Australasian Journal of Herpetology Australasian Journal of Herpetology 22:9-59. ISSN 1836-5698 9(Print) Published 1July 2014. ISSN 1836-5779 (Online) A logical new taxonomy for the Asian subfamily Draconinae based on obvious phylogenetic relationships and morphology of species (Squamata: Sauria: Agamidae: Draconinae). RAYMOND T. HOSER 488 Park Road, Park Orchards, Victoria, 3134, Australia. Phone: +61 3 9812 3322 Fax: 9812 3355 E-mail: [email protected] Received 10 November 2013, Accepted 1 June 2014, Published 1 July 2014. ABSTRACT A number of recent molecular studies have highlighted divergences between south-east Asian agamid lizards within the Draconinae previously thought to be sufficiently close as to be placed in the same genera. Consolidating recently published studies and further investigations into relevant taxa, incorporating available molecular, morphological and geological evidence some genera as presently recognized are re-arranged to better reflect the relationships of species. For some of these taxa, names are available and as a result of this review they are formally resurrected from synonymy. Where names for taxa don’t exist, the species groups are formally named and defined herein according to the Zoological Code (Ride et al. 1999). Well established genera that are in effect split up include the following: Gonocephalus, Kaup, 1825 is divided into three genera. The nominate genus is also split into five subgenera, all named for the first time except for (Dilophyrus Gray, 1845) and the nominate subgenus. Japalura Gray, 1853 is divided into three genera, one of which is named for the first time. Two genera are further divided into two subgenera. Calotes Daudin, 1802 is divided into three genera, two also divided into subgenera. The taxon originally described as “Calotes andamanensis Boulenger, 1891” is also placed in a new monotypic genus as it also clearly sits ouside the two genera in which it has been recently placed (Calotes and Pseudocalotes Kaup, 1827). Ceratophora Gray, 1835, is divided in three, with the two most divergent taxa, C. aspera Günther, 1864 and C. karu Pethiyagoda and Manamendra-Arachchi, 1998 each placed in a new monotypic genus. The remaining Ceratophora are split into two obvious subgenera. Bronchocela Kaup, 1827 is divided into two subgenera. The taxon currently known as Bronchocela cristatella Kuhl, 1820 from the island of Halmahera, Indonesia is formally described as a new species. Other potential new species are identified. Phoxophrys Hubrecht, 1881 is divided into three subgenera, one formally named for the first time. Aphaniotis Peters, 1864 is subdivided into two subgenera as is Ptyctolaemus Peters, 1864. Salea Gray, 1845 is divided into two subgenera for which a names are already available. Draco Linnaeus, 1758 is herein conservatively divided nine ways into subgenera of which five are formally named for the first time. It is likely that other taxonomists may treat these divisions as full genera. Furthermore, Draconinae taxonomy and nomenclature at the level between subfamily and genus is tidied up. The result is the formal erection of ten tribes and the addition six subtribes formally defined and named. As a result of this scientific reorganisation of the subfamily, this paper presents a list of all recognized Draconinae species in correct tribes, subtribes, genera and subgenera. Keywords: Taxonomy; nomenclature; genera; Gonocephalus; Japalura; Cophotis; Calotes; Ceratophora; Diploderma; Aphaniotis; Phoxophrys; Bronchocela; Lophocalotes; Dendragama; Otocryptis; Ptyctolaemus; Mictopholis; Salea; Sitana; Otocryptis; Pseudocalotes; Paracalotes; Draco; Dilophyrus; Oriotiaris; Pelturagonia; Lyriocephalus; Coryphophylax; Ptyctolaemus; Mantheyus; Acanthosaura; new genera; Daraninagama; Doongagama; Maxhoseragama; Crottyagama; Skrijelus; Notacalotes; Pethiyagodaus; Manamendraarachchius; subgenera; Dracontoides; Rhacodracon; Pterosaurus; new subgenera; Honlamagama; Mantheysaurus; Denzeragama; Eksteinagama; Jamesschulteus; Rubercalotes; Ghatscalotes; Laccadivecalotes; Ceyloncalotes; Tamilnaducalotes; Freudcalotes; Khasicalotes; Amboncalotes; Ferebronchocela; Olorenshawagama; Proboscisagama; Mindatagama; Macguiredraco; Philippinedraco; Engannodraco; Somniadraco; Spottydraco; new species; Harradineus. new tribes; Dracoiini; Maxhoseragamiini; Crottyagamiini; Daraninagamaiini; Pethiyagodaiini; Japaluraiini; Lophocalotesiini; Phoxophryiini; Mantheyiini; Dendragamaiini; new subtribes; Maxhoseragamiina; Sitanaiina; Acanthosauriina; Saleaiina; Pethiyagodaiina; Hoser 2014 - Australasian Journal of Herpetology 22:9-59. Doongagamaiina. Available online at www.herp.net Copyright- Kotabi Publishing - All rights reserved 10 Australasian Journal of Herpetology INTRODUCTION subgenus is