Abstract
Ulodidae is a small family of saproxylic and fungus feeding beetles restricted to New Zealand, Australia, Chile and New Caledonia. The phylogeny of this family is presented for the first time, based on a cladistic analysis of 53 adult characters from 16 ulodid genera, rooted with Parahelops Waterhouse (Promecheilidae). The topology shows Arthopus Sharp at the base of the tree and confirms the placement of Meryx Latreille as a member of Ulodidae and closely related to the Chilean genus Trachyderas Philippi & Philippi. The extinct New Zealand genus Waitomophylax Leschen & Rhode was placed among a clade consisting of Brouniphylax Strand, Exohadrus Broun, and Pteroderes Germain. Two new genera and two new species are described: Ulobostrichus gen. n. (type species: Ulobostrichus monteithi sp. n.) and Ulocyphaleus gen. n. (type species: Cyphaleus valdivianus Philippi & Philippi, 1864, now U. valdivianus (Philippi & Philippi) n. comb.; U. laetus sp. n.). Dipsaconia pyritosa Pascoe is designated as the type species of Dipsaconia Pascoe and a lectotype was designated for C. valdivianus. A fully illustrated key to the genera and a checklist of the 16 genera and 42 species is included. Based on the phylogeny, the following characters are derived in the family: tuberculate body surface and the presence of scales and /or encrustations. The presence of pore-fields in the abdominal cuticle has evolved at least three times in Meryx Latreille (Australia), Syrphetodes Pascoe (New Zealand) and Trachyderastes Kaszab (New Caledonia).
References
Beutel, R.G. & Friedrich, F. (2005) Comparative study of larvae of Tenebrionoidea (Coleoptera: Cucujiformia). European Journal of Entomology, 102 (2), 241–264.
http://dx.doi.org/10.14411/eje.2005.037Blackburn, T. (1891) Further notes on Australian Coleoptera, with descriptions of new species, ix. Transactions of the Royal Society of South Australia, 14, 65–153.
Blackburn, T. (1902) Further notes on Australian Coleoptera, with descriptions of new genera and species. Part XXXI. Transactions of the Royal Society of South Australia, 26, 288–321.
http://dx.doi.org/10.5479/si.03629236.185.3Blackwelder, R.E. (1945) Checklist of the coleopterous insects of Mexico, Central America, the West Indies, and South America. Bulletin of the United States National Museum, 185 (part 3), 343–550.
Broun, T. (1880) Manual of the New Zealand Coleoptera, Part I. Government Printer, Wellington.
Broun, T. (1893) Manual of the New Zealand Coleoptera, Parts V–VII. Government Printer, Wellington.
Broun, T. (1895) Descriptions of new Coleoptera from New Zealand. The Annals and Magazine of Natural History, (6) 15 (1895), 67–88, 194–203, 234–245, 405–419.
http://dx.doi.org/10.1080/00222939508677849Broun, T. (1903) Descriptions of new genera and species of New Zealand Coleoptera. Annals and Magazine of Natural History (series 7), 12, 69–86.
http://dx.doi.org/10.1080/00222930308678830Broun, T. (1904) Descriptions of new genera and species of New Zealand Coleoptera. Annals and Magazine of Natural History (series 7), 14, 41–59.
http://dx.doi.org/10.1080/03745480409442977Broun, T. (1912) Descriptions of new genera and species of New Zealand Coleoptera. Transactions and Proceedings of the New Zealand Institute, 44 (1911), 379–440.
Broun, T. (1914) Descriptions of new genera and species of Coleoptera. Part III. Bulletin of the New Zealand Institute, 1, 143–266.
Broun, T. (1917) Descriptions of new genera and species of New Zealand Coleoptera. Bulletin of the New Zealand Institute, 1 (5), 347–472.
Broun, T. (1921) Descriptions of new genera and species of New Zealand Coleoptera. Bulletin of the New Zealand Institute, 1 (6), 475–590.
Carter, H.J. (1926a) Revision of the Australasian species of Anilara (Fam. Buprestidae) and Helmis (Fam. Dryopidae), with notes, and descriptions of other Australian Coleoptera. Proceedings of the Linnean Society of New South Wales, 51, 50–71.
Carter, H.J. (1926b) Revision of Athemistus and Microtragus (Fam. Cerambycidae) with notes, and descriptions of other Australian Coleoptera. Proceedings of the Linnaean Society of New South Wales, 51 (4), 492–516.
Carter, H.J. (1926c) A check list of the Australian Tenebrionidae. The Australian Zoologist, 4 (3), 117–63
Carter, H.J. (1928) Revision of the Australian species of the genera Curis, Neocuris and Trachys, together with notes and descriptions of new species of other Coleoptera. Proceedings of the Linnaean Society of New South Wales, 53 (3), 270–290.
