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«© CSIRO 2006 10.1071/IS05020_AC ISSN 1445-5226 Invertebrate Systematics, 2006, 20(3), 305–365. Revision of Nicrophorus in part: new species and ...»

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© CSIRO 2006 10.1071/IS05020_AC

ISSN 1445-5226

Invertebrate Systematics, 2006, 20(3), 305–365.

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. SikesA,D, Ronald B. MadgeB and Stephen T. TrumboC


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: dsikes@ucalgary.ca Abstract. 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.

2 Museum and collections from which specimens were borrowed coden Name and address AMNH American Museum of Natural History, Central Park West at 79th St. New York, NY 10024-5192, USA ANIC Australian National Insect Collection CSIRO Division of Entomology, GPO Box 1700, Canberra, ACT 2601, AUSTRALIA BMNH Department of Entomology, The Natural History Museum, Cromwell Rd, London, SW7 5BD, UNITED KINGDOM BPBM Bishop Museum, 1525 Bernice St, Honolulu, HI 96817-0916, USA CASC California Academy of Sciences, Department of Entomology, San Francisco, CA 94118, USA CMNC Canadian Museum of Nature, Insects-Collection Division, PO Box 3443, Station “D”Ottawa, ON K1P 6P4, CANADA CNCI Canadian National Collection of Insects, Crop Protection Program, Eastern Cereal and Oilseeds Research Centre, K. W. Neatby Bldg, C.E.F. Ottawa, K1A OC6, CANADA CUIC Cornell University Insect Collections, Dept. of Entomology, Cornell University, Ithaca, NY 14853, USA DSSC Private collection of Derek S. Sikes, Dept. Biological Sciences, University of Calgary, Calgary, AB, T2N 1N4 CANADA EIHU [=SEHU] Systematic Entomology, Faculty of Agriculture, Hokkaido University, Kita 9 Nisi 9, Kitaku, Sapporo 060, JAPAN EMUS Department of Biology, Utah State University, College of Science, Logan, UT 84322USA FMNH The Field Museum of Natural History, Roosevelt Rd at Lake Shore Drive Chicago, IL 60605-2496, USA HNHM Coleoptera Collection, Department of Zoology, Hungarian Natural History Museum, HBudapest, Baross u. 13, HUNGARY.

ITLJ Laboratory of Insect Systematics, National Institute of Agro-Environmental Sciences, Kannondai 3-1-1, Tsukuba Ibaraki, 305, JAPAN IZCAS [IOZ-CAS] Chinese Academy of Sciences, Institute of Zoology, Beijing, CHINA JHC private collection of Jirí Háva, Branická 13, CZ-147 00, Praha 4, CZECH REPUBLIC JRC private collection of Jan Růžička, Pod Královkou 7, CZ-169 00 Praha 6, Praha, CZECH REPUBLIC JSC private collection of Jan Schneider, Lipová 15, CZ-120 00, Praha 5, CZECH REPUBLIC MCPM Milwaukee Public Museum 800 West Wells St., Milwaukee, WI 53233, USA MCZC Museum of Comparative Zoology Harvard University, 26 Oxford St. Cambridge, MA 02138, USA MNC private collection of Masaaki Nishikawa, 27-1-115, Higashi-kashiwagaya 1, Ebina 243JAPAN MNHN Museum National D’Histoire Naturelle, Laboratoire D’Entomologie, 45 rue Buffon, FParis, FRANCE MOC Systematic Entomology, Faculty of Agriculture, Hokkaido University, Kita 9 Nisi 9 Kitaku, Sapporo 060, JAPAN MSUC Center for Insect Diversity Study, Museum Department of Entomology 243 Natural Science, Michigan State University, East Lansing, MI 48824, USA MVMA Dept. of Natural Sciences Museum of Victoria, 71 Vicotria Crescent, Abbotsford, Victoria, 3067, AUSTRALIA MZHF Zoological Museum, PO Box 17, FIN-00014 University of Helsinki, Helsinki, FINLAND MZLU Museum of Zoology, Department of Zoology, Division of Systematics Lund University, Helgonavagen 3, S-233 62 Lund, SWEDEN NHMW Naturhistorisches Museum Wien, Zweite Zoologische Abteilung; Insekten,Burgring 7 AWien, AUSTRIA NHRS Naturhisoriska Riksmuseet of Natural History, Sektion För entomologi Box 50007, 104 05 Stockholm, SWEDEN NKUM Dept. of Biology, Nankai University, Tianjin 300071, P. R. CHINA 3 NMNH [=USNM] National Museum of Natural History, Smithsonian Institution, Dept.