also tentative, noting mixed results emerging from A number of recent molecular studies have highlighted deep cited molecular and morphological studies. divergences between relatively well-known south-east Asian Ceratophora Gray 1835, is divided in three, with the divergent agamid lizards. These lizards were thought to be sufficiently taxa, C. aspera Günther, 1864 and C. karu Pethiyagoda and close until now as to be placed in the same genera. While Manamendra-Arachchi, 1998 each placed in a new monotypic general affinities have not been disputed, it has become clear genus. The remaining Ceratophora is split into two obvious that some species placed within given genera should be subgenera. removed and placed elsewhere. This situation for some species Bronchocela Kaup, 1827 is divided into two subgenera, the new is not just a case of being within the same genus group, but one named Ferebronchocela subgen. nov... slightly divergent, but more often than not because the relevant The taxon currently referred to as Bronchocela cristatella Kuhl, species are not remotely related to the type species of the 1820 from the island of Halmahera, Indonesia is formally genus they have been placed in (e.g. Gonocephalus Kaup, 1825 described herein as a new and closely related species. as presently defined by most authors or Japalura Gray, 1853 as presently defined by most authors). Phoxophrys Hubrecht, 1881 is divided into three subgenera, one formally named for the first time. In a limited number of cases, genera with widely divergent species in need of a break up have also been identified (e.g. Aphaniotis Peters, 1864 is subdivided into two subgenera as is Ceratophora Gray 1835 or Calotes Daudin, 1802). Ptyctolaemus Peters, 1864. Following on from a number of molecular studies that have Salea Gray, 1845 is divided into two subgenera for which a indicated splits of genera are warranted on a molecular basis, I name is already available. audited all genera within the southern and south-east Asian Draco Linnaeus, 1758 is herein divided nine ways into agamid subfamily Draconinae to determine which needed to be subgenera of which five are formally named for the first time. divided. Furthermore, the taxonomy and the nomenclature at the level The material and methods of the audit involved a thorough between subfamily and genus is tidied up for the Draconinae. review of the published literature with a view to seeing if there The result is the formal erection of ten tribes with some of these were significant morphological differences between species as in turn divided into subtribes (6 subtribes being named). indicated by recently available molecular data. Added to this Unlike authors such as Wallach et al. (2009), Baig et al. 2012., was direct inspection of several hundred live and dead Bates et al. (2013), Reynolds et al. (2013a, 2013b and 2014), specimens and photos of the relevant taxa with known and Hedges et al. (2014) and Thomas et al. (2014), I do not feel the accurate locality data. need to dishonestly rehash other people’s previously published Genera, in the context of how they are recognized by most data and present it as new to justify the taxonomic decisions zoologists in 2013, as indicated by Pyron et al. (2013) found herein or falsely claim the scientific basis underpinning these worthy of formal division included the following: Gonocephalus, decisions is new. Kaup, 1825; Japalura Gray, 1853; Calotes Daudin, 1802; Unlike these authors just cited and others in the group known as Ceratophora Gray 1835; Bronchocela Kaup, 1827 and Draco the Wüster gang, I do not find a need to step outside of the Linnaeus, 1758. Zoological Code (Ride et al. 1999) to rename previously named For some of these taxa, and new genera to be defined, names taxa based on the research of others. are available and they are formally resurrected herein. In this and all previous papers published by myself, I have Where names for these taxa are not available, they are formally always respected and used originally proposed names that have named herein according to the Zoological Code (Ride et al. been named in a code compliant way, no matter how little 1999). Invariably these new genera, or at times herein more evidence was provided as justification for the original conservatively placed subgenera are readily defined using the descriptions (and in some cases this was none!). significant and generally well-known morphological differences Where names have been properly proposed, even if unused to between relevant species-level taxa. date, they are used herein if appropriate and in total compliance Well established genera that are in effect split up, include the with the established rules of zoology (Ride et al. 1999). In terms following: of newly defined taxa