A Re-evaluation of the Eucnemidae larval characters (Coleoptera)

Muona, Jyrki; Teräväinen, Marianna

doi:10.11606/1807-0205/2020.60.special-issue.28

Jyrki Muona ²2E-mail: jyrki.muona@helsinki.fi (corresponding author)

Finnish Museum of Natural History, Zoological Museum. Helsinki, Finland

http://orcid.org/0000-0003-2771-1171

Marianna Teräväinen ³3E-mail: marianna.teravainen@helsinki.fi

Finnish Museum of Natural History, Zoological Museum. Helsinki, Finland

http://orcid.org/0000-0002-4624-011X

²E-mail: jyrki.muona@helsinki.fi (corresponding author)

³E-mail: marianna.teravainen@helsinki.fi

Edited by: Sônia A. Casari / Gabriel Biffi

EUCNEMIDAE
	Perothopinae		Perothops sp. [*Lawrence et al.,* 2007**Lawrence, J.F.; Muona, J.; Teräväinen, M.; Ståhls, G. & Vahtera, V. 2007. Anischia, Perothops and the phylogeny of Elateroidea (Coleoptera: Elateriformia). Insect Systematics & Evolution, 38: 205-239. DOI]
	Phyllocerinae		Phyllocerus sp. [Gilyarov, 1979Gilyarov, M.S. 1979. A larva of Phyllocerus and its position in the system of Elateroidea. (in Russian). Zoologicheskii Zhurnal, 58: 655-663.; Klausnitzer, 1997Klausnitzer, B. 1997. Zur Kenntnis der Larve von Phyllocerus Lepeletier & Serville (Coleoptera, Eucnemidae, Phyllocerinae). Berichte des Naturwissenschaftlich-medizinischer Verein Innsbruck, 84: 371-378.]
	Pseudomeninae		Pseudomenes bakewelli (Bonvouloir)*, Australia, ACT
	Schizophilinae		Schizophilus subrufus (Randall)* [Otto & Young, 1998Otto, R. & Young, D. 1998. Description of the Larva of Schizophilus subrufus (Randall) (Coleoptera: Eucnemidae: Pseudomeninae), with Notes on Its Natural History. The Coleopterists Bulletin, 52: 306-312.]
	Palaeoxeninae		Palaeoxenus dohrni (Horn)* [Böving & Craighead, 1931Böving, A.G. & Craighead, F.C. 1931. An illustrated synopsis of the principal larval forms of the order Coleoptera. Entomologica Americana, 11: 1-351.; *Chang et al.,* 2016**Chang, H.; Muona, J.; Pu, H.; Xu, L.; Wang, C.; Teräväinen, M.; Ren, D.; Yang, Q.; Zhang, X. & Jia, S. 2016. Chinese Cretaceous larva exposes a southern Californian living fossil (Insecta, Coleoptera, Eucnemidae). Cladistics, 32(2): 211-214. DOI]
	Anischiinae		Anischia kuscheli*Lawrence [Lawrence et al., 2007**Lawrence, J.F.; Muona, J.; Teräväinen, M.; Ståhls, G. & Vahtera, V. 2007. Anischia, Perothops and the phylogeny of Elateroidea (Coleoptera: Elateriformia). Insect Systematics & Evolution, 38: 205-239. DOI]
	Melasinae
		Melasini	Isorhipis melasoides (Castelnau)* [Mamaev, 1976Mamaev, B.M. 1976. Morphological types of xylophagous beetle larvae (Coleoptera, Eucnemidae) and their evolutionary importance. In: Mamaev, B.M. (Ed.). Evolutionary Morphology of Wood-Boring Larvae. Moscow, Nauka. p. 136-155.]; I. obliqua (Say)* [Van Horn, 1909Van Horn, R.W. 1909. Notes on some of the Eucnemidae of the eastern states. (Coleoptera, Eucnemidae). Proceedings of the entomological Society Washington, 11: 54-62.]; Melasis buprestoides (Fabricius)* [Leiler, 1976Leiler, T.E. 1976. Zur Kenntnis der Entwicklungsstadien und der Lebensweise nord- und mitteleuropäischer Eucnemiden (Col.). Entomologische Blätter für Biologie und Systematik der Käfer, 72: 10-50.]; Melasis pectinicornis Melsheimer*
