Species delimitations, endemism, and invasions in ...

1 downloads 0 Views 36MB Size Report
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