CLASSIFICATION AND PHYLOGENY OF SOUTH AMERICAN GNAPHALIEAE (ASTERACEAE) FROM MORPHOLOGY AND MOLECULAR DATA
Susana E. Freire
Instituto Darwinion, San Isidro, Argentina. 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. 22 genera • Gamochaeta Wedd., c. 60 spp 15 • Gamochaetopsis Anderb. & S.E. Freire, 1 sp • Jalcophila Dillon & Sagástegui, exclusively • Loricaria Wedd., 19 spp from South • Lucilia Cass. (incl. Oligandra Less.), 8 or 12 spp America • Luciliocline Anderb. & S.E. Freire, 5 or 13-16 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.
S.E.Freire SOUTH AMERICAN GNAPHALIEAE (ASTERACEAE) 5/12/2011 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
S.E.Freire SOUTH AMERICAN GNAPHALIEAE (ASTERACEAE) 5/12/2011 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
S.E.Freire SOUTH AMERICAN GNAPHALIEAE (ASTERACEAE) 5/12/2011 V “Lucilia Group” Merxmüller, Leins & Roessler 1977
• 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 13 géneros • 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 1986 • Novenia S.E. Freire, 1 sp
S.E.Freire SOUTH AMERICAN GNAPHALIEAE (ASTERACEAE) 5/12/2011
SUBTRIBES OF GNAPHALIEAE Anderberg, 1991
LORICARIINAE
RELHANIINAE
CASSINIINAE
ANGIANTHIINAE
GNAPHALIINAE
S.E.Freire SOUTH AMERICAN GNAPHALIEAE (ASTERACEAE) 5/12/2011 “Lucilia Group” sensu Anderberg, 1991
• Belloa J. Rémy, 12 spp • Berroa Beauverd, 1 sp • Chevreulia Cass., 5 spp • Cuatrecasasiella H. Rob., 2 spp 7 géneros • Facelis Cass., 3 spp Anderberg, • Lucilia Cass., 11 spp. 1991 • Jalcophila Dillon & Sagástegui, 2 spp
S.E.Freire SOUTH AMERICAN GNAPHALIEAE (ASTERACEAE) 5/12/2011 V “Lucilia Group” Merxmüller et al. 1977/Anderberg 1991
• Belloa J. Rémy • Berroa Beauverd • Chevreulia Cass • Cuatrecasasiella H. Rob. 1985 (= Luciliopsis Wedd. ) • Facelis Cass. • Lucilia Cass. • Jalcophila Dillon & Sagástegui • Leucopholis Gardn. (= Chionolaeana DC.) – Freire 1993 • Oligandra Less. (= Lucilia Cass.) – Freire 1989
• Chionolaena DC. CASSINIINAE • Gnaphaliothamnus Kirp. “ • Raouliopsis S.F. Blake • Mniodes (A. Gray) Benth. LORICARIINAE • Loricaria Wedd. • Novenia S.E. Freire ASTEREAE
S.E.Freire SOUTH AMERICAN GNAPHALIEAE (ASTERACEAE) 5/12/2011 “Lucilia Group” sensu Anderberg, 1991/Anderberg & Freire, 1991
• Belloa J. Rémy • Berroa Beauverd • Chevreulia Cass. • Cuatrecasasiella H. Rob. 7 géneros 9 • Facelis Cass. Anderberg, géneros 1991 • Lucilia Cass. Anderberg & Freire, 1991 • Jalcophila Dillon & Sagástegui, 2 spp 3
• Gamochaetopsis Anderberg & S.E. Freire, 1 sp • Luciliocline Anderberg & S.E. Freire, 5 spp
S.E.Freire SOUTH AMERICAN GNAPHALIEAE (ASTERACEAE) 5/12/2011
GNAPHALIINAE - “Lucilia Group”, Anderberg & Freire 1991
SYNAPOMORPHIES OF “LUCILIA GROUP” • Style branches dorsally pubescent
• Pappus bristles connate
• Florets purple
S.E.Freire SOUTH AMERICAN GNAPHALIEAE (ASTERACEAE) 5/12/2011 “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
S.E.Freire SOUTH AMERICAN GNAPHALIEAE (ASTERACEAE) 5/12/2011