		Hylocharini	Hylochares cruentatus (Gyllenhal)* [Brüstle & Muona, 2009Brüstle, L. & Muona, J. 2009. Life-history studies versus genetic markers - the case of Hylochares cruentatus (Gyllenhal, 1808) (Col., Eucnemidae). Journal of Zoological Systematics and Evolutionary Research, 42: 100-107. DOI]; H. harmandi Fleutiaux [Mamaev, 1976Mamaev, B.M. 1976. Morphological types of xylophagous beetle larvae (Coleoptera, Eucnemidae) and their evolutionary importance. In: Mamaev, B.M. (Ed.). Evolutionary Morphology of Wood-Boring Larvae. Moscow, Nauka. p. 136-155.]; H. nigricornis (Say)* [Peterson, 1960Peterson, A. 1960. Larvae of Insects. Coleoptera, Diptera, Neuroptera, Siphonaptera, Mecoptera, Trichoptera. Columbus, A. Paterson. part II, 416p.]; H. populi Brüstle & Muona* [Brüstle & Muona, 2009Brüstle, L. & Muona, J. 2009. Life-history studies versus genetic markers - the case of Hylochares cruentatus (Gyllenhal, 1808) (Col., Eucnemidae). Journal of Zoological Systematics and Evolutionary Research, 42: 100-107. DOI]
		Calyptocerini	Otho sphondyloides Germar* [Leiler, 1976Leiler, T.E. 1976. Zur Kenntnis der Entwicklungsstadien und der Lebensweise nord- und mitteleuropäischer Eucnemiden (Col.). Entomologische Blätter für Biologie und Systematik der Käfer, 72: 10-50.]
		Xylobiini	Agalba sp., Australia, Queensland; Proxylobius gardneri* (Fleutiaux) [Gardner, 1935Gardner, J.C.M. 1935. Immature stages of Indian Coleoptera (17) (Eucnemidae). Indian Forest Records (New Series) Entomology, 1: 79-93, plates I-II.]: Xylophilus corticalis (Paykull)* [Leiler, 1976Leiler, T.E. 1976. Zur Kenntnis der Entwicklungsstadien und der Lebensweise nord- und mitteleuropäischer Eucnemiden (Col.). Entomologische Blätter für Biologie und Systematik der Käfer, 72: 10-50.]; X. ussurienisMamaev [Mamaev, 1976Mamaev, B.M. 1976. Morphological types of xylophagous beetle larvae (Coleoptera, Eucnemidae) and their evolutionary importance. In: Mamaev, B.M. (Ed.). Evolutionary Morphology of Wood-Boring Larvae. Moscow, Nauka. p. 136-155.]
		Epiphanini	Hylis cariniceps (Mannerheim)* [Leiler, 1976Leiler, T.E. 1976. Zur Kenntnis der Entwicklungsstadien und der Lebensweise nord- und mitteleuropäischer Eucnemiden (Col.). Entomologische Blätter für Biologie und Systematik der Käfer, 72: 10-50.]; H. foveicollis (Mannerheim) [Leiler, 1976Leiler, T.E. 1976. Zur Kenntnis der Entwicklungsstadien und der Lebensweise nord- und mitteleuropäischer Eucnemiden (Col.). Entomologische Blätter für Biologie und Systematik der Käfer, 72: 10-50.]; H. procerulus (Mannerheim)* [Leiler, 1976Leiler, T.E. 1976. Zur Kenntnis der Entwicklungsstadien und der Lebensweise nord- und mitteleuropäischer Eucnemiden (Col.). Entomologische Blätter für Biologie und Systematik der Käfer, 72: 10-50.]; H. olexai (Mannerheim) [Lucht, 1981Lucht, W. 1981. Die Präimaginalstadien von Hypocoelus olexai Palm (Col., Eucnemidae) nebst Bestimmungstabelle der Larven Nord- und Mitteleuropäischer Hypocoelus -Arten. Entomologische Blätter für Biologi und Systematik der Käfer, 77: 61-74.]
