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Home , Balgus, Brachypsectridae, Click beetle, Dystaxia (beetle), Histeroidea, Stenocladius

4.

Introduction, Phylogeny labrum in the , and were considered to be even more diffi cult to separate on adult features. The superfamily Cantharoidea was John F. Lawrence, Ladislav Bocak, Milada Bocakova, considered to be the most likely group to be merged Rolf G. Beutel and Jyrki Muona with Elateroidea. Lawrence & Newton (1982) followed Crowson The constitution of has varied over in considering Artematopodidae, Brachypsec- time, as discussed in detail by Beutel & Leschen tridae, Elateroidea and Cantharoidea to form a (2005) (see 1–14). The series was fi rst proposed by monophylum, and Lawrence (1988) formally rec- Crowson (1960) for his Dascilliformia (Crowson ognized an expanded Elateroidea to include all of 1955) minus the family (which was these groups. The position of Rhinorhipus Lawrence combined with to form the series at the base of the elateroid clade was considered to Scarabaeiformia) and the families , be tentative because of lack of information on the and , which were placed in a larva, combined with the fact that there are six free superfamily Eucinetoidea. This classifi cation was Malpighian tubules (instead of four as in all other also used in Crowson (1981) except that a series Euci- members of the group). Furthermore, in clado- netiformia was recognized and Rhipiceridae was grams produced by Lawrence et al. (1995), Rhinor- added to Scarabaeiformia-, based on hipus usually formed a clade with Dascillus Latreille Crowson (1971). In all of Crowson’s classifi cations, (Dascillidae), Sandalus Knoch (Rhipiceridae) and the superfamily Elateroidea included Perothopidae, Dystaxia LeConte ( or ) , , Cebrionidae, Elateridae, and was never placed within the elateroid-cantha- and , although the last was omitted in roid group. error from the 1981 work. In cladograms produced by Beutel (1995) and Lawrence & Newton (1982) followed Crowson’s based on larval characters, Elateroidea (sensu lato) classifi cation in most respects. They did not defi ne was always monophyletic, but this was true of series as such, but considered all Eucinetiformia, neither Elateroidea (sensu stricto) nor Cantharoidea. Scarabaeiformia and Elateriformia as belonging to Most cantharoid families plus Brachypsec- an “Elateriform lineage”. Although Scarabaeoidea tridae formed a clade sister to Cerophytidae + was tentatively included in this “lineage”, some Throscidae + Eucnemidae, while Cantharidae doubt was expressed about the relationship formed a clade with Artematopodidae and Elat- of the group to Dascilloidea. The families nor- eridae. The non-monophyly of the Cantharoidea mally included in Elateroidea and Cantharoidea, was also supported by Bocakova et al. (2007) in plus the Artematopodidae and Brachypsectridae cladograms based on nuclear and mitochondrial were considered to form a monophyletic group. gene sequences. While Elateroidea (sensu lato) was In the fi rst cladistic analysis of Elateriformia, strongly supported in all cladograms, the soft-bod- Lawrence (1988) excluded Scarabaeoidea alto- ied groups usually placed in Cantharoidea never gether, while Eucinetoidea were included as an formed a monophyletic group. The major clusters outgroup in some analyses. The monophyly of were formed by 1) Lampyridae (including Ototreti- Elateroidea + Cantharoidea + Artematopodidae + nae) + Cantharidae, 2) Elateridae (including Drilidae Brachypsectridae was confi rmed in analyses based and usually Omalisidae) + (including on both adult and larval characters, with the family ), 3) and 4) Eucnemi- Rhinorhipidae (known from adult characters only) dae. The positions of the genera Drilonius, Telegeusis, at the base of this clade. Trixagus and sometimes Omalisus varied with type Elateroidea was restricted by Crowson (1955) of alignment and analysis: 1) Drilonius, Telegeusis to those taxa the adults of which have more or less and Trixagus formed a clade with rounded procoxae with concealed trochantins, no and outside Elateroidea; 2) Drilonius and Telegeusis transverse metakatepisternal suture, contiguous