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A Re-Evaluation of the Crocodile Skinks, Genus Tribolonotus Duméril and Bibron, 1839 Sensu Lato Including

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A Re-Evaluation of the Crocodile Skinks, Genus Tribolonotus Duméril and Bibron, 1839 Sensu Lato Including Australasian Journal of Herpetology 33 Australasian Journal of Herpetology 32:33-39. ISSN 1836-5698 (Print) Published 1 August 2016. ISSN 1836-5779 (Online) A re-evaluation of the Crocodile Skinks, genus Tribolonotus Duméril and Bibron, 1839 sensu lato including the division of the genus into three, description of three new species, a new subspecies and the placement of all within a new tribe. RAYMOND T. HOSER 488 Park Road, Park Orchards, Victoria, 3134, Australia. Phone: +61 3 9812 3322 Fax: 9812 3355 E-mail: snakeman (at) snakeman.com.au Received 20 April 2016, Accepted 28 May 2016, Published 1 August 2016. ABSTRACT The Lygosominae, Crocodile Skinks genus Tribolonotus Duméril and Bibron, 1839 as currently recognized consists of eight described species. A related genus, Fojia Greer and Simon, 1982 consists of a single species Fojia bumui Greer and Simon, 1982. Molecular studies have shown the some of the known species of Tribolonotus (type species Zonurus novaeguineae Schlegel, 1834), to be significantly divergent from one another in spite of morphological similarities and distributional proximity (Austin et al. 2010). As a result of these studies, a re-assessment of the morphology of the relevant species and comparison with the geographical and geological records of the Solomon Islands, it is clear that the assemblage needs to be divided (as do some other species complexes in the area). At the genus level Tribolonotus needs to be divided in order to maintain relative parity as compared to other skink genera from the Australo-Papuan region in terms of their relative divergences. Therefore, Tribolonotus as currently recognized is herein divided into three genera, namely Tribolonotus, for the New Guinea species, Quazitribolonotus gen. nov. for T. blanchardi Burt, 1930 and two similar species formally described herein, namely Q. frankanthonyi sp. nov. and Q. tomlonsdalei sp. nov., and the genus Feretribolonotus gen. nov. for the rest of the species formerly within Tribolonotus. Feretribolonotus gen. nov. is further divided into two subgenera, the other named Propetribolonotus subgen. nov. to accommodate the divergent species “Tribolonotus brongersmai Cogger, 1972”. All of these and Fojia are in turn placed in a newly named tribe Tribolonotiini tribe nov.. Fojia is also placed into a subtribe Fojiina subtribe nov.. The species described as Tribolonotus pseudoponceleti Greer and Parker, 1968, is divided into two based on criteria set out by Greer and Parker (1968) and (Austin et al. 2010). The Buka Island population is formally named Feretribolonotus greeri sp. nov.. Kar Kar Island and Huon Peninsula specimens of Tribolonotus gracilis de Rooij, 1909 being significantly different in form to the nominate race, and divergent genetically are herein described as a new subspecies, T. gracilis karkarensis sp. nov.. Keywords: Taxonomy; lizards; Tribe; new tribe; Tribolonotiini; new subtribe; Fojiina; genus; genera; Tribolonotus; Solomon Islands; Solomons; Guadalcanal; Bougainville; Nggela; new genus; Quasitribolonotus; Feretribolonotus; new subgenus; Propetribolonotus; new species; Buka Island; greeri; frankanthonyi; tomlonsdalei; new subspecies; karkarensis. INTRODUCTION genera. Molecular studies have shown that some of the known The so-called Crocodile Skinks Tribolonotus Duméril and Bibron, species of Tribolonotus to be significantly divergent from one 1839 are found in New Guinea and islands to the north, another in spite of morphological similarities and distributional including the Bismarck Archipelago, and the Solomon Islands. proximity (Austin et al. 2010). The genus Tribolonotus as currently recognized consists of eight Pyron et al. (2013) found much the same in their more wide- described species all of which superficially at least appear to be ranging squamate phylogeny and showed the depth of morphologically similar. A related genus, Fojia Greer and divergence between the relevant species-level taxa to be Simon, 1982 consists of a single species Fojia bumui Greer and significant. Simon, 1982, but as noted in the original description, clearly As a result of these studies and a re-assessment of the shares affinities with Tribolonotus to the exclusion of all other morphology of the relevant species, it is clear that at the genus Hoser 2016 - Australasian Journal of Herpetology 32:33-39. Available online at www.herp.net Copyright- Kotabi Publishing - All rights reserved 34 Australasian Journal of Herpetology level, the assemblage needs to be divided in order to maintain populations is the general lower Sepik River basin, which is an relative parity as compared to other skink genera from the area of lowlands and swamps and clearly unsuitable habitat for Australo-Papuan region in relation to divergences and generic the species and so the absence of records can be reliably placements. attributed to an absence of specimens, as opposed to a mere This is most easily seen when comparing this