		Dirhagini	Adelothyreus bengalensis Fleutiaux [Gardner, 1935Gardner, J.C.M. 1935. Immature stages of Indian Coleoptera (17) (Eucnemidae). Indian Forest Records (New Series) Entomology, 1: 79-93, plates I-II.]; Arrhipis ?orientalis Fleutiaux [Gardner, 1935Gardner, J.C.M. 1935. Immature stages of Indian Coleoptera (17) (Eucnemidae). Indian Forest Records (New Series) Entomology, 1: 79-93, plates I-II.]; A. balwanti Fleutiaux [Gardner, 1935Gardner, J.C.M. 1935. Immature stages of Indian Coleoptera (17) (Eucnemidae). Indian Forest Records (New Series) Entomology, 1: 79-93, plates I-II.]; Dirrhagofarsus attenuatus (Mäklin)* [Burakowski, 1989Burakowski, B. 1989. Hypermetamorphosis of Rhocopus attenuatus (Mäklin) (Coleoptera, Eucnemidae). Annales Zoologici, Warszawa, 42: 165-180.]; D. lewisi (Fleutiaux)* [Ford & Spilman, 1979Ford, E.J. & Spilman, T.J. 1979. Biology and immature stages of Dirhagofarsus lewisi, a species new to the United States (Coleoptera, Eucnemidae). The Coleopterists Bulletin, 33: 75-83.], D. ernae*Otto, Muona & McClarin [Otto et al., 2014*Otto, R.; Muona, J. & McClarin, J. 2014. Description of Dirrhagofarsus ernae n. sp. with a key to the known Dirrhagofarsus species (Coleoptera: Eucnemidae). Zootaxa, 3878: 179-184.]; Entomophthalmus rufiolus* (LeConte) [Otto, 2012bOtto, R. 2012b. Eucnemid Larvae of the Nearctic Region. Part II: Description of the Mature Larva of Deltometopus amoenicornis (Say, 1836) (Coleoptera: Eucnemidae: Macraulacinae: Macraulacini), with notes on its biology. The Coleopterists Bulletin, 66(3): 285-288.] Farsus dubius (Piller & Mitterpacher) [Nemeth & Otto, 2016Nemeth, T. & Otto, R. 2016. Notes on the bionomics of Farsus dubius (Piller & Mitterpacher, 1783) (Coleoptera: Eucnemidae: Melasinae), with observations on its hypermetamorphic development. Elateridarium, 10: 133-144.; Microrhagus audax Horn [Otto, 2015Otto, R. 2015. Eucnemid larvae of the Nearctic region. Part V: Fifth instar larval descriptions for eight species of Microrhagus Dejean, 1833 (Coleoptera: Eucnemidae: Melasinae: Dirhagini), with descriptions of four new species and notes on their biology. Insecta Mundi, 421: 1-43.]; M. breviangularis Otto [Otto, 2015Otto, R. 2015. Eucnemid larvae of the Nearctic region. Part V: Fifth instar larval descriptions for eight species of Microrhagus Dejean, 1833 (Coleoptera: Eucnemidae: Melasinae: Dirhagini), with descriptions of four new species and notes on their biology. Insecta Mundi, 421: 1-43.]; M. brunneus Otto [Otto, 2015Otto, R. 2015. Eucnemid larvae of the Nearctic region. Part V: Fifth instar larval descriptions for eight species of Microrhagus Dejean, 1833 (Coleoptera: Eucnemidae: Melasinae: Dirhagini), with descriptions of four new species and notes on their biology. Insecta Mundi, 421: 1-43.]; M. carinicollis Otto [Otto, 2015Otto, R. 2015. Eucnemid larvae of the Nearctic region. Part V: Fifth instar larval descriptions for eight species of Microrhagus Dejean, 1833 (Coleoptera: Eucnemidae: Melasinae: Dirhagini), with descriptions of four new species and notes on their biology. Insecta Mundi, 421: 1-43.]; M. emyi Rouget, Sweden; M. lecontei* Otto [Otto, 2015Otto, R. 2015. Eucnemid larvae of the Nearctic region. Part V: Fifth instar larval descriptions for eight species of Microrhagus Dejean, 1833 (Coleoptera: Eucnemidae: Melasinae: Dirhagini), with descriptions of four new species and notes on their biology. Insecta Mundi, 421: 1-43.]; Microrhagus