cassini tribe heliantheae (compositae) - fapesp
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Cassini tribe Heliantheae (Compositae)
Today an alliance
Dr. Mara Angelina Galvão Magenta Universidade Santa Cecília - UNISANTA, Department of Biology, Santos, SP – Brazil, [email protected]
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CASSINI (1819)
“Hélianthées”: New Word Sunflowers - many characters
Key characters opposite leaves Yellow flowers Involucre uni to biseriate
Helianthus divaricatus
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Eupatoriées
Vernoniées
Ambrosiées
Hélianthées
“Affinités”: Eupatorieae near to Vernonieae, but with characters common to the Heliantheae
“Synanthérées” – Cassini (1817)
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“Hélianthées” in sections: ovary shape / pappus structure
1917 1819, 1826, 1829
Cassini
Pappus of bristles or reduced to scales
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LESSING (1832)
The most obvious characters
• Helianthées, Ambrosiées e Tagétinées de Cassini: subtribes of “Asteroideae” and “Senecioideae”
• System followed by Candolle (1836) with minor changes
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BENTHAM (1873) Restoring the characteristics employed by Cassini
“Helianthoideae”
Exclusion of groups with epaleate receptacles and with pappus reduced to scales
- Helenoideae (excluding Tagaetinae), adjacent to Heliantheae: Helenieae unnatural X natural Heliantheae
Exclusion of genera with bristly pappus
- Senecionideae
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Helianthoideae 140 genera, 1100 species
Key characters • Opposite leaves
• Receptacle with paleae subtending
individual florets
• Pappus of a few rigid awns or paleae
Viguiera dentata
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HELIANTHOIDEAE (10 subtribes, Including “Ambrosieae ” e “Coreopsideae” )
Helianthoideae
(Heliantheae)
with paleae
Subtribe 10: Madiae
(Madiinae)
partially with paleae
Helenioideae
(Helenieae)
without paleae
CONNECTIONS
Helianthoideae
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BENTHAM (1873)
System perpetuated by Hoffmann (1894)
USED FOR MORE THAN A CENTURY Exception: Helenieae, further treated as part of Heliantheae or other tribes
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CRONQUIST (1955, 1977)
• Suggestion to reinclude Helenieae in Heliantheae
• Heliantheae pointed out as the most basal
Helianthus annuus Helianthus grosseseratus
oppositae leaves multiflorous capitula, a few per plant involucral bracts in many series yellow corollas
Helenium flexuosum
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STUESSY (1977) Symposium on the Biology and Chemistry of the Compositae
Synthesis of Heliantheae Habit, floral characters and chromosome number
15 subtribes
Dismissed 1. Turner and Powell (1977): tribe Coreopsidae 2. Nordenstam (1977): re-inclusion (provisional) of the taxa without paleae and with pappus of bristles in Heliantheae (Tagaetae Cass. and Helenieae Benth.)
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STUESSY (1977)
• Bahiinae and Gaillardinae (Heleniinae of *Helenieae) provisionally in Heliantheae.
• Madiinae in Heliantheae, but connecting with Senecioneae
• Ambrosiinae and Coreopsidinae as subtribes
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Cladistic methodology
Evidence
for
relationships
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ROBINSON (1981)
Revolutionary review : a large number of characters
• Micro-morphology
• Cytology
• Chemistry
Significance of carbonized layer (phytomelanin)
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ROBINSON (1981)
Primary characters that separate certain groups are inconstant: recognition of Heliantheae sensu lato
Position of leaves and presence of paleae – minor features
Taxa without paleae (Senecioneae or Helenieae Bentham)
14 subtribes
*Senecioneae with carbonizes cypselae: transferred to Heliantheae
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ROBINSON (1981)
• Madiinae bridges the gap between Heliantheae s.s. and Helenieae
• Discarding of tribal names Madieae, Coreopsideae, Tagateae and
Helenieae
Heliantheae into 35 subtribes: 260 genera, 3000 species
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Heterogamous heads Polarized thickenings on endothecial cells Anther appendage with basal constriction
homogamous heads Radial thickenings on endothecial cells Expanded style appendages
Ancestral: carbonized achene walls, anther appendages without constricted bases
EUPATORIEAE HELIANTHEAE
INULAE
ROBINSON (1981)
Phytomelanin Base of style with hairs American distribution
Helianthus
Eupatorium
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Phylogenetic analysis
Improvement of Heliantheae alliance
hypotheses
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Reduced phylogenetic trees (Bremer et al. 1992)
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KARIS (1993) morphological data
• Inclusion of Athroisma DC. in the analysis
Athroisma latifolium (Eriksson 1995)
Eriksson (1990, 1991): Athroisma, Blepharispermum e Leuciblepharis from Inulae to Heliantheae (receptacle with pales and carbonized cypselas)
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KARIS (1993) Athroisma
Taxa without paleae: Sisters of the core group Helianthoide Members of Eupatorieae: no resolution or low resolution among lineages helenioides ...