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
S.E.Freire SOUTH AMERICAN GNAPHALIEAE (ASTERACEAE) 5/12/2011 TAXA Belloa (4) Berroa (1) Chevreulia (1) Cuatrecasasiella (1) Facelis (3) In group Gamochaetopsis (1) Jalcophila (1) Lucilia (4) Luciliocline (3)
GNAPHALINEAS: Achyrocline, Edmondia, Gamochaeta, Gnaphalium Helichrysum, Lasiopogon, Leontopodium, Logfia, Pseudognaphalium, Stuckertiella, Stuartina, Vellereophyton,
LORICARINEAS: Mniodes, Loricaria
CASSININEAS: Acanthocladium, Antennaria, Anaphalis, Out group Anaxeton, Cassinia, Ewartia, Langebergia
RHELANINEAS: Oedera, Relhania
ASTEREAS: Novenia, Solidago, Chiliotrichium, Chiliophyllum, Baccharis, Aster
S.E.Freire SOUTH AMERICAN GNAPHALIEAE (ASTERACEAE) 5/12/2011
Combined trnL-F,matK, ITS and ETS
Novenia, 1 sp - Argentina, Bolivia, Peru
Tribe Astereae Tribe Gnaphalieae Lucilia Group
S.E.Freire SOUTH AMERICAN GNAPHALIEAE (ASTERACEAE) 5/12/2011
Combined trnL-F,matK, ITS and ETS
CHEVREULIA , 5 spp - Argentina, Bolivia, Brasil, Chile, Paraguay, Uruguay, Falkland Islands
Tribe Gnaphalieae CUATRECASASIELLA, 2 spp Lucilia Group Ecuador to Argentina
S.E.Freire SOUTH AMERICAN GNAPHALIEAE (ASTERACEAE) 5/12/2011
Combined trnL-F,matK, ITS and ETS
Jalcophila. boliviensis
Tribe Gnaphalieae Lucilia Group JALCOPHILA, 3 spp - Ecuador, Peru, Bolivia
S.E.Freire SOUTH AMERICAN GNAPHALIEAE (ASTERACEAE) 5/12/2011
Combined trnL-F,matK, ITS and ETS
Tribe Gnaphalieae Lucilia Group LORICARIA, 19 spp - Colombia to Bolivia
S.E.Freire SOUTH AMERICAN GNAPHALIEAE (ASTERACEAE) 5/12/2011
Combined trnL-F,matK, ITS and ETS
Luciliocline subspicata
LUCILIOCLINE , 5 spp - Peru, Bolivia, Argentina
Tribe Gnaphalieae Belloa Lucilia Group schultzii
BELLOA, 9 spp – Venezuela to Chile
S.E.Freire SOUTH AMERICAN GNAPHALIEAE (ASTERACEAE) 5/12/2011
Combined trnL-F,matK, ITS and ETS
MNIODES, 4 spp – Peru, Bolivia
Tribe Gnaphalieae Lucilia Group BELLOA, 9 spp – Venezuela to Chile
S.E.Freire SOUTH AMERICAN GNAPHALIEAE (ASTERACEAE) 5/12/2011
Combined trnL-F,matK, ITS and ETS
STUCKERTIELLA, 1 sp - Argentina, Bolivia, Peru
GAMOCHAETA c. 60 spp-New World
Tribe Gnaphalieae GAMOCHAETOPSIS 1sp - Argentina, Chile Lucilia Group
S.E.Freire SOUTH AMERICAN GNAPHALIEAE (ASTERACEAE) 5/12/2011
Combined trnL-F,matK, ITS and ETS
Belloa chilensis– Argentina, Chile
Tribe Gnaphalieae FACELIS, 3 spp - Argentina, Bolivia, Lucilia Group Brazil, Chile, Paraguay, Peru, Uruguay
S.E.Freire SOUTH AMERICAN GNAPHALIEAE (ASTERACEAE) 5/12/2011
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
S.E.Freire SOUTH AMERICAN GNAPHALIEAE (ASTERACEAE) 5/12/2011 Combined Molecular and Morphological Evidence
Tribu Astereae Tribu Gnaphalieae Lucilia Group
S.E.Freire SOUTH AMERICAN GNAPHALIEAE (ASTERACEAE) 5/12/2011 Combined Molecular and Morphological Evidence
Tribu Gnaphalieae Lucilia Group
S.E.Freire SOUTH AMERICAN GNAPHALIEAE (ASTERACEAE) 5/12/2011 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.