lepidus (Rosenhauer)* [Leiler, 1976Leiler, T.E. 1976. Zur Kenntnis der Entwicklungsstadien und der Lebensweise nord- und mitteleuropäischer Eucnemiden (Col.). Entomologische Blätter für Biologie und Systematik der Käfer, 72: 10-50.]; M. pectinatus LeConte* [Otto, 2015Otto, R. 2015. Eucnemid larvae of the Nearctic region. Part V: Fifth instar larval descriptions for eight species of Microrhagus Dejean, 1833 (Coleoptera: Eucnemidae: Melasinae: Dirhagini), with descriptions of four new species and notes on their biology. Insecta Mundi, 421: 1-43.]; M. pygmaeus (Fabricius)* [Leiler, 1976Leiler, T.E. 1976. Zur Kenntnis der Entwicklungsstadien und der Lebensweise nord- und mitteleuropäischer Eucnemiden (Col.). Entomologische Blätter für Biologie und Systematik der Käfer, 72: 10-50.]; M. subsinuatus LeConte* [Otto, 2015Otto, R. 2015. Eucnemid larvae of the Nearctic region. Part V: Fifth instar larval descriptions for eight species of Microrhagus Dejean, 1833 (Coleoptera: Eucnemidae: Melasinae: Dirhagini), with descriptions of four new species and notes on their biology. Insecta Mundi, 421: 1-43.]; M. triangularis (Say)* [Otto, 2015Otto, R. 2015. Eucnemid larvae of the Nearctic region. Part V: Fifth instar larval descriptions for eight species of Microrhagus Dejean, 1833 (Coleoptera: Eucnemidae: Melasinae: Dirhagini), with descriptions of four new species and notes on their biology. Insecta Mundi, 421: 1-43.]; ?Prodirrhagus cairnesensis (Lea), Australia, Queensland; Rhacopus sahlbergi* (Mannerheim)* [Leiler, 1976Leiler, T.E. 1976. Zur Kenntnis der Entwicklungsstadien und der Lebensweise nord- und mitteleuropäischer Eucnemiden (Col.). Entomologische Blätter für Biologie und Systematik der Käfer, 72: 10-50.]; Rhagomicrus bonvouloiri (Say)* [Otto, 2012aOtto, R. 2012a. Eucnemid larvae of the Nearctic Region. Part I: Description of the larva of Rhagomicrus bonvouloiri (Horn, 1886) (Coleoptera: Eucnemidae: Melasinae: Dirrhagini), with notes on its biology. The Coleopterists Bulletin, 66(3): 219-223.)
	Eucneminae
		Mesogenini	Stethon pectorosus LeConte [Otto & Gruber, 2016Otto, R. & Gruber, J. 2016. Eucnemid larvae of the Nearctic region. Part VI: Descriptions of the fifth instar and prepupal larval stages Stethon pectorosus LeConte, 1866 (Coleoptera: Eucnemidae: Eucneminae: Mesogenini), with notes on their biology. Insecta Mundi, 474: 1-11.]
		Eucnemini	Eucnemis capucinus Ahrens* [Palm, 1955Palm, T. 1955 Weitere Beiträge zur Kenntnis der schwedischen Hypocoelus-Arten (Col. Eucnemidae). Opuscula Entomologica, 20: 211-217.; Lundberg, 1962Lundberg, 1962. Något om svenska eucnemider. Entomologisk Tidsskrift, 83: 89-94.; Leiler, 1976Leiler, T.E. 1976. Zur Kenntnis der Entwicklungsstadien und der Lebensweise nord- und mitteleuropäischer Eucnemiden (Col.). Entomologische Blätter für Biologie und Systematik der Käfer, 72: 10-50.]; E. zaitzevi*Mamaev [Mamaev, 1976*Mamaev, B.M. 1976. Morphological types of xylophagous beetle larvae (Coleoptera, Eucnemidae) and their evolutionary importance. In: Mamaev, B.M. (Ed.). Evolutionary Morphology of Wood-Boring Larvae. Moscow, Nauka. p. 136-155.]; Poecilochrus bengalensis* Fleutiaux [Gardner, 1935Gardner, J.C.M. 1935. Immature stages of Indian Coleoptera (17) (Eucnemidae). Indian Forest Records (New Series) Entomology, 1: 79-93, plates I-II.]