Carter, H.J. (1937) Some new Tenebrionidae in the South Australian Museum, together with notes and descriptions of other Australian Coleoptera. Transactions and Proceedings of the Royal Society of South Australia, 61, 121–144.
Carter, H.J. & Zeck, E.H. (1937) A monograph of the Australian Colydiidae. Proceedings of the Linnean Society of New South Wales, 62, 181–208, pls. VIII–IX.
Champion, G.C. (1894) On the Tenebrionidae collected in Australia and Tasmania by Mr James J. Walker, R.N., F.L.S., during the voyage of H.M.S. “Penguin” with descriptions of new genera and species. Transactions of the Entomological Society of London, 35, 351–408, pl. VIII.
Crowson, R.A. (1953) The classification of the families of British Coleoptera [continued]. The Entomologist’s Monthly Magazine, 89, 37–59.
Doyen, J.T. & Lawrence, J.F. (1979) Relationships and higher classification of some Tenebrionidae and Zopheridae (Coleoptera). Systematic Entomology, 4, 333–377.
http://dx.doi.org/10.1111/j.1365-3113.1979.tb00619.xErichson, W.F. (1842) Beitrag zur Insecten-Fauna von Vandiemensland, mit besonderer Berück- sichtigung der geographischen Verbreitung der Insecten. Archiv für Naturgeschichte, 8 (1), 83–287, pls. 4–5.
Fairmaire, L. & Germain, P. (1860) Coleoptera Chilensia. Typographie F. Malteste et cie. Part 2, Section 1, 8 pp.
Förster, A. (1869) Synopsis der Familien und Gattungen der Ichneumonen. Verhandlungen des Naturhistorischen Vereins der Preussischen Rheinlande und Westfalens, 25 (1868), 135–221.
Gebien, H. (1911) Coleopterorum Catalogus Pars 28. Tenebrionidae III. Pp. 355–585. W. Junk, Berlin.
Gebien, H. (1939) Katalog der Tenebrioniden. Teil II (part). Mitteilungen der Münchener Entomologischen Gesellschaft, 29, 443–474 (466–497), 739–770 (498–529).
Germain, P. (1894) Apuntes sobre los insectos de Chile. Anales de la Universidad de Chile, 87, 445–457, 1 plate.
Gerstaecker, A. (1862) Bericht über die wissenshaftlichen Leistungen im Gebiete der Entomologie wahrend des Jahres 1861. Archiv für Naturgeschichte, 28 (2), 273–571.
Gunter, N. L., Levkaničová, Z., Weir, T. H., Ślipiński, A., Cameron, S.L. & Bocak, L. (2014) Towards a phylogeny of the Tenebrionoidea (Coleoptera). Molecular Phylogenetics and Evolution, 79, 305–312.
http://dx.doi.org/10.1016/j.ympev.2014.05.028Hope, F.W. (1843) Continuation of a memoir containing descriptions of new species of Coleoptera from Port Essington, in New Holland. Annals and Magazine of Natural History, 12, 357–361.
Hudson, G.V. (1934) New Zealand Beetles and Their Larvae. Ferguson & Osborn: Wellington. 236 pp +17 pls.
Kaszab, Z. (1982) Die Tenebrioniden Neukaledoniens und der Loyauté-Inseln (Coleoptera). Folia Entomologica Hungarica, 28, 1–294.
Kaszab, Z. (1986) Tenebrioniden (Coleoptera) aus Neukaledonien. Annales Historico-Naturales Musei Nationalis Hungarici, 78, 151–175.
Kulzer, H. (1964) Über neue Tenebrionidenarten (Col.). 27. Beitrag zur Kenntnis der Tenebrioniden. Entomologische Arbeiten der Museum Georg Frey, 15, 221–276.
Latreille, P.A. (1802) Histoire naturelle, générale et particulière des crustacés et des insectes. Ouvrage faisant suite à l’histoire naturelle générale et particulière, composée par Leclerc de Buffon, et rédigée par C.S. Sonnini, membre de plusieurs sociétés savantes. Familles naturelles des genres. Tome troisième. F. Dufart, Paris, xii + 13–467 + [1] pp.
