PP, NAP, HWS, RGL, MS, NSK, SII, and CS collected samples and contributed to fieldwork; NAP, MS, and AVT generated molecular data and conducted phylogenetic analyses; PP, HWS, RGL, JLL, MS and NAP examined specimens and contributed photographs; JLL, HWS, RGL and NAP generated figures; JLL, NAP and HWS wrote the original draft; all authors read, reviewed and revised the final version of the manuscript.