S.E.Freire SOUTH AMERICAN GNAPHALIEAE (ASTERACEAE) 5/12/2011
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
S.E.Freire SOUTH AMERICAN GNAPHALIEAE (ASTERACEAE) 5/12/2011 Combined ETS-ITS
Belloa chilensis – Argentina, Chile
S.E.Freire SOUTH AMERICAN GNAPHALIEAE (ASTERACEAE) 5/12/2011 Combined ETS-ITS/ ETS ETS
ETS-ITS
STUCKERTIELLA, 1 sp Argentina, Bolivia, Peru
S.E.Freire SOUTH AMERICAN GNAPHALIEAE (ASTERACEAE) 5/12/2011 Combined ETS-ITS/ ETS ETS-ITS
ETS
GAMOCHAETOPSIS ,1 sp Argentina - Chile
S.E.Freire SOUTH AMERICAN GNAPHALIEAE (ASTERACEAE) 5/12/2011 Combined ETS-ITS/ ETS
Gamochaeta nivalis - Argentina, Chile
GAMOCHAETOPSIS ,1 sp Argentina - Chile
S.E.Freire SOUTH AMERICAN GNAPHALIEAE (ASTERACEAE) 5/12/2011 Combined ETS-ITS
G. axillaris G. filaginea G. nigrevestis G. platensis Gamochaeta filaginea, Argentina-Brazil- G. subfalcata Uruguay
S.E.Freire SOUTH AMERICAN GNAPHALIEAE (ASTERACEAE) 5/12/2011 Combined ETS-ITS
Gamochaeta purpurea América del Norte
Gamochaeta neuquensis Chile-Argentina
S.E.Freire SOUTH AMERICAN GNAPHALIEAE (ASTERACEAE) 5/12/2011 Combined ETS-ITS
Gamochaeta lulioana - Bolivia
Gamochaeta longepedicellata Argentina, Bolivia
S.E.Freire SOUTH AMERICAN GNAPHALIEAE (ASTERACEAE) 5/12/2011 Combined ETS-ITS
Gamochaeta grazielae - Brazil
Gamochaeta stachydifolia - Argentina, Brazil
S.E.Freire SOUTH AMERICAN GNAPHALIEAE (ASTERACEAE) 5/12/2011 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.
S.E.Freire SOUTH AMERICAN GNAPHALIEAE (ASTERACEAE) 5/12/2011
6. Molecular Evidence: Two clades
Cuatrecasasiella Chevreulia * Jalcophila Loricaria Luciliocline “Belloas” Mniodes Gamochaeta *Stucke rtiella* Gamochaetopsis Belloa Facelis * Berroa Lucilia *
S.E.Freire SOUTH AMERICAN GNAPHALIEAE (ASTERACEAE) 5/12/2011 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
S.E.Freire SOUTH AMERICAN GNAPHALIEAE (ASTERACEAE) 5/12/2011
AGRADECIMIENTOS ACKNOWLEDGMENTS
• • • Finnacial support was provided by Consejo Nacional de InvestigacionesCientíficas y Técnicas (CONICET) Argentina- PIP 112-200801-02196.
• 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.
S.E.Freire SOUTH AMERICAN GNAPHALIEAE (ASTERACEAE) 5/12/2011