group with other absence of collecting. The Sepik valley barrier would have been species and genera on the Pyron et al. (2013) figures. formed and maintained as the northern landmasses accreted to Therefore, Tribolonotus as currently recognized is herein divided the New Guinea landmass in the last 5 million years as into three genera and one of these into subgenera in described by Hall (2002) and adapted by Austin et al. (2010). accordance with the International Code of Zoological Furthermore the geologically recently created hilly regions in Nomenclature (Ride et al. 1999). northern New Guinea which is where the species occurs is not All of these and Fojia are in turn placed in a newly named tribe connected to the central cordillera of New Guinea, these being Tribolonotiini tribe nov.. Fojia is also placed into a subtribe called separated by a wide area of lowlands and so specimens from Fojiina subtribe nov.. the east and west populations do not have an obvious bridge by which to get from one population to another. Quazitribolonotus gen. nov. is the new genus name applied to Well established geological evidence suggests a division of the the T. blanchardi (Burt, 1930) complex. Until now this has been populations by more than a million years (somewhere under 4 treated as a single species, but is herein divided three ways. million years) and this divergence is of the order recognized by Q. frankanthonyi sp. nov. and Q. tomlonsdalei sp. nov. are the many authorites as being worthy of taxonomic recognition (e.g. two newly named species and occur in biologically distinct land Keogh et al., 2003). zones within the Solomon Islands. Keogh et al., (2003), found populations of Stephens Banded Feretribolonotus gen. nov. is the generic name now applied to Snakes H. stephensi Krefft, 1869 to have diverged some the following species: T. annectens Zwiefel, 1966, T. 800,000 years ago and went on to state that “managers should brongersmai Cogger, 1972, T. ponceleti Kinghorn, 1937, T. treat the Queensland and NSW populations of H. stephensi as pseudoponceleti Greer and Parker, 1968 as well as separate conservation units”. Feretribolonotus greeri sp. nov. (described below) and T. In the absence of known intermediates and a divergence greater schmidti Burt, 1930. The divergent species T. schmidti Burt, than a million years, it is appropriate that the two populations of 1930 is unique among the species in the tribe in being a live T. gracilis be afforded taxonomic recognition. The unnamed bearer, as opposed to laying eggs and is placed in the subgenus taxon is therefore described herein as T. gracilis karkarensis Proptribolonotus subgen. nov.. subsp. nov.. Tribolonotus Duméril and Bibron, 1839 is herein restricted to the The species T. blanchardi Burt, 1930 as recognized to date, is two New Guinea species, namely Tribolonotus novaeguineae clearly the most divergent in the group from the Solomon Islands (Schlegel, 1834) (the type species) and T. gracilis De Rooij, and as mentioned already is herein placed in the genus 1909. Quazitribolonotus gen. nov.. The relevant divisions outlined above also have a robust The known populations clearly sit within three currently known morphological basis to support them as a further reason why I well-defined groups. These are a largely montane dwelling form have no hesitation in breaking up the genus as currently from Bougainville, the nominate form known from Choisuel and recognized. a third form known from Nggela and Guadalcanal. Based on The species T. pseudoponceleti Greer and Parker, 1968 (herein their obvious differences in form (including colour variants) and placed in the new genus Feretribolonotus gen. nov.) was shown habits, they are formally divided into species. by both Greer and Parker (1968) and (Austin et al. 2010) to Molecular evidence, not available at present, will I anticipate consist of two distinct populations. support my position. This is formally divided into two species based on criteria set out This can be pre-empted with a high degree of probability (but not by Greer and Parker (1968) and (Austin et al. 2010). certainty) on the following grounds. In the case of these taxa, I have married the molecular data of The three populations of Quazitribolonotus blanchardi (Burt, Austin et al. (2010), with the morphological data of Greer and 1930), herein divided are significantly divergent in habits, that Parker (1968) to identify the unnamed taxon and provide a cannot be merely explained by location, which is both proximal description of it in compliance with the rules of the International and in the recent geological past connected by land bridges. Code of Zoological Nomenclature (Ride et al. 1999). The allopatry of species is based on a demonstrated lack of The Buka Island population is formally named Feretribolonotus dispersal in these lizards as shown by the current distribution of greeri sp. nov.. the tribe. Inspection of a large number of T. gracilis de Rooij, 1909 from Even at times of reduced sea levels, during glacial maxima, the general region they occur (northern New Guinea), showed potentially emergent land bridges between the relevant islands the presence of two distinct geographically disjunct morphs.
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