Entomology, Collections Management Unit, NHB MRC 165; 10th & Constitution Ave.

N. W. Washington, D. C. 20560, USA NMNS National Museum of Natural Science1, Kuan-Chein Rd., Taichung, Taiwan, R.O.C.

TAIWAN NMPC Národní Muzeum, Entomologicke Odd. PM, Golcova 1, Kunratice 148 00 Praha 4,


NSMT National Science Museum, Hyakunin-cho 3-23-1, Shinjuku-ku, Tokyo, JAPAN SBPC private collection of Stewart Peck, Department of Biology, Carleton University, 1125 Colonel By Drive, Ottawa, Ontario K1S 5B6 Ottawa, CANADA SEMC Snow Entomological Museum, Natural History Museum, University of Kansas, Lawrence, KS 66045, USA SMFD [=SMFC] Forschungsinstitut Senckenberg, Entomologie I, Senckenberganlage 25 DFrankfurt /M., F. R. GERMANY SMNS Staatliches Museum für Naturkunde in Stuttgart, Rosenstein 1, D-70191, Stuttgart 1, F.

R. GERMANY SMTD Staatliches Museum für Tierkunde, Augustusstraße 2, D-01067 Dresden, F. R.

GERMANY TAMU [=UWFC] Insect Collection, Dept. of Entomology, Texas A&M University, College Station, TX 77843, USA TARI Insect Collection, Taiwan Agricultural Research Institute, 189 Chungcheng Rd. Wufeng, Taichung Hsien, Taiwan 431,Taiwan, R. O. CHINA UCDC R. M. Bohart Museum of Entomology, University of California, 1124 Academic Surge, Davis, California, 95616, USA UMRM Wilbur R. Enns Entomology Museum, Dept. of Entomology, University of Missouri, Columbia, MO 65211, USA WBC private collection of Wolfgang Barries. Kinzerplatz 10-11/3/3/9, A; 1 2 1 0, Wien, AUSTRIA ZMAN Instituut voor Taxonomische Zoölogie, Zoölogisch Museum, Universiteit van Amsterdam, Afdeling Entomologie, Plantage Middenlaan 64, 1018 DH, Amsterdam, The


ZMAS Russian Academy of Sciences, Zoological Institute, 199034, Universitetskaya nab., 1, St.

Petersburg, RUSSIA ZMHB Museum für Naturkunde der Humboldt-Universität zu Berlin, D-10115 Berlin, Invalidenstr. 43, Berlin, F. R. GERMANY ZMUC Zoologisk Museum, Universitetsparken 15, DK 2100 Kobenhavn O., Kobenhavn, DENMARK ZSMC Zoologische Staatssammlung Münchhausenstraße 21, D-81247, München, München, F.

R. GERMANY 4 Complete lists of Material examined Asterisks (**) indicate material found at the same higher locality. The country for some label data localities could not be determined, these are listed as ‘country not found’

–  –  –

Material examined 29 ♂, 30 ♀, 17 sex undetermined: PHILIPPINES: Mindanao, Bukidnon, Mt. Katanglad, elev.

1480 m [8.1°N, 124.9°E] 27–31.x.1959; 1 ♀ (Honolulu [BPBM]); Bukidnon Prov., MT.

Kitanglad Range, 17km S, 7km E Baungon, elev. 1550 m, 8°10′N, 124°45′E, 1–31.v.1992 (coll.

L. Heaney); 3 ♀ 2 ♂ (Chicago [FMNH]); Davao Prov.: Baclayan, E. slope of Mt. Apo, elev. 1981 m [8°N, 125°E] 1–30.xi.1946 (coll. H. Hoogstraal); 1 paratype ♂ (Chicago [FMNH]); same locality, elev. 1981 m [8°N, 125°E] 12–18.xi.1946 (coll. H. Hoogstraal); 1 paratype ♀ (Chicago [FMNH]) [Davao Prov.] Baclayan, Mt. Apo, elev. 1650 m [8°N, 125°E] 13–15.xi.1946 (coll. D.

Davis); 1 ♀ (Washington D.C. [USNM]); Davao Prov.: E. slope of Mt. McKinley, elev. 914 m [8°N, 125°E] 29.iix.1946 (coll. F. G. Werner); 1 paratype ♂ (Chicago [FMNH]) 1 paratype ♀ (Washington D.C. [USNM]); same locality, elev. 1981 m [8°N, 125°E] 3.ix.1946 (coll. H.