formed a clade sister to Elateroidea and Trixagus was metacoxae, hind wing with an apically truncate sister to Elateroidea minus Drilonius and Telegeusis; wedge cell, acutely projecting hind pronotal 3) Drilonius was in Eucnemidae, Telegeusis sister to angles, head without a distinct frontoclypeal Elateroidea minus Eucnemidae, and Trixagus sister suture, trilobate aedeagus with freely articulated to Lycidae; or 4) Drilonius and Telegeusis formed a parameres, and 4 free Malpighian tubules, while clade sister to remaining elateroids, and Trixagus larvae lack a free labrum or epicranial stem and and Omalisus formed a clade within Eucnemidae. have simple, non-channeled mandibles. Artema- Similar results were published by Sagegami-Oba topodidae (then in Dryopoidea) were considered et al. (2007) and Bocak et al. (2008). to be separable from elateroids on little more The Elateroidea, as here delimited, exhibit than exposed trochantins in the adult and a free several major evolutionary trends which deserve 36 John F. Lawrence, Ladislav Bocak, Milada Bocakova, Rolf G. Beutel and Jyrki Muona further mention: 1) development of a type of defen- Leptolycinae (Lycidae) is not a result of the long evo- sive behavior known as “clicking” in adults of the lutionary history, but a consequence of relatively families Cerophytidae, Eucnemidae, Throscidae recent modifi ed function of the endocrine system. and Elateridae, 2) reduction in sclerotization These events potentially led to homoplasious mod- of the cuticle, often accompanied by chemical ifi cations of morphology. The resulting similarity defense mechanisms and aposematic color patterns of soft-bodied or neotenic lineages is therefore dif- in adults of various families formerly included in fi cult to interpret in morphology based analyses. Cantharoidea, and 3) retention of larval features Crowson (1972) postulated that some neoten- (neoteny) in adults of at least some of these families; ous groups, specifi cally the Southeast Asian lycid 4) the evolution of in both adults genera Duliticola and Lyropaeus, are members of and larvae; 5) the occurrence of an elateroid type of primitively neotenous lineages and that fully meta- ecdysis associated with biforous spiracles and the morphosed winged forms re-developed from neo- loss of the spiracular closing apparatus in larvae; tenic ancestors. Similar scenarios of evolutionary and 6) consolidation of the larval maxillae and ‘re-imaginalisation’ were proposed for Lycidae by labium to form a maxillolabial complex. Kazantsev (2005), and equally for the closely related The cuticular and muscular modifi cations which Lampyridae by Cicero (1988). Bocak et al. (2008) make the clicking maneuver possible have been hypothesized that soft-bodiedness represents a discussed by Evans (1972, 1973) for Elateridae, fi rst level of incomplete metamorphosis. Soft-bod- but precursors of these conditions are exhibited ied adults of both sexes are known in Telegeusidae, by members of various families of Dascilloidea, , Cantharidae, Lycidae, Lampyridae, and . The evolution of a Phengodidae, Rhagophthalmidae, Drilidae, and pro-mesothoracic interlocking device involving Omalisidae. Some adult females within these projections and concavities or crenulate edges at families are neotenic, i. e., they maintain appar- the posterior end of the prothorax, anterior ends of ently juvenile features resulting in incomplete the elytra, scutellum and/or mesanepisterna, com- metamorphosis and, in extreme cases, the lack of bined with a mesoventral cavity for reception of the adult stages. The neotenic development of females prosternal process, allow these to combine is obligatory in all Omalisidae, Drilidae, Phengodi- mobility with structural integrity, by the unlock- dae and Rhagophthalmidae, and in many lineages ing or locking of this device. The transformation of Lampyridae and Lycidae. The modifi cations of this condition to form the clicking mechanism include females with vestigial wings, but adult- involves the enlargement of the prothorax, increase like thorax (Omalisidae, Lampyridae part), wing- in the mass of the M4 muscle (Larsén 1966), reduc- less