catalogue of biphyllidae (coleoptera) of the world

A N N A L E S Z O O L O G I C I (Warszawa), 2015, 65(3): 409-471

CATALOGUE OF BIPHYLLIDAE (COLEOPTERA) OF THE WORLD PIOTR WĘGRZYNOWICZ Institute of Forest Sciences, University of Łódź, Branch in Tomaszów Mazowiecki, ul. Konstytucji 3 Maja 65/67, 97-200 Tomaszów Mazowiecki, Poland; e-mail: [email protected]

Abstract.— A catalogue of all known species of the family Biphyllidae is presented. All described extant and fossil species have been included, with synonyms, taxonomic notes, bionomy and distribution. Two new synonyms are proposed: Diphyllus lunatus var. inornatus Reitter, 1888 syn. nov. of Dermestes lunata Fabricius, 1787, and Thallestus typhaeoides var. obscuricollis Wollaston, 1865 syn. nov. of Thallestus typhaeoides Wollaston, 1862. Twenty three new combinations are proposed: Biphyllus alluaudi (Grouvelle, 1906), B. bolivari (Grouvelle, 1905), B. brunnescens (Reitter, 1877), B. canaliculatus (Grouvelle, 1906), B. concolor (Grouvelle, 1906), B. convexus (Reitter, 1878), B. dohrni (Reitter, 1877), B. escalerae (Grouvelle, 1905), B. histrio (Grouvelle, 1906), B. liliputanus (Reitter, 1877), B. maculatus (Grouvelle, 1906), B. madagascariensis (Fairmaire, 1898), B. magnus (Grouvelle, 1906), B. minimus (Grouvelle, 1905), B. obscurus (Reitter, 1877), B. parvulus (Grouvelle, 1906), B. sicardi (Grouvelle, 1906), B. sordidus (Grouvelle, 1906), B. subfasciatus (Reitter, 1878), B. undulatus (Grouvelle, 1906), B. v-notatus (Grouvelle, 1906), B. variegatus (Grouvelle, 1906), and B. wollastoni (Reitter, 1878). All above-mentioned species are being transferred from Diphyllus Berthold, 1827, which is an unjustified emendation of the name Biphyllus. Type species for three genera are designated: Anobocoelus championi Sharp, 1902 for Anobocoelus Sharp, 1902; Gonicoelus unicornis Sharp, 1900: 605 for Gonicoelus Sharp, 1900: 604; and Euderopus regularis Sharp, 1900 for Euderopus Sharp, 1900.
Key words.— Coleoptera, Cucujoidea, Biphyllidae, distribution, biology, new synonym, new combination, type species designation.

Contents Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Material and methods . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Catalogue . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Family Biphyllidae LeConte, 1861 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Genus Althaesia Pascoe, 1860 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Genus Anchorius Casey, 1900 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Genus Biphyllus Dejean, 1821 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Genus Diplocoelus Guérin-Méneville, 1844 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Genus Euderopus Sharp, 1900 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Genus Eurhanius Reitter, 1887 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Genus Gonicoelus Sharp, 1900 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Fossil taxa . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Acknowledgements . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Index . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Biphyllidae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Insecta. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Fungi and plants . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . PL ISSN 0003-4541 © Fundacja Natura optima dux doi: 10.3161/00034541ANZ2015.65.3.002

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INTRODUCTION Biphyllidae are a family of beetles belonging to the superfamily Cucujoidea. They are rather poorly known, perhaps because of their small size, inconspicuous appearance, and clandestine way of life. Currently, there are 185 extant species and 1 fossil species classified into 7 genera. The first named (as Dermestes lunata) representative of this group was Biphyllus lunatus (Fabricius, 1787) from Sweden, but most species were described in the mid to late 1800’s and early 1900’s which coincided with tropical and subtropical area becoming better explored and the resulting natural history specimens deposited back in major European collections. Antoine Grouvelle undoubtedly put the greatest contribution to the knowledge of this family describing 59 new species from around the world in 1898–1919. LeConte (1861: 105) was the first coleopterist to distinguished Biphyllidae as a separate group of beetles – under the name Diphyllidae, as the subfamily (sic!) of Mycetophagidae. Later most often species of Biphyllidae was included into family Cryptophagidae (Redtenbacher 1858: LXXXII; Reitter 1887: 12; Casey 1900: 78), Erotylidae (Ganglbauer 1899: 651; Kuhnt 1911: 81), and even Byturidae (Falcoz 1926: 69). The first author who distinguished Biphyllidae as an independent family was Arrow (1929: 306). Crowson (1952b: 118) started modern studies on the systematic position and phylogeny of Biphyllidae, and they have been continued by many authors (Abdullah and Abdullah 1966: 63; Abdullah 1971: 504; Węgrzynowicz 2002: 449; Leschen 2003: 17; Hughes et al. 2006: 270; Hunt et al. 2007: 1914; Leschen and Buckley 2007: 100; Lawrence et al. 2011: 9). Biphyllidae are distributed on all continents except Antarctica, with greatest diversity (66 species) in the Neotropics. The biology and immature stages are poorly known. Most taxa within the family are likely mycetophagous, developing on dendrophilous fungi.

MATERIAL

AND METHODS

All taxa are listed in alphabetical order. The following format is followed for each taxonomic entry: Name derivation. Etymology of the names of genera and species. Gender. Grammatical gender of the genus name. Type species. Type species of the genus and way of its designation. Type locality. The locality from where the holotype or lectotype of the species originated. If neither (in the original publication) the holotype nor (later) the lectotype has been designated, and the author mentioned more than one locality for the type material, all of them (separated by semicolons: “;”) have been listed.

All type localities have been quoted in the original spelling, with – if needed – comments and additions in square brackets: “[]”. Type material. Collections in which type specimens are hosted, their kind and number, according to the original publication, secondary sources (with reference), or author’s own remarks in collections (pers. obs.). Collection abbreviations are explained below. A question mark “?” before the abbreviation means that depository of the type material has not been specified in the original description or secondary sources but probably the type material is preserved there. References . Publications concerning particular taxa are provided in chronological order. Quotations include names of the authors, year of the publication, relevant pages, and (if any) illustration(s). Each reference is followed (in parentheses) by: • taxon name – only if different from that accepted in the catalogue, i.e.: the synonym considered valid by the author of cited publication (with remark: species bona or genus bonum), or only the genus name if the author classified the species in a different genus; • type of the information to be found in the publication, e.g. distribution, biology, etc. Immature stages. References to publications concerning immature stages of the taxon. Biology. Data on the biology of particular taxa with the respective references. Number of species. Number of species described in the given group. Distribution. Geographical distribution of particular taxa: list of countries, and for those of large territories their subdivisions are included as well. Note. Additional relevant notes and remarks. References have been carefully checked against original publications. Publications have been dated as exactly as possible based on analysis of original works as well as on secondary (bibliographic) sources. In the latter case, the source has been quoted. Abbreviations for types depositories: AMSA – Australian Museum, Sydney, Australia; CUMZ – Cambridge University, Museum of Zoology, Cambridge, United Kingdom; DEIC – Deutsches Entomologisches Institut im Leibniz Zentrums für Agrarlandschaftsforschung, Müncheberg, Germany; FVC – Francesco Vitali’s Collection, Genova, Italy; HNHM – Hungarian Natural History Museum, Budapest, Hungary; IBSS – nstitute of Biology and Soil Sciences Russian Academy of Sciences, Vladivostok, Russia; IRSNB – Institut Royal des Sciences Naturelles de Belgique, Brussels, Belgium; KUEC– Kyûshû University, Fukuoka, Japan; MACN – Museo Argentina de Ciencias Naturales, Buenos Aires, Argentina;

CATALOGUE OF BIPHYLLIDAE (COLEOPTERA) OF THE WORLD

MCZC – Museum of Comparative Zoology, Cambridge, Massachusetts, USA; MIZPAN – Muzeum i Instytut Zoologii PAN, Warszawa, Poland; MNHN – Muséum National d’Histoire Naturelle, Paris, France; MNSC – Museo Nacional de Historia Natural, Santiago de Chile, Chile; NHML – The Natural History Museum, London, United Kingdom; NHRS – Naturhistoriska Riksmuseet, Stockholm, Sweden; QMBA – Queensland Museum, Brisbane, Australia; SAMA – South Australian Museum, Adelaide, Australia; SEHU – Hokkaido University Museum, Sapporo, Japan; SMTD – Staatlishes Museum für Tierkunde, Dresden, Germany; USNM – National Museum of Natural History, Washington, USA; ZMAS – Russian Academy of Sciences, Zoological Institute, St. Petersburg, Russia; ZMHB – Museum für Naturkunde der Humboldt Universität, Berlin, Germany; ZMUM – Moscow State University, Moscow, Russia.

CATALOGUE Family Biphyllidae LeConte, 1861 Diphyllidae LeConte, 1861: 105.

Type genus. Diphyllus Berthold, 1827: 393 (= Biphyllus Dejean, 1821: 102). Note. Diphyllus is an unjustified emendation of the name Biphyllus only, so LeConte is the author of the family-group name Biphyllidae (Pakaluk et al. 1994: 228 and 243). References. Redtenbacher 1858: LXXXII (key for the identification of European genera); Jacquelin du Val 1859: 114 [catalogue] (catalogue of European species); Kiesenwetter 1862: 416 (systematic position); Thomson 1863: 239 (descriptions – adult and immature stages, Scandinavian species); Marseul 1866: 49 (catalogue of European species); Gemminger and Harold 1868: 881, 908 (world catalogue); Stein 1868: 49 (catalogue of European species); Thomson 1868: 61 (description, Scandinavian species); Kraatz 1869: 28 (catalogue of German species); Stierlin and Gautard 1869: 151 (catalogue of Helvetian species, biology); Seidlitz 1872: 45 [Gattungen] (key for the identification of European genera), 183 (description of European species); Crotch 1873: 44 (catalogue of North American species);

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Redtenbacher 1874: LXXXIX (key for the identification of Austrian genera); Stein and Weise 1877: 76 (catalogue of European species); Lewis 1879: 11 (catalogue of Japanese species); Rupertsberger 1879: 219 (list of described larvae of European species); Reitter 1880b: 86 [16] (key for the identification of European genera); LeConte and Horn 1883: 140 (classification of North American species); Reitter in Heyden et al. 1883: 78 (catalogue of European species); Dohrn 1884: 117 (description in key); Reitter 1885: 20 (key for the identification of European species); Masters 1886: 667 (catalogue of Australian species); Reitter 1887: 12 (key for the identification of Palearctic genera); Reitter 1888b: 12 (key for the identification of Palearctic genera); Schilsky 1888: 54 (catalogue of Central European species); Schönfeldt 1888: 47 (catalogue of Japanese species, additions); Seidlitz 1888: 66 (key for the identification of European genera); Seidlitz 1889: 66 (key for the identification of European genera); Haly 1890: 103 (catalogue of Sri Lanka species); Reitter in Heyden et al. 1891: 151 (catalogue of European species); Schoenfeldt 1891: 257 (catalogue of Japanese species, additions); Ragusa 1892: 131 (catalogue of Sicilian species, biology); Ganglbauer 1899: 651 (key for the identification of Central European species); Alluaud 1900: 133 (catalogue of Madagascan species); Casey 1900: 78 (review of genera and species from North America, keys for species identification); Sahlberg 1900: 75 (catalogue of Finnish species); Fauvel 1903: 314 (key for the identification of New Caledonian species, catalogue); Jakobson 1905: 936 (catalogue of Palearctic species); Reitter in Heyden et al. 1906: 338 (catalogue of European species); Kuhnt 1909: 116 (world catalogue, key for the identification of genera); Schilsky 1909: 91 (catalogue of Central European species); Kuhnt 1911: 81 (world catalogue); Reitter 1911: 73 (key for the identification of European species); Kuhnt 1912: 530 (description, key for the identification of German genera); Bruch 1914: 385 (catalogue of Argentinean species); Jakobson 1915: 936 (key for the identification of Palearctic genera; catalogue of Palearctic species); Schaufuss 1916: 483 (description, key for the identification of European species); Kuthy 1918: 88 (catalogue of Hungarian species); Leng 1920: 202 (catalogue of North American species); Falcoz 1926: 69 (systematic position); Mader 1926: 718–721 (catalogue of Palearctic species); Arrow 1929: 306 (classification); Portevin 1931: 193 (key for the identification of French species, biology); Miwa 1931: 76 (catalogue of Taiwanese species); Leng and Mutchler 1933: 33 (supplement to catalogue of North American species); Schenkling 1934: 1–7 (supplement to world catalogue); Roubal 1936: 161 (catalogue of Slovak species, biology); Blackwelder 1945: 245 (catalogue of the Central and South American species); Hellen 1947: 43 (catalogue of Finnish and Swedish species);

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Crowson 1952a: 70 (description in key); Crowson 1952b: 118 (systematic position); Crowson 1956: 36 (description in key); Rymer Roberts 1958: 276 (immature stages); Crowson 1960: 127 (systematic position); Abdullah and Abdullah 1966: 63 (systematic position); Crowson 1967: 106 (systematic position, biology); Vogt 1967: 104 (key for the identification of European genera); Damoiseau 1968: 28 (type specimens in the collection of IRSNB); Abdullah 1969: 684 (systematic position); Abdullah 1971: 504 (phylogeny, morphology, immature stages); Arnett 1971: 789 (description, key for the identification of USA genera, systematic position, biology); Bangsholt et al. 1979: 49 (catalogue of Scandinavian species); Camousseight 1980: 5 (list of types in the collection of MNSC); Wallace and Fox 1980: 638, figs. 125–128 (morphology, wings, systematic position); Lawrence and Newton 1982: 282 (systematic position); Burakowski et al. 1986: 208 (catalogue of Polish species, biology); Pal and Lawrence 1986: 195, fig 57 (morphology, systematic position); Lucht 1987: 187 (catalogue of Central European species); Basset 1988: 216, fig. 3 (biology, Australia); Costa et al. 1988: 202 (immature stages, biology, Brasil); Hirashima et al. 1989: 379 (catalogue of Japanese species); Maetô 1990: 383 (biology); Campbell 1991: 241 (catalogue of Canadian species); Lawrence 1991: 475 (immature stages, biology); Goodrich and Springer 1992: 361 (revision of the North American species); Hornig 1992: 70 (list of types in collection of SMTD); Nikitsky 1992: 279 (review of species from Russian Far East); Allison 1993: 17 (biology, tree canopies, Papua New Guinea); Jelínek 1993: 102 (catalogue of Czechoslovakian species); Pakaluk et al. 1994: 228 and 243 (nomenclature); Lawrence and Newton 1995: 880 (classification, nomanclature); Poggi 1995: 1, 7 (catalogue of Italian species); Alexandrovitch et al. 1996: 41 (catalogue of Belarusian species); Arnett 1997: 346 (North American species, biology); Zitani et al.1997: 180 (biology); Köchler and Klausnitzer 1998: 111 (catalogue of German species); Peck and Thomas 1998: 94 (catalogue of Florida species); Machado and Oromí 2000: 63 (catalogue of Canarian species); Azuma et al. 2002: 224 (catalogue of Ryukyu Islands species); Goodrich 2002: 356 (fauna of North America, catalogue, morphology, adults, immature stages, keys, biology); Węgrzynowicz 2002: 449 (morphology, phylogeny); Leschen 2003: 17, tab. 2 (morphology, phylogeny); Leschen et al. 2005: 28 (systematic position); Peck 2005: 134 (catalogue of Cuban species); Suh et al. 2005: 261, fig. 2 (biology, gut, yeasts); Hughes et al. 2006: 270 (genetics, phylogeny); Gormley et al. 2007: 134, tab. 1 (forests fragmentation, Costa Rica); Grimbacher and Stork 2007: 80, tab. 1 (Australia, ecology, rain forests); Hunt et al. 2007: 1914, tab. 1 (phylogeny, biology); Jelínek 2007: 547–548 (catalogue of Palearctic species); Leschen and Buckley 2007: 100, 103, figs. 4 and 5 (phylogeny); Grimbacher et

al. 2008: 77, appendix 2 (Australia, biology); Fedorenko 2009: 247 (hind wings, morphology, evolution); Hirao et al. 2009: 270, tab. 2 (Japan, ecology); Ricarte et al. 2009: 596 (biology); Cline and Shockley 2010: 306 (review, morphology, taxonomy, biology); Grebennikov and Leschen 2010: 90 (morphology, exoskeletal cavities); Ohsawa 2010: 483, tab. 3 (Japan, ecology, Malaise trap); Recalde Irurzun and San Martín Moreno 2010: 152 (distribution, Iberian fauna); Stone et al. 2010: 313, appendix 1 (Australian taxa, biology); Bouchard et al. 2011: 356 (nomenclature); Lawrence et al. 2011: 9 (phylogeny); Slipinski et al. 2011: 206 (classification); Park et al. 2012: 185 (revision of South Korean species, keys for identification, descriptions, distribution, biology, illustrations); Bousquet et al. 2013: 6, tab. 1 (catalogue of Canadian and Alaskan species, distribution); Lawrence and Ślipiński 2013: 299, figs.73A–B, 84G, and 86H (morphology, immature stages, biology, distribution, Australia); Quinto et al. 2013: 770, tab. 1 (biology); Hendrich et al. 2014: 4, tab. 1 (barcoding). Immature stages . Thomson 1863: 239 (larva, description); Rupertsberger 1879: 219 (catalogue of described larvae of European species); Crowson 1952a: 70 (larva, description in key); Rymer Roberts 1958: 276 (larva, morphology); Lawrence 1991: 475 (immature stages, biology); Goodrich 2002: 356 (description); Cline and Shockley 2010 (family overview of immature morphology and biology); Lawrence and Ślipiński 2013: 300, fig. 84G (larva, morphology, biology). Biology. Mycetophagous beetles, developing on dendrophilous fungi. An unidentified species from Australia was collect on the rotting flower stalks of Xanthorrhoea sp. (Xanthorrhoeaceae) and in rotting cycad cones, where they were likely feeding on molds or other fungi (Lawrence 1991: 475). Number of species. 185 extant and 1 fossil. Distribution. Worldwide, except Antarctica.

Genus Althaesia Pascoe, 1860 Althaesia Pascoe, 1860: 117.

Name derivation. Greek name for natural mallows (genus Althaea L.) – itself probably a derivation of altho (gr.) – I cure, I heal, or althos (gr.) – medicine, the ending -ia (lat.) means similarity; the name given to allude to the dense setation of both these plants (e.g. the European Althaea officinalis L.) and the beetles belonging to this genus. Gender. Feminine. Type species. Althaesia pilosa Pascoe, 1860: 117, by monotypy. References. Gemminger and Harold 1868: 909 (world catalogue); Arrow 1929: 308 (key for the identification of species – contains typographical errors –

CATALOGUE OF BIPHYLLIDAE (COLEOPTERA) OF THE WORLD

see Arrow (1937: 106)); Schenkling 1934: 3 (supplement to the world catalogue); Arrow 1937: 106 (corrected version of Arrow’s previous key for the identification of species). Immature stages. Unknown. Biology. Unknown. Number of species. 6. Distribution. Australia, New Guinea, Indonesia (Maluku Province: Island Larat).

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year of the description of this species erroneously given as 1903); Arrow 1929: 308 (description in key, systematic position); Schenkling 1934: 3 (distribution); Arrow 1937: 106 (description in key). Distribution. Australia (New South Wales).

Althaesia maxima (Lea, 1921) Diplocoelus maximus Lea, 1921b: 236 (in Althaesia – Arrow 1929: 309).

Althaesia acuminata Arrow, 1929 Althaesia acuminata Arrow, 1929: 309.

Name derivation. Acuminata, -us (lat.) – pointed; likely due to the denticles, one just at apex of each elytron. Type locality. Dorey. Type material. NHML. Note. In the original description the number of typespecimens is not provided. References. Schenkling 1934: 3 (distribution); Arrow 1937: 106 (description in key, distribution, discussion). Distribution. New Guinea.

Name derivation. Maximus (lat.) – the biggest, very big, enormous; the name alludes to the large size (8.0–9.0 mm) of the beetle, one of the biggest representatives of the family. Type locality. Cairns district, Queensland. Type material. SAMA (holotype) and QMBA (paratype). References. Arrow 1929: 309 (description in key, systematic position); Schenkling 1934: 3 (distribution); Arrow 1937: 106 (description in key). Distribution. Australia (Queensland).

Althaesia pilosa Pascoe, 1860 Althaesia pilosa Pascoe, 1860: 117, pl. VI, fig. 4.

Althaesia arrowi Grouvelle, 1914 Althaesia Arrowi Grouvelle, 1914a: 497.

Name derivation. Specific epithet honours English coleopterologist Gilbert John Arrow (1873–1948). Type locality. île Larat [= Island Larat]. Type material. NHML (2 syntypes – male and female). References. Arrow 1929: 309 (description in key); Schenkling 1934: 3 (distribution); Arrow 1937: 106 (description in key). Distribution. Indonesia (Maluku Province: Island Larat).

Name derivation. Pilosa, -us (gr./lat.) – hairy, comose; likely due to the dense body setation. Type locality. Dorey. Type material. NHML. Note. In the original description the number of typespecimens is not provided. References. Gemminger and Harold 1868: 909 (distribution); Arrow 1929: 308 (description in key); Schenkling 1934: 3 (distribution); Arrow 1937: 106 (description in key). Distribution. New Guinea.

Althaesia sericea (Lea, 1921) Althaesia leai (Blackburn, 1894) Diplocoelus Leai Blackburn, 1894: 204 (in Althaesia – Arrow 1929: 308).

Name derivation. Specific epithet honours Australian coleopterologist Arthur Mills Lea (1868–1932), who had collected the type material. Type locality. N.[ew] S.[outh] Wales. Type material. NHML and ?SAMA. Note. In the original description the number of typespecimens is not provided. References. Kuhnt 1909: 120 (Diplocoelus, distribution); Kuhnt 1911: 83 (Diplocoelus, distribution –

Diplocoelus sericeus Lea, 1921b: 237 (in Althaesia – Arrow 1929: 309).

Name derivation. Sericeus (lat.) – silky, covered with glossy pubescence (from medieval word for silk – sericum). According to the original description, the name was given because: “the elytra have a curious mottled appearance, owing to numerous sericeouslooking patches, which alter their positions with the point of view” (Lea, 1921b: 237). Type locality. Coen River, Queensland. Type material. SAMA (holotype) and QMBA (paratype).

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Note. When preparing the description, Lea had two more specimens which probably belonged to the same species but differed somewhat from the type, and so he did not include them in the type-series. References. Arrow 1929: 309 (description in key, systematic position); Schenkling 1934: 3 (distribution); Arrow 1937: 106 (description in key); Lawrence and Ślipiński 2013: 264, fig. 73A (adult, illustration). Distribution. Australia (Queensland).

Name derivation. Arcanus (lat.) – secretive, clandestine, mysterious; so named because of the dark colouration of this beetle or due to the secretive biology. Type locality. Loja. Type material. MNHN (holotype). References. Schenkling 1934: 4 (Anobocoelus, distribution); Blackwelder 1945: 425 (Anobocoelus, distribution); Cline and McHugh 2010: 98 (distribution, systematic position). Distribution. Ecuador.

Genus Anchorius Casey, 1900 Anchorius Casey, 1900: 79.

Name derivation. Anchor (lat.) – anchor. Gender. Masculine. Type species. Anchorius lineatus Casey, 1900: 80, by monotypy. Anobocoelus Sharp, 1902: 625. Synonymized by Cline and McHugh 2010: 98.

Name derivation. Anobioo (gr.) – revive, resuscitate – refers to the name of the genus Anobium (Anobiidae), whose representatives show general similarity to the beetles discussed here, and coelus (lat.) (gr. koilos – empty, hollow), alluding to the name Diplocoelus. Gender. Masculine. Type species. Anobocoelus championi Sharp, 1902: 626, here designated Note. Sharp (1902: 625) described Anobocoelus for two species, A. championi and A. optatus, but he did not designated a type species. References. Kuhnt 1909: 120 (description, world catalogue); Kuhnt 1911: 83 (world catalogue); Leng 1920: 202 (catalogue of North American species); Schenkling 1934: 4 (Anobocoelus, supplement to world catalogue); Blackwelder 1945: 245 (Anobocoelus, catalogue of Central and South American species); Arnett 1971: 790 (description in key); Goodrich and Springer 1992: 373 (description, distribution); Arnett 1997: 346 (distribution); Goodrich 2002: 356 (description, biology), 357 (catalogue of North American species); Peck 2005: 134 (Anobocoelus, catalogue of Cuban species); Cline and McHugh 2010: 98 (catalogue, systematic position). Immature stages. Lawrence 1991: figs. 34.554a-c (larva, illustration). Biology. Adults come strongly to light (Goodrich 2002: 356). Number of species. 5. Distribution. Neotropical and Nearctic Region.

Anchorius arcanus (Grouvelle, 1914) Anobocoelus arcanus Grouvelle, 1914b: 53 (in Anchorius – Cline and McHugh 2010: 98).

Anchorius championi (Sharp, 1902) Anobocoelus championi Sharp, 1902: 626, pl. XVIII, fig. 26 (in Anchorius – Cline and McHugh 2010: 98).

Name derivation. Specific epithet honours George Charles Champion (1851–1927), renowned British coleopterologist, who greatly contributed – as both collector (including the holotype of this species) and author – to the monumental series “Biologia Centrali Americana”. Type locality. Zapote [= El Zapote, Escuintla], Guatemala. Type material. NHML (holotype). Note. The type-specimen is worn and somewhat damaged (Sharp 1902: 626). References. Schenkling 1934: 4 (Anobocoelus, distribution); Blackwelder 1945: 425 (Anobocoelus, distribution); Cline and McHugh 2010: 98 (distribution, systematic position). Distribution. Guatemala.

Anchorius costulatus (Chevrolat, 1864) Diplocoelus costulatus Chevrolat, 1864: 615 (in Anchorius – Cline and McHugh 2010: 98).

Name derivation. Costula (lat.) – small rib (here: costa), and -atus (lat.) – suffix meaning -ate; costulate, adorned with costae. Type locality. Havane [= Havana]. Type material. MNHN (Coll. A. Grouvelle) (2 syntypes). Note. The species described from two specimens from the collection of Auguste Chevrolat (Chevrolat 1864: 615). Later, Grouvelle (1919: 66) stated that the type belonged to his own collection; now in MNHN. References. Gemminger and Harold 1868: 909 (Diplocoelus, distribution); Kuhnt 1909: 120 (Diplocoelus, distribution); Kuhnt 1911: 83 (Diplocoelus, distribution); Grouvelle 1919: 66 (Anobocoelus, systematic position); Schenkling 1934: 4 (Anobocoelus, distribution); Blackwelder 1945: 425 and 469 (Anobocoelus, distribution); Peck 2005: 134 (Anobocoelus,

CATALOGUE OF BIPHYLLIDAE (COLEOPTERA) OF THE WORLD

distribution); Cline and McHugh 2010: 98 (distribution, systematic position). Distribution. Cuba.

Anchorius lineatus Casey, 1900 Anchorius lineatus Casey, 1900: 80.