Lawrence, J.F. (1991) Zopheridae (Tenebrionoidea). In: Stehr, F.W. (Ed.), Immature Insects. Vol. 2. Kendall/Hunt Publishing Co., Dubuque, Iowa. pp. 518–519
Lawrence, J.F. (1994) The larva of Sirrhas variegata, sp. nov., with notes on the Perimylopidae, Ulodidae (stat. nov.), Zopheridae and Chalcodryidae (Coleoptera: Tenebrionoidea). Invertebrate Taxonomy, 8, 329–349.
http://dx.doi.org/10.1071/IT9940329Lawrence, J.F. & Britton, E.B. (1991) Coleoptera (Beetles). In: CSIRO Division of Entomology (Ed.), Insects of Australia: a Textbook for Students and Research Workers, Second Edition. Vol. 2. Melbourne University Press, Carlton, Victoria. pp. 543–683.
Lawrence, J.F., Escalona, H.E., Leschen, R.A.B. & Ślipiński, S.A. (2014) Review of the genera of Mycetophagidae (Coleoptera: Tenebrionoidea) with descriptions of new genera and a world generic key. Zootaxa, 3826 (1), 195–229.
http://dx.doi.org/10.11646/zootaxa.3826.1.6Lawrence, J.F., Hastings, A.M., Dallwitz, M.J., Paine, T.A. & Zurcher, E.J. (1999a) “Beetle Larvae of the World: Descriptions, Illustrations, Identification, and Information Retrieval for Families and Subfamilies.” CD-ROM, Version 1.1 for MS-Windows. CSIRO Publishing, Melbourne.
Lawrence, J.F., Hastings, A.M., Dallwitz, M.J., Paine, T.A. & Zurcher, E.J. (1999b) “Beetles of the World: A Key and Information System for Families and Subfamilies.” CD-ROM, Version 1.0 for MS-Windows. CSIRO Publishing, Melbourne.
Lawrence, J.F. & Hlavac, T.F. (1979) Review of the Derodontidae (Coleoptera: Polyphaga) with new species from North America and Chile. The Coleopterists Bulletin, 33, 369–414.
Lawrence, J.F. & Newton, A.F. Jr. (1995) Families and subfamilies of Coleoptera (with selected genera, notes and references, and data on family-group names). In: Pakaluk, J. & Ślipiński, S.A. (Eds.), Biology, Phylogeny, and Classification of Coleoptera: Papers Celebrating the 80th Birthday of Roy A. Crowson. Muzeum i Instytut Zoologii PAN, Warszawa, pp. 779–1006.
Lawrence, J.F., Ślipiński, S.A. & Elgueta, D.M. (2010) Promecheilidae Lacordaire 1859. In: Part 38. Coleoptera, beetles. Volume 2: Morphology and systematics (Elateroidea, Bostrichiformia, Cucujiformia partim). In: Leschen R.A.B., Beutel R.G., Lawrence J.F., Ślipiński S.A. (Eds) Handbook of zoology. Volume IV. Arthropoda: Insecta. W. de Gruyter, New York and Berlin, pp. 563–567.
Lawrence, J.F., Ślipiński, A., Seago, A.E., Thayer, M.K., Newton, A.F. & Marvaldi, A.E. (2011) Phylogeny of the Coleoptera based on morphological characters of adults and larvae. Annales Zoologici, 61 (1), 1–217.
http://dx.doi.org/10.3161/000345411X576725Leschen, R.A.B. 2003. Erotylidae (Insecta: Coleoptera: Cucujoidea): phylogeny and review. Fauna of New Zealand, 47, 1–108.
Leschen, R.A.B. & Buckley, R.T. (2015) Revision and phylogeny of Syrphetodes (Coleoptera: Ulodidae): implications for biogeography, alpinization and conservation. Systematic Entomology, 40, 143–168.
http://dx.doi.org/10.1111/syen.12094Leschen, R.A.B., Marris, J.W.M., Emberson, R.M., Nunn, J., Hitchmough, R.A. & Stringer, I.A.N. (2012) The conservation status of New Zealand Coleoptera. New Zealand Entomologist, 35, 91–98.
http://dx.doi.org/10.1080/00779962.2012.686311Leschen, R.A.B. & Rhode, B.E. (2002) A new genus and species of large extinct Ulodidae (Coleoptera) from New Zealand. New Zealand Entomologist, 25, 57–64.
http://dx.doi.org/10.1080/00779962.2002.9722095Leschen, R.A.B. & Rhode, B.E. (2005) A replacement name for Archaeophylax Leschen and Rhode (Coleoptera: Ulodidae). New Zealand Entomologist, 27, 125.
http://dx.doi.org/10.1080/00779962.2004.9722134Leschen, R.A.B. & Ślipiński, S.A. (2010) Chapter 11.10. Ulodidae Pascoe, 1869. In: Leschen, R.A.B., Beutel, R.G. & Lawrence, J.F. (Eds.), Handbook of Zoology, Coleoptera Volume 2: Morphology and Systematics (Elateroidea, Bostrichformia, Cucujiformia partim). Walter de Gruyter, Berlin, pp. 559–563.
