invasions in Madagascar - an iterative taxonomic approach to tackle a hyper-diverse ant genus in the Malagasy region. Francisco Hita Garcia & Brian Fisher.
Species delimitations, endemism, and invasions in Madagascar - an iterative taxonomic approach to tackle a hyper-diverse ant genus in the Malagasy region Francisco Hita Garcia & Brian Fisher Entomology Department California Academy of Sciences San Francisco, U.S.A.
Madagascar
Madagascar
Madagascar
Malagasy ant fauna Malagasy ant inventory (B. Fisher, 1992 to 2010): 8 subfamilies 67+ genera 1300+ species
ca. 98% endemism
less than 500 species described more than 60% undescribed
Malagasy ant fauna
Problematic genera: Camponotus 150+ species Pheidole 150+ species Tetramorium 150+ species
Malagasy ant fauna
Problematic genera: Camponotus 150+ species Pheidole 150+ species Tetramorium 150+ species
Taxonomic foundation Modern regional revisions Bolton, 1976, 1977, 1979, 1980 (Afrotropical, Malagasy, Oriental, IndoAustralian, New World)
Barry Bolton (BMNH)
Palearctic: Csösz et al, 2007, 2010; Steiner et al. 2010 Africa: Hita Garcia et al. 2010a, 2010b, 2010c New World: Vásquez-Bolaños, 2007; Vásquez-Bolaños et al., 2011
Diversity & distribution
Palearctic ca. 70
New World ca. 15
Afrotropical ca. 230 Malagasy 39
Oriental ca. 80 Indo-Australian 23
Diversity & distribution
Palearctic ca. 70
New World ca. 15
Afrotropical ca. 230 + 150 Malagasy 39 + 120
Oriental ca. 80 + 30
300+ undescribed species
Indo-Australian 23 + 20
Project goals 1. taxonomic revision of the genus for the Malagasy region (description of 100+ new species in 2 years) 2. identification tools: - species group key - species level keys - DNA barcodes for all species - Automontage images for all species (AntWeb) 3. framework for further research: - ecology - biogeography - phylogeny
Material Malagasy ant inventory (1992 to 2010): 6,000+ leaf litter samples 4,000+ pitfall traps 1,000+ malaise traps 9,000+ hand collection events
Material Mounted material: 43 drawers 30,000+ specimens 9529 databased
Initial species counts: 39 valid species 122 morphospecies 161 in total
Methodology morphology morphometry
Morphometry PH
HL
PTH
WL HW
PPH
EL
SL
PSL
PW
PTW PTL
14 measurements 13 indices
PPW PPL
Methodology morphology morphometry DNA barcoding habitat microhabitat geography
Why use DNA barcoding for taxonomy?
1. future species identification for non-taxonomists 2. test species groups and species hypotheses 3. reveal cryptic diversity 4. associate queens & males with their respective workers
Methodology morphology morphometry DNA barcoding
Why use species groups? 1. establishment of operational subunits - morphologically well-defined - easily identifiable (morphology & mtDNA) - tentatively monophyletic
2. work on global scale & allow zoogeographic comparisons
3. working hypotheses for future phylogenetic research
Tetramorium species group overview Afrotropical Oriental & Indo-Australian New World Palearctic
Malagasy (2010)
24 groups 20 groups 6 groups 6 groups
9 groups
Tetramorium species group overview Afrotropical Oriental & Indo-Australian New World Palearctic
24 groups 20 groups 6 groups 6 groups
Malagasy (2010) Malagasy (2011)
9 groups 19 groups
all 19 morphologically well defined
sericeiventre (2) tosii (2) simillimum (5)
New species groups 2011
marginatum (6) plesiarum (2)
Neighbor-joining tree CO1-5P (648bp)
kelleri (1) ranarum (1) dysalum (10)
18 species groups 127 species 1500 specimens
tortuosum (15) dysalum (4) schaufussii (30) naganum (4) bessonii (6) bonibony (6) (6 spp.) pleganon (2) bonibony (2) ranarum (20) plesiarum (3) severini (1) weitzeckeri (1) bicarinatum (2) obesum (1)
sericeiventre (2) tosii (2) simillimum (5) marginatum (6) plesiarum (2) kelleri (1) ranarum (1) dysalum (10)
14 well supported groups
tortuosum (15) dysalum (4) schaufussii (30) naganum (4) bessonii (6) bonibony (6) (6 spp.) pleganon (2) bonibony (2) ranarum (20) plesiarum (3) severini (1) weitzeckeri (1) bicarinatum (2) obesum (1)
sericeiventre (2) tosii (2) simillimum (5) marginatum (6) plesiarum (2) kelleri (1) ranarum (1) dysalum (10)
4 difficult groups
tortuosum (15) dysalum (4) schaufussii (30) naganum (4) bessonii (6) bonibony bonibony (6(6)spp.) pleganon (2) bonibony (2) ranarum (20) plesiarum (3) severini (1) weitzeckeri (1) bicarinatum (5) obesum (1)
Easy case: T. marginatum group
T. marginatum Forel
T. fulgidum sp. n.
T. silvicolum sp. n.
T. echinatum sp. n.
T. norvigi sp. n.
T. shamshir sp. n.
T. bessonii group
T. bessonii Forel
T. MG04
T. orientale Forel stat. n.
T. MG05
T. bessonii group
T. MG05
3 species? 44 specimens no material from T. orientale (only holotype) T. bessonii
T. MG04
T. bessonii group
T. MG05
3 or 5 species 44 specimens no material from T. orientale (only holotype) T. bessonii
T. MG04
T. bessonii group
T. bessonii Forel
T. orientale Forel stat. n.
T. MG04a
T. MG05a
T. MG04b
T. MG05b
T. bessonii group
T. bessonii Forel
T. orientale Forel stat. n.
T. wardi sp. n.
T. malagasy sp. n.
T. australimum sp. n.
T. ryanphelanae sp. n.
Morphology vs. DNA barcoding
DNA barcoding data supports morphology & morphometry in most cases
DNA barcoding can reveal undetected species in few cases
If species or complexes of species are morphologically very difficult, then DNA barcoding data is inconclusive
Invasive or tramp species
T. caldarium (Roger)
T. bicarinatum (Nylander)
T. simillimum (Smith)
T. insolens (Smith)
T. lanuginosum Mayr
T. pacificum Mayr
6 global tramps
T. delagoense Forel
T. humbloti Forel
T. sericeiventre Emery
3 invaders from Africa
Zoogeographic overview 13 species groups endemic for Madagascar 3 species groups global tramps 1 species group of global distribution (no tramps) 2 species groups shared with Afrotropical region
152 species Malagasy endemics 3 species shared with Afrotropical region 6 global tramp species
ca. 95% of Malagasy Tetramorium fauna endemic
Conclusions
importance of subgroups for hyper-diverse genera
combination of morphology, morphometry, and mtDNA barcoding works very well for most groups and species
uniqueness of the Malagasy ant fauna
Acknowledgements Funding
Brian Fisher
Barry Bolton
CAS antlab Masashi Yoshimura Georg Fischer Michele Esposito Estella Ortega Shannon Hartman William Ericson Ryan Perry
Ant images AntWeb (www.antweb.org)
Map sources AntWeb, EU, Madagascar GIS