INTEGRATING MOLECULAR AND MORPHOLOGICAL METHODS FOR GREGARINE SPECIES DELIMITATION: A REVISION AND PHYLOGENY OF THE BLABERICOLIDAE WITH RECOGNITION OF NEW SPECIES OF BLABERICOLA AND PROTOMAGALHAENSIA INFECTING BLAPTICA DUBIA, THE GUYANA SPOTTED COCKROACH (BLATTODEA: BLABERIDAE).

Integrated morphological and molecular methods were used to reexamine, revise, and expand the gregarine species that are known members of the Blabericolidae infecting blaberid cockroaches of the subfamilies Blaberinae and Oxyhaloinae. Here, Protomagalhaensia serpicula n. sp. and Blabericola blaberae n. comb. are described from the Guyana spotted cockroach Blaptica dubia, and 3 new synonymies are recognized: Blabericola cubensis (=Blabericola princisi = Gregarina princisi) parasitizing Blaberus discoidalis and Blaberus boliviensis; Protomagalhaensia granulosae (=Protomagalhaensia vipera) parasitizing B. discoidalis and Eublaberus posticus; and Protomagalhaensia wolfi (=Protomagalhaensia richardsoni) parasitizing Nauphoeta cinerea and Henschoutedenia flexivitta. One new combination also is recognized, Blabericola cerastes n. comb. (=Protomagalhaensia cerastes) parasitizing Phoetalia pallida. This study revealed that gregarine species delimitation errors can be avoided by integrating morphological and molecular methods with multiple conspecifics as a reference framework. This approach is preferred for delimiting gregarine species. Blabericolidae and its component genera, Blabericola and Protomagalhaensia, are monophyletic groups with a likely Pangean origin whose members speciated as New World and Old World groups following the breakup of Gondwana. Patterns of speciation appear to be largely vicariant, but host switching through human introduction of pest host species may also have played a role in gregarine radiation. The gross alimentary anatomy of B. dubia is described, and the effects of gamont morphology and phylogeny on patterns of site specificity are discussed.