Research Article |
Corresponding author: Wolfgang Denzer ( wolfdenoxford@yahoo.co.uk ) Academic editor: Johannes Penner
© 2016 Wolfgang Denzer, Ulrich Manthey.
This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
Citation:
Denzer W, Manthey U (2016) Remarks on the taxonomy and nomenclature of the genus Hypsilurus Peters, 1867 (Reptilia, Agamidae, Amphibolurinae). Zoosystematics and Evolution 92(1): 103-110. https://doi.org/10.3897/zse.92.7469
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Specimens of Hypsilurus spp. corresponding to sequences deposited on GenBank were re-examined. The voucher specimens relating to GenBank sequences were tracked down and their species status confirmed. Sequences reported in earlier publication as H. “bruijnii” and H. “nigrigularis” turned out to be those of H. magnus and H. schultzewestrumi instead. Further confusion surrounded specimens of H. modestus, H. dilophus and H. papuensis. Based on these results a new phylogenetic tree was constructed and the genus name Lophosaurus Fitzinger, 1843 was resurrected.
Hypsilurus , resurrection of Lophosaurus, Lophosaurus dilophus comb. n., Lophosaurus boydii comb. n., Lophosaurus spinipes comb. n.
The Melanesian-Australian agamid genus Hypsilurus currently contains 20 species of which the majority (14 species) occurs on the main island of New Guinea (
The phylogeny of the genus Hypsilurus has been investigated within molecular studies concerned with agamid lizards in general by for example
godeffroyi group: Hypsilurus godeffroyi Peters, 1867, H. binotatus Meyer, 1874, H. bruijnii Peters & Doria, 1878, H. hikidanus Manthey & Denzer, 2006, H. longii (Macleay, 1877), H. macrolepis Peters, 1872, H. magnus Manthey & Denzer, 2006, H. ornatus Manthey & Denzer, 2006, H. papuensis (Macleay, 1877), H. schoedei (Vogt, 1932), H. schultzewestrumi (Urban, 1999) and H. tenuicephalus Manthey & Denzer, 2006. This group also includes the recently described H. capreolatus Kraus & Myers, 2012.
dilophus group comprising Hypsilurus dilophus (Duméril & Bibron, 1837), H. boydii (Macleay, 1884) and H. spinipes (A. Duméril in Duméril & Duméril, 1851)
nigrigularis group Hypsilurus nigrigularis Meyer, 1874, H. geelvinkianus (Peters & Doria, 1878), and H. auritus Meyer, 1874.
Hypsilurus modestus Meyer, 1874 was considered as the sole representative of the modestus group.
Currently GenBank (gb, http://www.ncbi.nlm.nih.gov/genbank/) holds sequences for eight species. However, some specimen identifications and assignments of museum vouchers to their respective sequence have been questioned by
In this paper we present the results of our investigation into the correct determination of the museum material and assignment of the corresponding GenBank sequences, followed by a phylogenetic analysis of the identified species and a comparison with our earlier morphological study. Finally, we will briefly discuss nomenclatural consequences of the results.
For specimen identification we compared photographs of preserved specimens with material, figures and descriptions given in
Gene sequences that have been used in the earlier studies by
Museum and GenBank collection / accession numbers and collection data of Hypsilurus species used in this study. Where the currently accepted nomenclature differs from the species name provided on GenBank this is indicated below the gb accession number.
Species | Catalog No. | Genbank No. | Locality | Coordinates |
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Hypsilurus boydii |
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AY133013 | Mt. Boolbun, South Queensland, Australia | 15°55’S, 145°9’E |
Hypsilurus dilophus | AMS R122449 | AF128466 | Namosado, Southern Highlands, Papua New Guinea | 6°15’S, 142°47’E |
Hypsilurus magnus | AMS R122474 | AY133014 H. bruijnii | Fogamayiu [=Fogomaiu on Google Maps], Southern Highlands, Papua New Guinea | 6°31’S, 143°05’E |
Hypsilurus modestus | AMS R115478 | AY133015 | Yuro, Chimbu District, Papua New Guinea | 6°32’S, 144°51’E |
Hypsilurus papuensis |
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AY133017 | Wau, Morobe Province, Papua New Guinea | 7°20’S, 146°43’E |
Hypsilurus schultzewestrumi |
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AY133016 H. nigrigularis | Kaironk Village, ~10 km NW Simbai, Papua New Guinea | 5°16’S, 144°32’E |
Hypsilurus spinipes |
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AY133018 | Nana Creek Area, N(orth) of Coffs Harbour, New South Wales, Australia | 30°12’S, 152°57’E |
Initially material used in earlier studies (
Photographs of re-determined specimens. A: Hypsilurus magnus (AMS R122474, previously identified as H. bruijnii). Please note the number of small scale rows (n=4) between infralabialia and enlarged submandibular scales. Insert: Lateral view of the dorsum showing crossbands. Photos: Cecilie Beatson. B: Hypsilurus schultzewestrumi (
Some of the confusion surrounding the identification (or rather misidentification) of Hypsilurus nigrigularis specimens in museum collections most probably resulted from two earlier published figures, both of which did not depict “Gonyocephalus nigrigularis” [= H. nigrigularis] as written in the figure captions. In fact
The specimen of H. dilophus (gb AF128466; AMS R122449) is currently catalogued as H. magnus (OZCAM). This specimen is also erroneously (pers. comm. F. Kraus) listed in
Additionally there were inconsistencies with respect to the GenBank sequence of Hypsilurus papuensis (gb AY133017) and its corresponding voucher specimen. In an earlier publication (
Finally there also exists some confusion around Hypsilurus modestus on the OZCAM online database.