females (Lampyridae part) or females with only tion of the size of the exposed portion of the pro- mouthparts and head adult-like (Drilidae, Lampy- coxa, enclosure of the trochantin and (except in ridae part). Lineages affected by neoteny to the Cerophytidae) its fusion to the notum, enlarge- highest degree are found in Lycidae where females ment and deepening of the mesoventral cavity lack both pupal and adult stages and retain a larvae- combined with the formation of a prosternal rest like morphology after the last ecdysis (Wong 1996). and an oblique slide at the anterior end of the cav- Some neotenic lycids reach body sizes of fi ve centi- ity. Based on the topology given by Bocakova et al. meters and more and are frequently referred to as (2007), this condition could have arisen indepen- ‘trilobite larvae’ due to their appearance (Gravely dently from three to fi ve times in the Elateroidea. 1915; Mjöberg 1925). The corresponding males are Vahtera et al. (2009), however, suggested that, given regularly fully metamorphosed and only seldom the complexity of the clicking mechanism, it could brachelytrous (Alyculus in Lycidae and Phosphaenus have evolved at the base of the elateroid clade and in Lampyridae). been subsequently lost on numerous occasions, Several elateroid groups (Lampyridae, Phengo- usually in association with the development of didae, Rhagophthalmidae and two independent soft-bodiedness (see below). groups of Elateridae (: Cantharoidea were defi ned mainly on the shared and Thylacosterninae: ) are known for their morphological traits resulting from soft-bodied- bioluminescence. Previous morphological studies ness (generally reduced body sclerotisation and (Crowson 1972, Beutel 1995) often suggested close a soft, fl exible abdomen with extensive interseg- relationships of cantharoid luminescent lineages mental membranes reminiscent of those in the lar- (Lampyridae, Phengodidae). Latest molecular anal- vae). The molecular phylogenies mentioned above yses (Bocakova et al. 2007; Sagegami-Oba et al. 2007) ( Bocakova et al. 2007; Sagegami-Oba et al. 2007; showed that bioluminescent groups have arisen at Bocak et al. 2008) rejected monophyly of Cantha- least four times in Elateroidea. Likewise, recent roidea and proposed multiple origin of both soft- morphological analysis (Branham and Wenzel bodiedness and probably closely related female 2001, 2003) supported several independent origina- neoteny of some groups within broadly defi ned tions of bioluminescence in Elateroidea. Although Elateroidea. The hypothesis of frequent shifts to superfi cially similar, molecular conclusions differ neotenic development opens a possibility that the substantially. While morphological study sepa- morphological disparity suggesting establishment rates Rhagophthalmidae from Phengodidae, and of families like Drilidae or some subfamilies like Drilaster and Stenocladius from Lampyridae, neither Elateroidea 37 of these conclusions were confi rmed in the molecu- larvae is strongly modifi ed, with posterior arms lar studies, hence upholding the traditional view of very strongly developed, cranially directed and the constitution of Phengodidae and Lampyridae completely detached from the tentorial bridge. (Crowson 1972; Lawrence et al. 1995). Conversely, The dorsal and anterior parts of the tentorium latest molecular analyses found cantharoid lumi- are reduced. Interestingly, again a similar condi- nescent groups Lampyridae and Phengodidae tion is found in cleroid larvae and in some groups deeply separated which is also supported by the of (Beutel & S´ lipin´ ski 2001). Appar- structural and biochemical differences of the lucif- ently this condition is linked with the formation erases in either group (Viviani 2002). of a maxillolabial complex. A set of features dis- Vahtera et al. (2009) presented an hypothesis tinctly separating Elateroidea from Byrrhoidea is connecting the clicking mechanism with the evolu- the presence of a strongly developed lateral tento- tion of bioluminescence. If the clicking mechanism riohypopharyngeal muscle, a dense, preoral fi lter evolved in ancestors of the entire elateroid complex, formed by long