Hoogstraal & G. Alcasid); 1 paratype ♂ (Washington D.C. [USNM]); same locality, elev. 1981 m [8°N, 125°E] 7.ix.1946 (coll. H. Hoogstraal & G. Alcasid); 1 paratype ♀ 1 paratype ♂ (Chicago [FMNH]) 1 paratype ♀ 1 paratype ♂ (London [BMNH]) 1 ♀ 1 ♂ (San Francisco [CASC]) 2 paratype ♀ (Washington D.C. [USNM]); same locality, elev. 1951 m [8°N, 125°E] 10.ix.1946 (coll. H. Hoogstraal); 2 paratype ♀ 4 paratype ♂ (Chicago [FMNH]); same locality, elev. 1768 m [8°N, 125°E] 17.ix.1946 (coll. H. Hoogstraal); 4 paratype ♀ 3 paratype ♂ (Chicago [FMNH]);

same locality, elev. 1006 m [8°N, 125°E] 24.ix.1946 (coll. H. Hoogstraal); 1 paratype ♀ (Chicago [FMNH]); Davao Prov.: Meran, E. slope of Mt. Apo, elev. 1829 m [8°N, 125°E] 8.xi.1946 (coll.

H. Hoogstraal); 1 paratype ♀ 1 paratype ♂ (Washington D.C. [USNM]); **Misamis Or. MT.

Balatukan, 15km SW of Gingoog, elev. 1000–2000 m [8.83°N, 125.27°E] 27–30.iv.1960 (coll. H.

Torrevillas); 1 ♀ (Washington D.C. [USNM]); Mt Apo [8°N, 125°E](coll. E A Mearns); 1 allotype ♂ 1 paratype ♂ (Washington D.C. [USNM]); Mt Apo, Mindanao, Crampton [6.99°N,

125.27°E]; 1 ♀ 1 ♂ (Cambridge [MCZC]); Mt. Apo; Western Mindanao; Brgy.; Ilomabis;

Kidapawan City; North Cotabato, elev. 1750 m, 1–30.xi.2004 (coll. D. Mohagan); 1 DNA voucher 08 adult, 1 DNA voucher 09 adult, 1 DNA voucher 10 adult, 1 DNA voucher 11 adult, 1 DNA voucher 12 adult, 1 DNA voucher 13 adult, 1 DNA voucher 14 adult, 1 DNA voucher 15 adult, 1 DNA voucher 16 adult, 1 DNA voucher 17 adult (Calgary [DSSC]); ****probably 5 Mindanao Mts. [8°N, 125°E](coll. H. Hoogstraal ); 1 paratype ♀ (Chicago [FMNH]); South Cotabato, Kidapawan City, brgy : dlomabis, Mt. Apo, elev. 1700 m [6.98°N, 125.27°E]

2.vii.2004 (coll. D. Mohagan); 1 DNA voucher 01 adult, 1 DNA voucher 02 adult, 1 DNA voucher 03 adult, 1 DNA voucher 04 adult, 1 DNA voucher 05 adult, 1 DNA voucher 06 adult, 1 DNA voucher 07 adult, 7 ♀ 10 ♂ (Calgary [DSSC]).

Nicrophorus charon Sikes & Madge, sp. nov.

Material examined Holotype. ♀ labelled “INDONESIA: Sulawesi, Tengah: Mt. Tambusisi, elev. 1219 m, 1°39′S, 121°21′E, 3–13.iv.1980 (coll. M. J. D. Brendell) BMNH006420Nic.” (London [BMNH]).

Paratypes. 10 ♂, 7 ♀: INDONESIA: Sulawesi, Utara, Gng. Amang F. R. nr.

Kotamobagu, elev. 1200 m [1°N, 123°E] 4.ii.1985; 3 ♂: BMNH83802Nic, BMNH83803Nic, BMNH83804Nic; same locality, 2.vi.1985; 1 ♂: BMNH83805Nic; Utara Dumoga-Bone N.P., lower montane, 2000 m pitfall trap / R. Ent. Soc. Lond., PROJECT WALLACE, B.M. 1985-10, 1985 (coll. P. Whilton); 2 ♂: BMNH122043Nic, BMNH122045Nic; Utara, Gng. Ambang F. R., Gng. Muajak, nr. Kotamobagu, 1780 m, 1–30.xi.1985; 1 ♀: BMNH122046Nic; 30 km NW