Maddison, W.P. & Maddison, D.R. (2011) Mesquite: a modular system for evolutionary analysis. Version 2.75. Avaliable from: http://mesquiteproject.org (Accessed 14 Jul. 2016)
Matthews, E.G., Lawrence, J.F., Bouchard, P., Steiner, W.E. & Ślipiński, S.A. (2010) Tenebrionidae Latreille, 1802 [pp. 574–659]. In: Leschen RAB, Beutel RG, Lawrence JF (Eds.), Hand book of Zoology. Coleoptera, Beetles. Volume 2: Morphology and systematics (Elateroidea, Bostrichiformia partim). De Gruyter, Berlin, New York, xiii + 786 pp.
McKenna, D.D., Wild, A.L., Kanda, K., Bellamy, C.L., Beutel, R.G., Caterino, M.S., Farnum, C.W., Hawk, D.C., Ivie, M.A., Jameson, M.L., Leschen, R.A.B., Marvaldi, A.E., McHugh, J.V., Newton, A.F., Robertson, A.A., Thayer, M.K., Whiting, M.F., Lawrence, J.F., Ślipiński, A., Maddison, D.R. & Farrel, B.D. (2015) The beetle tree of life reveals Coleoptera survived end Permian mass extinction to diversify during the Cretaceous terrestrial revolution. Systematic Entomology, 40, 835–880.
http://dx.doi.org/10.1111/syen.12132Mosely, M.E. & Kimmins, D.E. (1953) The Trichoptera (caddis-flies) of Australia and New Zealand. British Museum (Natural History), London. 550 pp.
Newman, E. (1842) List of insects collected at Port Philipp, South Australia, by Edmund Thomas Higgins, Esq. Entomologist, London 1 (25), 401–405.
Pascoe, F.P. (1860) Notices of new or little-known genera and species of Coleoptera. Journal of Entomology, 1 (2), 98–132.
Pascoe, F.P. (1862) Notices of new or little-known genera and species of Coleoptera. Part II. Journal of Entomology, 1, 98–132, plates V–VIII.
Pascoe, F.P. (1869) Descriptions of new genera and species of Tenebrionidae from Australia and Tasmania. Annals and Magazine of Natural History, (4) 3, 29–45, plate X.
Pascoe, F.P. (1875) Descriptions of new genera and species of New Zealand Coleoptera. -Part I. Annals and Magazine of Natural History, (4) 16 (93), 210–223, plate V.
Peña G., L.E. (1966a) Catalogo de los Tenebrionidae (Coleoptera) de Chile. Entomologische Arbeiten aus dem Museum G. Frey, 17, 397–453.
Peña G., L.E. (1966b) A preliminary attempt to divide Chile into entomofaunal regions, based on the Tenebrionidae (Coleoptera). Postilla, 97, 1–17.
Philippi, R.A. & Philippi, F.H.E. (1864) Beschreibung einiger neuen Chilenischen Käfer. Stettiner Entomologische Zeitung, 25 (10–12), 313–406.
Sharp, D. (1876) Descriptions of some new genera and species of New Zealand Coleoptera. Entomologist's Monthly Magazine, 13, 20–28, 70–77.
http://dx.doi.org/10.5962/bhl.part.22816Sharp, D. (1886) On New Zealand Coleoptera with descriptions of new genera and species. Transactions of the Royal Dublin Society, 2, 351454 + pls. xii & xiii.
Ślipiński, S.A., Leschen, R.A.B. & Lawrence, J.F. (2011) Order Coleoptera Linnaeus, 1758. In: Zhang, Z.-Q. (Ed.), Animal biodiversity: An outline of higher-level classification and survey of taxonomic richness. Zootaxa, 3148, 203–208.
Strand, E. (1943) Miscellanea nomenclatorica zoologica et paleontologica. Folia zoologica et hydrobiologica, 12, 94–114.
Swofford, D.L. (2002) PAUP*: Phylogenetic analysis using parsimony (* and other methods). Version 4. Sinauer Associates, Sunderland, Massachusetts.
Watt, J.C. (1974) A revised subfamily classification of Tenebrionidae (Coleoptera). New Zealand Journal of Zoology, 1, 381–452.
http://dx.doi.org/10.1080/03014223.1974.9517846Watt, J.C. (1982) 1981 presidential address. New Zealand beetles. New Zealand Entomologist, 7, 213–221.
http://dx.doi.org/10.1080/00779962.1982.9722384