microtrichia, the immobilisation the bifunctional role of the pre-luciferase enzyme of the labrum, and the loss of the labral muscles. in combination with the high-energy demand of A labrum separated from the clypeal region is pre- the pronotal muscle were the preadaptive features served only in Artematopodidae and Brachypsectri- for the luminescence to evolve in the prothorax. dae. The preoral fi lter is apparently an adaptation Lineages evolving away from the compact elaterid- to liquid feeding. A similar condition has evolved type body structure retained the predisposition for in Carabidae and (Beutel 1993, 1999). luminescence, once a suitable luciferin was avail- Unusual modifi cations of the mandibular appara- able. The sources of luciferin type compounds in tus are characteristic for larvae of most Eucnemidae beetles, whether of symbiotic origin or not, facili- and Throscidae, where mandibles may be fi xed or tated the pronotal light spots at the muscular exodont. Another specifi c modifi cation is the pres- attachment points as well as the fat body region. ence of mandibular sucking channels occurring This scenario predicts that the source for luciferin in Brachypsectridae, Lampyridae, and a few other is most likely external and after becoming available groups. An unusual feature apparently linked for any elateroid clade could be picked up repeat- with highly specialised liquid feeding habits is the edly. It also explains why this feature is restricted origin of very strongly developed extrinsic maxil- to this one group beetles – the clicking mechanism lary muscles of the sclerotised ventral wall of the being unique within beetles. hypopharynx. A somewhat similar condition has Larval head structures of Elateroidea are quite evolved in and some cucujoid groups, characteristic, even though, as pointed out in Beu- where an anterior bundle of M. tentoriostipitalis tel (1995), several derived features are also found originates from the ventral prepharyngeal wall or in larvae of all or most groups presently assigned from the posteriormost hypopharynx (Beutel & to Byrrhoidea (see 1–2). A tendency to concentrate S´ lipin´ ski 2001). or reduce the stemmata is found in both lineages. Well separated stemmata occur in so me groups of Byrrhoidea (e. g., , , Heterocer- Literature idae) but in others (e. g., Psephenidae, Ptilodactyli- dae) they form tight clusters and in Eulichadidae Beutel, R. G. (1993): Phylogenetic analysis of there is a single large lens beneath which are two (Coleoptera) based on characters of the larval head. to fi ve pigment spots. In Elateroidea there is never – Systematic Entomology 18: 127–147. more than a single stemma on each side. As in all – (1995): Phylogenetic analysis of Elateriformia Byrrhoidea, elateroid larvae lack a basal mandibu- (Coleoptera: ) based on larval characters. lar mola, and as in Byrrhoidea excl. Byrrhidae the – Journal of Zoological Systematics and Evolutionary head is distinctly prognathous. Both conditions Research 33: 145–171. have evolved independently in different lineages – (1999): Morphology and evolution of the larval of Coleoptera, notably in groups with predacious head of and Histeroidea (Coleop- larvae (e. g., Adephaga, Hydrophiloidea, Cleroidea tera: ). – Tijdschrift voor Entomologie 142: 9–30. [see 1–7, 1–10, 2–9]). A characteristic feature found Beutel, R. G. & Leschen, R. A. B. (2005): 14. Elateri- in larvae of Elateroidea (and Byrrhoidea excluding formia Crowson, 1960. Introduction, Phylogeny. Byrrhidae, some and Eulichadidae) Pp. 427–429 in Handbuch der Zoologie/Handbook of is a maxillolabial complex, with closely connected Zoology. Band/Volume IV Arthropoda: Insecta Teilband/ labium and maxillae (Beutel 1995). The ventral Part 38. Coleoptera, Beetles. Volume 1: Morphology and mouthparts are moved only as a structural unit Systematics (, Adephaga, , vertically. The extrinsic tentoriomaxillary muscles Polyphaga partim). W. DeGruyter, Berlin. are vertically arranged. Similar conditions have Beutel, R. G. & S´ lipin´ ski, S. A. (2001): Comparative evolved independently in Cleroidea and in some study of larval head structures of and supposedly related groups of Cucujoidea (Beu- (Cucujoidea, Coleoptera). – Euro- tel & S´ lipin´ ski 2001). As in most byrrhoid groups pean Journal of Entomology 98: 219–232. (excluding Byrrhidae, Ptilodactylidae, Eulichadi- Bocak, L., Bocakova, M., Hunt, T. & Vogler, A. P. (2008): dae and Callirhipidae) the tentorium of elateroid Multiple ancient origins of neoteny in Lycidae 38 John F. Lawrence