Based on these findings we conducted a phylogenetic analysis the results of which are depicted in Figure
Phylogenetic analysis of Hypsilurus spp. and some Australian agamid lizards belonging to the subfamily Amphibolurinae. A: Most parsimonious tree (PHYLIP/Dnapars); B: Maximum likelihood tree (PhyML). Branch length scale represents number of substitutions/site. Branch support values are given above or next to the branch. Both trees clearly show that Hypsilurus is paraphyletic and forms two distinct clades.
With respect to species of the genus Hypsilurus both trees clearly support different clades. Our maximum likelihood analysis yields a clade containing the two Australian species Hypsilurus spinipes and H. boydii as well as the wide ranging H. dilophus as a sister group to the other studied Australian Amphibolurinae. The remaining four Melanesian species are well supported in an apparently monophyletic clade but still with well supported branches differentiating between H. modestus and H. magnus, H. papuensis, H. schultzewestrumi. In our parsimony analysis a clade containing Hypsilurus spinipes, H. boydii and H. dilophus is formed that also contains the closely related species Moloch horridus and Chelosania brunnea as a sister group. This clade is nested between the Melanesian species of Hypsilurus and the remaining Australian amphibolurine lizards. The branch supports in our maximum likelihood and parsimony analyses for the Moloch / Chelosania clade are comparatively weak. None of the resulting topologies is sufficiently supported to present a clear case for either phylogenetic position of these two genera. Again, our parsimony analysis produces a node separating the branch containing only H. modestus (100% bootstrap support) from the branch comprising the other Melanesian species of Hypsilurus. In summary the two resulting Hypsilurus clades are well supported by molecular genetics and well-defined by morphology (see
First, we would like to note that, if sequences are retrieved from databases or databases are consulted for identifying museum material, it may be necessary and appropriate to re-determine the voucher specimens. Great care has to be taken to ensure correct species identification or otherwise misleading phylogenies are published that do not reflect the true intra- and intergeneric relationships between species in accordance with their morphology based taxonomy. For an in-depth discussion on issues related to GenBank see
With respect to Hypsilurus species our analysis corroborates the results of earlier published phylogenetic studies by
Already in the original description of Gonyocephalus (Arua) inornatus [= H. modestus fide Boulenger 1885 and
The clade containing H. magnus, H. papuensis and H. schultzewestrumi is supported by morphological data and these species are members of the godeffroyi group as defined by
The clade containing the species Hypsilurus spinipes, H. boydii and H. dilophus was recognised as a species group by
We currently consider the clade containing Hypsilurus modestus and all Melanesian species of Hypsilurus (apart from H. dilophus) as monophyletic. We suggest to leave these species in Hypsilurus sensu lato until additional material becomes available and further biomolecular studies can be conducted that include additional species of Hypsilurus s.l. Morphologically Hypsilurus s.l. can be divided into a godeffroyi species group, a nigrigularis species group and the monotypic modestus species group (s.
Lophyrus dilophus Duméril & Bibron, 1837.
On p. 419
Medium to large sized, arboreal amphibolurine lizard without femoral or precloacal pores (present in all Australian agamid lizards apart from Chelosania and Moloch); no spines on the body (present in Moloch), no frill around the neck (present in Chlamydosaurus); a transverse gular fold (absent in Chelosania), dorsal scales heterogeneous in size (homogeneous in all Hypsilurus s. l.); TL/HBL < 2.3, typically < 2; lacrimal bone present (absent in all Australian Amphibolurinae apart from Intellagama and Chelosania)
Lophosaurus dilophus (Duméril & Bibron, 1837)
Distribution: New Guinea and adjacent islands
Lophosaurus boydii (Macleay, 1884)
Distribution: Australia (Northeastern Queensland)
Lophosaurus spinipes (Duméril & Bibron, 1851)
Distribution: Australia (Southeastern Queensland, northeastern New South Wales)
1a | Median line of gular pouch without lanceolate scales | L. spinipes |
1b | Median line of gular pouch with lanceolate scales | 2 |
2a | Diameter of conical scales below tympanum > ½ diameter of the tympanum | L. boydii |
2b | Diameter of conical scales below tympanum << ½ diameter of the tympanum | L. dilophus |
We are grateful to Travis LaDuc and David Cannatella (both
Genbank Accession Numbers and corresponding museum specimen data (for Hypsilurus specimens only): Caimanops (=Diporiphora) amphiboluroides (AF128472), Chelosania brunnea (AF128465), Chlamydosaurus kingii (EF090421), Ctenophorus adelaidensis (AF128471), Ctenophorus maculatus (AF375628), Diporiphora winneckei (AY133012), Lophognathus gilberti (AY133019), Moloch horridus (AF128467), Physignathus (=Intellagama) lesueurii (AF128463), Pogona nullarbor (AY133025), Rankinia diemensis (KF791202), Tympanocryptis lineata (AF128475).