Name derivation. Lineatus (lat.) – lineate; from linea (lat.) – line, and -atus (lat.) – suffix -ate; name referring to longitudinal ridges on pronotum. Type locality. Arizona. Type material. USNM (holotype) (Goodrich and Springer 1992: 373). References. Kuhnt 1909: 120 (distribution); Kuhnt 1911: 83 (distribution); Leng 1920: 202 (distribution); Arnett 1971: 790 (distribution); Lawrence 1991: 475, figs. 34.554a-c (biology, larva). Goodrich and Springer 1992: 373, figs. 1, 6, 11, 14, 17, and 20 (description, distribution); Arnett 1997: 346 (distribution); Goodrich 2002: 356 (biology), 357 (distribution), fig. 2.89 (first abdominal sternite); Cline and McHugh 2010: 98 (distribution, systematic position); Cline and Shockley 2010: 306 (as Anchorius sp., biology); Lawrence et al. 2011: 161, fig. 80H (mandible). Immature stages. Lawrence 1991: figs. 34.554a-c (larva, illustration – general view, head, and mandible). Biology. Found under bark of trees, also fermenting, especially frequent on mesquite – Prosopis sp. (Fabaceae) (Lawrence 1991: 475; Goodrich and Springer 1992: 373; Goodrich 2002: 356; Cline and Shockley 2010: 306). Often comes to light (Goodrich and Springer 1992: 373). Distribution. USA (Arizona, California), Mexico.

Anchorius optatus (Sharp, 1902) Anobocoelus optatus Sharp, 1902: 625 (in Anchorius – Cline and McHugh 2010: 98).

Name derivation. Optatus (lat.) – desired. Type locality. Vera Cruz. Type material. NHML (2 syntypes). Note. Both syntypes disintegrated (Sharp 1902: 625). References. Schenkling 1934: 4 (Anobocoelus, distribution); Blackwelder 1945: 425 (Anobocoelus, distribution); Cline and McHugh 2010: 98 (distribution, systematic position). Distribution. Mexico.

Genus Biphyllus Dejean, 1821 Biphyllus Dejean, 1821: 102.

415

Name derivation. Bi (lat.) – double, and phyllon (gr.) – leaf; the name refers to dimerous antennal club in this genus. Rather objectionable combination of Latin numeral with Greek stem, which provoked the later emendation (see remarks on Diphyllus). Gender. Masculine. Type species. Dermestes lunata [sic!] Fabricius, 1787: 378, by monotypy. Diphyllus Berthold, 1827: 393.

Name derivation. Dis (gr.) – twice, and phyllon (gr.) – leaf; the result of emendation of the hybrid (Latin/Greek) name Biphyllus. Note. Despite linguistic correctness this emendation is not justified according to the rules of the Code of Zoological Nomenclature, and so must be treated as a junior objective synonym of Biphyllus – according to article 19.1 (ICZN 1999). Sphaeriophagus Kirby in Stephens, 1829: 88 (nomen nudum).

Name derivation. Sphaeria Haller (a group of fungi belonging to Ascomycota), and phagein (gr.) – eat, devour; the genus contains mycetophagous beetles which, according to the author of this name, feed on the fruiting bodies of fungi. Note. Name mentioned among synonyms of Biphyllus. Thallestus Wollaston, 1862: 153. Synonymized with Diphyllus by Reitter 1880b: 86 [16].

Name derivation. Thallos (gr.) – branch, and estos (gr.) – amusing oneself. Gender. Masculine. Type species. Thallestus subellipticus Wollaston, 1862: 155, subsequent designation by Sasaji 1985a: 1. References. Stephens 1830: 87 (description); Westwood 1838: 14 (description); Shuckard, 1839: 178 (description); Schaum 1840: 105 (Diphyllus, systematic position); Redtenbacher 1849: 188 (Diphyllus, description); Jacquelin du Val 1857: 102 (Diphyllus, systematic position); Redtenbacher 1858: 358 (Diphyllus, description); Jacquelin du Val 1859: 213 (description), 114 [catalogue] (catalogue of European species); Schaum 1859: XXIV (Diphyllus, systematic position); Schaum 1860: XXII (Diphyllus, systematic position); Kiesenwetter 1862: 416 (Thallestus, systematic position); Perris 1862: 196 (Diphyllus, systematic position); Thomson 1863: 239 (Diphyllus, description, Scandinavian species); Marseul 1866: 49 (catalogue of European species); Gemminger and Harold 1868: 881 (Thallestus, world catalogue), 909 (Diphyllus, world catalogue); Stein 1868: 49 (Diphyllus, catalogue of European species); Kraatz 1869: 28 (Diphyllus, catalogue of German species); Redtenbacher 1874: 392

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(Diphyllus, description); Stein and Weise 1877: 76 (Diphyllus, catalogue of European species); Lewis 1879: 11 (Thallestus, catalogue of Japanese species); Reitter 1880b: 86 [16] (Diphyllus, key for the identification of European species); Reitter in Heyden et al. 1883: 78 (Diphyllus, catalogue of European species); Reitter 1885: 20 (Diphyllus, key for the identification of European species); Schilsky 1888: 54 (Dipyllus, catalogue of Central European species); Schönfeldt 1888: 47 (Thallestus, catalogue of Japanese species); Reitter 1889a: 301 (Diphyllus, key for the identification of Japanese species); Reitter in Heyden et al. 1891: 151 (Diphyllus, catalogue of European species); Schoenfeldt 1891: 257 (Diphyllus, catalogue of Japanese species); Ragusa 1892: 131 (Diphyllus, catalogue of Sicilian species); Ganglbauer 1899: 651 (Diphyllus, description, key for the identification of Central European species); Casey 1900: 79 (discussion on nomenclature); Grouvelle 1906: 137 (Diphyllus, key for the identification of Madagascan species); Reitter in Heyden et al. 1906: 338 (Diphyllus, catalogue of European species); Kuhnt 1909: 117 (Diphyllus, description, world catalogue); Schilsky 1909: 91 (Diphyllus, catalogue of Central European species); Kuhnt 1911: 81 (Diphyllus, world catalogue); Reitter 1911: 73 (Diphyllus, description, European species); Jakobson 1915: 936 (catalogue of Palearctic species); Grouvelle 1916: 48 (Diphyllus, key for the identification of Indian species); Schaufuss 1916: 483 (Diphyllus, description), 484 (description of larva); Schenkling 1923: 16 (Thallestus, world catalogue); Mader 1926: 718 (Diphyllus, catalogue of Palearctic species) and 721 (Thallestus, catalogue of Palearctic species – incorrectly listed species that previously, on pages 718–719, were included in the genus Diphyllus); Portevin 1931: 193 (Diphyllus, key for the identification of French species); Falcoz 1926: 73 (Diphyllus and Thallestus – both genera bonum, systematic position); Miwa 1929: 121 (Diphyllus, systematic position); Miwa 1931: 76 (Diphyllus, catalogue of Taiwanese species); Schenkling 1934: 1 (supplement to the world catalogue); Blair 1934: 95 (Diphyllus, biology); Hellen 1947: 43 (Diphyllus, catalogue of Finnish and Swedish species); Vogt 1967: 108 (Diphyllus, description, key for the identification of Central European species); Bangsholt et al. 1979: 49 (catalogue of Scandinavian species); Nikitsky 1983: 695 (description) and 698 (key for the identification of East Palearctic species); Sasaji 1983: 28 (review of Japanese species) and 36 (key for the identification of Japanese species); Sasaji 1985a: 1–2 (catalogue and distribution of Japanese species); Burakowski et al. 1986: 208 (catalogue of Polish species); Lucht 1987: 187 (Diphyllus, catalogue of Central European species); Hirashima et al. 1989: 379 (catalogue of Japanese species); Basset 1991: 174 (biology, Australia); Nikitsky 1992: 280 (key for identification of

Russian Far East species); Jelínek 1993: 102 (catalogue of Czechoslovak species); Poggi 1995: 7 (catalogue of Italian species); Alexandrovitch et al. 1996: 41 (catalogue of Belarusian species); Köchler and Klausnitzer 1998: 111 (catalogue of German species); Machado and Oromí 2000: 63 (catalogue of Canarian species); Węgrzynowicz, 2002: 449, table 1, figs. 140–143 (morphology, phylogeny); Jelínek 2007: 547 (catalogue of the Palearctic species, distribution); Park et al. 2012: 185 (revision of South Korean species, keys for identification, descriptions, distribution, biology, illustrations). Immature stages. Perris 1851b: 42 (description); Chapuis and Candèze 1853: 431, pl . II, fig. 7 (description, illustration); Rupertsberger 1879: 219; Letzner 1886: 280 (description); Ganglbauer 1899: 653 (description); Reitter 1911: fig. 24 (description); Schaufuss 1916: 484 (description); Nikitsky 1983: 695 (description) Rymer Roberts 1958: 277, fig. 47 (description, illustration). Biology. Mycetophagous beetles, developing on dendrophilous fungi. Unidentified species (close to Biphyllus minutus (Grouvelle)) belonging to this genus had been collected in the Dark Cave (Selangor) in July, 2–1600 ft. (= 0,7–488 m.) from the mouth and in the same cave in November, 600–1200 ft. (= 185–365 m.) from the mouth (Blair 1929: 381). Unidentified species had been collected in Queensland from traps on Argyrodendron actinophyllum (Bailey) Edlin (Malvaceae) (Basset 1991: 174). Number of species. 89. Distribution. Old world.

Biphyllus aequalis (Reitter, 1889) Diphyllus aequalis Reitter, 1889a: 300 (in Biphyllus – Jakobson, 1915: 936).

Name derivation. Aequalis (lat.) – uniform; likely due to the unicolorous dorsal habitus. Type locality. Japan. Type material. ?HNHM (holotype). References. Schoenfeldt 1891: 257 (Diphyllus, distribution); Kuhnt 1909: 118 (Diphyllus, distribution); Kuhnt 1911: 82 (Diphyllus, distribution); Jakobson 1915: 936 (distribution); Mader 1926: 719 (Diphyllus, distribution); Nakane 1963: 198 (description, distribution), pl. 99, fig. 19 (habitus, colour); Sasaji 1983: 29 (distribution) and 38 (description in key); Nikitsky 1983: 700, fig. 1 (3) (description in key); Hirashima et al. 1989: 379 (distribution); Sasaji 1985a: 1 (distribution); Nikitsky 1992: 282, fig. 127.7 (description in key); Sasaji 1998: 208, 211, pl. 33, fig. 25 (description in key, distribution); Anonym 2000: 272 (distribution); Nomura et al. 2000: 225 (distribution, biology); Nomura et al.

CATALOGUE OF BIPHYLLIDAE (COLEOPTERA) OF THE WORLD

2006: 213 (distribution, biology); Jelínek 2007: 547 (distribution). Biology. The holotype of this species was collected at the beginning of June (Reitter 1889a: 300). On Honshû, the species was collected in May (Nomura et al. 2006: 213) and October (Nomura et al. 2000: 225), on Kyûshû in May and July (Sasaji 1983: 29). Distribution. Japan (Honshû; Kyûshû – Fukuoka Prefecture).

Biphyllus africanus (Grouvelle, 1914)

417

Biphyllus andrewesi (Grouvelle, 1916) Diphyllus Andrewesi Grouvelle, 1916: 33 (in Biphyllus – Schenkling 1934: 1).

Name derivation. Name honouring British coleopterologist Herbert Edward Andrewes (1863–1950). Type locality. Nilgiri Hills, Inde [= India]. Type material. NHML (Coll. H. E. Andrewes) and MNHN (Coll. A. Grouvelle). Note. In the original description the number of typespecimens is not provided. References. Schenkling 1934: 1 (distribution). Distribution. India (Nilghiri Hills).

Diphyllus africanus Grouvelle, 1914c: 179 (in Biphyllus – Schenkling 1934: 1).

Biphyllus bolivari (Grouvelle, 1905) comb. nov. Name derivation. Africanus (lat.) – African; name refers to the distribution of the species. Type locality. Sénégal. Type material. MNHN (Coll. A. Grouvelle) (holotype). References. Schenkling 1934: 1 (distribution). Distribution. Senegal.

Biphyllus alluaudi (Grouvelle, 1906) comb. nov. Diphyllus Alluaudi Grouvelle, 1906: 128.

Name derivation. Specific epithet honours French coleopterologist, explorer and collector Charles Alluaud (1861–1949), especially meritorious in studies of beetles of Madagascar and surrounding islands. Type locality. Forèt de Tanala [= Forest Tanala]; Andrangoloaka, près [= near] Tananarive. Type material. MNHN (Colls. A. Grouvelle and Ch. Alluaud). Note. In the original description the number of typespecimens is not provided. References. Kuhnt 1911: 82 (Diphyllus, distribution). Distribution. Madagascar.

Biphyllus amabilis (Grouvelle, 1916) Diphyllus amabilis Grouvelle, 1916: 30 (in Biphyllus – Schenkling 1934: 1).

Name derivation. Amabilis (lat.) – kind, lovable. Type locality. Pegu [= Bago]. Type material. MNHN (Coll. A. Grouvelle) (holotype). References. Schenkling 1934: 1 (distribution). Distribution. Burma.

Diphyllus Bolivari Grouvelle, 1905a: 256.

Name derivation. Most likely, the specific epithet honours Spanish entomologist Ignacio Bolívar y Urrutia (1850–1944). Type locality. Cabo San Juan, Biafra [= Bight of Biafra (now Bight of Bonny), Spanish Guinea]; Gabon. Type material. MNHN (Coll. A. Grouvelle). Note. In the original description the number of typespecimens is not provided. References. Kuhnt 1909: 119 (Diphyllus, distribution); Kuhnt 1911: 82 (Diphyllus, distribution). Distribution. Equatorial Guinea, Gabon.

Biphyllus brevis (Grouvelle, 1914) Diphyllus brevis Grouvelle, 1914c: 174 (in Biphyllus – Schenkling 1934: 1).

Name derivation. Brevis (lat.) – short, small; likely due to the diminutive size (~ 1.0 mm in length) of the beetle. Type locality. Afrique orientale [= East Africa]. Type material. MNHN (Coll. A. Grouvelle) (holotype). References. Schenkling 1934: 1 (distribution). Distribution. “East Africa”.

Biphyllus brunnescens (Reitter, 1877) comb. nov. Thallestus brunnescens Reitter, 1877: 138.

Name derivation. Brunneus (lat.) – brown, and -escens (lat.) – becoming, changing into; likely due to the colouration of the beetle. Type locality. India orientalis. Type material. MIZPAN (Coll. C. A. Dohrn). Note. In the original description number of specimens was not provided.

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References. Schenkling 1923: 16 (Thallestus, distribution). Distribution. “East India”.

Biphyllus canaliculatus (Grouvelle, 1906) comb. nov. Diphyllus canaliculatus Grouvelle, 1906: 131.

Name derivation. Canaliculus (lat.) – fine channel, and -atus (lat.) – suffix meaning -ate; likely due to pronotal sculpture being in the form of longitudinal sulci. Type locality. Nord du pays Androy [= north of Androy Country]; Diego Suarez; Suberbieville. Type material. MNHN (Colls. Ch. Alluaud, S. A. Sicard, L. Fairmaire, and A. Grouvelle) (pers. obs.). References. Kuhnt 1911: 82 (Diphyllus, distribution). Distribution. Madagascar.

Type locality. Malvern, Natal. Type material. NHML. Note. In the original description the number of typespecimens is not provided. References. Schenkling 1934: 1 (distribution). Biology. The type material was collected in February (Arrow 1929: 312). Distribution. Republic of South Africa (Natal).

Biphyllus complexus Sasaji, 1983 Biphyllus complexus Sasaji, 1983: 31, figs. 4A and D.

Name derivation. Specific epithet honours père [= father] Cardon, who collected holotype of this species. Type locality. Barway, Inde [= India]. Type material. IRSNB (holotype only). References. Schenkling 1934: 1 (distribution). Distribution. India (Uttarakhand).

Name derivation. Complexus (lat.) – embracing, encircling; likely due to the complicated pattern of elytral setation: “Elytral pubescens [...] arranged in very complex currents as figured, strongly turning laterally at several parts” (Sasaji 1983: 31). Type locality. Mt. Hikosan, Fukuoka Pref., Kyûshû. Type material. KUEC (holotype – male, and 3 paratypes – male and 2 females). References. Sasaji 1985a: 1 (distribution); Hirashima et al. 1989: 379 (distribution); Nikitsky 1992: 280 (description in key); Sasaji 1998: 208, 211, pl. 33, fig. 29 (description in key, distribution); Jelínek 2007: 547 (distribution). Biology. The type material was colected between May and August (Sasaji 1983: 33–34). Distribution. Japan (Honshû – Prefectures Tôkyô and Kanagawa; Shikoku – Tokushima Prefecture; Kyûshû – Fukuoka Prefecture).

Biphyllus centromaculatus (Grouvelle, 1916)

Biphyllus concolor (Grouvelle, 1906) comb. nov.

Biphyllus cardoni (Grouvelle, 1916) Diphyllus Cardoni Grouvelle, 1916: 39 (in Biphyllus – Schenkling 1934: 1).

Diphyllus centromaculatus Grouvelle, 1916: 49 (in Biphyllus – Schenkling 1934: 1).

Name derivation. Centrum (lat.) – middle, and maculatus (lat.) – spotted; likely due to the differently coloured setal patch at the middle of the elytra. Type locality. Usambara. Type material. MNHN (Coll. A. Grouvelle). Note. In the original description the number of typespecimens is not provided. References. Schenkling 1934: 1 (distribution). Distribution. Tanzania (Usambara Mts.).

Diphyllus concolor Grouvelle, 1906: 130.

Name derivation. Concolor (lat.) – of equal colouration; likely due to the uniformly dark-brown colouration and homogeneous yellow pubescence. Type locality. Andrangoloaka, près [= near] Tananarive. Type material. MNHN (Coll. A. Grouvelle). Note. In the original description the number of typespecimens is not provided. References. Kuhnt 1911: 82 (Diphyllus, distribution). Distribution. Madagascar.

Biphyllus clavatus Arrow, 1929 Biphyllus clavatus Arrow, 1929: 312.

Biphyllus convexus (Reitter, 1878) comb. nov. Thallestus convexus Reitter, 1878: 185.

Name derivation. Clavatus (lat.) – club-shaped, from clava (lat.) – club, and -atus (lat.) – suffix -ate; likely due to the unusually wide antennal club.

Name derivation. Convexus (lat.) – convex, hunched; likely due to the convex body shape.

CATALOGUE OF BIPHYLLIDAE (COLEOPTERA) OF THE WORLD

Type locality. Ceylon [= Sri Lanka]. Type material. Unknown. References. Haly 1890: 103 (Thallestus, distribution); Schenkling 1923: 16 (Thallestus, distribution). Distribution. Sri Lanka.

Biphyllus corpulentus Arrow, 1929 Biphyllus corpulentus Arrow, 1929: 312.

Name derivation. Corpus (lat.) – body, and -lentus (lat.) – suffix meaning abundance, inclinationy. Type locality. Entebbe. Type material. NHML. Note. In the original description the number of typespecimens is not provided. References. Schenkling 1934: 1 (distribution). Biology. The type material was collected in February (Arrow 1929: 313). Distribution. Uganda.

Biphyllus decoratus (Grouvelle, 1916) Diphyllus decoratus Grouvelle, 1916: 31 (in Biphyllus – Schenkling 1934: 1).

Name derivation. Decoro (lat.) – decorate, and -atus (lat.) – suffix -ated; decorated, in view of the ornate elytral pattern. Type locality. Shembaganur. Type material. MNHN (Coll. A. Grouvelle). Note. In the original description the number of typespecimens is not provided. References. Schenkling 1934: 1 (distribution). Distribution. India (Shembaganur).

Biphyllus distinctus (Grouvelle, 1916) Diphyllus distinctus Grouvelle, 1916: 37 (in Biphyllus – Schenkling 1934: 1).

Name derivation. Distinctus (lat.) – distinct, differentiated; likely due to the distinctive elytral pattern. Type locality. Nilgiri Hills [= Nilghiri Hills]. Type material. NHML (Coll. H. E. Andrewes) (holotype). References. Schenkling 1934: 1 (distribution). Distribution. India (Nilghiri Hills).

Biphyllus dohrni (Reitter, 1877) comb. nov. Thallestus Dohrni Reitter, 1877: 136.

Name

derivation. Specific epithet honours

419

German coleopterologist and collector Carl August Dohrn (1806–1892). Type locality. India orientalis [= East India]. Type material. MIZPAN (Coll. C. A. Dohrn). Note. In the original description the number of typespecimens is not provided. References. Schenkling 1923: 16 (Thallestus, distribution). Distribution. “East India”.

Biphyllus dollmani Arrow, 1929 Biphyllus dollmani Arrow, 1929: 310.

Name derivation. Name honouring the collector of the type material, Hereward Chune Dollman (1888– 1919). Type locality. Mwengwa, N. W. Rhodesia [now Zambia]. Type material. MNHL. Note. In the original description the number of typespecimens is not provided. References. Schenkling 1934: 1 (distribution). Biology. The type material was collected in November (Arrow 1929: 311). Distribution. Zambia.

Biphyllus egens (Grouvelle, 1916) Diphyllus egens Grouvelle, 1916: 45 (in Biphyllus – Schenkling 1934: 1).

Name derivation. Egens (lat.) – poor; likely due to indistinctive body colouration. Type locality. Nilgiri Hills [= Nilghiri Hills]. Type material. NHML (Coll. H. E. Andrewes) (syntype) and MNHN (Coll. A. Grouvelle) (syntype). References. Schenkling 1934: 1 (distribution). Distribution. India (Nilghiri Hills).

Biphyllus elegans Arrow, 1929 Biphyllus elegans Arrow, 1929: 311.

Name derivation. Elegans (lat.) – showy, elegant; likely due to the beauty and elegance of the beetle. Type locality. Port St. John, Pondoland, Cape Prov. Type material. NHML. Note. In the original description the number of typespecimens is not provided. References. Schenkling 1934: 1 (distribution). Biology. The type material was collected in September (Arrow 1929: 311).

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Distribution. Republic of South Africa (Cap Colony).

Biphyllus escalerae (Grouvelle, 1905) comb. nov. Diphyllus Escalerae Grouvelle, 1905a: 257.

Name derivation. Specific epithet honours Manuel Martinez de Escalera (1867–1936). Type locality. Cabo San Juan, Biafra [= Bight of Biafra (now Bight of Bonny)]. Type material. ?MNHN (Coll. A. Grouvelle). Note. In the original description the number of typespecimens is not provided. References. Kuhnt 1909: 119 (Diphyllus, distribution); Kuhnt 1911: 82 (Diphyllus, distribution). Distribution. Equatorial Guinea.

Biphyllus euphorbiae (Peyerimhoff, 1923) Diphyllus euphorbiae Peyerimhoff, 1923: 51 (in Biphyllus – Schenkling 1934: 1).

Name derivation. Euphorbiae (gr./lat.) – related to spurge; the type series was collected on branches of spurge (Euphorbia L.), which is the basis for the specific epithet. Type locality. Agadir, Vallé de Sous. Type material. ?MNHN. Note. In the original description the number of typespecimens is not provided. References. Mader 1926: 719 (Diphyllus, distribution); Schenkling 1934: 1 (distribution); Kocher 1956: 56 (distribution, biology); Jelínek 2007: 547 (distribution). Biology. The type material was collected in April on dead branches of Euphorbia beaumierana Hooker et Cosson (= E. officinarum L.) (Euphorbiacae) (Peyerimhoff 1923: 52). Reported generally from Euphorbia sp. (Kocher 1956: 56). Distribution. Morocco.

Biphyllus fastidiosus (Grouvelle, 1916) Diphyllus fastidiosus Grouvelle, 1916: 42 (in Biphyllus – Schenkling 1934: 1).

Name derivation. Fastidiosus (lat.) – hideous, abominable. Type locality. Barway. Type material. IRSNB (holotype). References. Schenkling 1934: 1 (distribution). Distribution. India (Uttarakhand).

Biphyllus flavonotatus (Lea, 1921) Diphyllus flavonotatus Lea, 1921a: 364 (in Biphyllus – Schenkling 1934: 1).

Name derivation. Flavus (lat.) – yellow, nota (lat.) – mark, and -atus (lat.) – suffix of past participle, -ate; altogether meaning “marked with yellow”, likely due to the yellow spots on elytra. Type locality. Ourimbah. Type material. SAMA (holotype). References. Schenkling 1934: 1 (distribution). Distribution. Australia (New South Wales).

Biphyllus flexuosus (Reitter, 1889) Diphyllus flexuosus Reitter, 1889a: 300 (in Biphyllus – Jakobson 1915: 936).

Name derivation. Flexuosus (lat.) – wavy, zigzaggy; likely due to the pale zigzag spots on the elytra. Type locality. Hiogo [= Hyôgo]; Oyama, Sapporo. Type material. ?HNHM (Coll. E. Reitter). Note. In the original description the number of typespecimens is not provided. References. Schoenfeldt 1891: 257 (Diphyllus, distribution); Kuhnt 1909: 118 (Diphyllus, distribution); Kuhnt 1911: 82 (Diphyllus, distribution); Grouvelle 1914d: 67 (Diphyllus, distribution); Jakobson, 1915: 936 (distribution); Mader 1926: 719 (Diphyllus, distribution); Miwa 1931: 76 (Diphyllus, distribution); Nakane 1963: 198 (description, distribution), pl. 99, fig. 21 (habitus, colour); Sasaji 1983: 31 (distribution), 38 (description in key); Nikitsky 1983: 702, fig. 1 (12) (description in key); Sasaji 1985a: 1 (distribution); Nakane 1988: 81 (description); Hirashima et al. 1989: 379 (as Biphyllus flexiosus [sic!] – error typographicum, distribution); Nikitsky 1992: 284, fig. 127.14 (description in key, biology); Sasaji 1998: 208, 211, pl. 33, fig. 28 (Biphyllus flexiosus [sic!] – error typographicum, description in key, illustration, distribution); Hua 2002: 109 (distribution); Jelínek 2007: 547 (distribution). Immature stages. Nikitsky 1983: 704 (description of larva), fig. 2 (1–7) (illustrations: 1 – head, 2 – antenna, 3 – mandible, 4 – labium and maxilla, 5 – hypopharynx, 6 – leg, 7 – VIII–X abdominal segments). Biology. Larvae are known from the periderm or cork layer of birches (Betula sp., Betulaceae), elms (Ulmus sp., Fagaceae), and Maack’shagberry (Prunus maacki (Rupr.) Komarov, Rosaceae), on surfaces covered with ascomycetous fungi. On elms they were observed, like larvae of Corticeus colydioides Lewis (Tenebrionidae, Coleoptera) and Europs sp. (Monotomidae, Coleoptera) (Nikitsky 1983: 704; Nikitsky 1992: 284) – in galleries of the bark-beetle Xyleborus

CATALOGUE OF BIPHYLLIDAE (COLEOPTERA) OF THE WORLD

orientalis Eggers (Curculionidae, Coleoptera) within the outer layers of wood. Some type material was collected in Honshû at the end of June (Reitter 1889a: 300). Distribution. Russia (Southern Primorye), Japan (Hokkaidô; Honshû – Prefectures Nagano, Gumma, and Fukui; Kyûshû – Prefectures Fukuoka and Kumamoto), Taiwan.

Biphyllus formosianus (Grouvelle, 1914) Diphyllus formosianus Grouvelle, 1914d: 67 (in Biphyllus – Schenkling 1934: 1).

