg 1991 Belloa J. Rémy Berroa Beauverd Chevreulia Cass Cuatrecasasiella H. Rob (= Luciliopsis Wedd. ) Facelis Cass. Lucilia Cass. Jalcophila Dillon & Sagástegui Leucopholis Gardn. (= Chionolaeana DC.) Freire 1993 Oligandra Less. (= Lucilia Cass.) Freire 1989 Chionolaena DC. Gnaphaliothamnus Kirp. Raouliopsis S.F. Blake Mniodes (A. Gray) Benth. Loricaria Wedd. Novenia S.E. Freire CASSINIINAE LORICARIINAE ASTEREAE

9 Lucilia Group sensu Anderberg, 1991/Anderberg & Freire, 1991 Belloa J. Rémy Berroa Beauverd Chevreulia Cass. Cuatrecasasiella H. Rob. Facelis Cass. Lucilia Cass. Jalcophila Dillon & Sagástegui, 2 spp 3 7 géneros Anderberg, géneros Anderberg & Freire, 1991 Gamochaetopsis Anderberg & S.E. Freire, 1 sp Luciliocline Anderberg & S.E. Freire, 5 spp

10 GNAPHALIINAE - Lucilia Group, Anderberg & Freire 1991 SYNAPOMORPHIES OF LUCILIA GROUP Style branches dorsally pubescent Pappus bristles connate Florets purple

11 Lucilia Group sensu Dillon, 2000, 2003 Belloa de 9 1 sp Berroa 1 sp Chevreulia 5 spp Cuatrecasasiella 2 spp Facelis 3 spp Lucilia de 8 12 spp Jalcophila de 3 2 spp Gamochaetopsis 1 sp Luciliocline de 8 12 spp

12 METHODS 4 molecular markers: trnl-f and matk (chloroplast), ITS and ETS (from the nucleus) 2 Phylogenetic analyses: Molecular matrix composed of 4 markers Molecular matrix + morphological matrix (36 caracteres) Molecular matrices were aligned using the online application MAFFT Phylogenetic analyses were performed using the software TNT Branch support was evaluated using jackknife expressed as absolute values and as frequency differences (GC) Gaps were treated as missing data

13 TAXA In group Belloa (4) Berroa (1) Chevreulia (1) Cuatrecasasiella (1) Facelis (3) Gamochaetopsis (1) Jalcophila (1) Lucilia (4) Luciliocline (3) GNAPHALINEAS: Achyrocline, Edmondia, Gamochaeta, Gnaphalium Helichrysum, Lasiopogon, Leontopodium, Logfia, Pseudognaphalium, Stuckertiella, Stuartina, Vellereophyton, LORICARINEAS: Mniodes, Loricaria Out group CASSININEAS: Acanthocladium, Antennaria, Anaphalis, Anaxeton, Cassinia, Ewartia, Langebergia RHELANINEAS: Oedera, Relhania ASTEREAS: Novenia, Solidago, Chiliotrichium, Chiliophyllum, Baccharis, Aster

14 Combined trnl-f,matk, ITS and ETS Novenia, 1 sp - Argentina, Bolivia, Peru Tribe Astereae Tribe Gnaphalieae Lucilia Group

15 Combined trnl-f,matk, ITS and ETS CHEVREULIA, 5 spp - Argentina, Bolivia, Brasil, Chile, Paraguay, Uruguay, Falkland Islands Tribe Gnaphalieae Lucilia Group CUATRECASASIELLA, 2 spp Ecuador to Argentina

16 Combined trnl-f,matk, ITS and ETS Jalcophila. boliviensis Tribe Gnaphalieae Lucilia Group JALCOPHILA, 3 spp - Ecuador, Peru, Bolivia

17 Combined trnl-f,matk, ITS and ETS Tribe Gnaphalieae Lucilia Group LORICARIA, 19 spp - Colombia to Bolivia

18 Combined trnl-f,matk, ITS and ETS Luciliocline subspicata LUCILIOCLINE, 5 spp - Peru, Bolivia, Argentina Tribe Gnaphalieae Lucilia Group Belloa schultzii BELLOA, 9 spp Venezuela to Chile

19 Combined trnl-f,matk, ITS and ETS MNIODES, 4 spp Peru, Bolivia Tribe Gnaphalieae Lucilia Group BELLOA, 9 spp Venezuela to Chile

20 Combined trnl-f,matk, ITS and ETS STUCKERTIELLA, 1 sp - Argentina, Bolivia, Peru GAMOCHAETA c. 60 spp-new World Tribe Gnaphalieae Lucilia Group GAMOCHAETOPSIS 1sp - Argentina, Chile

