Taxonomy and species-groups of the subgenus Crematogaster ( Orthocrema) in the Malagasy region (Hymenoptera, Formicidae).

The species-level taxonomy of the subgenus Crematogaster (Orthocrema) in the Malagasy region is evaluated with both morphological data from worker and queen ants, and genetic data from three nuclear markers (long wavelength rhodopsin, arginine kinase and carbamoylphosphate synthase). These two types of data support the existence of six Orthocrema species: Crematogaster madecassa Emery, Crematogaster rasoherinae Forel, Crematogaster telolafysp. n., Crematogaster razanasp. n., Crematogaster volamenasp. n. and Crematogaster mpanjonosp. n.. Two new synonyms of Crematogaster rasoherinae Forel are recognized, Crematogaster rasoherinae brunneola Emery, syn.n. and Crematogaster voeltzkowi Forel, syn. n., as these were not supported as distinct taxa by the data. A neotype is designated for Crematogaster rasoherinae; lectotypes are designated for Crematogaster madecassa, Crematogaster rasoherinae brunneola and Crematogaster voeltzkowi. Species descriptions, images, distribution maps and identification keys based on worker and queen ants are given for all six species. A diagnosis of the subgenus Orthocrema in the Malagasy region is presented for both workers and queens. Within the Malagasy Orthocrema, three distinct phylogenetic lineages are suggested by molecular and morphological data. Newly defined monophyletic species-groups are thus the Crematogaster madecassa-group (Crematogaster madecassa, Crematogaster telolafy and Crematogaster razana) and the Crematogaster volamena-group (Crematogaster volamena and Crematogaster mpanjono); Crematogaster rasoherinae represents an isolated lineage in the Malagasy region and its closest relatives remain unclear. Other interesting biological findings are the presence of an intermediate caste between workers and queens in Crematogaster rasoherinae and Crematogaster madecassa, and unusually large workers in Crematogaster volamena resembling a major caste.