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CLASSIFICATION AND PHYLOGENY OF SOUTH AMERICAN
GNAPHALIEAE (ASTERACEAE)
FROM MORPHOLOGY AND MOLECULAR DATA
Susana E. Freire
Instituto Darwinion, San Isidro, Argentina.
• 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 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.
22 genera
15
exclusively
from South
America
V Southamerican Gnaphalieae
S.E.Freire SOUTH AMERICAN GNAPHALIEAE (ASTERACEAE) 5/12/2011
PHYLOGENY OF GAMOCHAETA
PHYLOGENY OF “LUCILIA GROUP”
S.E.Freire SOUTH AMERICAN GNAPHALIEAE (ASTERACEAE) 5/12/2011
• To test the monophyly of “Lucilia Group”
• To test the monophyly of each genus of “Lucilia Group”
• To test the monophyly of Gamochaeta and to
establish the relationships among its species
USING MOLECULAR AND MORPHOLOGICAL
EVIDENCE:
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
• 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
V “Lucilia Group” Merxmüller, Leins & Roessler 1977
1986
S.E.Freire SOUTH AMERICAN GNAPHALIEAE (ASTERACEAE) 5/12/2011
LORICARIINAE
RELHANIINAE
CASSINIINAE
ANGIANTHIINAE
GNAPHALIINAE
SUBTRIBES OF GNAPHALIEAE Anderberg, 1991
S.E.Freire SOUTH AMERICAN GNAPHALIEAE (ASTERACEAE) 5/12/2011
• 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.
• Jalcophila Dillon & Sagástegui, 2 spp
“Lucilia Group” sensu Anderberg, 1991
7
géneros Anderberg,
1991
S.E.Freire SOUTH AMERICAN GNAPHALIEAE (ASTERACEAE) 5/12/2011
• Belloa J. Rémy
• Berroa Beauverd
• Chevreulia Cass
• Cuatrecasasiella H. Rob. 1985 (= Luciliopsis Wedd. )
• Facelis Cass.
• Lucilia Cass.
• Jalcophila Dillon & Sagástegui
• Novenia S.E. Freire
• Chionolaena DC.
• Gnaphaliothamnus Kirp.
• Raouliopsis S.F. Blake
• Mniodes (A. Gray) Benth.
• Loricaria Wedd.
• Leucopholis Gardn. (= Chionolaeana DC.) – Freire 1993
• Oligandra Less. (= Lucilia Cass.) – Freire 1989
“
CASSINIINAE
LORICARIINAE
ASTEREAE
V “Lucilia Group” Merxmüller et al. 1977/Anderberg 1991
S.E.Freire SOUTH AMERICAN GNAPHALIEAE (ASTERACEAE) 5/12/2011
• Belloa J. Rémy
• Berroa Beauverd
• Chevreulia Cass.
• Cuatrecasasiella H. Rob.
• Facelis Cass.
• Lucilia Cass.
• Jalcophila Dillon & Sagástegui, 2 spp
• Gamochaetopsis Anderberg & S.E. Freire, 1 sp
• Luciliocline Anderberg & S.E. Freire, 5 spp
9
géneros Anderberg &
Freire, 1991 3
“Lucilia Group” sensu Anderberg, 1991/Anderberg & Freire, 1991
7
géneros Anderberg,
1991
S.E.Freire SOUTH AMERICAN GNAPHALIEAE (ASTERACEAE) 5/12/2011
SYNAPOMORPHIES OF “LUCILIA GROUP”
GNAPHALIINAE - “Lucilia Group”, Anderberg & Freire 1991
S.E.Freire SOUTH AMERICAN GNAPHALIEAE (ASTERACEAE) 5/12/2011
• Style branches dorsally
pubescent
• Pappus bristles connate
• Florets purple
• 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
“Lucilia Group” sensu Dillon, 2000, 2003
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)
• 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
2 Phylogenetic analyses:
Molecular matrix composed of 4 markers
Molecular matrix + morphological matrix (36 caracteres)
S.E.Freire SOUTH AMERICAN GNAPHALIEAE (ASTERACEAE) 5/12/2011
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
CASSININEAS: Acanthocladium, Antennaria, Anaphalis,
Anaxeton, Cassinia, Ewartia, Langebergia
RHELANINEAS: Oedera, Relhania
ASTEREAS: Novenia, Solidago, Chiliotrichium, Chiliophyllum,
Baccharis, Aster
In group
Out group
TAXA
S.E.Freire SOUTH AMERICAN GNAPHALIEAE (ASTERACEAE) 5/12/2011
Combined trnL-F,matK, ITS and ETS
Tribe Astereae
Tribe Gnaphalieae
Lucilia Group
S.E.Freire SOUTH AMERICAN GNAPHALIEAE (ASTERACEAE) 5/12/2011
Novenia, 1 sp - Argentina, Bolivia, Peru
Combined trnL-F,matK, ITS and ETS
Tribe Gnaphalieae
Lucilia Group
S.E.Freire SOUTH AMERICAN GNAPHALIEAE (ASTERACEAE) 5/12/2011
CHEVREULIA , 5 spp - Argentina, Bolivia, Brasil,
Chile, Paraguay, Uruguay, Falkland Islands