Name derivation. Grouvelle gave this name because of the occurrence of the species on the island of Formosa [= Taiwan]. Type locality. Kosempo. Type material. DEIC (Coll. W. Horn) and MNHN (Coll. A. Grouvelle) (totally 5 syntypes). References. Miwa 1931: 76 (Diphyllus, distribution); Schenkling 1934: 1 (distribution); Hua 2002: 109 (distribution); Jelínek 2007: 547 (distribution). Distribution. Taiwan.

Biphyllus frater (Aubé, 1850) Diphyllus frater Aubé, 1850: 330 (in Biphyllus – Jacquelin du Val 1859: 114 [catalogue]).

Name derivation. frater (lat.) – brother; according to the author of the original description this species is similar to Diplocoelus fagi, and the name likely alludes to this similarity. Type locality. île de Sardaigne [= Sardinia]. Type material. MNHN (Coll. Ch. Aubé) (2 syntypes). References. Jacquelin du Val 1859: 114 [catalogue] (as Biphyllus Fraater [sic!], distribution); Marseul 1866: 49 (distribution); Gemminger and Harold 1868: 909 (Diplocoelus, distribution); Stein 1868: 49 (distribution); Kraatz 1869: 28 (Diplocoelus, distribution); Stein and Weise 1877: 76 (Diplocoelus, distribution); Reitter 1880b: 86 [16] (Diphyllus, description, distribution); Reitter in Heyden et al. 1883: 78 (Diphyllus, distribution); Frivaldszky 1884: 286 (Diphyllus, distribution); Reitter 1885: 20 (Diphyllus, description, distribution); Seidlitz 1888: 270 (Diphyllus, distribution); Seidlitz 1889: 289 (Diphyllus, distribution); Reitter in Heyden et al. 1891: 151 (Diphyllus, distribution); Ganglbauer 1899: 654 (Diphyllus, description, distribution); Reitter in Heyden et al. 1906: 338 (Diphyllus, distribution); Kuhnt 1909: 118 (Diphyllus, distribution); Kuhnt 1911: 82 (Diphyllus, distribution); Jakobson 1915: 936 (distribution); Schaufuss 1916: 484

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(Diphyllus, distribution); Kuthy 1918: 88 (Diphyllus, distribution); Mader 1926: 718 (Diphyllus, distribution); Portevin 1931: 193 (Diphyllus, description, distribution); Roubal 1936: 161 (Diphyllus, distribution, biology); Horion 1960: 214 (Diphyllus, distribution); Vogt 1967: 109 (Diphyllus, description in key, distribution); Lucht 1987: 187 (Diphyllus, distribution); Jelínek 1993: 102 (distribution); Poggi 1995: 7 (distribution); Kremer 1996: 94 (biology); Zúber et al. 1996: 122 (distribution, biology); Brustel and Rogé 1998: 206, fig 4 (Diphyllus, distribution, biology, illustration); Gräf 1998a: 9 (biology); Gräf 1998b: 113 (biology); Brustel and Clary 2000: 366 (distribution, biology); Rogé 2001: 234, fig. 2 (distribution, biology); Jelínek 2007: 547 (distribution); Recalde Irurzun and San Martín Moreno 2010: 152, fig. 1e (description, illustration, distribution, Iberian fauna, biology); Rusakov and Kalabkina 2013: 26, tab. 1 (distribution); Recalde Irurzun and San Martín Moreno 2014: 178 (distribution, biology). Biology. Collected in May on dead, trunks of deciduous trees in sunny places (Kremer 1996: 94). Under the bark (Roubal 1936: 191) or on trunks of Quercus sp. (Fagaceae) (Recalde Irurzun and San Martín Moreno 2010: 152). In the Czech Republic, specimens were collected on Turkey oak (Quercus cerris L., Fagaceae) (Gräf 1998a: 9; Gräf 1998b: 113) and on the stem of Aesculus sp. (Sapindaceae) – together with Synchita separanda (Reitter), S. mediolanensis (Villa et Villa), Cicones undatus (Guérin-Ménéville), and Colobicus hirtus (Rossi) (Colydiidae, Coleoptera) (Zúber et al. 1996: 122). In France and Spain, specimens were collected on pubescent oak (Quercus pubescens Willd. [= Q. humilis Mill.], Fagaceae) (Brustel and Rogé 1998: 206; Recalde Irurzun and San Martín Moreno 2014: 178). In the Czech Republic and France, specimens were collected from May to August (Zúber et al. 1996: 122; Brustel and Rogé 1998: 206; Brustel and Clary 2000: 366; Rogé 2001: 234), in Spain in May and June (Recalde Irurzun and San Martín Moreno 2010: 152; Recalde Irurzun and San Martín Moreno 2014: 178). Distribution. South Europe (Czech Republic, Slovakia, France, Austria, Hungary, Italy, Romania, Bosnia Herzegovina, Croatia, Slovenia, southern parts of European Russia and Ural Mts., Azerbaijan), Algeria.

Biphyllus frequens (Grouvelle, 1916) Diphyllus frequens Grouvelle, 1916: 35 (in Biphyllus – Schenkling 1934: 2).

Name derivation. Frequens (lat.) – frequent. Type locality. Nilgiri Hills [= Hilghiri Hills]. Type material. NHML (Coll. H. E. Andrewes) and MNHN (Coll. A. Grouvelle) (syntypes).

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Note. In the original description the number of typespecimens is not provided. References. Schenkling 1934: 2 (distribution). Distribution. India (Nilghiri Hills).

Biphyllus fulvus (Grouvelle, 1910) Diphyllus fulvus Grouvelle, 1910: 328 (in Biphyllus – Arrow 1929: 307).

Name derivation. Fulvus (lat.) – yellow; likely due to the dense yellow pubescence of the dorsal side. Type locality. Kibonoto, Kilimandjaro. Type material. ?NHRS (holotype). References. Arrow 1929: 307 (systematic position); Schenkling 1934: 2 (distribution). Biology. The holotype has been found in “cultivated areas” (Grouvelle 1910: 329). Distribution. Tanzania.

Biphyllus hildebrandti (Grouvelle, 1913) Diphyllus Hildebrandti Grouvelle, 1913b: 417 (in Biphyllus – Schenkling 1934: 2).

Name derivation. Specific epithet honours German botanist and traveller Jahann Maria Hildebrandt (1847–1881), meritorious explorer of Africa and Madagascar. Type locality. Madagascar. Type material. ZMHB. Note. In the original description the number of typespecimens is not provided. References. Schenkling 1934: 2 (distribution). Distribution. Madagascar.

Biphyllus histrio (Grouvelle, 1906) comb. nov. Diphyllus histrio Grouvelle, 1906: 135.

Name derivation. Histrio (lat.) – actor. Type locality. Diego Suarez. Type material. MNHN (Colls. S. A. Sicard and A. Grouvelle) (pers. obs.). Note. In the original description the number of typespecimens is not provided. References. Kuhnt 1911: 82 (Diphyllus, distribution). Distribution. Madagascar.

Name derivation. Humerus (lat.) – shoulder, and -alis (lat.) – suffix -al; the name refers to rusty spots on the elytral humeri. Type locality. Miyanoshita; Kiga; Yuyama. Type material. HNHM (syntype in Coll. E. Reitter, but subsequently labeled as holotype and not published (pers. obs.). Similar phenomenon was observed by Kamiński (2015) for Platynotina darkling beetles. Note. Number of type specimens has not been specified in the original publication. However, description implies that there were few individuals investigated (Reitter 1889a: 299). References. Schoenfeldt 1891: 257 (Diphyllus, distribution); Kuhnt 1909: 118 (Diphyllus, distribution); Kuhnt 1911: 82 (Diphyllus, distribution); Jakobson 1915: 936 (distribution); Mader 1926: 719 (Diphyllus, distribution); Sasaji 1983: 29 (distribution), 37 (description in key); Nikitsky 1983: 702, fig. 1 (9) (description in key); Sasaji 1985a: 1 (distribution); Hirashima et al. 1989: 379 (distribution); Nikitsky 1992: 283, fig. 127.12 (description in key); Sasaji 1998: 208, 211, pl. 33, fig. 26 (description in key, distribution); Narukawa 2001: 15 (description); Jelínek 2007: 547 (distribution). Distribution. Japan (Honshû – Prefectures Fukui and Hyôgo; Shikoku – Kôchi Prefecture; Kyûshû – Fukuoka Prefecture). Biphyllus inaequalis (Reitter, 1889) Diphyllus inaequalis Reitter, 1889a: 299 (in Biphyllus – Jakobson 1915: 936).

Name derivation. Inaequalis (lat.) – uneven. Type locality. Japan. Type material. ?HNHM (holotype). References. Schoenfeldt 1891: 257 (Diphyllus, distribution); Kuhnt 1909: 118 (Diphyllus, distribution); Kuhnt 1911: 82 (Diphyllus, distribution); Jakobson 1915: 936 (distribution); Mader 1926: 719 (Diphyllus, distribution); Sasaji 1983: 30 (distribution, morphological variability), 37 (description in key); Nikitsky 1983: 699, fig. 1 (5) (description in key); Sasaji 1985a: 1 (distribution); Nakane 1988: 80 (description); Hirashima et al. 1989: 379 (distribution); Sasaji 1998: 211, pl. 34, fig. 2 (description in key, distribution); Jelínek 2007: 547 (distribution). Biology. Collected from late May to early August (Sasaji 1983: 30). Distribution. Japan (Honshû – Prefectures Fukui and Wakayama; Shikoku – Prefectures Kanagawa and Ehime).

Biphyllus humeralis (Reitter, 1889)

Biphyllus infans (Grouvelle, 1914)

Diphyllus humeralis Reitter, 1889a: 299 (in Biphyllus – Jakobson 1915: 936).

Diphyllus infans Grouvelle, 1914c: 177 (in Biphyllus – Schenkling 1934: 2).

CATALOGUE OF BIPHYLLIDAE (COLEOPTERA) OF THE WORLD

Name derivation. Infans (lat.) – weak, slim; likely due to the small size and elongate body. Type locality. Afrique orientale [= East Africa]. Type material. MNHN (Coll. A. Grouvelle) (holotype). References. Schenkling 1934: 2 (distribution). Distribution. East Africa.

Biphyllus inops (Grouvelle, 1916) Diphyllus inops Grouvelle, 1916: 38 (in Biphyllus – Schenkling 1934: 2).

Name derivation. Inops (lat.) – lean, poor; likely due to distinctly elongate body. Type locality. Nilgiri Hills [= Nilghiri Hills]. Type material. NHML (Coll. H. E. Andrewes) and MNHN (Coll. A. Grouvelle). Note. In the original description the number of typespecimens is not provided. References. Schenkling 1934: 2 (distribution). Distribution. India (Nilghiri Hills).

Biphyllus insignis (Grouvelle, 1914) Diphyllus insignis Grouvelle, 1914c: 166 (in Biphyllus – Schenkling 1934: 2).

Name derivation. Insignis (lat.) – distinctive; name refers to the striking orange spots on elytra. Type locality. Afrique australe [=South Africa]. Type material. MNHN (Coll. A. Grouvelle) (holotype). References. Schenkling 1934: 2 (distribution). Distribution. South Africa.

Biphyllus japonicus Sasaji, 1983 Biphyllus japonicus Sasaji, 1983: 34.

Name derivation. Japonicus (lat.) – Japanese; from the country where it was collected. Type locality. Mt. Hoonji, Fukui Pref., Honshû. Type material. KUEC (holotype – male, and 30 paratypes). References. Sasaji 1985a: 2 (distribution); Nakane 1988: 81 (description); Hirashima et al. 1989: 379 (distribution); Nikitsky 1992: 280 (description in key); Sasaji 1998: 208, 211, pl. 33, fig. 27 (description in key, distribution); Narukawa 2001: 15 (description); Jelínek 2007: 547 (distribution). Biology. The type material was collected between late April and August (Sasaji 1983: 35). Distribution. Japan (Honshû – Prefectures Fukui and Huôgo).

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Biphyllus kasuganus Nakane, 1988 Biphyllus kasuganus Nakane, 1988: 81, pl. 9, fig. 8.

Name derivation. Derivation of Kasuga, where the species was collected. Type locality. Kasuga, Nara [Pref.]. Type material. SEHU (holotype – female). References. Hirashima et al. 1989: 379 (distribution); Narukawa 2001: 15 (distribution, biology), fig. (habitus); Jelínek 2007: 547 (distribution). Biology. Collected in June (Nakane 1988: 81; Narukawa 2001: 15). Collected from beech dead branches (Narukawa 2001: 15). Distribution. Japan (Honshû). Biphyllus kolosovi Nikitsky, 1983 Biphyllus kolosovi Nikitsky, 1983: 703, figs. 1, 2, 7, and 14.

Name derivation. Named in honour of the zoologist Andrei M. Kolosov. Type locality. V[illage] Kamenushka, Ussurysky region, Southern Primorye. Type material. ZMAS (holotype – male and paratype), ZMUM (paratype), and IBSS (2 paratypes). References. Nikitsky 1992: 282, figs. 127.4–6 (description in key); Jelínek 2007: 547 (distribution). Biology. Collected in June and August (Nikitsky 1983: 703). Distribution. Russia (Southern Primorye). Biphyllus kuzurius Sasaji, 1985 Biphyllus kuzurius Sasaji, 1985b: 11, fig. 1.

Name derivation. Derivation from the river Kuzuryû (meaning nine-headed dragon) in the Fukui Prefecture on the Japanese island Honshû where the type series was collected. Type locality. Izumi-mura [= village Izumi], Washikura-dake [= Mount Washikura], Fukui Prefecture, Honshû. Type material. KUEC (holotype – male, and 4 paratypes – females). References. Hirashima et al. 1989: 379 (distribution); Sasaji 1997: 27 (distribution, biology); Jelínek 2007: 547 (distribution). Biology. Collected in the end of April and early July (Sasaji, 1997: 27). Distribution. Japan (Honshû – Fukui Prefecture). Biphyllus lanuginosus (Grouvelle, 1914) Diphyllus lanuginosus Grouvelle, 1914c: 172 (in Biphyllus – Schenkling 1934: 2).

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Name derivation. Lanugo (gen. lanuginis) (lat.) – fluff, down, and -osus (lat.) – adjectival suffix meaning “full of”; likely due to the dense tomentose dorsal pubescence. Type locality. Afrique orientale [=South Africa]. Type material. MNHN (Coll. A. Grouvelle) (2 syntypes). References. Schenkling 1934: 2 (distribution). Distribution. East Africa.

Biphyllus latipes (Grouvelle, 1916) Diphyllus latipes Grouvelle, 1916: 34 (in Biphyllus – Schenkling 1934: 2).

Name derivation. Latus (lat.) – broad, and pes (lat.) – foot; likely due to the widened anterior tarsi. Type locality. Calcutta. Type material. MNHN (Coll. A. Grouvelle) (holotype). References. Schenkling 1934: 2 (distribution). Distribution. India.

Biphyllus lewisi (Reitter, 1889) Diphyllus Lewisi Reitter, 1889a: 299 (in Biphyllus – Jakobson 1915: 936).

Name derivation. Named in honour of the British coleopterologist George Lewis (1839–1926), who significantly contributed to the study of the Japanese fauna. Type locality. Japan. Type material. HNHM (Coll. E. Reitter – see Note below). Note. Original publication does not contain information on the depository and number of types – it might be a single specimen (Reitter 1889: 299). The Budapest Museum has one type-specimen of this species (pers. obs.). References. Schoenfeldt 1891: 257 (Diphyllus Lewisii [sic!], distribution); Kuhnt 1909: 118 (Diphyllus, distribution); Kuhnt 1911: 82 (Diphyllus, distribution); Jakobson 1915: 936 (distribution); Mader 1926: 719 (Diphyllus, distribution); Nakane 1963: 198 (description, distribution), pl. 99, fig. 18 (habitus, colour); Sasaji 1969: 13 (distribution, biology); Sasaji 1983: 30 (distribution, morphological variability), 38 (description in key); Nikitsky 1983: 701 (description in key); Sasaji 1985a: 2 (distribution); Nakane 1988: 80 (description); Hirashima et al. 1989: 379 (distribution); Nikitsky 1992: 283 (description in key); Sasaji 1998: 208, 211, pl. 33, fig. 24 (description in key, distribution); Iwai et al. 2001: 14 (distribution, biology); Jelínek 2007: 547 (distribution); Ohmomo et al. 2011: 85 (distribution, biology); Park et al. 2012: 187 (description,

distribution, biology), figs. 1a (pronotum), 2a (mesoand metasternite), 4a, d (male genitalia), 5a (female genitalia), 6a (habitus), 7a (elytron), 8a (antenna). Biology. The type material was collected by Reinhold Hiller in Japan at the beginning of November (Reitter 1889a: 299). On Honshû the species was found in the end of April until the begining of June (Iwai et al. 2001: 14; Ohmomo et al. 2011: 85), on Tsushima – in July (Sasaji 1969: 13). In South Korea it was collected from April to June (Park et al. 2012: 188). Distribution. Japan (Honshû – Prefectures Gumma, Gifu, and Fukui; Shikoku – Kôchi Prefecture; Kyûshû – Prefectures Fukuoka and Nagasaki (Tsushima Island)), South Korea.

Biphyllus liliputanus (Reitter, 1877) comb. nov. Thallestus liliputanus Reitter, 1877: 137.

Name derivation. The name means “Lilliputian” and derives from the fictive island of Lilliput, conceived by Jonathan Swift in the novel “Gulliver’s Travels” (1726); the island was described as inhabited by small (less than 6 inches) people, thence “lilliput” (optional spelling – “liliput”) means dwarf, midget. Edmund Reitter apparently wished to point out the small body size of this species, which measures ca. 1.5 mm in length. Type locality. India orientalis [= East India]. Type material. MIZPAN (Coll. C. A. Dohrn). Note. In the original description the number of typespecimens is not provided. References. Schenkling 1923: 16 (Thallestus, distribution). Distribution. “East India”.

Biphyllus Sasaji, 1991 Biphyllus loochooanus Sasaji, 1991: 12, figs. 1A–F.

Name derivation. From the Loochoo [= Ryukyu] Archipelago; the adjectival suffix -anus (lat.) denotes origin or affiliation. Type locality. Mt. Urabu, Yonaguni Island. Type material. KUEC (holotype – male, paratypes – 5 males and 2 females). References. Azuma et al. 2002: 224 (distribution); Jelínek 2007: 548 (distribution). Biology. The type material was collected at the end of December and beginning of January (Sasaji 1991: 12). Distribution. Japan (Ruykyus – Yonaguni Island).

Biphyllus lunatus (Fabricius, 1787) Dermestes lunata [sic!] Fabricius, 1787: 378 (in Biphyllus – Dejean 1821: 102).

CATALOGUE OF BIPHYLLIDAE (COLEOPTERA) OF THE WORLD

Name derivation. Lunatus (lat.) – semilunar; alludes to the distinctive pale semilunar elytral spots. Type locality. Suecia [= Sweden]. Type material. ZMHB (syntype) (pers. obs.). Note. Fabricius mentioned in the original description that the type-material came from the collection of Gustav Paykull. Biphyllus sphaeriae Stephens, 1829: 88 (nomen nudum).

Note. In the synonymy Stephens (1829) included the name Biphyllus lunatus. Diphyllus lunatus var. inornatus Reitter, 1888a: 170, syn. nov.

Name derivation. In- (lat.) – prefix meaning negation, and ornatus (lat.) – adorned; likely due to this taxon lacking elytral pale spots characteristic of the nominate form. Type locality. [Caucas.[us] or.[ientalis] [= East Caucasus], Circassien (pers. obs. – see Note below)]; Samara. Type material. HNHM (Coll. E. Reitter). Note. The single syntype in HNHM has been labelled as the holotype (pers. obs.), but Reitter had certainly possessed more than one specimen when preparing the description, and he did not designate a holotype in the original paper (Reitter 1888a: 170). References. Gmelin 1790: 1600 (Dermestes, description, biology); Dillwyn 1829: 23 (biology); Stephens 1830: 87 (description, distribution, biology), pl. XVII – fig. 3 (adult, colour illustration); Dejean 1835: 311 (catalogue); Dejean 1836: 336 (catalogue); Shuckard, 1839: 179 (distribution); Redtenbacher 1849: 188 (Diphyllus, description); Perris 1851a: 566 (Diphyllus, biology); Perris 1851b: 42, pl. 2, figs. 10–16 (Diphyllus, immature stages, biology); Chapuis and Candèze 1853: 431 (Diphyllus, immature stages); Dohrn 1853: 61 (Diphyllus, systematic position); Elditt 1854: 191 (Diphyllus, immature stages, catalogue); Redtenbacher 1858: 359 (Diphyllus, description); Jacquelin du Val 1859: 114 [catalogue] (distribution); Thomson 1863: 240 (Diphyllus, description, distribution); Wollaston 1864: 134 (Diphyllus, distribution, biology); Wollaston 1865: 159 (Diphyllus, distribution, biology); Marseul 1866: 49 (distribution); Gemminger and Harold 1868: 909 (Diphyllus, distribution); Stein 1868: 49 (Diphyllus, distribution); Kraatz 1869: 28 (distribution); Stierlin and Gautard 1869: 151 (distribution, biology); Seidlitz 1872: 183 (Diphyllus, description, distribution); Redtenbacher 1874: 392 (Diphyllus, description); Stein and Weise 1877: 76 (Diphyllus, distribution); Rupertsberger 1879: 219 (Diphyllus, larva); Reitter 1880b: 86 [16] (Diphyllus, description, distribution); Reitter in Heyden et al. 1883: 78 (Diphyllus, distribution); Reitter 1885: 20 (Diphyllus, description, distribution); Letzner 1886: 280 (Diphyllus, immature

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stages); Oertzen 1887: 231 (Diphyllus, distribution); Reitter 1888a: 170 (biology); Schilsky 1888: 54 (distribution); Seidlitz 1888: 270 (Diphyllus, description, distribution); Seidlitz 1889: 286 (Diphyllus, description, distribution); Reitter in Heyden et al. 1891: 151 (Diphyllus, distribution); Ragusa 1892: 131 (Diphyllus, distribution); Ganglbauer 1899: 653 (Diphyllus, larva, description, biology), 654 (description); Sahlberg 1900: 75 (Diphyllus, distribution); Jakobson 1905: pl. 23, fig. 14 (adult, colour illustration); Reitter in Heyden et al. 1906: 338 (Diphyllus, distribution); Kuhnt 1909: 118 (Diphyllus, distribution); Schilsky 1909: 91 (distribution); Kuhnt 1911: 82 (Diphyllus, distribution); Reitter 1911: 73, fig. 24 (larva), pl. 89, fig. 15 (habitus) (Diphyllus, description in key); Falcoz 1912: 38 (Diphyllus, biology); Kuhnt 1912: 530 (Diphyllus, description), figs. 4a (antenna) and 12 (habitus); Morley 1912: 4 (Diphyllus, distribution, biology); Jakobson 1915: 936 (distribution); Schaufuss 1916: 484 (Diphyllus, description, distribution, biology); Kuthy 1918: 88 (Diphyllus, distribution); Mader 1926: 718 (Diphyllus, distribution); Trella 1929: 131 (Diphyllus, biology); Portevin 1931: 193, fig. 210 (Diphyllus, description, illustration, biology); Schenkling 1934: 2 (distribution); Donisthorpe 1935: 25 (Diphyllus, biology); Siegemund 1935: 226 (Diphyllus, distribution, biology); Eagles 1936: 31 (Diphyllus, biology); Roubal 1936: 161 (Diphyllus, distribution, biology); Coulson 1938a: 37 (Diphyllus, biology); Coulson 1938b: 76 (Diphyllus, biology); Coulson 1939: 41 (Diphyllus, biology); Hellen 1947: 43 (Diphyllus, distribution); Buck 1949: 223 (distribution, biology); Duffy 1950: 76 (distribution, biology); Benick 1952: 127 (biology); Crowson 1952a: 69, fig. 113 (wing), 71, fig.121 (first abdominal ventrite); Crowson 1952b: 118 (biology); Hawkins 1952: 7 (Diphyllus, distribution, biology); Carlier 1954: 22 (Diphyllus, biology); Jacobs 1955: 92 (Diphyllus, distribution, biology); Crowson 1956: 35, fig. 91, and 38, fig. 99 (illustrations); Eagles 1956: 80 (distribution, biology); Kocher 1956: 56 (distribution); Rymer Roberts 1958: 277, fig. 47 (Diphyllus, larva, morphology); Palm 1959: 262 (Diphyllus, biology); Horion 1960: 212 (Diphyllus, distribution, biology); Gardiner 1965: 33 (distribution, biology); Crowson 1967: 92, (fig. 113 – wing), 94 (fig. 121 – first abdominal ventrite), 106 (biology); Vogt 1967: 109 (Diphyllus, description in key, distribution); Hingley 1971: 26 (biology); Bangsholt et al. 1979: 49 (distribution); Silfverberg 1992: 57 (distribution); Huddleston 1980: 53 (biology); Allen 1981: 90 (distribution, biology); McClenaghan 1982: 43 (distribution, biology); Sokoloff 1985: 6 (Biphylus [sic!], biology); Burakowski et al. 1986: 208 (distribution, biology); Jäch 1986: 29 (distribution, biology); Palm, 1986: 61 (morphology, distribution); Lucht 1987: 187 (Diphyllus, distribution); Alexander 1988: 128 (distribution, biology); Kirby and Lambert 1990:

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127 (distribution, biology); Lawrence 1991: 475 (biology); Halstead 1992: 73 (distribution, biology); Jelínek 1993: 102 (distribution); Alexander 1993: 98 (biology); Alexandrovitch et al. 1996: 41 (distribution); Poggi 1995: 7 (distribution); Hornig 1998: 9, tab.3 (B. lunulatus [sic!], distribution); Jones 1998: 172 (biology); Köchler and Klausnitzer 1998: 111 (distribution); Jones 1999: 201 (biology); Machado and Oromí 2000: 63 (distribution); Alexander 2002: 52 (distribution, biology); Grossa-Silva 2002: 52 (distribution); Theodorides et al. 2002: 468 (genetics); Węgrzynowicz, 2002: 449 (morphology, phylogeny); Hughes et al. 2006: 270, tabs. 1 and 2, figs. 2 and 3 (genetics, phylogeny); Shatters et al. 13 (genetics); 2006: Jelínek 2007: 548 (Biphyllus nornatus [sic!], distribution); Fedorenko 2009:330, fig. A188 (hind wing, morphology); Guéorguiev et al. 2010: 364 (distribution, biology); Recalde Irurzun and San Martín Moreno 2010: 152 (distribution); Vitali 2010: fig. 4d (antennal club); Tamutis et al. 2011: 254 (distribution); Lassauce et al. 2013: 407, appendix A (distribution, biology); Rusakov and Kalabkina 2013: 26, tab. 1 (distribution). Immature stages. Perris 1851b: 42, pl.2, figs. 10–16 (description of larva, illustrations); Chapuis and Candèze 1853: 431, pl . II, fig. 7 (description of larva, illustration – body, lateral view); Rupertsberger 1879: 219; Letzner 1886: 280 (description of larva); Ganglbauer 1899: 653 (description of larva); Reitter 1911: fig. 24 (larva); Rymer Roberts 1958: 277 (description of larva), fig. 47 (larva, mandible). Biology. Develops in Daldinia concentrica (Bolton) Cesati et Notaris (Xylariaceae) (Perris 1851b: 44; Wollaston 1865: 159 [on Sphaeria fraxinea Sibth. (= D. concentrica)]; Gangelbauer 1899: 654; Reitter 1911: 73; Falcoz 1912: 38; Morley 1912: 4; Schaufuss 1916: 484; Portevin 1931: 193; Donisthorpe 1935: 25; Siegemund 1935: 226; Eagles 1936: 31; Roubal 1936: 191; Coulson 1938a: 37; Buck 1949: 223; Benick 1952: 127; Crowson 1952b: 118; Carlier 1954: 22; Palm 1959: 262; Horion 1960: 213; Crowson 1967: 106; Hingley 1971: 26; Allen 1981: 90; Nikitsky 1983: 695; Sokoloff 1985: 6; Burakowski et al. 1986: 209; Lawrence 1991: 475; Halstead 1992: 73; Alexander 1993: 98; Jones 1999: 201; Jones 2000: 177; Alexander 2002: 52) and black fungus (Hypoxylon sp.) (Coulson 1938b: 76), infesting trunks of ash (Fraxinus excelsior L., Oleaceae) (Perris 1851b: 44; Carlier 1954: 22; Hingley 1971: 26; Alexander 1993: 98; Alexander 2002: 52) or other deciduous trees (Kuhnt 1909: 118; Palm 1959: 262). Reportedly occurs in black sooty spores of the ‘sooty bark disease’, i.e. fungus Cryptostroma corticale (Ellis et Everh.) Greg. et Waller, on Acer pseudoplatanus L. (Jones 1998: 172; Jones 2000: 178). Pupation in fructifications of host fungi (Hingley 1971: 26; Alexander 2002: 52) or in rotten, fungus-infested trunks of e.g. limetrees (Trella 1929: 131). Pupation in soil was reported