		Galbitini	Arisus bituberculatus Fleutiaux [Gardner, 1935Gardner, J.C.M. 1935. Immature stages of Indian Coleoptera (17) (Eucnemidae). Indian Forest Records (New Series) Entomology, 1: 79-93, plates I-II.]; Galbimorpha agastoceroides Fleutiaux [Gardner, 1935Gardner, J.C.M. 1935. Immature stages of Indian Coleoptera (17) (Eucnemidae). Indian Forest Records (New Series) Entomology, 1: 79-93, plates I-II.]; Galbites fulvus Fleutiaux [Gardner, 1935Gardner, J.C.M. 1935. Immature stages of Indian Coleoptera (17) (Eucnemidae). Indian Forest Records (New Series) Entomology, 1: 79-93, plates I-II.]
	Macraulacinae
		Euryptychini	Euryptychus lewisi Fleutiaux [Mamaev, 1976Mamaev, B.M. 1976. Morphological types of xylophagous beetle larvae (Coleoptera, Eucnemidae) and their evolutionary importance. In: Mamaev, B.M. (Ed.). Evolutionary Morphology of Wood-Boring Larvae. Moscow, Nauka. p. 136-155.]; ?Euryptychus sp.*, Panama
		Macraulacini	Asiocnemis ?mcclayi Muona, USA, California; A. morawitzi* (Semenov) [Mamaev, 1976Mamaev, B.M. 1976. Morphological types of xylophagous beetle larvae (Coleoptera, Eucnemidae) and their evolutionary importance. In: Mamaev, B.M. (Ed.). Evolutionary Morphology of Wood-Boring Larvae. Moscow, Nauka. p. 136-155.]; Deltometopus amoenicornis (Say) [[Otto, 2012Otto, R. 2012b. Eucnemid Larvae of the Nearctic Region. Part II: Description of the Mature Larva of Deltometopus amoenicornis (Say, 1836) (Coleoptera: Eucnemidae: Macraulacinae: Macraulacini), with notes on its biology. The Coleopterists Bulletin, 66(3): 285-288.b]; Dromaeoloides parvulus (Bonvouloir), Australia, NSW, reared; Dromaeoloides* spp., New Caledonia, Fiji, both reared; D. ?badius* (Melsheimer), USA, Idaho; D. barnabita* (Villa)* [Leiler, 1976Leiler, T.E. 1976. Zur Kenntnis der Entwicklungsstadien und der Lebensweise nord- und mitteleuropäischer Eucnemiden (Col.). Entomologische Blätter für Biologie und Systematik der Käfer, 72: 10-50.]; ?Dromaeolus lugubris Lea, Australia, NSW; ?Dromaeolus punctatus* (LeConte), USA, Georgia; Fornax distinguendus* Bonvouloir [Gardner, 1935Gardner, J.C.M. 1935. Immature stages of Indian Coleoptera (17) (Eucnemidae). Indian Forest Records (New Series) Entomology, 1: 79-93, plates I-II.]; F. hisamatsui Mamaev [Mamaev, 1976Mamaev, B.M. 1976. Morphological types of xylophagous beetle larvae (Coleoptera, Eucnemidae) and their evolutionary importance. In: Mamaev, B.M. (Ed.). Evolutionary Morphology of Wood-Boring Larvae. Moscow, Nauka. p. 136-155.]; F. gardneri Fleutiaux [Gardner, 1926Gardner, J.C.M. 1926. (a) Description of the early stages of Fornax gardneri Fleut. (Melasidae, Col.). (b) Description of the larva of Atractocerus emarginatus Cast. (Lymexylonidae, Col.). Indian Forest Records, 12: 273-286.]; F. holmi Leiler [Leiler, 1990Leiler, T.E. 1990. Beschreibung einiger malaysischer Eucnemiden-Arten, ihrer Larven un Lebensweise (Col.). Entomologische Blätter für Biologie und Systematik der Käfer, 86: 145-158.]; F. raja Leiler* [Leiler, 1990Leiler, T.E. 1990. Beschreibung einiger malaysischer Eucnemiden-Arten, ihrer Larven un Lebensweise (Col.). Entomologische Blätter für