Rantepao, Bulu-Bulu, elev. 1600 m [2.98°S, 119.9°E] 9–15.v.1966 (coll. R. Straatman); 1 ♀:

BPBM003389Nic; ~30 km NE of Enrekang. Gn. Rantemario. Borong Tangga Camp., 3°24′S, 120°0′E, 5.xi.1993 (coll. J. P. & M. J. Duffels); 2 ♀: ZMAN003702Nic, ZMAN003703Nic;

Latimodionggeb., elev. 2200 m, 23.vi.1930 (coll. G. Heinrich); 1 ♂: BMNH006421Nic; Palu Central-celebes [0.9°S, 119.86°E] 1–30.iv.1991; 1 ♂: WBC080867Nic; Tengah Lore Lindu N.P., 10 km SE Poloka, Stat. 55 Disturbed lower montane forest ML–light, canopy, elev. 1900 m,

25.iii.1985 (coll. J.P. & M. J. Duffels); 2 ♂: ZMAN003740Nic, ZMAN003741Nic; Tengah Lore Lindu N.P., 10 km SE Poloka, Stat. 57 Disturbed lower montane forest ML–light, canopy, elev.

1900 m, 26.iii.1985 (coll. J.P. & M. J. Duffels); 1 ♀: ZMAN003690Nic; Tengah Lore lindu N.P., Rano Rano 10km NE Gimpu, Stat 41 lower montane forest MV–light, elev. 1600 m [1.6°S,

120.03°E] 14.iii.1985 (coll. J.P. & M. J. Duffels); 2 ♀: ZMAN003688Nic, ZMAN003689Nic.


Nicrophorus herscheli Sikes & Madge, sp. nov.

Material examined Holotype. ♀, labelled “[INDONESIA]: Sumatra, N. Sumatra Gng Leuser NP Mt.

Mamas, elev. 1630 m, 3°33′N, 97°39′E, 7–10.viii.1983 (coll. H. Räisänen); MCZC73701Nic.” (Cambridge [MCZC]).

Paratypes. 3 ♂, 5 ♀. [INDONESIA]: Sumatra, N. Sumatra Gng Leuser NP Mt.

Mamas, elev. 1630 m, 3°33′N, 97°39′E, 24–28.vii.1983 (coll. H. Räisänen); 2 ♀:

MCZC73703Nic, MCZC73705Nic, 1 ♂: MCZC73699Nic; **same locality, 28 vii; 7.viii.1983; 1 ♂: MZHF73698Nic; **same locality, 7–10.viii.1983; 1 ♀: MCZC73704Nic, 1 ♂:

MCZC73700Nic; **same locality, elev. 1650–1680 m, 3°20′N, 97°46′E, 10–17.ix.1983 (coll. H.

Räisänen); 1 ♀: MZHF73702Nic; **N. Sumatra Tele [southwest of lake Toba & Samosir Island;

southwest of Mt. Belirang, ca. 1700m elev.] [2°33′N, 98°35′E] 23–24.iii.1984 (coll. G. Hangay);

1 ♀: SMNS003361Nic.

Nicrophorus heurni Portevin, 1926: 209

Material examined 75 ♂, 85 ♀, 56 sex undetermined: INDONESIA: Irian Jaya, Abmisibl, 18.ix.1985 (coll. H. V.

M.); 1 ♂ (Amsterdam [ZMAN]); Maoke Mountains, road Wamena, Jayaputa, km 48, Primarurwald, elev. 2100 m [4°S, 138°E] 4.xii.1997 (coll. K. Carny); 1 ♀ (Praha, priv.coll [JSC]); Star; Mountains Abmisibil, 9–14.xi.1990 (coll. H. J. G. v. Mastrigt); 2 ♀ (Amsterdam [ZMAN]); NEW GUINEA: Adelberts [4.717°S, 145.25°E] 1–31.iii.1974 (coll. H. Ohlmus); 1 ♀ 1 ♂ (Canberra [ANIC]); Bosavi [6.583°S, 142.833°E] 1–30.xi.1974 (coll. H. Ohlmus); 1 ♀ (Canberra [ANIC]); Dormanpadbivak, elev. 1410 m, 1–31.x.1920 (coll. W. C. van Huern); 1 paratype ♀ 1 paratype ♂ (Paris [MNHN]); **Ekuti Range north slope 2400 m. south Wau [7.33°S, 146.5°E] 18.xii.1977 (coll. R. Harmsen & P. Hebert); 2 ♀ 1 ♂ (Ottawa [CNCI]);

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