(Perris 1851b: 44; Ganglbauer 1899: 654; Reitter 1911: 73; Falcoz 1912: 38; Portevin 1931: 193; Palm 1959: 262; Burakowski et al. 1986: 209), but put in question by Trella (1929: 131). Adults were found under the bark of ash (Fraxinus sp., Oleaceae) (Dillwyn 1829: 23; Stephens 1830: 88) and – less frequently – birch (Betula sp., Betulaceae) (Coulson 1938a: 37; Coulson, 1939: 41; Palm 1959: 262; Sokoloff 1985: 6), oak (Quercus sp., Fagaceae) – on mould or in company of Siagonium humerale Frivaldszky in Germar (Staphylinidae, Coleoptera) and Cerylon sp. (Cerylonidae, Coleoptera) (Roubal 1936: 191), beeches (Fagus sp., Fagaceae) (Schaufuss 1916: 484; Roubal 1936: 191), Platanus orientalis L. (Platanaceae) (Guéorguiev et al. 2010: 364), and alders (Alnus sp., Betulaceae) (Burakowski et al. 1986: 209). Collected on trunks of old laurel (Laurus sp., Lauraceae) (Wollaston 1864: 134), as well as fungal covered birch (Hawkins 1952: 7). Found in galleries of Hylesinus crenatus (F.) (Scolytidae, Coleoptera) (Burakowski et al. 1986: 209). Collected on leaves of beech (Fagus sp., Fagaceae) (Reitter 1888a: 170). Swept from grass under ash (Allen 1981: 90). Attacked and consumed by Thanasimus formicarius L. (Cleridae, Coleoptera) (Carlier 1954: 22), parasitized by Meteorus sp. (Hingley 1971: 26) and Meteorus vexator (Haliday) (Braconidae, Hymenoptera) (Morley 1912: 4) – Huddleston (1980 : 53) and Stigenberg et Ronquist (2011: 75) question the reliability of these data, because this species parasitizes Psychidae (Lepidoptera). Distribution. Europe (Austria, Belarus, Belgium, Bosnia Hercegovina, Croatia, Czech Republic, Denmark, Finland, France, Germany, Great Britain, Greek, Hungary, Italy, Macedonia, The Netherlands, Norway, Poland, Portugal, Romania, southern parts of European Russia and Ural Mts., Serbia and Montenegro, Slovakia, Slovenia, Spain, Sweden, Ukraine), Canary Islands (La Palma, La Gomera, Gran Canaria), Morocco, Algeria, Libya, Madeira Archipelago, Iran.

Biphyllus maculatus (Grouvelle, 1906) comb. nov. Diphyllus maculatus Grouvelle, 1906: 130.

Name derivation. Macula (lat.) – spot, and -tus (lat.) – adjectival suffix; maculatus means spotted; likely due to the distinct pale elytral setal spots. Type locality. Andrangoloaka, près [= near] Tananarive. Type material. MNHN (Coll. A. Grouvelle). Note. In the original description the number of typespecimens is not provided. References. Kuhnt 1911: 82 (Diphyllus, distribution); Grouvelle 1913b: 417 (Diphyllus, distribution). Distribution. Madagascar.

CATALOGUE OF BIPHYLLIDAE (COLEOPTERA) OF THE WORLD

Biphyllus madagascariensis (Fairmaire, 1898) comb. nov. Triphyllus madagascariensis Fairmaire, 1898: 438.

Name derivation. Madagascariensis (lat.) – Madagascarian; name refers to the distribution of the species. Type locality. Tsarasotra. Type material. MNHN (Coll. A. Grouvelle). Note. In the original description the number of typespecimens is not provided. Note. Grouvelle (1906b: 137) stated that the species was erroneously included in the genus Triphyllus, but quotes himself as the author of the description. The cited publication, however, was written by Fairmaire, who does mention in the original description that it was based on a specimen (or specimens) from Grouvelle’s collection, but nothing more. Grouvelle as the author of the name was mistakenly quoted also by other authors – Alluaud (1900: 133) and Kuhnt (1911: 82). References. Alluaud 1900: 133 (Triphyllus, distribution); Grouvelle 1906: 137 (Diphyllus, systematic position, description in key); Kuhnt 1911: 82 (Diphyllus, world catalogue). Distribution. Madagascar.

Biphyllus magnus (Grouvelle, 1906) comb. nov. Diphyllus magnus Grouvelle, 1906: 129.

Name derivation. Magnus (lat.) – great, large; the species is distinctive among congeners by its large size. Type locality. Diego Suarez. Type material. MNHN (Colls. Ch. Alluaud, A. Grouvelle, and S. A. Sicard). Note. In the original description the number of typespecimens is not provided. References. Kuhnt 1911: 82 (Diphyllus, distribution). Distribution. Madagascar.

Biphyllus maindroni (Grouvelle, 1903) Diphyllus Maindroni Grouvelle, 1903a: 342, fig. 1 (in Biphyllus – Schenkling 1934: 2).

Name derivation. Specific epithet honours Maurice Maindron (1857–1911), French entomologist, popularizer of biological knowledge and outstanding expert in Carabidae (Coleoptera) systematics. Type locality. Coonoor, Nilghiris [= Nilghiri Hills]. Type material. MNHN (holotype). References. Kuhnt 1909: 119 (Diphyllus, distribution); Kuhnt 1911: 82 (Diphyllus, distribution);

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Grouvelle 1916: 48 (Diphyllus, description in key); Schenkling 1934: 2 (distribution). Biology. Holotype was collected at the end of July, at 1500 m. a.s.l, in a laceration of a tree-trunk (Grouvelle 1903a: 342). Distribution. India (Nilghiri Hills).

Biphyllus marmoratus (Reitter, 1889) Diphyllus marmoratus Reitter, 1889a: 300 (in Biphyllus – Jakobson 1915: 936).

Name derivation. Marmor (lat.) – marble, and -atus (lat.) – adjectival suffix meaning “-like”, “making appearance of”; likely due to “marbled” elytral pattern of multicoloured setulae. Type locality. Kiga; Higo. Type material. HNHM (Coll. E. Reitter). Note. Number of type specimens is not provided in the original publication (Reitter 1889a: 300); the single specimen preserved in HNHM has been secondarily marked as holotype (pers. obs.). References. Schoenfeldt 1891: 257 (Diphyllus, distribution); Kuhnt 1909: 118 (Diphyllus, distribution); Kuhnt 1911: 82 (Diphyllus, distribution); Jakobson 1915: 936 (distribution); Mader 1926: 719 (Diphyllus, distribution); Nakane 1963: 198 (description, distribution), pl. 99, fig. 20 (habitus, colour); Sasaji 1983: 31 (distribution), 38 (description in key); Nikitsky 1983: 702, fig. 1 (8 and 11) (description in key); Sasaji 1985a: 2 (distribution); Hirashima et al. 1989: 379 (distribution); Nikitsky 1992: 284, figs. 127.10 and 127.11 (description in key); Inoue and Sasaji 1998: 3 (distribution, biology); Sasaji 1998: 211, pl. 34, fig. 1 (description in key, distribution); Jelínek 2007: 548 (distribution); Park et al. 2012: 188 (description, distribution), figs. 1b (pronotum), 2b (meso- and metasternite), 3 (first abdominal sternite), 4b, e (male genitalia), 5b (female genitalia), 6b (habitus), 7b (elytron), 8b (antenna). Biology. In Japan collected in June and July (Reitter 1889a: 300; Sasaji 1983: 31; Inoue and Sasaji 1998: 3), in South Korea in May and June (Park et al. 2012: 189). Distribution. Russia (Southern Primorye), Japan (Honshû; Kyûshû – Prefectures Fukuoka and Kumamoto), South Korea.

Biphyllus marshalli (Grouvelle, 1911) Diphyllus Marshalli Grouvelle, 1911: 237 (in Biphyllus – Arrow 1929: 307).

Name derivation. The specific epithet honours Gay Anstruther Knox Marshall (1871–1959), collector of the type series.

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Type locality. Chirinda, Mashonaland. Type material. NHML and MNHN (Coll. A. Grouvelle) (totally 3 syntypes). References. Arrow 1929: 309 (distribution, systematic position); Schenkling 1934: 2 (distribution). Distribution. Zimbabwe (Mashonaland, Southern Rhodesia), Republic of South Africa (Zululand, Pondoland).

Biphyllus medius (Grouvelle, 1914) Diphyllus medius Grouvelle, 1914d: 72 (in Biphyllus – Schenkling 1934: 2).

Name derivation. Medius (lat.) – middle, medium. Type locality. Taihorin. Type material. DEIC (Coll. W. Horn) and/or MNHN (Coll. A. Grouvelle) (totally 2 syntypes). References. Miwa 1931: 77 (Diphyllus, distribution); Schenkling 1934: 2 (distribution); Hua 2002: 109 (distribution); Jelínek 2007: 548 (distribution). Distribution. Taiwan.

Biphyllus micros (Grouvelle, 1900) Diphyllus micros Grouvelle, 1900: 266 (in Biphyllus – Schenkling 1934: 2).

Name derivation. Mikros (gr.) – small; likely due to the small (ca. 1.5 mm) body size. Type locality. Indrapoera, Sumatra. Type material. ?MNHN. Note. In the original description the number of typespecimens is not provided. References. Kuhnt 1909: 119 (Diphyllus, distribution); Kuhnt 1911: 82 (Diphyllus, distribution); Schenkling 1934: 2 (distribution). Distribution. Indonesia (Sumatra).

Biphyllus minimus (Grouvelle, 1905) comb. nov. Diphyllus minimus Grouvelle, 1905a: 259.

Name derivation. Minimus (lat.) – very small, the smallest; name alluding to the small size of the beetle. Type locality. Cabo San Juan, Biafra [= Bight of Biafra (now Bight of Bonny)]. Type material. ?MNHN. Note. In the original description the number of typespecimens is not provided. References. Kuhnt 1909: 119 (Diphyllus, distribution); Kuhnt 1911: 82 (Diphyllus, distribution). Distribution. Equatorial Guinea.

Biphyllus minutus (Grouvelle, 1902) Diphyllus minutus Grouvelle, 1902b: 485 (in Biphyllus – Schenkling 1934: 2).

Name derivation. Minutus (lat.) – small; Grouvelle gave this name because of small size of the beetle. Type locality. Nalanda. Type material. DEIC (Coll. W. Horn) and MNHN (Coll. A. Grouvelle) (totally 3 syntypes). References. Kuhnt 1909: 119 (Diphyllus, distribution); Kuhnt 1911: 82 (Diphyllus, distribution); Schenkling 1934: 2 (distribution). Distribution. Sri Lanka.

Biphyllus molestus (Grouvelle, 1913) Diphyllus molestus Grouvelle, 1913a: 114 (in Biphyllus – Schenkling 1934: 2).

Name derivation. Molestus (lat.) – arduous. Type locality. Sadiya [river], N. E. Assam. Type material. ?MNHN (Coll. A. Grouvelle) (holotype). References. Schenkling 1934: 2 (distribution). Biology. Holotype was collected under bark (Grouvelle 1913a: 115). Distribution. India (Assam).

Biphyllus obscuronotatus (Lea, 1922) Diphyllus obscuronotatus Lea, 1922: 297 (in Biphyllus – Schenkling 1934: 2).

Name derivation. Obscurus (lat.) – indistinct, nota (lat.) – marking, and -atus (lat.) – past participle ending -ate, altogether means “indistinctly marked”; likely due to the two inconspicuous elytral spots. Type locality. Cairns district, Queensland. Type material. SAMA (holotype). References. Schenkling 1934: 2 (distribution); Grove 2000: 158, appendix A (distribution, biology); Grove 2002: 164 (distribution, biology). Biology. Collected in rainforest (Grove 2000: 158, appendix A; Grove 2002: 164). Distribution. Australia (Queensland).

Biphyllus obscurus (Reitter, 1877) comb. nov. Thallestus obscurus Reitter, 1877: 137.

Name derivation. Obscurus (lat.) – dark; likely due to the dark coloured body of this species. Type locality. India orientalis [= East India]. Type material. MIZPAN (Coll. C. A. Dohrn).

CATALOGUE OF BIPHYLLIDAE (COLEOPTERA) OF THE WORLD

Note. In the original description the number of typespecimens is not provided. References. Schenkling 1923: 16 (Thallestus, distribution). Distribution. “East India”.

Biphyllus odiosus (Grouvelle, 1913) Diphyllus odiosus Grouvelle, 1913a: 112 (in Biphyllus – Schenkling 1934: 2).

Name derivation. Odiosus (lat.) – hateful, abominable. Type locality. Kobo. Type material. ?MNHN (Coll. A. Grouvelle) (holotype). References. Schenkling 1934: 2 (distribution). Biology. Holotype was collected at 400 ft. (= 122 m.) a.s.l. (Grouvelle 1913a: 113). Distribution. India (Assam).

Biphyllus ornatellus (Blackburn, 1903) Diphyllus ornatellus Blackburn, 1903: 156 (in Biphyllus – Stone 1993: 293, tab. 2).

Name derivation. Ornatus (lat.) – adorned, and -ellus (lat.) – diminutive suffix; likely due to the small spots on the elytra. Type locality. Dividing Range, Victoria. Type material. NHML and ?SAMA. Note. In the original description the number of typespecimens is not provided. References. Kuhnt 1909: 119 (Diphyllus, distribution); Kuhnt 1911: 82 (Diphyllus, distribution); Stone 1993: 293, tab. 2 (distribution, biology); Grove 2000: 158, appendix A (distribution, biology). Biology. The type material was collected from under bark of Eucalyptus sp. (Myrtacae) (Blackburn 1903: 157) and on billets of loblolly pine (Pinus taeda L., Pinaceae) (Stone 1993: 293, tab. 2). Collected in rainforest (Grove 2000: 158, appendix A). Distribution. Australia (Queensland, New South Wales, Victoria).

Biphyllus oshimanus Nakane, 1988 Biphyllus oshimanus Nakane, 1988: 80, pl. 9, fig. 7.

Name derivation. Specific epithet derived from Amami Ôshima Island, where the species was collected. Type locality. Hatsuno, Amami-Oshima. Type material. SEHU (holotype – male and 4 paratypes – 2 females and 2 specimens with unknown sex).

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References. Hirashima et al. 1989: 379 (distribution); Sasaji 1991: 12 (distribution, biology); Azuma et al. 2002: 224 (distribution); Jelínek 2007: 548 (distribution). Biology. Species collected in March, May, July, August, and October (Nakane 1988: 80; Sasaji 1991: 12). Distribution. Japan (Kyûshû and Ryukyus: Amami Ôshima Island, Ishigaki Island, Iriomote Island).

Biphyllus parvulus (Grouvelle, 1906) comb. nov. Diphyllus parvulus Grouvelle, 1906: 133.

Name derivation. Parvus (lat.) – small, and -ulus (lat.) – diminutive suffix; likely due to the small body size (ca. 1.5 mm). Type locality. Fort-Dauphin. Type material. MNHN (Coll. Ch. Alluaud). Note. In the original description the number of typespecimens is not provided. References. Kuhnt 1911: 82 (Diphyllus, distribution). Distribution. Madagascar.

Biphyllus rosti (Grouvelle, 1916) Diphyllus Rosti Grouvelle, 1916: 41 (in Biphyllus – Schenkling 1934: 2).

Name derivation. Specific epithet honours German entomologist, Karl Rost (1859–1918). Type locality. Barway. Type material. IRSNB (holotype only). References. Schenkling 1934: 2 (distribution). Distribution. India (Uttarakhand).

Biphyllus rufopictus (Wollaston, 1874) Thallestus rufopictus Wollaston, 1874: 172 (in Biphyllus – Jakobson 1915: 936).

Name derivation. Rufus (lat.) – red, and pictus (lat.) – painted; refers to the red elytral pattern. Type locality. Japan. Type material. NHML. Note. In the original description the number of typespecimens is not provided. References. Lewis 1879: 11 (Thallestus, distribution); Schönfeldt 1888: 47 (Thallestus, distribution); Reitter 1889a: 301 (Diphyllus, description in key); Schoenfeldt 1891: 257 (Diphyllus, distribution); Kuhnt 1909: 119 (Diphyllus, distribution); Kuhnt 1911: 82 (Diphyllus, distribution); Jakobson 1915: 936

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(distribution); Schenkling 1923: 16 (Thallestus, distribution); Mader 1926: 719 (Diphyllus, distribution) and 721 (Thallestus, distribution); Nakane 1950: 1083 (description, distribution), fig. (habitus); Nakane 1963: 197 (description, distribution), pl. 99, fig. 15 (habitus, colour); Sasaji 1983: 28 (distribution, morphological variability), 36 (description in key); Nikitsky 1983: 701, fig. 1 (10) (description in key); Sasaji 1985a: 2 (distribution); Hirashima et al. 1989: 379 (distribution); Nikitsky 1992: 283, fig. 127.13 (description in key); Sasaji 1998: 208, 211, pl. 33, fig. 30 (description in key, distribution); Sawada et al. 1999: 166 (biology, distribution); Anonym 2000: 272 (distribution); Nomura et al. 2000: 225 (distribution, biology), fig. 73 (habitus); Iwai et al. 2001: 14 (distribution, biology); Nomura and Hirano 2005: 202 (distribution, biology); Nomura et al. 2006: 214 (distribution, biology); Jelínek 2007: 548 (distribution); Ohmomo et al. 2011: 85 (distribution, biology); Park et al. 2012: 189 (description, distribution, biology), figs. 1c (pronotum), 2c (meso- and metasternite), 4c, f (male genitalia), 5c (female genitalia), 6c (habitus), 7c (elytron), 8c (antenna). Biology. In Japan, the species was collected from April to September (Nomura et al. 2000: 225; Nomura et al. 2006: 214; Iwai et al. 2001: 14; Ohmomo et al. 2011: 85), in South Korea – from April to July (Park et al. 2012: 189). In Japan, the species was collected in forests where oriental white oak (Quercus aliena Blume, Fagaceae) is dominant (Sawada et al. 1999: 166). Distribution. Japan (Honshû – Prefectures Gumma, Gifu, Fukui, and Wakayama; Shikoku – Kôchi Prefecture; Kyûshû – Prefectures Fukuoka, Nagasaki, and Kagoshima), South Korea.

Biphyllus satsumanus Nakane, 1988 Biphyllus satsumanus Nakane, 1988: 79, pl. 9, fig. 6.

Name derivation. Specific epithet derived from Satsuma – old province of Japan that is now the western half of Kagoshima Prefecture, where the species was collected. Type locality. Kurino, Kagoshima Pref., Kyûshû. Type material. SEHU (holotype – female). References. Hirashima et al. 1989: 379 (distribution); Jelínek 2007: 548 (distribution). Biology. Holotype was collected in May (Nakane 1988: 80). Distribution. Japan (Kyûshû).

Biphyllus sauteri (Grouvelle, 1914) Diphyllus Sauteri Grouvelle, 1914d: 70 (in Biphyllus – Schenkling 1934: 2).

Name derivation. Specific epithet honours Hans Sauter (1871–1943), explorer of Formosa (now Taiwan). Type locality. Taihorin; Combeh. Type material. DEIC (Coll. W. Horn) and MNHN (Coll. A. Grouvelle) (totally 5 syntypes – 4 from Taihorin and single from Combeh). References. Miwa 1931: 77 (Diphyllus, distribution); Schenkling 1934: 2 (distribution); Hua 2002: 109 (distribution); Jelínek 2007: 548 (distribution). Distribution. Taiwan.

Biphyllus schenklingi (Grouvelle, 1914) Diphyllus Schenklingi Grouvelle, 1914d: 68 (in Biphyllus – Schenkling 1934: 2).

Name derivation. Name honouring the eminent coleopterologist, originator and editor of the monumental series Coleopterorum Catalogus, Sigmund Schenkling (1865–1946). Type locality. Kosempo. Type material. ?DEIC (holotype). References. Miwa 1931: 76 (Diphyllus, distribution); Schenkling 1934: 2 (distribution); Hisamatsu 1965: 139 (distribution, diagnostic characters); Hua 2002: 109 (distribution); Jelínek 2007: 548 (distribution). Biology. Collected at the end of June (Hisamatsu 1965: 139). Distribution. Taiwan.

Biphyllus semifuscus (Grouvelle, 1914) Diphyllus semifuscus Grouvelle, 1914c: 175 (in Biphyllus – Schenkling 1934: 2).

Name derivation. Semi (lat.) – half, and fuscus (lat.) – dark, brown; likely due to the indistinctive body colouration. Type locality. Afrique orientale [= East Africa]. Type material. MNHN (Coll. A. Grouvelle) (holotype). References. Schenkling 1934: 2 (distribution). Distribution. East Africa.

Biphyllus sicardi (Grouvelle, 1906) comb. nov. Diphyllus Sicardi Grouvelle, 1906: 136.

Name derivation. Specific epithet honours French coleopterologist Siméon Albert Sicard (1864–1930) who, during his stay on Madagascar as army doctor, gathered a large collection of beetles inhabiting the island.

CATALOGUE OF BIPHYLLIDAE (COLEOPTERA) OF THE WORLD

Type locality. Diego Suarez. Type material. MNHN (Colls. S. A. Sicard and A. Grouvelle) (3 syntypes – pers. obs). References. Kuhnt 1911: 82 (Diphyllus, distribution). Distribution. Madagascar.

Biphyllus sjostedti (Grouvelle, 1910) Diphyllus Sjöstedti Grouvelle, 1910: 329 (in Biphyllus – Arrow 1929: 307).

Name derivation. Specific epithet honours Swedish naturalist Bror Yngwe Sjöstedt (1866–1948), organizer of expeditions to Africa. Type locality. Kiboscho, Kilimandjaro. Type material. ?NHRS (2 syntypes). References. Arrow 1929: 307 (systematic position); Schenkling 1934: 2 (distribution). Biology. Type specimens were collected at 3000– 4000 m. a.s.l. (Grouvelle 1910: 332). Distribution. Tanzania.

Biphyllus sordidus (Grouvelle, 1906) comb. nov. Diphyllus sordidus Grouvelle, 1906: 133.

Name derivation. Sordidus (lat.) – dirty; likely due to the densely pubescent body surface. Type locality. Fort-Dauphin. Type material. MNHN (Colls. A. Grouvelle and Ch. Alluaud). Note. In the original description the number of typespecimens is not provided. References. Kuhnt 1911: 82 (Diphyllus, distribution). Distribution. Madagascar.

Biphyllus subellipticus (Wollaston, 1862) Thallestus subellipticus Wollaston, 1862: 155, pl. VII, fig. 4 (in Biphyllus – Jakobson 1915: 936).

Name derivation. Sub- (lat.) – prefix curtailing or reducing the meaning of a word, here “almost”, and ellipticus (lat.) – elliptic; likely due to the nearly elliptic body outline. Type locality. Mountains above Santa Cruz, Teneriffe. Type material. NHML. Note. In the original description the number of typespecimens is not provided. References. Wollaston 1864: 134 (Thallestus, distribution, biology); Wollaston 1865: 158 (Thallestus,

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distribution, biology); Gemminger and Harold 1868: 881 (Thallestus, distribution); Kuhnt 1909: 118 (Diphyllus, distribution); Kuhnt 1911: 82 (Diphyllus, distribution); Jakobson 1915: 936 (distribution); Schenkling 1923: 16 (Thallestus, distribution); Peyerimhoff 1923: 49 (biology); Mader 1926: 718 (Diphyllus, distribution) and 721 (Thallestus, distribution); Machado and Oromí 2000: 63 (distribution); Jelínek 2007: 548 (distribution). Biology. Syntypes were collected by Wollaston in February, in the decaying stem of Euphorbia canariensis L. (Euphorbiacae) (Wollaston 1862: 155; Wollaston 1864: 134; Wollaston 1865: 158). Specimens were also found on arborescent forms of Euphorbia sp. (Euphorbiacae) (Peyerimhoff 1923: 49). Distribution. Canary Islands (Tenerife).

Biphyllus subfasciatus (Reitter, 1878) comb. nov. Thallestus subfasciatus Reitter, 1878: 185.

Name derivation. Sub (lat.) – curtailing prefix, fascia (lat.) – band and -atus (lat.) – adjectival suffix -ate; likely due to the indefinite elytral bands. Type locality. Mexico. Type material. ?MNHN (Coll. A. Grouvelle). Note. In the original description the number of typespecimens is not provided. References. Schenkling 1923: 16 (Thallestus, distribution); Blackwelder, 1945: 428 (Thallestus, distribution). Note. Systematic position of this species is uncertain. Distribution. Mexico.

Biphyllus substriatus (Grouvelle, 1916) Diphyllus substriatus Grouvelle, 1916: 44 (in Biphyllus – Schenkling 1934: 2).

Name derivation. Sub- (lat.) – restrictive prefix, and stria (lat.) – line; likely due to the regular but inconspicuous rows of elytral punctures. Type locality. Nilgiri Hills. Type material. MNHN (Coll. A. Grouvelle) (holotype). References. Schenkling 1934: 2 (distribution). Distribution. India (Nilghiri Hills).

Biphyllus suffusus (Wollaston, 1874) Thallestus suffusus Wollaston, 1874: 171 (in Biphyllus – Jakobson 1915: 936).

