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Systematics, phylogeny and biogeography

Revision of Nicrophorus in part: new species and inferred phylogeny of the nepalensis-group based on evidence from morphology and mitochondrial DNA (Coleoptera : Silphidae : Nicrophorinae)

Derek S. Sikes A D , Ronald B. Madge B and Stephen T. Trumbo C
+ Author Affiliations
- Author Affiliations

A Department of Biological Sciences, University of Calgary, 2500 University Drive NW, Calgary, Alberta, T2N 1N4, Canada.

B 1637 16 Street S. E., Calgary, Alberta, T2G 3P6, Canada.

C Department of Ecology and Evolutionary Biology, University of Connecticut, Waterbury, Connecticut, 06710, USA.

D Corresponding author. Email:

Invertebrate Systematics 20(3) 305-365
Submitted: 13 May 2005  Accepted: 22 February 2006   Published: 20 June 2006


Carrion beetles of the genus Nicrophorus Fabricius, 1775 (Silphidae) are well known for their biparental brood care and monopolisation of small vertebrate carcasses in subterranean crypts. Although the taxonomy of New World species has received modern attention, the fauna of Asia, primarily of the nepalensis-group of species, has not. Herein we revise this species-group and describe as new the following seven species: Nicrophorus charon Sikes & Madge (Sulawesi), Nicrophorus herscheli Sikes & Madge (Sumatra), Nicrophorus insignis Sikes & Madge (Flores Island), Nicrophorus melissae Sikes & Madge (Nepal, Bhutan), Nicrophorus reticulatus Sikes & Madge (Guadalcanal), Nicrophorus schawalleri Sikes & Madge (Gansu, Shaanxi, Sichuan Province) and Nicrophorus trumboi Sikes & Madge (Nepal, Bhutan). We obtained a preliminary phylogeny using morphology and mtDNA (COII). This was inferred using maximum likelihood and Bayesian methods with the Mkv and GTR+I+G models (parsimony was rejected by the Akaike information criterion for being excessively parameter-rich). The phylogenetic signal in the morphological dataset was not strong and results were confounded by a ‘long-branch’ species, N. reticulatus. The signal was stronger in the combined dataset and the COII-only dataset. The molecular phylogeny supported the new status of species N. trumboi and N. melissae. Support was found for a mainland origin of the group with subsequent radiations into the Malay Archipelago.

Additional keywords: Bayesian phylogenetic inference, China, Himalayas, Malay Archipelago, maximum likelihood, Mkv model, Nepal.


We thank the multitude of curators and collections managers who assisted with loans of study material including: N. Berti (MNHN), M. Brendell & M. Barclay (BMNH), R. W. Brooks (SEMC), B. Brugge (ZMAN), M.-L. Chan (NMNS), L.-y. Chou (TARI), R. Danielsson (MZLU), A. Davis (CNCI), F. Génier (CMNC), W. J. Hanson (EMUS), J. Háva (JHC), L. H. Herman (AMNH), E. R. Hoebeke (CUIC), G. N. House (NMNH, USNM), M. Jäch (NHMW), O. Jäger (SMTD), J. Jelínek (NMPC), D. Kavanaugh (CASC), A. G. Kireitshuk (ZMAS), K. Konishi (ITLJ), S. Krauth (IRCW), W. Wehling (MSUC), O. Martin (ZMUC), O. Merkl (HNHM), M. Nishikawa (MNC), S. Nomura (NSMT), G. R. Noonan (MCPM), M. Ôhara (EIHU), P. Parrillo (FMNH), S. B. Peck (SBPC), P. Perkins (MCZC), E. G. Riley (TAMU), G. A. Sammuelson (BPBM), W. Schawaller (SMNS), J. Schneider (JSC), H. Silfverberg (MZHF), R. W. Sites (UMRM), M. Uhlig (ZMHB), T. W. Pietsch (TAMU2), A. Vesmanis (SMFD), B. Viklund (NHRS), K. Walker (MVMA), T. Wier (ANIC), and L.-y. Zheng (NKUM). We thank those who helped us obtain specimens with preserved DNA: A. Riede, S. Suzuki, M. Kon, K. Araya, M. Maruyama, J. Horák, S. B. Peck, D. Mohagan, B. D. Gill. Numerous faculty in the Ecology and Evolutionary Biology department at the University of Connecticut have assisted this project in various ways, including Janine Caira, Paul Lewis, Chris Simon, Jim Slater, and David Wagner. Expeditions to Nepal and Japan were wonderfully successful thanks to the generosity and assistance of Dhruba Manandhar (Nepal) and Masahiro Kon, Seizi Suzuki, Masahiro Ôhara, Tomoyosi Nisimura, Munetoshi Maruyama, and Masahiro Nagano (Japan). DSS thanks his good friend Piotr Naskrecki for countless hours of taxonomic, computer, and photography related discussions (and for tolerating all the carrion). This research was enormously simplified by use of Piotr’s custom-built database MANTIS (A Manager of Taxonomic Information and Specimens (Naskrecki 2001)). Derek thanks his wonderful wife Melissa for her support and love. This project was supported by an Ernst Mayr grant and NSERC Discovery grant to Derek Sikes and a National Science Foundation Grant (DEB-9981381), a University of Connecticut Research Council grant, and a National Geographic Society grant to Stephen Trumbo.


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Appendix 1.  Specimens from which DNA was extracted for sequencing of the COII mitochondrial gene
DNA voucher specimens are stored in the first author’s collection at –80°C. Specimen ID codes correspond to specimen records in the project database and the labels associated with the specimens

Appendix 2.  Museums and collections from which specimens were borrowed
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