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KARIS (1993)
• Maybe Eupatoriae constitute a sister group of Heliantheae s.l.
• The status of the large Eupatorieae may have to be questioned
• Helenieae must be splitted into smaller units, with some names available: Madieae, Tageteae and Helenieae s.s.
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NEW DIVISION: HELIANTHEAE S.S. IN 10 SUBTRIBES
Eupatorieae + Althroisma: basal to a series of Helenieae lineages (provisional recognition of the tribe) finishing
with a monophyletic Heliantheae s.s.
KARIS & RYDING (1994) Karis (1993) studies formalized
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NEW
MOLECULAR
STUDIES
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KIM & JANSEN (1995)
ndhF
Helenieae
Eupatorieae
Difficult placement (Eup., Hel., Vern., Inul.)
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2. Helenium flexuosum (Helenieae)
4. Helianthus debilis (Heliantheae)
1. Athroisma stuhlmannii
(Athroisma group)
3. Eupatorium adenophorum (Euparorieae)
Images:1. Arne Anderberg (2009) 2. Lisa Brown (2007) 3. Sajam Subedi (2011) 4. Bill Rogers (2008)
PANERO et al. (2001)
(Heliantheae alliance)
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BALDWIN et al. 2002 Internal Trasncribed Spacer (ITS)
• Heliantheae s.s. and Eupatorieae: tradicional senses
• Eupatorieae derived from a heleniod lineage; Helenieae s.l. in six tribes: Helenieae s.s., Madieae and Tageteae (expanded), Bahieae, Chaenactideae, and Perityleae (new tribes )
• Helenieae s.s.: basal position
without phytomelanin epidermal cells with raphides leaves often in basal rosettes
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• Paleae in Heliantheae s.s. and Madiinae (Helenieae s.l.): homoplasious
• Pappus of bristles or reduced to scales: arisen in various lineages of Heliantheae s.l.
Features received too much weight in previous circumscriptions of suprageneric taxa
BALDWIN et al. 2002 ITS
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HELIANTHEAE ALLIANCE
12 TRIBES
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PANERO & FUNK (2002) Dna sequence with 10 genes or markers;
matrix of 13.380 pb
Modified Panero & Funk (2002, 2008)
SUBTRIBES TO TRIBES
PANERO & FUNK (2008) combined analyses of 10 cpDNA loci:
STRONG STATISTICAL SUPORT
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CARIAGA et al. (2008) nucleotides sequences ndhF
Inuleae Heliantheae sensu lato Feddea cubensis Athroismeae
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Supertribe Helianthodae H. Rob (2004)
Extracted from: Tree of Life web project (http://tolweb.org/Helianthodae/22899)
cpDNA Panero (2007)
Cariaga et al. (2008) /Panero and Funk (2008)
Inulae Helenieae Phytomelanic Cypsela Clade Feddeeae Athroismeae
Coreopsideae Bahieae Chaenactideae Tageteae Neurolaeneae Polymnieae Heliantheae Millerieae Madieae Peryleae Eupatorieae
HELIANTHEAE ALLIANCE: 13 TRIBES 5500 species
20%-25% of Compositae (Baldwin 2000)
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PANERO (2007) – Heliantheae s.s.
cpDNA: Galinsoginae, Melampodineae and Milleriinae sensu Robinson (1981): transferred to Millerieae
* New or recent subtribes
Modified from Panero (2008) - Heliantheae s.s. - Tree of Life web project (http://tolweb.org/Heliantheae/22924)
14 subtribes 13 genera 1500 spp
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HELIANTHEAE SENSU STRICTO • Leaves triplinerved • Receptacle with paleae
moderately to strongly conduplicate
• Ray florets with corollas trilobed (sometimes bilobed)
• Cypselas with phytomelanin
HELIANTHEAE ALLIANCE • Anthers usually blackened
with no tail
• styles usually hairy above the bifurcation; mature branches curved
• Most cypselae with phytomelanin
BALDWIN (2009) PANERO (2007)
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FUNK et al. (2005, 2009) MEGATREES
Madagascar, Tropical Africa
Central America , Caribbean
North and Central Andes
Modified from FUNK et al. (2009)
General Africa
North America, Mexico
Wildespread or ambiguous
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With the Cuban Feddea at the base of Heliantheae alliance: a direct dispersal must be taken seriously (Panero 2007, Funk 2009)
Bering land-bridge (Panero & Funk 2005)
Feddea