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Name derivation. Suf (= sub) (lat.) – restrictive prefix, and fusus (lat.) – fuzzy, diffuse; likely due to the red elytral spots that are smaller and less distinct than B. rufopictus. Type locality. Japan. Type material. NHML. Note. In the original description the number of typespecimens is not provided. References. Lewis 1879: 11 (Thallestus, distribution); Schönfeldt 1888: 47 (Thallestus, distribution); Reitter 1889a: 301 (Diphyllus, description in key); Schoenfeldt 1891: 257 (Diphyllus, distribution); Kuhnt 1909: 119 (Diphyllus, distribution); Kuhnt 1911: 82 (Diphyllus, distribution); Jakobson 1915: 936 (distribution); Schenkling 1923: 16 (Thallestus, distribution); Mader 1926: 719 (Diphyllus, distribution) and 721 (Thallestus, distribution); Nakane 1959: 70 (distribution); Nakane 1963: 198 (description, distribution), pl. 99, fig. 16 (habitus, colour); Sasaji 1983: 29 (distribution), 36 (description in key); Nikitsky 1983: 700 (description in key); Sasaji 1985a: 2 (distribution); Hirashima et al. 1989: 379 (distribution); Nikitsky 1992: 282 (description in key); Setsuda 1993: 18 (biology); Sasaji 1998: 208, 211, pl. 33, fig. 31 (description in key, distribution); Jelínek 2007: 548 (distribution). Biology. Collected on fruiting bodies of Fomes fomentarius (L.) Fr. (Polyporaceae) (Setsuda 1993: 18). Distribution. Japan (Honshû – Prefectures Gumma and Fukui; Kyûshû – Fukuoka Prefecture).

Biphyllus tenuis (Grouvelle, 1916) Diphyllus tenuis Grouvelle, 1916: 46 (in Biphyllus – Schenkling 1934: 2).

Name derivation. Tenuis (lat.) – lean, slender; likely due to the strongly elongate body. Type locality. Nilgiri Hills. Type material. NHML (Coll. H.-E. Andrewes) and MNHN (Coll. A. Grouvelle). Note. In the original description the number of typespecimens is not provided. References. Schenkling 1934: 2 (distribution). Distribution. India (Nilghiri Hills).

resemblance to Throscus Latreille (= Trixagus Kugellan) (Throscidae, Coleoptera). Type locality. Japan. Type material. NHML. Note. In the original description the number of typespecimens is not provided. References. Lewis 1879: 11 (Thallestus, distribution); Schönfeldt 1888: 47 (Thallestus, distribution); Reitter 1889a: 301 (Diphyllus, description in key); Schoenfeldt 1891: 257 (Diphyllus, distribution); Kuhnt 1909: 119 (Diphyllus, distribution); Kuhnt 1911: 82 (Diphyllus, distribution); Jakobson 1915: 936 (distribution); Schenkling 1923: 16 (Thallestus, distribution); Mader 1926: 719 (Diphyllus, distribution) and 721 (Thallestus, distribution); Nakane 1959: 70 (distribution); Nakane 1963: 198 (description, distribution), pl. 99, fig. 17 (habitus, colour); Sasaji 1983: 29 (distribution, biology), 30 (morphological variability), 36 (description in key); Nikitsky 1983: 701, fig. 1 (4) (description in key); Sasaji 1985a: 2 (distribution); Hirashima et al. 1989: 379 (distribution); Nikitsky 1992: 283, fig. 127.8 (description in key); Kato et al. 1993: 161 (biology); Sasaji 1997: 27 (distribution, biology); Sasaji 1998: 208, 211, pl. 33, fig. 32 (description in key, distribution); Sawada et al. 1999: 166 (biology, distribution); Jelínek 2007: 548 (distribution); Park et al. 2012: 190 (description, distribution, biology), figs. 1d (pronotum), 2d (meso- and metasternite), 5d (female genitalia), 6d (habitus), 7d (elytron), 8d (antenna). Biology. In Japan, the species has been collected from mid-April to early August (Sasaji 1983: 29–30; Kato et al. 1993: 161; Sasaji 1997: 27). In South Korea, the species has been collected in May and June (Park et al. 2012: 190). Adults have been collected in forests where oriental white oak (Quercus aliena Blume, Fagaceae) is dominant (Sawada et al. 1999: 166); and on flowers of Japanese rowan (Sorbus commixta Hedlung, Rosaceae) (Kato et al. 1993: 161). Distribution. Japan (Honshû – Prefectures Fukui and Hyôgo; Shikoku – Kôchi Prefecture; Kyûshû – Prefectures Fukuoka, Kumamoto, and Nagasaki (Tsushima Island)), South Korea.

Biphyllus turneri Arrow, 1929 Biphyllus turneri Arrow, 1929: 310.

Biphyllus throscoides (Wollaston, 1874) Thallestus throscoides Wollaston, 1874: 171 (in Biphyllus – Jakobson 1915: 936).

Name derivation. Throsko (gr.) – jump (but here derived from the genus-name Throscus), and -ides (gr.) – suffix denoting son or descendant, use to mark affinity or similarity; likely due to the superficial

Name derivation. Specific epithet honours Rowland Edwards Turner (1863–1945), who had collected the type material. Type locality. Port St. John, Pondoland, Cape prov. Type material. NHML. Note. In the original description the number of typespecimens is not provided.

CATALOGUE OF BIPHYLLIDAE (COLEOPTERA) OF THE WORLD

References. Schenkling 1934: 2 (distribution). Biology. The type material was collected in September (Arrow 1929: 310). Distribution. Republic of South Africa.

Biphyllus typhaeoides (Wollaston, 1862) Thallestus typhaeoides Wollaston, 1862: 155, pl. VII, figs. 6 and 6a-e (in Biphyllus – Jakobson 1915: 936).

Name derivation. Typhe (gr.) – reed-mace (Typha), and -ides (gr.) – suffix denoting son or descendant, use to mark affinity or similarity; Wollaston had in mind the dense golden-brown pubescence of the beetle to the “fluff” on reed-mace (Typha L.) (Typhaceae) inflorescences and fructifications. Type locality. Hills immediately to the north of San Sebastian, Gomera. Type material. NHML. Note. In the original description the number of typespecimens is not provided. Thallestus typhaeoides var. obscuricollis Wollaston, 1865: 158, syn. nov.

Name derivation. Obscurus (lat.) – dark, and collum (lat.) – neck; likely due to the dark pronotal colouration. Type locality. Gomera. Type material. NHML. Note. In the original description the number of typespecimens is not provided. References. Wollaston 1864: 135 (Thallestus, distribution, biology); Wollaston 1865: 158 (Thallestus, distribution, biology); Gemminger and Harold 1868: 881 (Thallestus, distribution); Kuhnt 1909: 118 (Diphyllus, distribution); Kuhnt 1911: 82 (Diphyllus, distribution); Jakobson 1915: 936 (distribution); Schenkling 1923: 16 (Thallestus, distribution); Peyerimhoff 1923: 49 (biology); Mader 1926: 718 (as Diphyllus typhoeoides [sic!], distribution); Franz 1996: 96 (Diphyllus, biology); Machado and Oromí 2000: 63 (distribution); Jelínek 2007: 548 (distribution). Biology. Type specimens were collected by Wollaston and Gray in February, on rotten branches of Euphorbia canariensis L. (Euphorbiacae) (Wollaston 1862: 156; Wollaston 1864: 135; Wollaston 1865: 158). Subsequently, collected in April and August from dead Euphorbia canariensis L. (Franz 1996: 96), and on arborescent forms of Euphorbia sp. (Euphorbiacae) (Peyerimhoff 1923: 49). Distribution. Canary Islands (El Hierro, La Gomera).

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Name derivation. Undulatus (lat.) – wavy; likely due to the pale elytral spots with wavy or undulating margins. Type locality. Diego Suarez. Type material. MNHN (Colls. Ch. Alluaud, A. Grouvelle, and S. A. Sicard). Note. In the original description the number of typespecimens is not provided. References. Kuhnt 1911: 82 (Diphyllus, distribution). Distribution. Madagascar.

Biphyllus uniformis (Grouvelle, 1914) Diphyllus uniformis Grouvelle, 1914c: 170 (in Biphyllus – Schenkling 1934: 2).

Name derivation. Unus (lat.) – one, and forma (lat.) – shape; likely due to the homogeneous body pubescence. Type locality. Afrique orientale [= East Africa]. Type material. MNHN (Coll. A. Grouvelle) (2 syntypes). References. Schenkling 1934: 2 (distribution). Distribution. East Africa.

Biphyllus ussuriensis Nikitsky, 1983 Biphyllus ussuriensis Nikitsky, 1983: 703, figs. 1 (1, 6, and 13).

Name derivation. Specific epithet derived from the Ussuri area in the Far East, where the species was collected. Type locality. V[illage] Kamenushka, Ussurysky region, Southern Primorye. Type material. ZMAS (holotype – male; 5 paratypes), ZMUM (5 paratypes), IBSS (2 paratypes). References. Nikitsky 1992: 280, figs. 127.1–3 (description in key, biology); Jelínek 2007: 548 (distribution). Biology. Specimens were collected in June and July on bark of recently dead maple (Acer sp., Aceraceae) covered with lichens (Lichenes) and ascomycetous fungi (Ascomycota); the trees were infested with bark-beetles Scolytoplatypus tycon Blandford (Curculionidae, Coleoptera) (Nikitsky 1983: 703; Nikitsky 1992: 280). Distribution. Russia (Southern Primorye).

Biphyllus v-notatus (Grouvelle, 1906) comb. nov. Diphyllus v notatus Grouvelle, 1906: 134, pl. 8, fig. 6.

Biphyllus undulatus (Grouvelle, 1906) comb. nov. Diphyllus undulatus Grouvelle, 1906: 128.

Name derivation. Nota (lat.) – mark, and -atus (lat.) – past participle suffix -ate; the elytra of this

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beetle are adorned with two distinct v-shaped, black bands. Type locality. Diego Suarez. Type material. MNHN (Colls. S. A. Sicard and A. Grouvelle). Note. In the original description the number of typespecimens is not provided. Diphyllus parvulus var. notatus Kuhnt, 1911: 82 (nomen nudum).

References. Kuhnt 1911: 82 (Diphyllus, distribution, mistakenly given in that paper as var. notatus of the species Biphyllus parvulus (Grouvelle). The description of Biphyllus parvulus is in Grouvelle’s paper just before that of Biphyllus v-notatus, and Kuhnt erroneously interpreted the name as v[arietas] notatus instead of “v notatus” [sic!]). Distribution. Madagascar.

Biphyllus variegatus (Grouvelle, 1906) comb. nov. Diphyllus variegatus Grouvelle, 1906: 132.

Name derivation. Variegatus (lat.) – speckled, spotted; likely due to the characteristic elytral pattern. Type locality. Madagascar. Type material. MNHN (Coll. A. Grouvelle). Note. In the original description the number of typespecimens is not provided. References. Kuhnt 1911: 82 (Diphyllus, distribution). Distribution. Madagascar.

Biphyllus weisei (Grouvelle, 1914) Diphyllus Weisei Grouvelle, 1914c: 168 (in Biphyllus – Schenkling 1934: 2).

Name derivation. Specific epithet honours Paul Weise (?–1902), untimely deceased collector of African fauna; Paul Weise was a son of the famous entomologist, expert in Chrysomelidae and Coccinellidae, Julius Weise (1844–1925). Type locality. Kwai, Afrique orientale [= East Africa]. Type material. MNHN (Coll. A. Grouvelle) (holotype). References. Schenkling 1934: 2 (distribution). Distribution. Botswana.

Biphyllus wollastoni (Reitter, 1878) comb. nov. Phallestus [sic!] (errore typographico) Wollastoni Reitter, 1878: 184.

Name derivation. Specific epithet honours British coleopterologist Thomas Vernon Wollaston (1822– 1878), who described, among others, the genus Thallestus. Note. The generic name Thallestus has been printed, probably mistakenly, as Phallestus. Type locality. Ceylon. Type material. ?MNHN (Coll. A. Grouvelle). Note. In the original description the number of typespecimens is not provided. References. Schenkling 1923: 16 (Thallestus, distribution). Distribution. Sri Lanka.

Genus Diplocoelus Guérin-Méneville, 1844 Diplocoelus Guérin-Méneville, 1844: 196.

Name derivation. diploos (gr.) – double, and koilos (gr.) – empty, hollow; likely due to the two deep foveae on the pronotal disk. Gender. Masculine. Type species. Triphyllus fagi Chevrolat, 1837: pl. 41, fig.7, subsequent designation by Jacquelin du Val 1859: 212. Marginus LeConte, 1861: 105. Synonymized by Reitter 1878: 189 (as subgenus of Diplocoelus).

Name derivation. Margo (lat.) – margin. Gender. Masculine. Type species. Marginus rudis LeConte, 1863: 73, subsequent monotypy by LeConte 1863: 73. Note. LeConte (1861: 105) described the genus without mentioning any species. Two years later, LeConte (1863: 73) diagnosed the genus together with single species. References. Jacquelin du Val 1857: 102 (systematic position); Redtenbacher 1858: 359 (description); Jacquelin du Val 1859: 212 (description), 114 [catalogue] (catalogue of European species); Schaum 1859: XXIV (systematic position); Schaum 1860: XXII (systematic position); Marseul 1866: 49 (catalogue of European species); Gemminger and Harold 1868: 908 (world catalogue), 909 (Marginus, world catalogue); Stein 1868: 49 (catalogue of European species); Kraatz 1869: 28 (catalogue of German species); Crotch 1873: 44 (Diplocoelus and Marginus – both genera bonum, catalogue of North American species); Redtenbacher 1874: 392 (description); Stein and Weise 1877: 76 (catalogue of European species); Reitter 1878: 190 (key for the identification of known species); Horn 1878: 606 (key for identification of North American species, description, distribution); Austin 1880: 20 (supplement to catalogue of North American species); Reitter 1880b: 86 [16] (key for identification of European species);

CATALOGUE OF BIPHYLLIDAE (COLEOPTERA) OF THE WORLD

Reitter in Heyden et al. 1883: 78 (catalogue of European species); Reitter 1885: 20 (key for identification of European species); Masters 1886: 667 (catalogue of Australian species); Redtenbacher 1886: 214 (wings, morphology); Philippi 1887: 678 (catalogue of Chilean species); Schilsky 1888: 55 (catalogue of Central European species); Reitter in Heyden et al. 1891: 151 (catalogue of European species); Ragusa 1892: 131 (catalogue of Sicilian species, biology); Ganglbauer 1899: 651 (as subgenus of Diphyllus, key for identification of Central European species); Casey 1900: 80 (key for identification of North American species); Blackburn 1903: 158 (key for identification of Australian species); Grouvelle 1903b: 195 (key the identification of New Caledonian species); Fauvel 1903: 314 (key for identification of New Caledonian species, catalogue); Reitter in Heyden et al. 1906: 338 (catalogue of European species); Kuhnt 1909: 119 (description of genus, world catalogue); Schilsky 1909: 91 (catalogue of Central European species); Blatchley 1910: 571 (key for identification of North American species); Kuhnt 1911: 83 (world catalogue); Reitter 1911: 73 (as subgenus of Diphyllus, description in key); Bruch 1914: 385 (catalogue of Argentinean species); Jakobson 1915: 936 (catalogue of Palearctic species); Schaufuss 1916: 484 (European species); Thaxter 1918: 226 (biology); Leng 1920: 202 (catalogue of North American species); Falcoz 1926: 69 (systematic position); Mader 1926: 719 (distribution); Portevin 1931: 193 (Diphyllus (Diplocoelus), description); Schenkling 1934: 2 (supplement to world catalogue); Hellen 1947: 43 (catalogue of Swedish species); Vogt 1967: 109 (description); Arnett 1971: 790 (description in key); Bangsholt et al. 1979: 49 (catalogue of Scandinavian species); Burakowski et al. 1986: 209 (catalogue of Polish species); Lucht 1987: 187 (catalogue of Central European species); Campbell 1991: 241 (catalogue of Canadian species); Goodrich and Springer 1992: 365 (revision of North American species); Jelínek 1993: 102 (catalogue of Czechoslovak species); Poggi 1995: 7 (catalogue of Italian species); Alexandrovitch et al. 1996: 41 (catalogue of Belarusian species); Arnett 1997: 346 (North American species); Köchler and Klausnitzer 1998: 111 (catalogue of German species); Peck and Thomas 1998: 94 (catalogue of Florida species); Barbosa and Marquet 2002: 300, tab. 1 (Chile, ecology, forests defragmentation); Goodrich 2002: 357 (description, biology); Jelínek 2007: 548 (catalogue of Palearctic species, distribution). Immature stages. Rymer Roberts, 1958: 277 (description) Costa et al. 1988: 203, pl. 98, figs. 1–12 (description and illustrations); Lawrence and Ślipiński 2013: 399, fig. 84G (illustration). Biology. Elytra of the Chilean specimens of this genus are inhabited by the fungus Cucujomyces diplocoeli Thaxter (Laboulbeniaceae, Laboulbeniales) (Thaxter 1918: 226). Adults come to light (Goodrich 2002: 356).

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Number of species. 37. Distribution. Europe, Near East, North and Northeast Africa, Sri Lanka, New Caledonia, Australia, and New World.

Diplocoelus amplicollis Reitter, 1878 Diplocoelus amplicollis Reitter, 1878: 188.

Name derivation. Amplus (lat.) – large, magnificent, and collum (lat.) – neck; likely due to the relatively broad pronotum. Type locality. Rio [de] Janeiro, Columbia. Note. The case of actual type-locality is difficult to clarify. Reitter in the original description indicated “Rio Janeiro, Columbia”. There are at least two localities named Rio de Janeiro in Columbia: one in the Department of Tolima, the other in Department of Caldas. Evidently later authors too hastily accepted for sure that “Rio de Janeiro” refers to the Brazilian town. That information was then repeated. Type material. ?MNHN (Coll. A. Grouvelle). Note. In the original description the number of typespecimens is not provided. References. Schenkling 1934: 2 (distribution); Blackwelder 1945: 425 (distribution); Costa et al. 1988: 203, pl. 98, figs. 1–12 (immature stages) and 12 (adult) (as Diplocoelus af. amplicollis, immature stages, distribution, biology); Mecke et al. 2000: 171, tab. IV (as Diplocoelus cf amplicollis, distribution, biology); Mecke et al. 2005: 11, annex (as Diplocoelus cf amplicollis, distribution, biology); Vitali 2010: 171 (biology). Immature stages. Costa et al. 1988: 203, pl. 98, figs. 1–12 (as Diplocoelus af. amplicollis, description and illustrations of larva and pupa, biology). Biology. Specimens tentatively identified as this species [as pointed out by the authors themselves and by Vitali (2010: 171)] were found in Brazil on ground cones of Araucaria angustifolia (Bert.) Kuntze (Araucariaceae) (Costa et al. 1988: 204; Costa et al. 1990: 5; Mecke et al. 2000: 171, tab. IV; Mecke et al. 2005: 11, annex). Laboratory studies showed that prior to pupation the larvae left the cones and constructed a prepupal chamber (Costa et al. 1988: 204; Costa et al. 1990: 5). Distribution. Colombia, ?Brazil.

Diplocoelus angustulus Blackburn, 1891 Diplocoelus angustulus Blackburn, 1891: 122.

Name derivation. Angustus (lat.) – narrow, and -ulus (lat.) – diminutive suffix; likely due to the slender body.

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Type locality. S. [outh] Australia, various localities (Blackburn, 1891: 123). Type material. NHML, IRSNB (single syntype – see Damoiseau 1968: 28), and ?SAMA. Note. In the original description the number of typespecimens is not provided. References. Blackburn 1903: 158 (description in key); Kuhnt 1909: 120 (distribution); Kuhnt 1911: 83 (distribution); Damoiseau 1968: 28 (type specimens in IRSNB); Grove 2009: 96 (distribution); Grove and Forster 2011: 62, appendix 2 (distribution, biology). Biology. Type specimens were found under bark of Eucalyptus sp. (Myrtaceae) (Blackburn 1891: 123). Also, collected on Eucalyptus obliqua L’Héritier (Myrtaceae) in Tasmania (Grove and Forster 2011: 62, appendix 2). Distribution. Australia (South Australia), Tasmania.

Diplocoelus apicicollis Lea, 1921 Diplocoelus apicicollis Lea, 1921a: 365.

Name derivation. Apex, apicis (lat.) – peak, and collum (lat.) – neck; likely due to the pronotum narrowed anterad. Type locality. Swan River; Donnybrook. Type material. SAMA. Note. In the original description the number of typespecimens is not provided. References. Schenkling 1934: 2 (distribution). Distribution. Australia (Western Australia).

Diplocoelus atomus Grouvelle, 1916 Diplocoelus atomus Grouvelle, 1916: 51.

Name derivation. Atomos (gr.) – indivisible, dot, here: something very small; likely due to the small size (ca. 0.7 mm). Type locality. St-Laurent du Maroni. Type material. MNHN (Coll. A. Grouvelle). Note. In the original description the number of typespecimens is not provided. References. Schenkling 1934: 2 (distribution); Blackwelder 1945: 425 (distribution). Biology. Commonly occurs on flowers (Grouvelle 1916: 53). Distribution. French Guyana.

Diplocoelus bicolor Sharp, 1900 Diplocoelus bicolor Sharp, 1900: 621.

Name derivation. Bis (lat.) – twice, and color (lat.) – colour; likely due to the characteristic bicolorous body pubescence. Type locality. Capetillo, Guatemala. Type material. NHML (2 syntypes). References. Schenkling 1934: 2 (distribution); Blackwelder 1945: 425 (distribution). Distribution. Guatemala.

Diplocoelus bombycinus Grouvelle, 1903 Diplocoelus bombycinus Grouvelle, 1903b: 194.

Name derivation. Bombyx (lat.) – silkworm, and -inus (lat.) – adjectival suffix; likely due to the dense and uniform body pubescence, which is similar in appearance to setae covering the body of silkworm moths. Type locality. Ile de Pins; Nouvelle-Calédonie [= New Caledonia]. Type material. IRSNB (Coll. A. Fauvel) and MNHN (Coll. A. Grouvelle). Note. In the original description the number of typespecimens is not provided. References. Fauvel 1903: 314 (description in key, distribution); Kuhnt 1909: 120 (distribution); Kuhnt 1911: 83 (distribution). Distribution. New Caledonia, Ile des Pins.

Diplocoelus brunneus LeConte, 1863 Diplocoelus brunneus LeConte, 1863: 73.

Name derivation. Brunneus (mediaeval lat.) – brown; likely due to the body colouration. Type locality. Middle States [of USA]. Type material. MCZC (holotype – according to Goodrich and Springer 1992: 367). Note. In the original publication Melsheimer and Ulke are mentioned as collectors of the type material, thus probably LeConte had more than one specimen at his disposal. Diplocoelus angusticollis Horn, 1878: 605. Synonymized by Goodrich and Springer 1992: 367.

Name derivation. Angustus (lat.) – narrow, and collum (lat.) – neck; likely due to the slender anterior part of pronotum (see note below). Type locality. Marquette, Michigan. Type material. MCZC (holotype). Note. The holotype of the species described by Horn is teratological (malformed pronotum) specimen of Diplocoelus brunneus LeConte (Goodrich and Springer 1992: 370). Despite Kuhnt’s (1911: 83) erro-

CATALOGUE OF BIPHYLLIDAE (COLEOPTERA) OF THE WORLD

neous information, Casey (1900: 80) is not the author of the description of this species. References. Gemminger and Harold 1868: 908 (distribution); Crotch 1873: 44 (distribution); Horn 1878: 605–606 (Diplocoelus angusticollis and D. brunneus – as separate species, description, distribution); Austin 1880: 20 (Diplocoelus angusticollis – species bona, distribution); Henshaw 1885: 53 (Diplocoelus angusticollis and D. brunneus – as separate species, distribution); Casey 1900: 80 (Diplocoelus angusticollis – species bona, description in key); Kuhnt 1909: 120 (Diplocoelus brunneus and D. angusticollis – as separate species, distribution); Blatchley 1910: 572 (Diplocoelus brunneus and D. angusticollis – as separate species, description, biology); Kuhnt 1911: 83 (Diplocoelus angusticollis Casey [sic!] – species bona, distribution); Leng 1920: 202 (Diplocoelus brunneus and Diplocoelus angusticollis – as separate species, catalogue of North American species); Beaulne 1923: 264 (distribution); Arnett 1971: 789, fig.1.91 (adult, illustration); Wallace and Fox 1980: 630, fig. 132 (wings, morphology); Campbell 1991: 241 (distribution); Goodrich and Springer 1992: 367, figs. 2, 4, 8, 9, 12, 15, and 18 (description, distribution); Peck and Thomas 1998: 94 (distribution); Goodrich 2002: 356, fig.1.89 (adult) and 357, fig. 3.89 (first abdominal sternite); Vitali 2010: fig. 4a (antennal club); Cline and Shockley 2010: 306 (biology). Biology. Adults can be found almost throughout the year (except January and August) on old (rather on dead but still standing than on fallen trunks) oaks (Quercus sp., Fagaceae) strongly infested by fungi of the genus Hypoxylon Bulliard (Xylariaceae, Ascomycota) (Goodrich and Springer 1992: 369; Cline and Shockley 2010: 306). They have been also sifted from dead leaves of maple (Acer sp., Aceraceae), beech (Fagus sp., Fagaceae) and other hardwood trees (Cline and Shockley 2010: 306). In summer, it was collected on fungi growing on beech and maple, and in spring among roots of these trees; probably adults overwinter (Blatchley 1910: 572). Distribution. Canada (New Brunswick, Ontario, Quebec), USA (New Hampshire, Michigan, Wisconsin, New York, New Jersey, Pennsylvania, Maryland, Virginia, Ohio, Indiana, Illinois, Missouri, Kentucky, North Carolina, Alabama, Oklahoma, Mississippi, Florida).

Diplocoelus consobrinus Grouvelle, 1905 Diplocoelus (Marginus) consobrinus Grouvelle, 1905b: 127.

Name derivation. Consobrinus (lat.) – nephew, relative; likely due to its similarity to congeners. Type locality. Buenos Ayres [= Buenos Aires], République Argentine [= Argentina]; Province de Tucuman.

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Type material. MACN (Coll. C. Bruch) and ?MNHN (Coll. A. Grouvelle). Note. In the original description the number of typespecimens is not provided. References. Kuhnt 1909: 120 (distribution); Kuhnt 1911: 83 (distribution); Bruch 1914: 385 (distribution); Blackwelder 1945: 469 (distribution). Distribution. Argentina (Tucuman Prov., Buenos Aires Prov.).

Diplocoelus decemlineatus Lea, 1921 Diplocoelus decemlineatus Lea, 1921a: 364.

Name derivation. Decem (lat.) – ten, linea (lat.) – line, and -atus (lat.) – past participle suffix -ate; likely due to the presence of ten longitudinal elevations on pronotum. Type locality. Lucindale, South Australia; Port Lincoln, South Australia; Ulverstone, Hobart, Tasmania (Lea 1922: 298). Type material. SAMA (a few syntypes – according to Lea (1922: 298)). Note. On the occasion of diagnosing Diplocoelus xanthorrhoeae Lea (1921a: 364) mentioned D. decemlineatus in one-sentence description and made this name formally available (probably unintentionally). One year later Lea (1922: 294) published detailed description of D. decemlineatus, with type locality and other details. References. Lea 1922: 298 (description); Schenkling 1934: 3 (distribution). Distribution. Australia (South Australia), Tasmania.

Diplocoelus dilataticollis Lea, 1921 Diplocoelus dilataticollis Lea, 1921a: 364.