CUATRECASASIELLA, 2 spp
Ecuador to Argentina
Combined trnL-F,matK, ITS and ETS
Tribe Gnaphalieae
Lucilia Group
S.E.Freire SOUTH AMERICAN GNAPHALIEAE (ASTERACEAE) 5/12/2011
JALCOPHILA, 3 spp - Ecuador, Peru, Bolivia
Jalcophila. boliviensis
Combined trnL-F,matK, ITS and ETS
Tribe Gnaphalieae
Lucilia Group
S.E.Freire SOUTH AMERICAN GNAPHALIEAE (ASTERACEAE) 5/12/2011
LORICARIA, 19 spp - Colombia to Bolivia
Tribe Gnaphalieae
Lucilia Group
S.E.Freire SOUTH AMERICAN GNAPHALIEAE (ASTERACEAE) 5/12/2011
LUCILIOCLINE , 5 spp - Peru, Bolivia, Argentina
Combined trnL-F,matK, ITS and ETS
BELLOA, 9 spp – Venezuela to Chile
Luciliocline subspicata
Belloa schultzii
Tribe Gnaphalieae
Lucilia Group
S.E.Freire SOUTH AMERICAN GNAPHALIEAE (ASTERACEAE) 5/12/2011
Combined trnL-F,matK, ITS and ETS
BELLOA, 9 spp – Venezuela to Chile
MNIODES, 4 spp – Peru, Bolivia
Tribe Gnaphalieae
Lucilia Group
S.E.Freire SOUTH AMERICAN GNAPHALIEAE (ASTERACEAE) 5/12/2011
Combined trnL-F,matK, ITS and ETS
GAMOCHAETOPSIS 1sp - Argentina, Chile
GAMOCHAETA
c. 60 spp-New World
STUCKERTIELLA, 1 sp - Argentina, Bolivia, Peru
Tribe Gnaphalieae
Lucilia Group
S.E.Freire SOUTH AMERICAN GNAPHALIEAE (ASTERACEAE) 5/12/2011
Belloa chilensis– Argentina, Chile
Combined trnL-F,matK, ITS and ETS
FACELIS, 3 spp - Argentina, Bolivia,
Brazil, Chile, Paraguay, Peru, Uruguay
Combined trnL-F,matK, ITS and ETS
Tribe Gnaphalieae
Lucilia Group
S.E.Freire SOUTH AMERICAN GNAPHALIEAE (ASTERACEAE) 5/12/2011
LUCILIA 8 spp -Argentina, Bolivia, Brasil,
Chile, Paraguay, Uruguay
BERROA 1 sp Argentina, Uruguay
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)
• 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
Phylogenetic analysis:
S.E.Freire SOUTH AMERICAN GNAPHALIEAE (ASTERACEAE) 5/12/2011
Taxon sampling: 27 species of Gamochaeta
Belloa chilensis – Argentina, Chile
S.E.Freire SOUTH AMERICAN GNAPHALIEAE (ASTERACEAE) 5/12/2011
Combined ETS-ITS
Combined ETS-ITS/ ETS
ETS-ITS
ETS
STUCKERTIELLA, 1 sp
Argentina, Bolivia, Peru
S.E.Freire SOUTH AMERICAN GNAPHALIEAE (ASTERACEAE) 5/12/2011
S.E.Freire SOUTH AMERICAN GNAPHALIEAE (ASTERACEAE) 5/12/2011
ETS-ITS
ETS
GAMOCHAETOPSIS ,1 sp Argentina - Chile
Combined ETS-ITS/ ETS
S.E.Freire SOUTH AMERICAN GNAPHALIEAE (ASTERACEAE) 5/12/2011
GAMOCHAETOPSIS ,1 sp Argentina - Chile
Combined ETS-ITS/ ETS
Gamochaeta nivalis - Argentina, Chile
Combined ETS-ITS
Gamochaeta filaginea, Argentina-Brazil-
Uruguay
S.E.Freire SOUTH AMERICAN GNAPHALIEAE (ASTERACEAE) 5/12/2011
G. axillaris
G. filaginea
G. nigrevestis
G. platensis
G. subfalcata
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 longepedicellata
Argentina, Bolivia
Gamochaeta lulioana - Bolivia
S.E.Freire SOUTH AMERICAN GNAPHALIEAE (ASTERACEAE) 5/12/2011
Gamochaeta stachydifolia - Argentina, Brazil
S.E.Freire SOUTH AMERICAN GNAPHALIEAE (ASTERACEAE) 5/12/2011
Combined ETS-ITS
Gamochaeta grazielae - Brazil
1. “Lucilia group” sensu Anderb. is paraphyletic.
CONCLUSIONS – Lucilia Group
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.
S.E.Freire SOUTH AMERICAN GNAPHALIEAE (ASTERACEAE) 5/12/2011
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.
Jalcophila
Cuatrecasasiella
Chevreulia *
Luciliocline
“Belloas”
Mniodes
Belloa
Berroa
Facelis *
6. Molecular Evidence: Two clades
S.E.Freire SOUTH AMERICAN GNAPHALIEAE (ASTERACEAE) 5/12/2011
Loricaria
Gamochaeta
* Stucke rtiella*
Gamochaetopsis
Lucilia *
PHYLOGENY OF GAMOCHAETA
PHYLOGENY OF “LUCILIA GROUP”
S.E.Freire SOUTH AMERICAN GNAPHALIEAE (ASTERACEAE) 5/12/2011
• To add Micropsis (Gnaphaliinae) and Chionolaeana
(Cassiniinae)
• To add species from Chile
• To combine molecular and morphological evidence
WORK IN PROGRESS:
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
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