Systematics and historical biogeography of the old world butterfly subtribe Mycalesina (Lepidoptera: Nymphalidae: Satyrinae).
BACKGROUND
Butterflies of the subtribe Mycalesina have radiated successfully in almost all habitat types in Africa, Madagascar, the Indian subcontinent, Indo-China and Australasia. Studies aimed at understanding the reasons behind the evolutionary success of this spectacular Old World butterfly radiation have been hampered by the lack of a stable phylogeny for the group. Here, we have reconstructed a robust phylogenetic framework for the subtribe using 10 genes from 195 exemplar taxa.


RESULTS
We recovered seven well supported clades within the subtribe corresponding to the five traditional genera (Lohora, Heteropsis, Hallelesis, Bicyclus, Mycalesis), one as recently revised (Mydosama) and one newly revised genus (Culapa). The phylogenetic relationships of these mycalesine genera have been robustly established for the first time. Within the proposed phylogenetic framework, we estimated the crown age of the subtribe to be 40 Million years ago (Mya) and inferred its ultimate origin to be in Asia. Our results reveal both vicariance and dispersal as factors responsible for the current widespread distribution of the group in the Old World tropics. We inferred that the African continent has been colonized at least twice by Asian mycalesines within the last 26 and 23 Mya. In one possible scenario, an Asian ancestor gave rise to Heteropsis on continental Africa, which later dispersed into Madagascar and most likely back colonised Asia. The second colonization of Africa by Asian ancestors resulted in Hallelesis and Bicyclus on continental Africa, the descendants of which did not colonise other regions but rather diversified only in continental Africa. The genera Lohora and Mydosama are derivatives of ancestors from continental Asia.


CONCLUSIONS
Our proposed time-calibrated phylogeny now provides a solid framework within which we can implement mechanistic studies aimed at unravelling the ecological and evolutionary processes that culminated in the spectacular radiation of mycalesines in the Old World tropics.
Department of Zoology, Radiating Butterflies Group, University of Cambridge, Downing Street, Cambridge, CB2 3EJ, UK. ka374@cam.ac.uk.
Kwaku
K
Aduse-Poku
Department of Zoology, Radiating Butterflies Group, University of Cambridge, Downing Street, Cambridge, CB2 3EJ, UK. ob269@cam.ac.uk.
Oskar
O
Brattström
Department of Zoology, Radiating Butterflies Group, University of Cambridge, Downing Street, Cambridge, CB2 3EJ, UK. ullasa@iisertvm.ac.in.|School of Biology, Indian Institute of Science Education and Research Thiruvananthapuram (IISER-TVM), CET campus, Sreekaryam, Thiruvananthapuram Kerala, 695016, India. ullasa@iisertvm.ac.in.
Ullasa
U
Kodandaramaiah
Department of Zoology, Radiating Butterflies Group, University of Cambridge, Downing Street, Cambridge, CB2 3EJ, UK. dclees@gmail.com.
David C
DC
Lees
Department of Zoology, Radiating Butterflies Group, University of Cambridge, Downing Street, Cambridge, CB2 3EJ, UK. pb499@cam.ac.uk.
Paul M
PM
Brakefield
Department of Biology, NSG, Laboratory of Genetics, University of Turku, Turku, 20014, Finland. niklas.wahlberg@utu.fi.
Niklas
N
Wahlberg
D000349:Africa; D000818:Animals; D001208:Asia; D001315:Australia; D005075:Biological Evolution; D002080:Butterflies / Q000145:classification* / Q000235:genetics*; D017753:Ecosystem; D010802:Phylogeny; D058974:Phylogeography