Name derivation. Dilato (lat.) – broaden, and collum (lat.) – neck; likely due to the distinctly widened pronotal margins that are wider than the elytral base. Type locality. Mount Tambourine, Queensland (Lea 1921b: 237). Type material. SAMA (holotype) and QMBA (paratype) (Lea 1921b: 237). Note. On the occasion of diagnosing Diplocoelus xanthorrhoeae Lea (1921a: 364) mentioned D. dilaticollis in one-sentence description and made this name formally available (probably unintentionally). Two days [sic!] later (see references) Lea (1921b: 237) published detailed description of D. dilaticollis, with type locality and other details.

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References. Lea 1921b: 237 (description); Schenkling 1934: 3 (distribution); Hugo et al. 2003: 355 (biology); Vitali 2010: 171 (biology). Biology. Larvae feed on fungi associated with damaged seeds of Australian blackwood (Acacia melanoxylon Brown, Fabaceae) (Hugo et al. 2003: 355; Vitali 2010: 171). Distribution. Australia (Queensland).

Diplocoelus exiguus Blackburn, 1891 Diplocoelus exiguus Blackburn, 1891: 123.

Name derivation. Exiguus (lat.) – small, tiny, miserable; likely due to the small body size (ca. 2.1 mm). Type locality. Near Port Lincoln, South Australia. Type material. NHML and ?SAMA. Note. In the original description the number of typespecimens is not provided. References. Blackburn 1903: 158 (description in key); Kuhnt 1909: 120 (distribution); Kuhnt 1911: 83 (distribution). Distribution. Australia (South Australia).

Diplocoelus fagi (Chevrolat, 1837) Mycetophagus [(]Triphyllus[)] fagi Chevrolat, 1833: 472 (nomen nudum). Triphyllus Serratus Dejean, 1835: 311 (nomen nudum, in synonyms i.a. name of Triphyllus fagi Chevrolat). Triphyllus Fagi Chevrolat in Dejean, 1835: 311 (nomen nudum, as synonym of Triphyllus Serratus Dejean). Triphyllus Pilicollis Parreyss in Dejean, 1835: 311 (nomen nudum, as synonym of Triphyllus Serratus Dejean). Triphyllus fagi Chevrolat, 1837: pl. 41, fig. 7 (in Diplocoelus – Guérin-Méneville, 1844: 196).

Name derivation. Fagus (lat.) – beech; specific epithet is derived from the genus name of the host plant. Type locality. Fontainebleau (Guérin-Méneville, 1844: 196). Type material. Unknown. Note. Number of type specimens unknown, but there were more than one (Guérin-Méneville, 1844: 196). Triphyllus serratus Dufour, 1843: 93. Synonymized by Bedel 1904: 235.

Name derivation. Serratus (lat.) – similar to the saw; likely due to the shape of the antennal club. Type locality. [Vallée d’Ossau]. Type material. MNHN (Coll. L. Dufour). Note. In the original description the number of typespecimens is not provided.

References. Chevrolat 1833: 472 (Mycetophagus, biology); Dejean 1835: 311 (Triphyllus, distribution); Dejean 1836: 336 (Triphyllus, distribution); Dufour 1843: (Triphyllus serratus – species bona, description, biology); Aubé 1850: 329 (Diphyllus, description, biology); Habelmann 1854: 28 (distribution, biology); Redtenbacher 1858: 359 (description); Jacquelin du Val 1859: 114 [catalogue] (distribution); Eichoff 1866: 294 (distribution, biology); Marseul 1866: 49 (distribution); Gemminger and Harold 1868: 909 (distribution); Stein 1868: 49 (distribution); Kraatz 1869: 28 (distribution); Stierlin and Gautard 1869: 151 (Diplocaelus [sic!], distribution, biology); Seidlitz 1872: 183 (description, distribution, biology); Brauns 1874: 127 (distribution); Redtenbacher 1874: 393 (description); Heyden 1877: 177 (distribution, biology); Stein and Weise 1877: 76 (distribution); Reitter 1878: 190 (description in key); Heyden 1879: 163 (distribution, biology); Reitter 1880b: 86 [16] (description, distribution); Buddeberg 1883: 71 (distribution); Reitter in Heyden et al. 1883: 78 (distribution); Reitter 1885: 20 (description, distribution); Redtenbacher 1886: 214 (wings, morphology); Oertzen 1887: 231 (distribution); Schilsky 1888: 55 (distribution); Seidlitz 1888: 270 (description, distribution); Rey 1889: 36 (biology); Seidlitz 1889: 289 (description, distribution); Favre 1890: 171 (Diplocaelus [sic!], biology); Reitter in Heyden et al. 1891: 151 (distribution); Ragusa 1892: 131 (distribution, biology); Ganglbauer 1899: 655 (Diphyllus (subgenus Diplocoelus), description, distribution); Kuhnt 1909: 119 (distribution); Schilsky 1909: 91 (distribution); Kuhnt 1911: 83 (distribution); Reitter 1911: 73, pl. 89, fig. 16 (Diphyllus (Diplocoelus), description in key); Kuhnt 1912: 530 (description), figs. 4b (antenna) and 13 (habitus); Jakobson 1915: 936 (distribution); Lüllwitz 1916: 233 (biology); Schaufuss 1916: 484 (description, distribution, biology); Kuthy 1918: 88 (distribution); Falcoz 1926: fig. 2 (mandibule), figs. 4 and 5 (male genitalia and genital segments); Mader 1926: 719 (distribution); Portevin 1931: 193 (Diphyllus (Diplocoelus), description); Donisthorpe 1935: 25 (biology); Roubal 1936: 161 (distribution, biology); Hellen 1947: 43 (distribution); Crowson 1952b: 118 (biology); Benick 1952: 127 (biology); Eagles 1955: 79(distribution, biology); Kocher 1956: 56 (distribution); Gould 1957: 30 (distribution); Jacobs 1957: 84 (distribution, biology); Rymer Roberts 1958: 277 (Diphyllus (Diplocoelus), larva, morphology); Palm 1959: 262 (biology); Horion 1960: 214 (distribution, biology); Gardner 1964: 22 (distribution, biology); Clifton 1965: 89 (distribution, biology); Crowson 1967: 106 (biology); Vogt 1967: 109 (description, distribution), fig. 6:1 (habitus); Bangsholt et al. 1979: 49 (distribution); Silfverberg 1992: 57 (distribution); Burakowski et al. 1986: 209 (distribution, biology); Drane 1987: 62 (biology); Lucht 1987: 187 (distribution); Lawrence 1991: 475 (biology); Borowiec et al.

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1992: 139 (distribution, protected areas, primary forests); Hoare and Hoare 1992: 74 (distribution, biology); Nikitsky 1993: fig. 10 (morphology); Jelínek 1993: 102 (distribution); Poggi 1995: 7 (distribution); Alexandrovitch et al. 1996: 41 (catalogue of Belarusian species); Jones 1996: 92 (biology); Cecchi and Bartolozzi 1997: 126 (biology); Hornig, 1998: 9, tab. 3 (distribution); Jones 1998: 172 (biology); Köchler and Klausnitzer 1998: 111 (distribution); Bakke 1999: 205, tab. 2 (distribution, biology); Schaffrath 1999: 28–29, tab. (biology); Jones 2000: 178 (biology); Merkl 2001: 206 (distribution); Schaffrath 2001: 27, tab. (distribution, biology); Alexander 2002: 52 (distribution, biology); Grossa-Silva 2002: 52 (distribution); Mitter 2004: 242, map 13 (distribution, biology); Bouget et al. 2005: 20 (biology); Jelínek 2007: 548 (distribution); Ricarte et al. 2009: 596 (biology); Löbl and Smetana 2010: 28 (correction to Palearctic catalogue); Recalde Irurzun and San Martín Moreno 2010: 152 (distribution); Schaffrath 2010: 365, tab. (distribution, biology); Vitali 2010: fig. 4c (antennal club); Horák 2011: 219, appendix A (distribution, biology); Przewoźny 2011: 39 (distribution, biology); Tamutis et al. 2011: 254 (distribution, distribution); Lassauce et al. 2012: 280, appendix A (distribution, biology); Lassauce et al. 2013: 407, appendix A (distribution, biology); Rusakov and Kalabkina 2013: 26, tab. 1 (distribution). Immature stages. Rymer Roberts, 1958: 277 (description of larva). Biology. The species inhabites deciduous forests (Burakowski et al. 1986: 210; Schaffrath 1999: 28–29; Schaffrath 2001: 27; Bouget et al. 2005: 17, 20; Ricarte et al. 2009: 596; Schaffrath 2010: 365; Horák 2011: 219, appendix A). Adults are collected on old beechs (Fagus silvatica L., Fagaceae) (Dufour 1843: 93; Rey 1889: 36; Ragusa 1892: 131; Hubenthal 1902: 274; Ricarte et al. 2009: 596; Przewoźny 2011: 39) (also under bark – Chevrolat 1833: 472; Aubé 1850: 330; Habelmann 1854: 28; Seidlitz 1872: 183; Heyden 1877: 177; Heyden 1879: 163; Reitter 1911: 73; Lüllwitz 1916: 233; Schaufuss 1916: 484; Roubal 1936: 191; Crowson 1952b: 118; Gardner 1964: 22; Hoare and Hoare 1992: 74; Cecchi and Bartolozzi 1997: 126; Jones 2000: 178; Alexander 2002: 52), sometimes on oaks (Quercus sp., Fagaceae) (Eichoff 1866: 294; Ricarte et al. 2009: 596; Lassauce et al. 2013: 407, appendix A) (also under bark – Hubenthal 1902: 274; Roubal 1936: 191 – with Staphylinidae [Thamiaraea sp. and Atheta euryptera (Stephens)], Nitidulidae [Epurea sp., Soronia sp., and Cryptarcha sp.], and Mycetophagidae [Triphyllus bicolor (F.)]), Turkey oak (Quercus cerris L.) (Roubal 1936: 191 – together with Rhinosimus aeneus Olivier, Salpingidae), Portuguese oak (Quercus faginea Lam.) (Ricarte et al. 2009: 596), Pyrenean oak (Quercus pyrenaica L.) (Ricarte et al. 2009: 596), cork oak (Quercus suber L.) (Ricarte et al. 2009: 596), narrow-

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leaft ash (Fraxinus angustifolia Vahl, Oleaceae) (Ricarte et al. 2009: 596), maples (Acer sp., Aceraceae) (Ricarte et al. 2009: 596) (sometimes frequently on Acer pseudoplatanus L. – Rey 1889: 36 and Denton 2007: 91; around London this is one of the most common beetles in June under bark of the same tree species – Jones 2000: 178, or later, in September – Jones 1996: 92), lime-trees (Tilia sp., Malvaceae) (Palm 1959: 263; Burakowski et al. 1986: 210; Bakke 1999: 205), aspens (Populus sp., Salicaceae) (Rey 1889: 36), planes (Platanus sp., Platanaceae) (Rey 1889: 36), and elms (Ulmus sp., Ulmaceae) (Burakowski et al. 1986: 210), rarely under bark of pine (Pinus sp., Pinaceae) (Drane 1987: 62) or common ivy (Hedera helix L., Araliaceae) (Stierlin and Gautard 1869: 151; Favre 1890: 171), or Quercus cerris L. (Fagaceae) (Roubal 1936: 191) or on spruce (Picea sp., Pinaceae) stump in May (Eagles 1955: 79). Also on Acer pseudoplatanus L., in black sooty spores of the “sooty bark disease” a fungus Cryptostroma corticale (Ellis et Everh.) Greg. et Waller (Ascomycota) (Jones 1998: 172; Alexander 2002: 52). Mycetophage, caught on dead leafless branches in April and May (Bouget et al. 2005: 20). The host is fungus – Nectria cinnabarina (Tode) Fr. (Nectriaceae) (Palm 1959: 263; Horion 1960: 214; Lawrence 1991: 475; Bakke 1999: 205; Alexander 2002: 52; Vitali 2010: 171 – all authors, exept Vitali (2010), reported fungus under the synonymous name, Tubercularia confluens Pers.), but also collected on Fomes fomentarius (L.) Fr. (Polyporaceae) (Donisthorpe 1935: 25; Benick 1952: 127), and Hypoxylon fuscum (Pers.) Fr. (Xylariaceae) (Donisthorpe 1935: 25; Benick 1952: 128). Larvae live under bark of deciduous trees (mainly beech, but also hornbeam, oak, lime, elm and maple) infested by fungi (Mitter 2004: 242); they winter, in spring pupate in the soil (Palm 1959: 263; Mitter 2004: 242) and as adults turn back to the host fungus in one year cycle (Mitter 2004: 242). Adults overwinter in deadwood, including oak (Quercus sp.) (Alexander 2002: 53). Distribution. Europe (Azerbaijan, Austria, Belarus, Belgium, Bosnia Hercegovina, Bulgaria, Croatia, Czech Republic, Denmark, France, Georgia, Germany, Great Britain, Greek, Hungary, Italy, Norway, Poland, Romania, Russia [southern part of Ural Mts.], Slovakia, Slovenia, Spain, Sweden, Switzerland, Ukraine, Serbia and Montenegro), Morocco, Algeria, Iran.

Diplocoelus fasciatus (MacLeay, 1871) Triphyllus fasciatus MacLeay, 1871: 170 (in Diplocoelus – Blackburn 1903: 157).

Name derivation. Fascia (lat.) – band, and -atus (lat.) – adjectival suffix -ate; likely due to the elytral black band.

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Type locality. [Gayndah]. Type material. AMSA. Note. In the original description the number of typespecimens is not provided. Note. Blackburn (1903: 157) wrote that he received from Lea a specimen of this species that was compared to the type, agreeing with the description, and certainly belonging to the genus Diplocoelus rather than Triphyllus. He added that this observation had not yet been published, though Lea in one of his papers mentioned Diplocoelus fasciatus (not quoting the author of the name). References. Masters 1886: 667 (Triphyllus, distribution); Blackburn 1903: 158 (description in key); Kuhnt 1909: 120 (distribution); Kuhnt 1911: 83 (distribution); Lawrence 1987: 361 (systematic position); Lawrence et al. 2011: 9 (phylogeny), fig. 28B (abdomen); Lawrence and Ślipiński 2013: 399, fig. 84G (immature stages). Immature stages. Lawrence and Ślipiński 2013: 399, fig. 84G (larva, illustration). Distribution. Australia.

Diplocoelus foveolatus Reitter, 1878 Diplocoelus foveolatus Reitter, 1878: 187.

Name derivation. Fovea (lat.) – pit, pitfall and suffixes: diminutive -olus (lat.), and participle -atus (lat.) -ate; likely due to the pronotal sculpture. Type locality. Chili [= Chile]. Type material. MNHN (Coll. A. Grouvelle). Note. In the original description the number of typespecimens is not provided. References. Philippi 1887: 678 (distribution); Schenkling 1934: 3 (distribution); Blackwelder 1945: 425 (distribution); Wallace and Fox 1980: 630, fig. 130 (wings, morphology); Elgueta and Arriagada 1989: 35 (distribution). Distribution. Chile.

Diplocoelus grandis Reitter, 1878 Diplocoelus grandis Reitter, 1878: 186.

Name derivation. Grandis (lat.) – large; likely due to the relatively large body (ca. 5.0–5.5 mm). Type locality. Mexico. Type material. MNHN (Coll. A. Grouvelle). Note. In the original description the number of typespecimens is not provided. References. Schenkling 1934: 3 (distribution); Blackwelder 1945: 425 (distribution). Distribution. Mexico.

Diplocoelus haagi Reitter, 1878 Diplocoelus Haagi Reitter, 1878: 186.

Name derivation. Specific epithet honours Johann Georg Haag-Rutenberg (1830–1879) (see also Kraatz 1880: 235). Type locality. Mexico. Type material. ?MNHN (Coll. A. Grouvelle). Note. In the original description the number of typespecimens is not provided. References. Kraatz 1880: 235 (Diplocaelus [sic!]); Schenkling 1934: 3 (distribution); Blackwelder 1945: 425 (distribution). Distribution. Mexico.

Diplocoelus humerosus Reitter, 1876 Diplocoelus humerosus Reitter, 1876: 293.

Name derivation. Humerus (lat.) – shoulder, and -osus (lat.) – suffix meaning “full of something”, “augmented”; name alluding to elytra distinctly widened behind humeri. Type locality. Mamudly; Kaukasus [= Caucasus] (Hornig 1992: 70). Type material. HNHM (type specimen marked as holotype in Coll. E. Reitter – pers. obs.); SMTD (however, Hornig 1992: 70 mentioned this collection with question mark). Note. Number of type specimens unknown, but it might be only the holotype (Reitter 1876: 293). References. Stein and Weise 1877: 76 (distribution); Reitter 1878: 190 (description in key); Reitter 1880b: 86 [16] (description, distribution); Reitter in Heyden et al. 1883: 78 (distribution); Reitter 1885: 20 (description, distribution); Reitter 1888a: 170 (description, biology); Seidlitz 1888: 270 (distribution); Seidlitz 1889: 289 (distribution); Reitter 1889b: 314 (description); Reitter in Heyden et al. 1891: 151 (distribution); Reitter in Heyden et al. 1906: 338 (distribution); Kuhnt 1909: 119 (world catalogue); Kuhnt 1911: 83 (distribution); Jakobson 1915: 936 (distribution); Schaufuss 1916: 484 (distribution); Mader 1926: 719 (distribution); Jelínek 2007: 548 (distribution). Biology. Found in October (Reitter 1876: 293) on leaf of beech (Reitter 1888a: 170). Distribution. Caucasus – Georgia, Southern regions of European Russia, Turkey.

Diplocoelus indicus Motschulsky, 1866 Diplocoelus indicus Motschulsky, 1866: 397.

Name derivation. Indicus (lat.) – Indian; the

CATALOGUE OF BIPHYLLIDAE (COLEOPTERA) OF THE WORLD

species was described from Sri Lanka, which in the past was included in India. Type locality. Montagnes de Nura-Ellia [= Nuwara Elija District]. Type material. ?ZMUM (Coll. V. Motschulsky). Note. In the original description the number of typespecimens is not provided. Distribution. Sri Lanka. Note. Species not recorded since its original description. The systematic position is uncertain.

Diplocoelus latus Lea, 1895 Diplocoelus latus Lea, 1895: 228.

Name derivation. Latus (lat.) – wide; likely due to the wide body. Type locality. Donnybrook. Type material. SAMA. Note. In the original description the number of typespecimens is not provided. References. Blackburn 1903: 158 (description in key); Kuhnt 1909: 120 (distribution); Kuhnt 1911: 83 (distribution); Lea 1921a: 364; Lawrence and Ślipiński 2013: 264, fig. 73B (adult, illustration). Distribution. Australia (Western Australia).

Diplocoelus mauritii Grouvelle, 1909 Diplocoelus Mauritii Grouvelle, 1909: 373.

Name derivation. Name derived from the latin form of the name Maurice – Mauritius, and have been given by Antoine Grouvelle in honour of Maurice de Rothschild (1881–1957), who was the organizer of the expedition to East Africa. Type locality. Confluent Akaki [= Akaki River catchment, near Addis Abeba], rive droite [= right bank of the river]. Type material. ?MNHN or ?NHML (holotype). Note. The type depository is unknown, maybe MNHN or NHML. References. Kuhnt 1911: 83 (distribution). Distribution. Ethiopia. Note. Kuhnt (1911: 83) having not mentioned Ethiopia, erroneously wrote that the species inhabited Mauritius. Probably it was a consequence of misinterpretation of the species name, which has nothing to do with Mauritius island.

Diplocoelus mus Reitter, 1878 Diplocoelus mus Reitter, 1878: 188.

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Name derivation. Mus (lat.) – mouse; likely due to the pubescence resembling the fur of a mouse. Type locality. Unknown. Note. According to the author of the original description the type specimens might have been collected in West India. Type material. ?DEIC (Coll. G. Kraatz) and ?MNHN (Coll. A. Grouvelle). Note. At least two syntypes exist. References. Horn 1878: 606 (distribution); Henshaw 1885: 53 (distribution); Leng 1920: 202 (distribution); Schenkling 1934: 3 (distribution); Blackwelder 1945: 425 (distribution). Distribution. ?West India. Note. Horn (1878: 606) suspected that this species is distributed in Mexico. Leng (1920: 202) wrote “West India or California?”.

Diplocoelus oblongus (Germain, 1855) Philophlaeus oblongus Germain, 1855: 395 (in Diplocoelus – Grouvelle 1919: 67).

Name derivation. Oblongus (lat.) – elongated; likely due to the relatively slender body. Type locality. Chile (Camousseight 1980: 5). Type material. MNSC (10 syntypes – Camousseight 1980: 5). Note. The genus Philophlaeus Germain, 1855: 395, nec Chaudoir, 1844: 472 (= Loberoschema Reitter, 1896: 160) had been erected for two Chilean species, one of them belongs to the family Erotylidae and has been designated by Leschen and Węgrzynowicz (1998: 231) as the type-species of that genus, while the other belongs to Biphyllidae. Diplocoelus tessellatus Reitter, 1878: 187. Synonymized by Grouvelle, 1919: 67.

Name derivation. Tesella (lat.) – mosaic and -atus (lat.) -ate; likely due to the small groups of white setae on the elytral intervals that form a mosaic-like pattern. Type locality. Chili [=Chile]. Type material. MNHN (Coll. A. Grouvelle). References. Philippi 1887: 676 (Philophlaeus oblongus, distribution), 678 (Diplocoelus tesselatus – species bona, distribution); Grouvelle 1919: 67 (systematic position); Schenkling 1934: 3 (distribution; both species, Diplocoelus oblongus and D. tesselatus are treated as species bona); Blackwelder 1945: 425 (distribution, both species, Diplocoelus oblongus and D. tesselatus are treated as species bona); Elgueta and Arriagada 1989: 35 (distribution, both species, Diplocoelus oblongus and D. tesselatus are treated as species bona). Distribution. Chile.

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Diplocoelus opacior Blackburn, 1903 Diplocoelus opacior Blackburn, 1903: 158.

Name derivation. Opacus (lat.) – mat, dim, and -ior (lat.) – suffix denoting comparative degree of adjectives; likely due to the conspicuous (especially pronotal) sculpture, making it more mat. Type locality. Glenelg R.[iver], Victoria. Type material. NHML and ?SAMA. Note. In the original description the number of typespecimens is not provided. References. Kuhnt 1909: 120 (distribution); Kuhnt 1911: 83 (distribution). Biology. The type material was collected under bark of Eucalyptus sp. (Myrtaceae) (Blackburn 1903: 158). Adults were found on leafs of Xanthorrhoea sp. (Xanthorrhoeaceae) (Lea 1921a: 364). Distribution. Australia (Victoria).

Diplocoelus ovatus MacLeay, 1871 Diplocoelus ovatus MacLeay, 1871: 170.

Name derivation. Ovatus (lat.) – egg-shaped; likely due to the ovate body outline. Type locality. [Gayndah]. Type material. AMSA. Note. In the original description the number of typespecimens is not provided. References. Masters 1886: 667 (distribution); Blackburn 1891: 123 (quotes this name as possible synonym of Diplocoelus piliger Reitter, 1880); Blackburn 1903: 157 (discussion of the systematic position); Kuhnt 1909: 120 (distribution); Kuhnt 1911: 83 (distribution). Distribution. Australia.

Diplocoelus parnoides Grouvelle, 1903 Diplocoelus parnoides Grouvelle, 1903b: 195.

Name derivation. Parnus, name of a beetle genus, itself derived from parnops (gr.) – a species of locust, and -ides (gr.) – suffix denoting son or descendant, use to mark affinity or similarity; likely due to the superficial similarity to, also conspicuously pilose, representatives of the genus Parnus (= Dryops) (Dryopidae, Coleoptera). Type locality. Coulée Boulari; Kanala; Nouméa; Tonghoué; Yahoué; forèt du Mont Dor; Bourail. Type material. IRSNB (Coll. A. Fauvel – 4 syntypes – Damoiseau 1968: 28) and MNHN (Colls. E. J.-B. Fleutiaux and A. Grouvelle).

Note. In the original description the number of typespecimens is not provided. References. Fauvel 1903: 314 (description in key, distribution, biology); Kuhnt 1909: 120 (distribution); Kuhnt 1911: 83 (distribution); Damoiseau 1968: 28 (type specimens in IRSNB). Biology. Collected under bark (Fauvel 1903: 314). Distribution. New Caledonia.

Diplocoelus parvus Sharp, 1900 Diplocoelus parvus Sharp, 1900: 622, pl. XVIII, fig. 24.

Name derivation. Parvus (lat.) – small; likely due to the small body size (ca. 2.0 mm in length). Type locality. Cordova [= Córdoba, Vera Cruz], Mexico; El Tumbador, Guatemala; Las Mercedes, Guatemala; Volcan de Chiriqui, Panama. Type material. NHML (7 syntypes). Note. Besides the type-series (seven syntypes), Sharp mentioned also four specimens from Cerro Zunil and Senahu (Guatemala), which he considered a variety (left unnamed). References. Schenkling 1934: 3 (distribution); Blackwelder 1945: 425 (distribution). Distribution. Mexico, Guatemala, Panama.

Diplocoelus piliger Reitter, 1880 Diplocoelus piliger Reitter, 1880a: 6.

Name derivation. Pilus (lat.) – hair (pl. pili – hairs), and gero (lat.) – I am carrying; likely due to the strongly setose body. Type locality. Australia. Type material. MNHN (Colls. E. Reitter and A. Grouvelle). Note. In the original description the number of typespecimens is not provided. References. Masters 1886: 667 (distribution); Blackburn 1903: 157 (discussion on the systematic position, possible synonym of Diplocoelus ovatus MacLeay, 1871); Kuhnt 1909: 120 (distribution, as synonym of Diplocoelus ovatus MacLeay, 1871); Schenkling 1934: 3 (distribution). Distribution. Australia. Diplocoelus pilinotatus Lea, 1922 Diplocoelus pilinotatus Lea, 1922: 297.

Name derivation. Pilus (lat.) – hair, nota (lat.) – mark, and -atus (lat.) – past participle ending, -ate, altogether: “marked with hairs”; likely due to the pattern of body pubescence.

CATALOGUE OF BIPHYLLIDAE (COLEOPTERA) OF THE WORLD

Type locality. Cairns district, Queensland. Type material. SAMA (holotype). References. Schenkling 1934: 3 (distribution). Distribution. Australia (Queensland).

Diplocoelus platysomus Lea,1922 Diplocoelus platysomus Lea,1922: 298.

Name derivation. Platys (gr.) – flat, and soma (gr.) – body; likely due to the characteristic flat body. Type locality. Lord Howe Island. Type material. SAMA (7 syntypes). References. Schenkling 1934: 3 (distribution). Distribution. Australia (Lord Howe Island).

Diplocoelus punctatus Lea, 1895 Diplocoelus punctatus Lea, 1895: 227.

Name derivation. Punctum (lat.) – prick, aperture, point, dot, and -atus (lat.) – suffix of past participle, meaning -ate; likely due to the rather coarse body punctation. Type locality. Richmond River. Type material. SAMA. Note. In the original description the number of typespecimens is not provided. References. Blackburn 1903: 157 (discussion on systematic position, possible synonym of Diplocoelus ovatus MacLeay, 1871); Kuhnt 1909: 120 (distribution); Kuhnt 1911: 83 (distribution); Lea 1921a: 364 (description); Lawrence and Ślipiński 2013: 401, fig. 86H (adult, ventral side, illustration). Distribution. Australia (New South Wales).