Biologie und Systematik der Käfer, 86: 145-158.]; F. silvestris Fleutiaux [Gardner, 1935Gardner, J.C.M. 1935. Immature stages of Indian Coleoptera (17) (Eucnemidae). Indian Forest Records (New Series) Entomology, 1: 79-93, plates I-II.]; F. vestitus Fleutiaux [Gardner, 1935Gardner, J.C.M. 1935. Immature stages of Indian Coleoptera (17) (Eucnemidae). Indian Forest Records (New Series) Entomology, 1: 79-93, plates I-II.]; Fornax sp. [*Costa et al.,* 2003*Costa, C.; Vanin, S.A.; Lawrence, J.F. & Ide, S. 2003. Systemtics and cladistics analysis of Cerophytidae (Elateroidea: Coleoptera). Systematic Entomology, 28: 375-407.]; Isarthrus rufipes* (Melsheimer) [Mosher, 1919Mosher, E. 1919. Notes on the immature stages of Deltometopus rufipes Mels. (Coleoptera, Eucnemidae). Annals of the Entomological Society of America, 12: 49-53, plate IV.]; Onichodon canadensis (Brown)* [Otto, 2013Otto, R. 2013. Eucnemid Larvae of the Nearctic Region. Part III: Mature Larval Descriptions for Three Species of Onichodon Newman, 1838 (Coleoptera: Eucnemidae: Macraulacini, with notes on their biology. The Coleopterists Bulletin, 67: 97-106.]; O. orchesides Newman [Otto, 2013Otto, R. 2013. Eucnemid Larvae of the Nearctic Region. Part III: Mature Larval Descriptions for Three Species of Onichodon Newman, 1838 (Coleoptera: Eucnemidae: Macraulacini, with notes on their biology. The Coleopterists Bulletin, 67: 97-106.]; O. rugicollis (Fall) [Otto, 2013Otto, R. 2013. Eucnemid Larvae of the Nearctic Region. Part III: Mature Larval Descriptions for Three Species of Onichodon Newman, 1838 (Coleoptera: Eucnemidae: Macraulacini, with notes on their biology. The Coleopterists Bulletin, 67: 97-106.]; Pseudoscython parvulus Fleutiaux [Gardner, 1935Gardner, J.C.M. 1935. Immature stages of Indian Coleoptera (17) (Eucnemidae). Indian Forest Records (New Series) Entomology, 1: 79-93, plates I-II.]
		Nematodini	Nematodes filum (Fabricius) [Leiler, 1976Leiler, T.E. 1976. Zur Kenntnis der Entwicklungsstadien und der Lebensweise nord- und mitteleuropäischer Eucnemiden (Col.). Entomologische Blätter für Biologie und Systematik der Käfer, 72: 10-50.]; Nematodes penetrans (LeConte) [Otto, 2017Otto, R. 2017. Eucnemid larvae of the Nearctic Region. Part VII: Description of the larvae of Nematodes penetrans (LeConte, 1852) (Coleoptera: Eucnemidae: Macraulacinae: Nematodini), with notes on its hypermetamorphic life cycle. Insecta Mundi, 545: 1-9.]; Nematodes sp., Fiji, Viti Levu, reared; Trigonopleurus rugulosus* Bonvouloir*, Australia, NSW, reared
CEROPHYTIDAE			Cerophytum elateroides (Latreille) [*Costa et al.,* 2003**Costa, C.; Vanin, S.A.; Lawrence, J.F. & Ide, S. 2003. Systemtics and cladistics analysis of Cerophytidae (Elateroidea: Coleoptera). Systematic Entomology, 28: 375-407.]
THROSCIDAE			Trixagus dermestoides (Linnaeus) [Burakowski, 1975Burakowski, B. 1975. Development, distribution and habits of Trixagus dermestoides (L.), with notes on the Throscidae and Lissomidae (Coleoptera, Elateroidea). Annales Zoologici, Warszawa, 32: 375-405.]

[1] ²E-mail: jyrki.muona@helsinki.fi (corresponding author)

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A Re-evaluation of the Eucnemidae larval characters (Coleoptera)

Abstract