21 Combined trnl-f,matk, ITS and ETS Belloa chilensis Argentina, Chile Tribe Gnaphalieae Lucilia Group FACELIS, 3 spp - Argentina, Bolivia, Brazil, Chile, Paraguay, Peru, Uruguay

22 Combined trnl-f,matk, ITS and ETS LUCILIA 8 spp -Argentina, Bolivia, Brasil, Chile, Paraguay, Uruguay Tribe Gnaphalieae Lucilia Group BERROA 1 sp Argentina, Uruguay

23 Combined Molecular and Morphological Evidence Tribu Astereae Tribu Gnaphalieae Lucilia Group

24 Combined Molecular and Morphological Evidence Tribu Gnaphalieae Lucilia Group

25 PHYLOGENY OF GAMOCHAETA (GNAPHALIEAE, ASTERACEAE) INFERRED FROM MORPHOLOGY AND MOLECULAR DATA E. Urtubey1, S. E. Freire1, A. López1, M. A. Chemisquy1, L. Paz Deble 2 & N. Bayón3 1 Instituto Darwinion, Argentina. 2 Universidade da Região da Campanha, RS, Brasil. 3 Universidad Nacional de La Plata, Argentina.

26 METHODS 2 molecular markers: ITS and ETS (from the nucleus) Taxon sampling: 27 species of Gamochaeta Phylogenetic analysis: Molecular matrices were aligned using the online application MAFFT Phylogenetic analyses were performed using the software TNT Branch support was evaluated using jackknife expressed as absolute values and as frequency differences (GC) Gaps were treated as missing data

27 Combined ETS-ITS Belloa chilensis Argentina, Chile

28 Combined ETS-ITS/ ETS ETS ETS-ITS STUCKERTIELLA, 1 sp Argentina, Bolivia, Peru

29 Combined ETS-ITS/ ETS ETS-ITS ETS GAMOCHAETOPSIS,1 sp Argentina - Chile

30 Combined ETS-ITS/ ETS Gamochaeta nivalis - Argentina, Chile GAMOCHAETOPSIS,1 sp Argentina - Chile

31 Combined ETS-ITS G. axillaris G. filaginea G. nigrevestis G. platensis G. subfalcata Gamochaeta filaginea, Argentina-Brazil- Uruguay

32 Combined ETS-ITS Gamochaeta purpurea América del Norte Gamochaeta neuquensis Chile-Argentina

33 Combined ETS-ITS Gamochaeta lulioana - Bolivia Gamochaeta longepedicellata Argentina, Bolivia

34 Combined ETS-ITS Gamochaeta grazielae - Brazil Gamochaeta stachydifolia - Argentina, Brazil

35 CONCLUSIONS Lucilia Group 1. Lucilia group sensu Anderb. is paraphyletic. 2. Molecular + Morphological evidence confirm the monophyly of 8 of 9 genera of Lucilia group sensu Anderb.: Berroa, Chevreulia, Cuatrecasasiella, Facelis, Gamochaetopsis, Jalcophila, Lucilia, Luciliocline sensu Anderb. & Freire. 3. Molecular analysis: Mniodes (Loricariinae) forms a monophyletic group with Belloa sensu Anderb. & Freire (excl. Type) and Luciliocline sensu Anderb. & Freire. 4. Molecular + Morphological characters: Jalcophila boliviensis belongs to genus Jalcophila. 5. Molecular and Molecular + Morphological confirm: Belloa as a monotypic genus.

36 6. Molecular Evidence: Two clades Cuatrecasasiella Chevreulia * Jalcophila Loricaria Luciliocline Belloas Mniodes Gamochaeta * Stucke rtiella* Gamochaetopsis Belloa Facelis * Berroa Lucilia *

37 WORK IN PROGRESS: PHYLOGENY OF LUCILIA GROUP To add Micropsis (Gnaphaliinae) and Chionolaeana (Cassiniinae) PHYLOGENY OF GAMOCHAETA To add species from Chile To combine molecular and morphological evidence

38 AGRADECIMIENTOS ACKNOWLEDGMENTS Finnacial support was provided by Consejo Nacional de InvestigacionesCientíficas y Técnicas (CONICET) Argentina- PIP To C. Ezcurra, (UNCOMA) and F. Zuloaga et al. (SI), for specimens of Belloa chilensis, and Belloa piptolepis, respectively. To J. M. Bonifacino (UDeLaR) for his photograph of Chevreulia sarmentosa.