Diplocoelus rudis (LeConte, 1863) Marginus rudis LeConte, 1863: 73 (in Diplocoelus – Reitter 1878: 189).

Note. Reitter did not formally introduce the name combination, but synonymized the genus of which rudis is the type-species. Name derivation. Rudis (lat.) – rough, unpolished; likely due to the coarse body punctation. Type locality. Middle and Southern States [of USA]. Type material. MCZC (holotype according to Goodrich and Springer 1992: 371 – see note). Note. The status of the type material presented by Goodrich and Springer (1992: 371) might be incorrect. LeConte had certainly analysed several specimens, and originally did not designated the holotype.

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Diplocoelus philothermoides Reitter, 1878: 189. Synonymized by Horn 1878: 606.

Name derivation. Philothermus – name of one of the genera of the family Cerylonidae, philos (gr.) – friend, therme (gr.) – warmth, and -ides (gr.) – suffix denoting affinity or similarity; likely due to the similarity with Philothermus (Cerylonidae, Coleoptera). Type locality. Nordamerica [= North America]. Type material. DEIC (Coll. G. Kraatz). Note. In the original description the number of typespecimens is not provided. Note. Goodrich and Springer (1992: 371) had not found type (or types) of Diplocoelus philothermoides Reitter in Edmund Reitter’s collection, but considered them as but smaller and more arched specimens within the range of variability of Diplocoelus rudis LeConte. References. Gemminger and Harold 1868: 909 (Marginus, distribution); Crotch 1873: 44 (Marginus, distribution); Horn 1878: 606 (description, distribution); Schwarz 1878: 445 (Marginus, distribution, biology); Austin 1880: 20 (distribution); Henshaw 1885: 53 (distribution); Casey 1900: 80 (description in key); Kuhnt 1909: 120 (world catalogue); Blatchley 1910: 572 (description); Kuhnt 1911: 83 (distribution); Leng 1920: 202 (distribution); Schenkling 1934: 3 (philothermoides – species bona, distribution); Goodrich and Springer 1992: 371, figs. 5, 10, 13, 16, and 19 (description, distribution); Peck and Thomas 1998: 94 (distribution); Ulyshen and Hanula 2009: 659, Appendix B (distribution, biology); Cline and Shockley 2010: 306 (biology); Grebennikov and Leschen 2010: 90, fig. 23 (exoskeletal cavities on head); Ulyshen and Hanula 2010: 189, tab.1 and 190, appendix 1 (biology); Ulyshen et al. 2010: 249, tab. 1 (biology); Lawrence et al. 2011: 9 (phylogeny). Biology. This species may be collected throughout the year under bark of trees, most frequently pines (Pinus sp., Pinaceae) (Cline and Shockley 2010: 306) (e. g. Pinus taeda L. – Ulyshen and Hanula 2009: 659, Appendix B; Ulyshen and Hanula 2010: 189, tab.1 and 190, appendix 1; Ulyshen et al. 2010: 249, tab. 1), but also under loose bark of oak (Quercus sp., Fagaceae) (Schwarz 1878: 445; Cline and Shockley 2010: 306), on logs of water oak (Quercus nigra L.) (Ulyshen and Hanula 2009: 659, Appendix B) or American sweetgum (Liquidambar styraciflua L., Altingiaceae) (Ulyshen and Hanula 2009: 659, Appendix B), and hickory (Carya sp., Juglandaceae) (Cline and Shockley 2010: 306); contrary to Diplocoelus brunneus LeConte which prefers fallen logs rather than standing (even if dead) trees. The species is often collected in pitfall traps with various baits (Goodrich and Springer 1992: 371). Distribution. USA (New Jersey, Indiana, Maryland, Virginia, Kentucky, Illinois, Missouri, North

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Carolina, South Carolina, Tennessee, Arkansas, Oklahoma, Georgia, Alabama, Mississippi, Louisiana, Texas, Florida).

Distribution. Azerbaijan.

Diplocoelus turbinatus Grouvelle, 1905 Diplocoelus similis Grouvelle, 1898 Diplocoelus (Marginus) similis Grouvelle, 1898: 44.

Name derivation. Similis (lat.) – similar; likely due to the resemblance with D. rudis LeConte. Type locality. Mount Gay Estate, Grenada. Type material. CUMZ. Note. In the original description the number of typespecimens is not provided. References. Kuhnt 1909: 120 (distribution); Kuhnt 1911: 83 (distribution); Blackwelder 1945: 469 (distribution). Distribution. Grenada. Diplocoelus subjectus Grouvelle, 1919 Diplocoelus (Marginus) oblongus Reitter, 1878: 189, nec (Germain, 1855: 395).

Name derivation. Oblongus (lat.) – elongated; likely due to the oblong body outline. Type locality. Brasilia [= Brazil]. Type material. MNHN (Coll. A. Grouvelle). Note. In the original description the number of typespecimens is not provided. Selenoderus oblongus Deyrolle in Reitter, 1878: 189 (nomen nudum). Diplocoelus subjectus Grouvelle, 1919: 67.

Name derivation. Subiectus (lat.) – lying under something, in this case – replacing. Note. Replacement name for Diplocoelus oblongus Reitter, 1878: 188, nec (Germain, 1855: 395). References. Schenkling 1934: 3 (distribution); Blackwelder 1945: 425 (distribution). Distribution. Brazil. Diplocoelus talyshensis Nikitsky, 1993 Diplocoelus talyshensis Nikitsky, 1993: 154, figs. 6–9.

Name derivation. From the type locality, eastern part of Talysh Mts. Type locality. S. [selo] [= village] Avrora. Type material. ZMUM (holotype – male, 3 paratypes – female and 2 of sex unknown). References. Löbl and Smetana 2008: 27 (distribution). Biology. Type specimens were collected in July (Nikitsky 1993: 154).

Diplocoelus turbinatus Grouvelle, 1905b: 127.

Name derivation. Turbinatus (lat.) – conical; likely due to the body outline. Type locality. Tucuman, République Argentine [= Argentina]. Type material. MACN (Coll. C. Bruch) and ?MNHN (Coll. A. Grouvelle). Note. In the original description the number of typespecimens is not provided. References. Kuhnt 1909: 120 (distribution); Kuhnt 1911: 83 (distribution); Bruch 1914: 385 (distribution); Blackwelder 1945: 469 (distribution). Distribution. Argentina (Tucuman Prov.).

Diplocoelus villosus Grouvelle, 1905 Diplocoelus villosus Grouvelle, 1905b: 126.

Name derivation. Villosus (lat.) – hirsute; likely due to the abundant body pilosity. Type locality. San Pedro, Paraguay. Type material. MACN (Coll. C. Bruch) and ?MNHN (Coll. A. Grouvelle). Note. In the original description the number of typespecimens is not provided. References. Kuhnt 1909: 120 (distribution); Kuhnt 1911: 83 (distribution); Bruch 1914: 385 (distribution); Blackwelder 1945: 469 (distribution). Distribution. Paraguay, Argentina.

Diplocoelus xanthorrhoeae Lea, 1921 Diplocoelus xanthorrhoeae Lea, 1921a: 364.

Name derivation. Specific epithet derived from the host plant belonging to the genus Xanthorrhoea Smith, from which type-specimens were collected. Type locality. Swan River, Darling Ranges – Western Australia; Sydney, New South Wales; Huon River, Tasmania. Type material. SAMA. Note. In the original description the number of typespecimens is not provided. References. Schenkling 1934: 3 (distribution); Hawkeswood 1985: 163 (biology, distribution). Biology. Imagines were collected on leaf-bases of several species of Xanthorrhoea sp. (Xanthorrhoeaceae) (Lea 1921a: 364) – e.g. X. johnsoni Lee (Hawkeswood 1985: 163). Collected in October and

CATALOGUE OF BIPHYLLIDAE (COLEOPTERA) OF THE WORLD

November, and it may breed in rotting vegetation in moist areas amongst leaves near the top of the caudex (Hawkeswood 1985: 163). Distribution. Australia (Western Australia, Queensland, New South Wales), Tasmania.

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Distribution. Panama.

Euderopus iteratus Sharp, 1900 Euderopus iteratus Sharp, 1900: 624.

Genus Euderopus Sharp, 1900 Euderopus Sharp, 1900: 622.

Name derivation. Eu (gr.) – amplifying prefix meaning “good, genuine”, deros (gr.) – skin, and pus (gr.) – leg. Gender. Masculine. Type species. Euderopus regularis Sharp, 1900: 624, here designated References. Schenkling 1934: 5 (supplement to world catalogue); Blackwelder 1945: 425 (catalogue of Central and South American species). Immature stages. Unknown. Biology. Unknown, except that most species were collected in mountainous regions (Sharp 1900: 623, 624; Sharp, 1902: 625). Number of species. 8. Distribution. Central America.

Euderopus brevipes Sharp, 1900 Euderopus brevipes Sharp, 1900: 622.

Name derivation. Brevis (lat) – short and pes (lat.) – foot; likely due to the relatively short tarsi. Type locality. Capetillo, Guatemala. Type material. NHML (holotype). Note. Holotype is in poor condition, as mentioned by the author of original description (Sharp 1900: 622). References. Schenkling 1934: 5 (distribution); Blackwelder 1945: 425 (distribution). Distribution. Guatemala.

Euderopus chaetophorus Sharp, 1902 Euderopus chaetophorus Sharp, 1902: 625.

Name derivation. Chaite (gr.) – bristle, and phoreo (gr.) – bear, carry; likely due to the ornate body setation. Type locality. Volcan de Chiriqui, Panama. Type material. NHML (5 syntypes). References. Schenkling 1934: 5 (distribution); Blackwelder 1945: 425 (distribution). Biology. The type material was collected on slopes of Chiriqui volcano at 2500–4000 ft. (= 760–1220 m.) a.s.l. (Sharp 1902: 625).

Name derivation. Iteratio (lat.) – repetition. Type locality. Volcan de Chiriqui, Panama. Type material. NHML (holotype). References. Schenkling 1934: 5 (distribution); Blackwelder 1945: 425 (distribution). Biology. Holotype was collected on slopes of Chiriqui volcano at 2500–4000 ft. (= 760–1220 m.) a.s.l. (Sharp 1900: 624). Distribution. Panama.

Euderopus microps Sharp, 1900 Euderopus microps Sharp, 1900: 623.

Name derivation. Mikros (gr.) – small, and opsis (gr.) – view, appearance; likely due to the small body size. Type locality. Panima, Vera Paz [= Baja Verapaz], Guatemala; Volcan de Chiriqui, Panama. Type material. NHML (3 syntypes). Note. Sharp treated the unique specimen from Volcan de Chiriqui as a variety, and did not name it. References. Schenkling 1934: 5 (distribution); Blackwelder 1945: 425 (distribution). Distribution. Guatemala, Panama. Note. Schenkling (1934: 5) did not mention Panama when describing the distribution.

Euderopus perbrevis Sharp, 1900 Euderopus perbrevis Sharp, 1900: 623.

Name derivation. Per- (lat.) – amplifying prefix, and brevis (lat.) – short; likely due to the relatively short and wide body. Type locality. Cordova [= Córdoba, Vera Cruz], Mexico; Senahu, Guatemala. Type material. NHML (2 syntypes). Note. Syntype originating from Mexico is in poor condition (Sharp 1900: 623). References. Schenkling 1934: 5 (distribution); Blackwelder 1945: 425 (distribution). Distribution. Mexico, Guatemala.

Euderopus regularis Sharp, 1900 Euderopus regularis Sharp, 1900: 624.

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Name derivation. Regula (lat.) – rule, and -aris (lat.) – suffix meaning -ar, regular; likely due to the regular body outline. Type locality. Volcan de Chiriqui, Panama. Type material. NHML (2 syntypes). References. Schenkling 1934: 5 (distribution); Blackwelder 1945: 425 (distribution). Biology. The type material was collected on slopes of the Chiriqui volcano at 2000–3000 ft. (= 610–915 m.) a.s.l. (Sharp 1900: 624). Distribution. Panama.

Type species. Eurhanius humeralis Reitter, 1887: 13, by monotypy. References. Reitter 1888b: 13 (description); Schoenfeldt 1891: 258 (catalogue of Japanese species, additions); Kuhnt 1909: 121 (description, world catalogue); Kuhnt 1911: 84 (world catalogue); Jakobson 1915: 936 (catalogue); Falcoz 1926: 73 (systematic position); Mader 1926: 719 (distribution); Miwa 1929: 121 (systematic position). Immature stages. Unknown. Biology. Unknown. Number of species. 1. Distribution. Japan.

Euderopus setosus Sharp, 1900 Euderopus setosus Sharp, 1900: 624.

Name derivation. Setosus (lat.) – setaceous; likely due to the erect setae on the body and in particular the head. Type locality. Volcan de Chiriqui, Panama. Type material. MNHL. Note. In the original description the number of typespecimens is not provided. References. Schenkling 1934: 5 (distribution); Blackwelder 1945: 425 (distribution). Biology. The type material was collected on slopes of the Chiriqui volcano at 2000–3000 ft. (= 610–915 m.) a.s.l. (Sharp 1900: 624). Distribution. Panama.

Euderopus unicolor Sharp, 1900

Eurhanius humeralis Reitter, 1887 Eurhanius humeralis Reitter, 1887: 13.

Name derivation. Humerus (lat.) – shoulder, and -alis (lat.) – suffix -al; likely due to the presence of spots on elytral humeri. Type locality. Japan. Type material. Unknown. Note. Depository of the type unknown, Reitter’s collection preserved in the Hungarian Natural History Museum in Budapest does not contain types of this species (O. Merkl – pers. obs.). References. Reitter 1888b: 12 (description, distribution); Reitter 1889a: 301 (distribution); Schoenfeldt 1891: 258 (distribution); Kuhnt 1909: 121 (distribution); Kuhnt 1911: 84 (distribution); Jakobson 1915: 936 (distribution); Mader 1926: 719 (distribution). Distribution. Japan.

Euderopus unicolor Sharp, 1900: 623.

Name derivation. Unus (lat.) – one, and color (lat.) – colour; likely due to the uniform body colouration. Type locality. Volcan de Chiriqui, Panama. Type material. NHML. Note. In the original description the number of typespecimens is not provided. References. Schenkling 1934: 5 (distribution); Blackwelder 1945: 425 (distribution). Biology. The type material was collected on slopes of the Chiriqui volcano at 2000–3000 ft. (= 610–915 m.) a.s.l. (Sharp 1900: 623). Distribution. Panama.

Genus Eurhanius Reitter, 1887 Eurhanius Reitter, 1887: 13.

Name derivation. Eu- (gr.) – amplifying prefix and rhanis (gr.) – drop. Gender. Masculine.

Genus Gonicoelus Sharp, 1900 Gonicoelus Sharp, 1900: 604.

Name derivation. Gonia (gr.) – angle, and coelus (lat.) [from gr. koilos – empty, hollow] (allusion to the name Diplocoelus). Gender. Masculine. Type species. Gonicoelus unicornis Sharp, 1900: 605, here designated Note. Sharp (1900: 604) described genus Gonicoelus for numerous new species, but did not designate the type species. References. Schenkling 1934: 3 (supplement to the world catalogue), Blackwelder 1945: 425 (catalogue of the Central and South American species); Lawrence 1991: 475 (biology); Chavarria 1994: 116 (biology); Weir and Rossi 2001: 174 (biology). Immature stages. Unknown. Biology. Unspecified representatives of this genus were found under fermenting bark (Lawrence 1991:

CATALOGUE OF BIPHYLLIDAE (COLEOPTERA) OF THE WORLD

475). Adults of undetermined species were observed in the nest of bumblebee Bombus ephippiatus Say (Apidae, Hymenoptera) in Costa Rica, but relations with these insects are not known (Chavarria 1994: 116). From the specimens of this genus collected on Trinidad, a parasitic fungus Cucujomyces gonicoeli Thaxter (Laboulbeniaceae, Laboulbeniales) (Thaxter 1931: 179) has been described, which was found also on a congeneric female from Bolivia (Weir and Rossi 2001: 174). Numberof species. 39. Distribution. Central and South America. Note. Single species, Gonicoelus deplanatus Sharp was intercepted to North America.

Gonicoelus affinis Grouvelle, 1902 Gonicoelus affinis Grouvelle, 1902a: 465.

Name derivation. Affinis (lat.) – relative. Type locality. Colombie [= Colombia]. Type material. ?MNHN (Coll. A. Grouvelle). Note. In the original description the number of typespecimens is not provided. References. Grouvelle and Léveillé 1903: 460, pl. VII, fig. 5 (illustration not included in original description); Schenkling 1934: 4 (distribution); Blackwelder 1945: 425 (distribution). Distribution. Colombia.

Gonicoelus angustus Sharp, 1900 Gonicoelus angustus Sharp, 1900: 615.

Name derivation. Angustus (lat.) – narrow; name likely alludes to the slender body of the beetle. Type locality. Cordova [= Córdoba, Vera Cruz], Mexico. Type material. NHML (holotype – male). References. Schenkling 1934: 4 (distribution); Blackwelder 1945: 425 (distribution). Distribution. Mexico.

Gonicoelus arduus Sharp, 1900

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References. Schenkling 1934: 4 (distribution); Blackwelder 1945: 425 (distribution). Biology. Type specimens were collected on slopes of the Chriqui volcano at 2500–4000 ft. (= 760–1220 m.) a.s.l. (Sharp 1900: 617). Distribution. Guatemala, Panama.

Gonicoelus bimargo Sharp, 1900 Gonicoelus bimargo Sharp, 1900: 616.

Name derivation. Bis (lat.) – twice, and margo (lat.) – margin; like the majority of representatives of the family Biphyllidae, this species shows conspicuous longitudinal costulae just at sides of pronotum, providing an appearance of an additional lateral margin. Type locality. Paraiso, Guatemala. Type material. NHML (holotype). References. Schenkling 1934: 4 (distribution); Blackwelder 1945: 425 (distribution). Distribution. Guatemala.

Gonicoelus brevicollis Sharp, 1900 Gonicoelus brevicollis Sharp, 1900: 607.

Name derivation. Brevis (lat.) – short, and collum (lat.) – neck; likely due to the strongly transverse pronotum. Type locality. Jalapa, Mexico; Tuxtla, Mexico. Type material. NHML (2 syntypes). References. Schenkling 1934: 4 (distribution); Blackwelder 1945: 425 (distribution). Distribution. Mexico.

Gonicoelus celatus Sharp, 1900 Gonicoelus celatus Sharp, 1900: 608.

Name derivation. Celate (lat.) – secretly. Type locality. Bugaba, Panama. Type material. NHML (3 syntypes). References. Schenkling 1934: 4 (distribution); Blackwelder 1945: 425 (distribution). Distribution. Panama.

Gonicoelus arduus Sharp, 1900: 617.

Name derivation. Arduus (lat.) – steep, high. Type locality. El Tumbador, Guatemala; Zapote [= El Zapote, Escuintla], Guatemala; Volcan de Chiriqui, Panama. Type material. NHML. Note. In the original description the number of typespecimens is not provided.

Gonicoelus championi Sharp, 1900 Gonicoelus championi Sharp, 1900: 613.

Name derivation. Specific epithet honours George Charles Champion (1851–1927), eminent British coleopterologist, who – both as collector (among others, of

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the type of this species) and as author – greatly contributed to the elaboration of the monumental series “Biologia Centrali Americana”. Type locality. Capetillo, Guatemala. Type material. NHML (holotype – male). References. Schenkling 1934: 4 (distribution); Blackwelder 1945: 425 (distribution). Distribution. Guatemala. Gonicoelus chontalenus Sharp, 1900 Gonicoelus chontalenus Sharp, 1900: 614.

Name derivation. Name derived from the department of Chontales in Nicaragua (Selander and Vaurie, 1962: 1), where Edward Mason Janson collected the only hitherto known specimen of this species. Type locality. Chontales, Nicaragua. Type material. NHML (holotype). References. Schenkling 1934: 4 (distribution); Blackwelder 1945: 425 (distribution). Distribution. Nicaragua. Gonicoelus concolor Sharp, 1900 Gonicoelus concolor Sharp, 1900: 618.

Name derivation. Concolor (lat.) – unicolorous; likely due to the uniformly rufous colouration of the body. Type locality. Volcan de Chiriqui, Panama. Type material. NHML (2 syntypes). References. Schenkling 1934: 4 (distribution); Blackwelder 1945: 425 (distribution). Biology. Type specimens were collected on slopes of Chriqui volcano at 6000–8000 ft. (= 1830–2440 m.) a.s.l. (Sharp 1900: 618). Distribution. Panama. Gonicoelus crispatus Sharp, 1900 Gonicoelus crispatus Sharp, 1900: 615.

Name derivation. Crispus (lat.) – curly, and -atus (lat.) – suffix -ate; likely due to the specific structure of male heads. Type locality. Volcan de Chiriqui, Panama. Type material. NHML. Note. In the original description the number of typespecimens is not provided. References. Schenkling 1934: 4 (distribution); Blackwelder 1945: 425 (distribution). Biology. Type specimens were collected on slopes of Chriqui volcano at 2500–4000 ft. (= 760–1220 m.) a.s.l. (Sharp 1900: 615).

Distribution. Panama.

Gonicoelus cultratus Sharp, 1900 Gonicoelus cultratus Sharp, 1900: 609.

Name derivation. Culter (lat.) – ploughshare, knife, and -atus (lat.) – suffix -ate; likely in reference to conspicuous blade-like extensions on the femora. Type locality. Volcan de Chiriqui, Panama. Type material. NHML (holotype – male). References. Schenkling 1934: 4 (distribution); Blackwelder 1945: 425 (distribution). Biology. Single male (holotype) has been collected on slopes of Chriqui volcano at 2500 ft. (= 760 m.) a.s.l. (Sharp 1900: 609). Distribution. Panama.

Gonicoelus deplanatus Sharp, 1900 Gonicoelus deplanatus Sharp, 1900: 620.

Name derivation. Deplano (lat.) – to level, and -atus (lat.) – suffix -ed; likely alluding to flattened body. Type locality. Cordova [= Córdoba, Vera Cruz], Mexico; Teapa, Mexico; Motzorongo, Mexico; Belize [City], British Honduras [= Belize]; Rio Hondo, British Honduras [= Belize]; San Juan, Guatemala; Senahu, Guatemala; Chacoj, Guatemala; Tamahu, Guatemala; Champerico, Guatemala; Zapote [= El Zapote, Escuintla], Guatemala; Chontales, Nicaragua; Volcan de Chiriqui, Panama; Bugaba, Panama. Type material. MNHL. Note. In the original description the number of typespecimens is not provided. References. Schenkling 1934: 4 (distribution); Blackwelder 1945: 425 (distribution); Mumford 1967: 24, tab. (distribution, biology, introduced species). Biology. Intercepted to USA (Florida) from Guatemala with leaf of Yucca sp. (Asparagaceae) (Mumford 1967: 24, tab.). Distribution. Mexico, British Honduras, Guatemala, Nicaragua, Panama, USA (Florida) (introduced from Guatemala).

Gonicoelus germanus Sharp, 1900 Gonicoelus germanus Sharp, 1900: 606.

Name derivation. Germanus (lat.) – relative, kin. Type locality. Mexico; Jalapa, Mexico; Chontales, Nicaragua. Type material. NHML (5 syntypes).

CATALOGUE OF BIPHYLLIDAE (COLEOPTERA) OF THE WORLD

References. Schenkling 1934: 4 (distribution); Blackwelder 1945: 425 (distribution). Distribution. Mexico, Nicaragua. Gonicoelus guatemalenus Sharp, 1900 Gonicoelus guatemalenus Sharp, 1900: 611.

Name derivation. From the place of collection, Guatemala, whose name is a derivative of quauhtlemallan, meaning “area with many trees/forests” in Middle-American language nahuatl. Type locality. El Tumbador, Guatemala. Type material. NHML (holotype – male). References. Schenkling 1934: 4 (distribution); Blackwelder 1945: 425 (distribution). Distribution. Guatemala. Gonicoelus hirtus Sharp, 1900 Gonicoelus hirtus Sharp, 1900: 608.

Name derivation. Hirtus (lat.) – pilose; referring to the abundant erect pilosity on the dorsal side of the beetle. Type locality. Volcan de Chiriqui, Panama. Type material. NHML (holotype). References. Schenkling 1934: 4 (distribution); Blackwelder 1945: 425 (distribution). Distribution. Panama. Gonicoelus humilis Sharp, 1900 Gonicoelus humilis Sharp, 1900: 616.

Name derivation. Humilis (lat.) – low, weak; probably alluding to flat front of males, which is devoid of any characteristic extension, protuberances or tubercles. Type locality. Jalapa, Mexico; Mexico. Type material. NHML (2 syntypes – males). References. Schenkling 1934: 4 (distribution); Blackwelder 1945: 425 (distribution). Distribution. Mexico. Note. Schenkling’s (1934: 4) report of the species from Guatemala was probably erroneous, as he did not mention Mexico; Blackwelder (1945: 425) listed both countries but was not certain that the information concerning Guatemala is true and put this name with a question mark.

Gonicoelus hystrix Sharp, 1900 Gonicoelus hystrix Sharp, 1900: 612.

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Name derivation. Hystrix (gr./lat.) – porcupine; refers to long erect setae on dorsal side of this species Type locality. Jalapa, Mexico; Cordova [= Córdoba, Vera Cruz], Mexico. Type material. NHML (11 syntypes). References. Schenkling 1934: 4 (distribution); Blackwelder 1945: 425 (distribution). Distribution. Mexico. Gonicoelus laticollis Sharp, 1900 Gonicoelus laticollis Sharp, 1900: 617.

Name derivation. Latus (lat.) – wide, and collum (lat.) – neck; likely due to the distinctly transverse pronotum. Type locality. Sinanja, Vera Paz [= Baja Verapaz], Guatemala. Type material. NHML (holotype – male). References. Schenkling 1934: 4 (distribution); Blackwelder 1945: 425 (distribution). Distribution. Guatemala. Gonicoelus latus Sharp, 1900 Gonicoelus latus Sharp, 1900: 611.

Name derivation. Latus (lat.) – wide; likely due to the wide body of the beetle. Type locality. Volcan de Chiriqui, Panama. Type material. NHML (holotype – female). References. Schenkling 1934: 4 (distribution); Blackwelder 1945: 425 (distribution). Distribution. Panama. Gonicoelus longicornis Sharp, 1900 Gonicoelus longicornis Sharp, 1900: 609.

Name derivation. Longus (lat.) – long, and cornu (lat.) – horn – in this case antenna; likely due to the exceptionally elongate antennae. Type locality. Volcan de Chiriqui, Panama. Type material. NHML (2 syntypes – females). References. Schenkling 1934: 4 (distribution); Blackwelder 1945: 425 (distribution). Distribution. Panama. Gonicoelus mediocris Sharp, 1900 Gonicoelus mediocris Sharp, 1900: 610.

Name derivation. Mediocris (lat.) – medium, average; likely due to the indistinctive appearance of this species.

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Type locality. Bugaba, Panama. Type material. NHML (holotype). References. Schenkling 1934: 4 (distribution); Blackwelder 1945: 425 (distribution). Distribution. Panama.

Gonicoelus mexicanus Sharp, 1900

Name derivation. Muticus (lat.) – mutilated; in reference to lack of characteristic dimorphic characters on head of male. Type locality. Jalapa, Mexico. Type material. NHML (4 syntypes). References. Schenkling 1934: 4 (distribution); Blackwelder 1945: 425 (distribution). Distribution. Mexico.

