Anischia, Perothops and the phylogeny of Elateroidea (Coleoptera: Elateriformia) JOHN F. LAWRENCE, JYRKI MUONA, MARIANNA TERÄVÄINEN, GUNILLA STÅHLS and VARPU VAHTERA Insect Syst.Evol. Lawrence, J.F., Muona, J., Teräväinen, M., Ståhls, G. and Vahtera, V.: Anischia, Perothops and the phylogeny of Elateroidea (Coleoptera: Elateriformia). Insect Syst. Evol. 38: 205-239. Co- penhagen, July, 2007. ISSN 1399-560X The larvae of Anischia Fleutiaux and Perothops Laporte are described. Cladistic analyses based on adult and larval morphology, as well as CO1 sequence data, place both genera in the Eucnemidae clade within the Elateroidea (sensu stricto). The subfamily Anischiinae Fleutiaux, 1936 is placed in the family Eucnemidae in a clade containing the more derived eucnemid subfamilies (Melasinae, Eucneminae and Macraulacinae), while Perothops and Phyllocerus Lepeletier & Serville represent subfamilies basal to the remaining eucnemid taxa. The genus Afranischia Basilewsky, 1955 is synonymized with Anischia Fleutiaux, 1896, and Anischia boliviana Fleutiaux is selected as the type species of the latter. Three new species are described: Anischia bicolor (New Caledonia), Anischia kuscheli (New Caledonia) and Anischia monteithi (NE Australia), and Anischia stupenda Fleutiaux, 1897 is recorded from Australia. Anischia crassicornis Champion, 1897 is synonymized with Anischia mexicana Fleutiaux, 1896. One new combination is made, Anischia ruandana (Basilewsky, 1955). J. F. Lawrence, CSIRO Entomology, GPO Box 1700, Canberra, ACT 2601, Australia *J. Muona, M. Teräväinen, G. Ståhls and V. Vahtera, Finnish Museum of Natural History, P. O. Box 17, 00014- University of Helsinki, Finland *Corresponding author ([email protected]) Introduction Perothops Laporte was placed in a separate sub- Although originally placed in Eucnemidae, Ani- family of Eucnemidae (Perothopsitae) by Lacord- schia Fleutiaux has been included in Cerophytidae aire (1857) and the genus is usually included in by many workers, based mainly on the absence of that family. Perothopidae was also recognized at metacoxal plates. Fleutiaux (1936) transferred the the family level by Horn (1878), Schenkling genus to Elateridae, placing it in a new subfamily. (1928), Crowson (1955), Arnett (1963) and a few Anischiinae was considered to be incertae sedis other workers. Muona (1993) suggested that within Elateridae by Lawrence and Newton (1995) Perothops might belong to either Eucnemidae or a but recognized as a distinct family by Lawrence et clade comprising Throscidae (sensu lato) and al. (1999). In cladograms produced by Muona Elateridae; however he chose to recognize the (1995) based on 25 elateroid genera and 70 char- group as the most basal of eucnemid subfamilies. acters (6 larval), Anischia was at or near the base A number of years ago, one of us (JFL) discovered of a clade comprising Elateridae and Throscidae in a large and unusual, eucnemid-like larva in the the sense of Crowson (1955) (including Lissom- collections of the National Museum of Natural inae and Thylacosterninae). The discovery of a History in Washington. Although not reared or unique, eucnemid-like larva associated with adult associated with adult beetles, the size, structure Anischia in New Caledonia led us to reconsider and locality of this soil-inhabiting larva were con- the affinities of the genus as part of a reanalysis of sidered in making the identification (see larval the elateroid complex. description below). © Insect Systematics & Evolution (Group 2, 11) 206 Lawrence, J. F. et al. INSECT SYST. EVOL. 38:2 (2007) The purpose of the present paper is to clarify the Taxon Sampling. – In addition to Perothops and position of these two unusual genera and deter- Anischia, 10 exemplar genera were chosen repre- mine their relationships to the families Eucnem- senting all subfamilies recognized by Muona idae, Cerophytidae, Throscidae and Elateridae. (1993) except Phlegoninae, three genera repre- senting each of the largest elaterid subfamilies Agrypninae, Dendrometrinae and Elaterinae, five Materials and Methods genera representing the disputed elaterid subfami- Morphological terms. – The terms mesoventrite lies Lissominae and Thylacosterninae, and one and metaventrite have been used in place of the genus each from the families Cerophytidae and misapplied terms mesosternite and metasternite Throscidae. In addition, the genera Brachypsectra following Lawrence 1999 and Lawrence et al. and Macropogon were chosen because the families 1999 (see also Beutel & Haas 2000). Wing vein Brachypsectridae and Artematopodidae were con- terminology follows that of Kukalová-Peck & sidered to be at or near the base of Elateroidea Lawrence (1993, 2004). (sensu lato) in Lawrence (1988), Lawrence et al. (1995) and Beutel (1995). In the larval cladogram Measurements and ratios. – BL = body length produced by Beutel (1995), Brachypsectra was (total length excluding head, or PL + EL); PL = basal to the cantharoid complex, and it occupied a median pronotal length; PW = greatest pronotal similar position in a cladogram based on adults width; EL = greatest elytral length; EW = greatest and larvae with Dascilloidea as an outgroup in elytral width. Lawrence (1988). This taxon was preferred over other more common cantharoid taxa, partly Image enhancement. – Images in Figs 1–10, 26–28 because of its basal position, but also because and 31 were enhanced using Auto-Montage soft- adult Brachypsectra lack most of the secondary ware version 4.00 (Synoptics Ltd., http//www.syn- cantharoid features associated with neoteny, short croscopy.com). adult life span, and the loss of cuticular strength- Specimen repositories. – The following abbrevia- ening mechanisms in favor of chemical defenses. tions have been used: ANIC: Australian National The three outgroups chosen represent 1) Dascill- Insect Collection, Canberra; BMH: Bishop Mu- idae, the most basal member of the series Ela- seum, Honolulu; CAS: California Academy of teriformia as used by Lawrence 1988 (Dascilli- Sciences, San Francisco; FMNH: Field Museum formia of Crowson 1955), Eucinetidae, a member of Natural History, Chicago; MNHN: Muséum of the Scirtoidea as used by Lawrence 2001 (Euci- National d’Histoire Naturelle, Paris; MZSP: Mu- netoidea of Crowson 1960) and one of the most seu de Zoologia de Univesidade de São Paulo; basal groups of Polyphaga, and 3) Scarabaeidae, a NMNH: National Museum of Natural History, member of the series Scarabaeiformia. Washington, D.C.; NZAC: New Zealand Arthro- pod Collection, Auckland; QMB: Queensland Mu- Specimens Examined. – The following list in- seum, Brisbane; UZMH: Finnish Museum of Na- cludes those taxa used for the coding of morpho- tural History, Helsinki. logical character states and for DNA extraction and representing exemplar genera in the cladistic Character Coding: – Elateroid relationships have analyses. Genera are arranged alphabetically with been studied quite intensively, but these studies their current family group placement in brackets. have yielded conflicting results (Calder et al. Ampedus Dejean (Elateridae: Elaterinae: Am- 1993; Muona 1995). It was clear to us that a very pedini). Morphology based on adults of A. san- thorough evaluation of the characters previously guineus (Linnaeus) and A. rubricollis (Herbst), used was needed. We hope to have removed all and descriptions and illustrations in Ôhira (1962) characters that could not be unambiguously coded and Dolin (1978). Sequence data: Ampedus nigro- from the present data set. flavus (Goeze); Finland, Uusimaa, Lehtisaari, v. The 118 morphological characters and character 1997, J. Muona leg. GenBank accession number states are listed in Appendix 1, along with notes on EF589366. the distribution of these states among the exemplar Anischia Fleutiaux (Elateridae: Anischiinae or taxa. The character matrix is included as Table 2. Anischiidae). Morphology based on adults of Anischia species described below and a larva of A. INSECT SYST. EVOL. 38:2 (2007) Phylogeny of the Elateroidea 207 kuscheli sp. nov. Sequence data: Anischia bicolor phology based on adults and larvae of Drapetes n. sp.; New Caledonia, Mts. Koegis, v. 1996, J. geminatus (Say) and descriptions and illustrations Muona leg. GenBank accession number of D. biguttatus (Piller) in Burakowski (1973). No EF589367. sequence data. Aulonothroscus Horn (Throscidae: Throscinae). Eucnemis Ahrens (Eucnemidae: Eucneminae: Morphology based on adults of several Aulono- Eucnemini). Morphology based on adults of E. throscus species from North America and Austra- capucina Ahrens and descriptions and illustrations lia, a putative Aulonothroscus larva from North in Muona (1993), Leiler (1976) and Mamaev America, and descriptions and illustrations in (1976). Sequence data: Eucnemis capucina Ah- Burakowski (1975, 1991) and Cobos (1967). Se- rens; Russia, Karelia, P. Martikainen leg. GenBank quence data: Aulonothroscus sp., Malesia, Sabah, accession number EF589373 N. Laurenne leg. GenBank accession number Galbites Fleutiaux (Eucnemidae: Eucneminae: EF589368. Galbitini). Morphology based on adults of Galbi- Austrelater Calder & Lawrence (Elateridae: tes spp. and descriptions and illustrations in Lissominae?). Morphology based on adults and Muona (1991, 1993) and Gardner (1935). Se- larvae of A. macphersonensis Calder and descrip- quence data: Galbites wallacei (Perroud & Mon- tions and illustrations in Calder et al. (1993). trouzier); Malaysia, Sznik leg. GenBank accession Brachypsectra LeConte (Brachypsectridae). number EF589374 Morphology based