Gonicoelus mexicanus Sharp, 1900: 607.

Name derivation. Mexicanus (lat.) – Mexican; specific epithet derived from where the species was collected. Type locality. Jalapa, Mexico. Type material. NHML (holotype – male). References. Schenkling 1934: 4 (distribution); Blackwelder 1945: 425 (distribution). Distribution. Mexico.

Gonicoelus mollis Sharp, 1900

Gonicoelus oopsis Sharp, 1900 Gonicoelus oopsis Sharp, 1900: 619.

Name derivation. Oon (gr.) – egg, and opsis (gr.) – appearance; likely due to the ovate body outline. Type locality. El Tumbador, Guatemala. Type material. NHML (2 syntypes). References. Schenkling 1934: 4 (distribution); Blackwelder 1945: 425 (distribution). Distribution. Guatemala.

Gonicoelus mollis Sharp, 1900: 621.

Gonicoelus parnoides Sharp, 1900 Name derivation. Mollis (lat.) – soft; for soft, silky pubescence of dorsal surface. Type locality. Capetillo, Guatemala; Cerro Zunil, Guatemala. Type material. NHML (7 syntypes). References. Schenkling 1934: 4 (distribution); Blackwelder 1945: 425 (distribution). Distribution. Guatemala.

Gonicoelus monticola Sharp, 1900 Gonicoelus monticola Sharp, 1900: 619.

Name derivation. Monticola (lat.) – mountaineer, from mons, montis (lat.) – mountain, and colo (lat.) – inhabit; likely due to the species’ occurrence in mountains. Type locality. Totonicapam [= Totonicapan], Guatemala. Type material. NHML (holotype). References. Schenkling 1934: 4 (distribution); Blackwelder 1945: 425 (distribution). Biology. The only known specimen of this species was collected at 8500–10500 ft. (= 2590–3200 m.) a.s.l. (Sharp 1900: 619). Distribution. Guatemala.

Gonicoelus muticus Sharp, 1900 Gonicoelus muticus Sharp, 1900: 618.

Gonicoelus parnoides Sharp, 1900: 610, pl. XVIII, fig. 23.

Name derivation. From beetle genus name Parnus, derived in turn from parnops (gr.) – a locust species, and -ides (gr.) – suffix meaning relative; name given probably because of superficial similarity to representatives of Parnus (= Dryops) (Dryopidae, Coleoptera). Type locality. Cordova [= Córdoba, Vera Cruz], Mexico. Type material. NHML (3 syntypes – male and 2 females). References. Schenkling 1934: 4 (distribution); Blackwelder 1945: 425 (distribution). Distribution. Mexico.

Gonicoelus planus Sharp, 1900 Gonicoelus planus Sharp, 1900: 614.

Name derivation. Planus (lat.) – flat; likely due to the distinctly flattened body. Type locality. Bugaba, Panama. Type material. NHML (holotype). Note. Poor condition of the holotype was mentioned by the author in the original description (Sharp 1900: 615). References. Schenkling 1934: 4 (distribution); Blackwelder 1945: 425 (distribution). Distribution. Panama.

CATALOGUE OF BIPHYLLIDAE (COLEOPTERA) OF THE WORLD

Gonicoelus relictus Sharp, 1900 Gonicoelus relictus Sharp, 1900: 606.

Name derivation. Relictus (lat.) – left. Type locality. Volcan de Chiriqui, Panama. Type material. NHML (holotype – female). References. Schenkling 1934: 4 (distribution); Blackwelder 1945: 425 (distribution). Biology. Holotype was collected on slopes of Chiriqui volcano at 3000 ft. (= 915 m.) a.s.l. (Sharp 1900: 606). Distribution. Panama. Gonicoelus rudis Sharp, 1900 Gonicoelus rudis Sharp, 1900: 606.

Name derivation. Rudis (lat.) – unpolished, crude; likely due to the strong setation of dorsal side that gives the beetle a somewhat rough appearance. Type locality. Caldera, Chiriqui, Panama. Type material. NHML. Note. In the original description the number of typespecimens is not provided. References. Schenkling 1934: 4 (distribution); Blackwelder 1945: 425 (distribution). Biology. Type specimens were caught at 1200 ft. (= 370 m.) a.s.l. (Sharp 1900: 606). Distribution. Panama. Gonicoelus securiger Sharp, 1900 Gonicoelus securiger Sharp, 1900: 612.

Name derivation. Securis (lat.) – axe, and gero (lat.) – carry; likely due to the distal joint of maxillary palpus markedly widened in the form of an axe, i.e. securiform shape. Type locality. Volcan de Chiriqui, Panama. Type material. NHML (2 syntypes). References. Schenkling 1934: 4 (distribution); Blackwelder 1945: 425 (distribution). Biology. Type specimens were caught by George Charles Champion on slopes of Chiriqui volcano at 3000 ft. (= 915 m.) a.s.l. (Sharp 1900: 612). Distribution. Panama. Gonicoelus segnis Sharp, 1900 Gonicoelus segnis Sharp, 1900: 618.

Name derivation. Segnis (lat.) – lazy. Type locality. Cordova [= Córdoba, Vera Cruz], Mexico; Jalapa, Mexico; Mexico. Type material. NHML.

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Note. In the original description the number of typespecimens is not provided. References. Schenkling 1934: 4 (distribution); Blackwelder 1945: 425 (distribution). Distribution. Mexico. Gonicoelus sellatus Sharp, 1900 Gonicoelus sellatus Sharp, 1900: 620.

Name derivation. Sella (lat.) – saddle, and -atus (lat.) – adjectival ending – ate; likely due to the black saddle-shaped spot at the elytral base. Type locality. Near the city Guatemala, Guatemala; Capetillo, Guatemala; Zapote [= El Zapote, Escuintla], Guatemala. Type material. NHML. Note. In the original description the number of typespecimens is not provided. References. Schenkling 1934: 4 (distribution); Blackwelder 1945: 425 (distribution). Distribution. Guatemala. Gonicoelus spheniscus Sharp, 1900 Gonicoelus spheniscus Sharp, 1900: 616.

Name derivation. Spheniskos (gr.) – small wedge; likely due to the relatively slender and convex body with tapering elytra. Type locality. Chiacam, Vera Paz [= Alta Verapaz], Guatemala. Type material. NHML (holotype). References. Schenkling 1934: 4 (distribution); Blackwelder 1945: 425 (distribution). Distribution. Guatemala. Gonicoelus subtilis Sharp, 1900 Gonicoelus subtilis Sharp, 1900: 613.

Name derivation. Subtilis (lat.) – delicate, fine. Type locality. Volcan de Chiriqui, Panama; Bugaba, Panama. Type material. NHML. Note. In the original description the number of typespecimens is not provided. Three specimens from Bugaba were treated by Sharp as a variety but he left it unnamed. References. Schenkling 1934: 4 (distribution); Blackwelder 1945: 425 (distribution). Biology. Syntypes were caught on slopes of Chiriqui volcano at 2000–4000 ft. (= 610–1220 m.) a.s.l. (Sharp 1900: 614). Distribution. Panama.

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Gonicoelus throscoides Sharp, 1900 Gonicoelus throscoides Sharp, 1900: 613.

Name derivation. Throsko (gr.) – jump (but in this case directly from the genus name Throscus) and -ides (gr.) – suffix meaning affinity or kinship; likely due to the superficial resemblance to representatives of Throscus Latr. (= Trixagus Kugell.) (Throscidae, Coleoptera). Type locality. Volcan de Chiriqui, Panama. Type material. NHML (holotype). References. Schenkling 1934: 4 (distribution); Blackwelder 1945: 425 (distribution). Distribution. Panama.

Gonicoelus tricornis Sharp, 1900

Name derivation. Vestitus (lat.) – dressed, adorned; likely due to the dense pubescence of the beetle. Type locality. Cordova [= Córdoba, Vera Cruz], Mexico; San Juan, Vera Paz [= Alta Verapaz], Guatemala. Type material. NHML (2 syntypes). References. Schenkling 1934: 4 (distribution); Blackwelder 1945: 425 (distribution). Distribution. Mexico, Guatemala.

Fossil taxa Unlike other small families of beetles (e. g. Dascillidae – see Jin et al. 2013a: 501 and Jin et al. 2013b: 551) the number of known fossil specimens of Biphyllidae is low.

Gonicoelus tricornis Sharp, 1900: 605.

Name derivation. Treis (gr.) – three, and cornu (lat.) – horn; likely due to the three outgrowths (one at anterior margin of pronotum and two on head) in males of this species. Type locality. Bugaba, Panama. Type material. NHML (2 syntypes – male and female). References. Schenkling 1934: 4 (distribution); Blackwelder 1945: 425 (distribution). Distribution. Panama.

Gonicoelus unicornis Sharp, 1900 Gonicoelus unicornis Sharp, 1900: 605, pl. XVIII, fig. 22.

Name derivation. Unus (lat.) – one, and cornu (lat.) – horn; likely due to the single sharp curcular extension resembling horn of unicorn. Type locality. Panima, Vera Paz [= Baja Verapaz], Guatemala; Volcan de Chiriqui, Panama; Boquete [= Bajo Boquete], Panama; Bugaba, Panama. Type material. NHML (23 syntypes). References. Schenkling 1934: 4 (distribution); Blackwelder 1945: 425 (distribution). Biology. Type specimens were collected on the slopes of Chiriqui volcano at 3000 ft. (= 915 m.) a.s.l. (Sharp 1900: 605). Distribution. Guatemala, Panama.

Gonicoelus vestitus Sharp, 1900 Gonicoelus vestitus Sharp, 1900: 611.

?Biphyllidae Kirejtshuk and Azar 2008: 15 (Lebanese amber).

Note. According to authors systematic position is uncertain.

Biphyllus sp. ?B Kuhnt 1909: 119 suggested that Triphyllus heeri Oustalet, 1874: 189, pl. 6, fig. 13 belongs to the genus Diphyllus (= Biphyllus). The species was described from a single specimen from Aix-en-Provence in France. The systematic position of this species is still unclear, but probably it doesn’t belong to Biphyllidae (pers. obs).

Diplocoelus sp. Klebs 1910: 239 (Baltic amber, according to author systematic position uncertain – see also Spahr 1981: 51); Bachofen-Echt 1949: 111 (Baltic amber); Spahr 1981: 51 (catalogue).

Diplocoelus probiphyllus Vitali, 2010 Diplocoelus probiphyllus Vitali, 2010: 168, figs. 1–2 (photos of holotype), 3 (reconstruction) and 4b (antennal club).

Name derivation. Pro (gr.) – before, and Biphyllus. Type locality. Lithuania, Baltic Coast, amber. Type material. FVC (holotype).

CATALOGUE OF BIPHYLLIDAE (COLEOPTERA) OF THE WORLD

ACKNOWLEDGEMENTS This work would never have been made without the help of many persons. First of all I would like to thank my wife, Monika, for her incessant, essential, and spiritual support. I am grateful to Dr. Otto Merkl (Hungarian Natural History Museum, Budapest) for information about the collections of the HNHM; to Prof. Dariusz Iwan and Dr. Marcin Kamiński (Muzeum i Instytut Zoologii PAN, Warszawa) for helpful advice concerning the project; to Prof. Masahiro Ohara (Hokkaido University Museum, Sapporo) and Dr. Adam Stroiński (Muzeum i Instytut Zoologii PAN, Warszawa) for assistance in the acquisition of important publications; and Dr. Roman Hołyński for help in translation of the text and relevant suggestions.

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CATALOGUE OF BIPHYLLIDAE (COLEOPTERA) OF THE WORLD

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Received: April 12, 2015 Accepted: August 26, 2015

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INDEX Biphyllidae acuminata, Althaesia 413 aequalis, Biphyllus 416 aequalis, Diphyllus 416 affinis, Gonicoelus 447 africanus, Biphyllus 417 africanus, Diphyllus 417 alluaudi, Biphyllus 417 Alluaudi, Diphyllus 417 Althaesia 412, 413 amabilis, Biphyllus 417 amabilis, Diphyllus 417 amplicollis, Diplocoelus 435 Anchorius 414, 415 andrewesi, Biphyllus 417 Andrewesi, Diphyllus 417 angusticollis, Diplocoelus 436, 437 angustulus, Diplocoelus 435 angustus, Gonicoelus 447 Anobocoelus 414, 415 apicicollis, Diplocoelus 436 arcanus, Anchorius 414 arcanus, Anobocoelus 414 arduus, Gonicoelus 447 arrowi, Althaesia 413 Arrowi, Althaesia 413 atomus, Diplocoelus 436 bicolor, Diplocoelus 436 bimargo, Gonicoelus 447 Biphyllidae, 410, 411, 441, 447, 452 Biphyllus 411, 415, 421–424, 427–434, 452 bolivari, Biphyllus 417 Bolivari, Diphyllus 417 bombycinus, Diplocoelus 436 brevicollis, Gonicoelus 447 brevipes, Euderopus 445 brevis, Biphyllus 417 brevis, Diphyllus 417 brunnescens, Biphyllus 417 brunnescens, Thallestus 417 brunneus, Diplocoelus 436, 437, 443 canaliculatus, Biphyllus 418 canaliculatus, Diphyllus 418 cardoni, Biphyllus 418 Cardoni, Diphyllus 418 celatus, Gonicoelus 447 centromaculatus, Biphyllus 418 centromaculatus, Diphyllus 418 chaetophorus, Euderopus 445 championi, Anchorius 414 championi, Anobocoelus 414 championi, Gonicoelus 447 chontalenus, Gonicoelus 448 clavatus, Biphyllus 418 complexus, Biphyllus 418 concolor, Biphyllus 418 concolor, Diphyllus 418 concolor, Gonicoelus 448

consobrinus, Diplocoelus 437 consobrinus, Diplocoelus (Marginus) 437 convexus, Biphyllus 418 convexus, Thallestus 418 corpulentus, Biphyllus 419 costulatus, Anchorius 414 costulatus, Diplocoelus 414 crispatus, Gonicoelus 448 Cucujoidea 410 cultratus, Gonicoelus 448 decemlineatus, Diplocoelus 437 decoratus, Biphyllus 419 decoratus, Diphyllus 419 deplanatus, Gonicoelus 447, 448 dilataticollis, Diplocoelus 437 Diphyllidae 410, 411 Diphyllus 411, 415–418, 420–422, 424–435, 438, 440, 452 Diplocaelus 438, 440 Diplocoelus 413, 414, 435, 438, 439, 441, 443, 452 distinctus, Biphyllus 419 distinctus, Diphyllus 419 dohrni, Biphyllus 419 Dohrni, Thallestus 419 dollmani, Biphyllus 419 egens, Biphyllus 419 egens, Diphyllus 419 elegans, Biphyllus 419 escalerae, Biphyllus 420 Escalerae, Diphyllus 420 Euderopus 445 euphorbiae, Biphyllus 420 euphorbiae, Diphyllus 420 Eurhanius 446 exiguus, Diplocoelus 438 fagi, Diplocoelus 421, 438 fagi, Mycetophagus (Triphyllus) 438 fagi, Triphyllus 434, 438 Fagi, Triphyllus 438 fasciatus, Diplocoelus 439, 440 fasciatus, Triphyllus 439 fastidiosus, Biphyllus 420 fastidiosus, Diphyllus 420 flavonotatus, Biphyllus 420 flavonotatus, Diphyllus 420 flexiosus, Biphyllus 420 flexuosus, Biphyllus 420 flexuosus, Diphyllus 420 formosianus, Biphyllus 421 formosianus, Diphyllus 421 foveolatus, Diplocoelus 440 Fraater, Biphyllus 421 frater, Biphyllus 421 frater, Diphyllus 421 frequens, Biphyllus 421 frequens, Diphyllus 421 fulvus, Biphyllus 422 fulvus, Diphyllus 422 germanus, Gonicoelus 448 Gonicoelus 446

CATALOGUE OF BIPHYLLIDAE (COLEOPTERA) OF THE WORLD

grandis, Diplocoelus 440 guatemalenus, Gonicoelus 449 haagi, Diplocoelus 440 Haagi, Diplocoelus 440 hildebrandti, Biphyllus 422 Hildebrandti, Diphyllus 422 hirtus, Gonicoelus 449 histrio, Biphyllus 422 histrio, Diphyllus 422 humeralis, Biphyllus 422 humeralis, Diphyllus 422 humeralis, Eurhanius 446 humerosus, Diplocoelus 440 humilis, Gonicoelus 449 hystrix, Gonicoelus 449 inaequalis, Biphyllus 422 inaequalis, Diphyllus 422 indicus, Diplocoelus 440 infans, Biphyllus 422 infans, Diphyllus 422 inops, Biphyllus 423 inops, Diphyllus 423 inornatus, Diphyllus lunatus var. 425 insignis, Biphyllus 423 insignis, Diphyllus 423 iteratus, Euderopus 445 japonicus, Biphyllus 423 kasuganus, Biphyllus 423 kolosovi, Biphyllus 423 kuzurius, Biphyllus 423 lanuginosus, Biphyllus 423 lanuginosus, Diphyllus 423 laticollis, Gonicoelus 449 latipes, Biphyllus 424 latipes, Diphyllus 424 latus, Diplocoelus 441 latus, Gonicoelus 449 leai, Althaesia 413 Leai, Diplocoelus 413 lewisi, Biphyllus 424 Lewisi, Diphyllus 424 Lewisii, Diphyllus 424 liliputanus, Biphyllus 424 liliputanus, Thallestus 424 lineatus, Anchorius 414, 415 longicornis, Gonicoelus 449 loochooanus, Biphyllus 424 lunata, Dermestes 410, 415, 424 lunatus, Biphyllus 410, 424, 425 lunulatus, Biphyllus 426 maculatus, Biphyllus 426 maculatus, Diphyllus 426 madagascariensis, Biphyllus 427 madagascariensis, Triphyllus 427 magnus, Biphyllus 427 magnus, Diphyllus 427 maindroni, Biphyllus 427 Maindroni, Diphyllus 427 Marginus 434, 443 marmoratus, Biphyllus 427 marmoratus, Diphyllus 427

marshalli, Biphyllus 427 Marshalli, Diphyllus 427 mauritii, Diplocoelus 441 Mauritii, Diplocoelus 441 maxima, Althaesia 413 maximus, Diplocoelus 413 mediocris, Gonicoelus 449 medius, Biphyllus 428 medius, Diphyllus 428 mexicanus, Gonicoelus 450 microps, Euderopus 445 micros, Biphyllus 428 micros, Diphyllus 428 minimus, Biphyllus 428 minimus, Diphyllus 428 minutus, Biphyllus 416, 428 minutus, Diphyllus 428 molestus, Biphyllus 428 molestus, Diphyllus 428 mollis, Gonicoelus 450 monticola, Gonicoelus 450 mus, Diplocoelus 441 muticus, Gonicoelus 450 Mycetophagus 438 nornatus, Biphyllus 426 notatus, Diphyllus parvulus var. 434 oblongus, Diplocoelus 441, 444 oblongus, Diplocoelus (Marginus) 444 oblongus, Philophlaeus 441 oblongus, Selenoderus 444 obscuricollis, Thallestus typhaeoides var. 433 obscuronotatus, Biphyllus 428 obscuronotatus, Diphyllus 428 obscurus, Biphyllus 428 obscurus, Thallestus 428 odiosus, Biphyllus 429 odiosus, Diphyllus 429 oopsis, Gonicoelus 450 opacior, Diplocoelus 442 optatus, Anchorius 415 optatus, Anobocoelus 414, 415 ornatellus, Biphyllus 429 ornatellus, Diphyllus 429 oshimanus, Biphyllus 429 ovatus, Diplocoelus 442, 443 parnoides, Diplocoelus 442 parnoides, Gonicoelus 450 parvulus, Biphyllus 429, 434 parvulus, Diphyllus 429 parvus, Diplocoelus 442 perbrevis, Euderopus 445 philothermoides, Diplocoelus 443 Pilicollis, Triphyllus 438 piliger, Diplocoelus 442 pilinotatus, Diplocoelus 442 pilosa, Althaesia 412, 413 planus, Gonicoelus 450 platysomus, Diplocoelus 443 probiphyllus, Diplocoelus 452 punctatus, Diplocoelus 443 regularis, Euderopus 445

469

470

relictus, Gonicoelus 451 rosti, Biphyllus 429 Rosti, Diphyllus 429 rudis, Diplocoelus 443 rudis, Gonicoelus 451 rudis, Marginus 434, 443 rufopictus, Biphyllus 429, 432 rufopictus, Thallestus 429 satsumanus, Biphyllus 430 sauteri, Biphyllus 430 Sauteri, Diphyllus 430 schenklingi, Biphyllus 430 Schenklingi, Diphyllus 430 securiger, Gonicoelus 451 segnis, Gonicoelus 451 sellatus, Gonicoelus 451 semifuscus, Biphyllus 430 semifuscus, Diphyllus 430 sericea, Althaesia 413 sericeus, Diplocoelus 413 serratus, Triphyllus 438 Serratus, Triphyllus 438 setosus, Euderopus 446 sicardi, Biphyllus 430 Sicardi, Diphyllus 430 similis, Diplocoelus 444 similis, Diplocoelus (Marginus) 444 sjostedti, Biphyllus 431 Sjöstedti, Diphyllus 431 sordidus, Biphyllus 431 sordidus, Diphyllus 431 Sphaeriae, Biphyllus 425 Sphaeriophagus 415 spheniscus, Gonicoelus 451 subellipticus, Biphyllus 431 subellipticus, Thallestus 415, 431 subfasciatus, Biphyllus 431 subfasciatus, Thallestus 431 subjectus, Diplocoelus 444 substriatus, Biphyllus 431 substriatus, Diphyllus 431 subtilis, Gonicoelus 451 suffusus, Biphyllus 431 suffusus, Thallestus 431 talyshensis, Diplocoelus 444 tenuis, Biphyllus 432 tenuis, Diphyllus 432 tessellatus, Diplocoelus 441 Thallestus 415, 416, 418, 419, 424, 429–434 throscoides, Biphyllus 432 throscoides, Gonicoelus 452 throscoides, Thallestus 432 tricornis, Gonicoelus 452 Triphyllus 427, 438, 440 turbinatus, Diplocoelus 444 turneri, Biphyllus 432 typhaeoides, Biphyllus 433 typhaeoides, Thallestus 433 typhoeoides, Diphyllus 433 undulatus, Biphyllus 433 undulatus, Diphyllus 433

P. WĘGRZYNOWICZ

unicolor, Euderopus 446 unicornis, Gonicoelus 446, 452 uniformis, Biphyllus 433 uniformis, Diphyllus 433 ussuriensis, Biphyllus 433 v notatus, Diphyllus 433 v-notatus, Biphyllus 433, 434 variegatus, Biphyllus 434 variegatus, Diphyllus 434 vestitus, Gonicoelus 452 villosus, Diplocoelus 444 weisei, Biphyllus 434 Weisei, Diphyllus 434 wollastoni, Biphyllus 434 Wollastoni, Phallestus 434 xanthorrhoeae, Diplocoelus 437, 444

Insecta aeneus, Rhinosimus 439 Anobiidae 414 Anobium 414 Apidae 447 bicolor, Triphyllus 439 Braconidae 426 Byturidae 410 Cerylon 426 Cerylonidae 426, 443 Cleridae 426 Coleoptera 420, 421, 426, 432, 433, 442, 443, 450, 452 Colydiidae 421 colydioides, Corticeus 420 crenatus, Hylesinus 426 Cryptarcha 439

Cryptophagidae 410 Cucujoidea 410 Curculionidae 421, 433 Dascillidae 452 Dermestes 425 Dryopidae 442, 450 Dryops 442, 450 ephippiatus, Bombus 447 Epurea 439 Erotylidae 410, 441 Europs 420 euryptera, Atheta 439 formicarius, Thanasimus 426 heeri, Triphyllus 452 hirtus, Colobicus 421 humerale, Siagonium 426 Hymenoptera 426, 447 Lepidoptera 426 Loberoschema 441 mediolanensis, Synchita 421 Meteorus 426 Monotomidae 420 Mycetophagidae 410, 439 Nitidulidae 439 orientalis, Xyleborus 420 Parnus 442, 450

CATALOGUE OF BIPHYLLIDAE (COLEOPTERA) OF THE WORLD

Philophlaeus 441 Philothermus 443 Psychidae 426 Salpingidae 439 Scolytidae 426 separanda, Synchita 421 Soronia 439 Staphylinidae 426, 439 Tenebrionidae 420 Thamiaraea 439 Throscidae 432, 452 Throscus 432, 452 Trixagus 432, 452 tycon, Scolytoplatypus 433 undatus, Cicones 421 vexator, Meteorus 426

Fungi and plants Acer 433, 437, 439 Aceraceae 433, 437, 439 actinophyllum, Argyrodendron 416 Aesculus 421 aliena, Quercus 430, 432 Alnus 426 Althaea 412 Altingiaceae 443 angustifolia, Araucaria 435 angustifolia, Fraxinus 439 Araliaceae 439 Araucariaceae 435 Ascomycota 415, 433, 437, 439 Asparagaceae 448 beaumierana, Euphorbia 420 Betula 420, 426 Betulaceae 420, 426 canariensis, Euphorbia 431, 433 Carya 443 cerris, Quercus 421, 439 cinnabarina, Nectria 439 commixta, Sorbus 432 concentrica, Daldinia 426 confluens, Tubercularia 439 corticale, Cryptostroma 426, 439 diplocoeli, Cucujomyces 435 Eucalyptus 429, 436, 442 Euphorbia 420, 431, 433 Euphorbiacae 420, 431, 433 excelsior, Fraxinus 426 Fabaceae 415 Fagaceae 420, 426, 430, 432, 437, 439, 443 faginea, Quercus 439 Fagus 426, 437 fomentarius, Fomes 432, 439

fraxinea, Sphaeria 426 Fraxinus 426 fuscum, Hypoxylon 439 gonicoeli, Cucujomyces 447 helix, Hedera 439 humilis, Quercus 421 Hypoxylon 426, 437 johnsoni, Xanthorrhoea 444 Juglandaceae 443 Laboulbeniaceae 435, 447 Laboulbeniales 435, 447 Lauraceae 426 Laurus 426 Lichenes 433 maacki, Prunus 420 Malvaceae 439 melanoxylon, Acacia 438 Myrtaceae 429, 436, 442 Nectriaceae 439 nigra, Quercus 443 obliqua, Eucalyptus 436 officinalis, Althaea 412 officinarum, Euphorbia 420 Oleaceae 426, 439 orientalis, Platanus 426 Picea 439 Pinaceae 429, 439, 443 Pinus 439, 443 Platanaceae 426, 439 Platanus 439 Polyporaceae 432, 439 Populus 439 Prosopis 415 pseudoplatanus, Acer 426, 439 pubescens, Quercus 421 pyrenaica, Quercus 439 Quercus 421, 426, 437, 439, 443 Rosaceae 432 Salicaceae 439 Sapindaceae 421 silvatica, Fagus 439 Sphaeria 415 styraciflua, Liquidambar 443 suber, Quercus 439 taeda, Pinus 429, 443 Tilia 439 Typha 433 Typhaceae 433 Ulmaceae 439 Ulmus 420, 439 Xanthorrhoea 412, 442, 444 Xanthorrhoeaceae 412, 442, 444 Xylariaceae 426, 437, 439 Yucca 448

471