(Coleoptera): consequences for ecology and mac- Sagegami-Oba, R., Takahashi, N., Oba, Y. (2007): roevolution. – Proceedings of the Royal Society B, 275: The evolutionary process of bioluminescence and 2015–2023. aposematism in cantharoid beetles (Coleoptera: Bocakova, M., Bocak, L., Hunt, T., Teraväinen, M. & Elateroidea) inferred by the analysis of 18S ribo- Vogler, A. P. (2007): Molecular phylogenetics of somal DNA. Gene 400 (1–2): 104–113. Elateriformia (Coleoptera): evolution of biolumin- Vahtera, V. Muna, J., Ståhls, G. & Lawrence, J. F. escence and neoteny. – Cladistics 23: 477–496. (2009): The phylogeny of Thylacosterninae beetles Branham, M. A. & Wenzel, J. W. (2001): The evolution (Coleoptera, Elateridae). – Cladistics (in press). of bioluminescence in cantharoids (Coleoptera: Viviani, V. R. (2002): The origin, diversity, and struc- Elateroidea). – Florida Entomologist 84: 565–586. ture function relationships of luciferases. – (2003): The origin of photic behavior and the evolu- Cellular and Molecular Life Sciences 59: 1833–1850. tion of sexual communication in fi refl ies (Coleoptera: Lampyridae). – Cladistics 19: 1–22. Cicero, J. M. (1988): Ontophylogenetics of cantharoid larviforms (Coleoptera: Cantharoidea). Coleopterists Bulletin 42: 105–151. Crowson, R. A. (1955): The Natural Classifi cation of the 4.1. Rhinorhipidae Lawrence, 1988 Families of Coleoptera. 187 pp. Nathaniel Lloyd, London. John F. Lawrence – (1960): The phylogeny of Coleoptera. – Annual Review of Entomology 5: 111–134. – (1971): Observations on the superfamily Dascilloi- Distribution. Rhinorhipus tamborinensis Lawrence dea (Coleoptera: Polyphaga), with the inclusion of has been collected in a few localities in southern Karumiidae and Rhipiceridae. – Zoological Journal of Queensland, Australia, all at higher elevations the Linnean Society 50: 11–19. in the vicinity of closed forest. It is likely that the – (1972): A review of the classifi cation of Cantharoi- species also occurs in montane regions in northern dea (Coleoptera), with the defi nition of two new New South Wales. families, Cneoglossidae and Omethidae. – Revista de la Universidad de Madrid 21: 35–77. Biology and Ecology. The largest series of adults – (1981): The Biology of Coleoptera. 802 pp. John were collected during the day on leaf surfaces of an Murray, London. introduced weed, Ageratina adenophora (Asteraceae), Evans, M. E. G. (1972): The jump of the click at the edge between rainforest and cleared areas. (Coleoptera, Elateridae) – a preliminary study. Unfortunately, this site has now become a suburb – Journal of Zoology, London 167: 319–336. – (1973): The jump of the (Coleoptera: and recent collecting expeditions have failed to Elateridae) — energetics and mechanics. – Journal of produce more specimens. At another locality, a few Zoology, London 169: 181–194. beetles were found in an open area on low vegetation Gravely, F. H. (1915): The larvae and pupae of some bordering a creek. When disturbed, the beetles beetles from Cochin. Records of the Indian Museum exhibited a death-feigning reaction, dropping to 11: 353–366. the ground. Males greatly outnumbered females in Kazantsev, S. V. (2005): Morphology of Lycidae with this habitat. It is likely that these clearings were the some considerations on evolution of the Coleop- sites of mating aggregations and that the beetles tera. Elytron 19: 49–226. fl ew to them from within the rainforest. Red mud Larsén, O. (1966): On the morphology and function of was present on a number of the specimens, which the locomotor organs of the Gyrinidae and other suggests that they either emerged from the Coleoptera. – Opuscula Entomologica Supplementum after eclosion or sheltered there. The structure of 30: 1–242. the metacoxae and hind legs also suggests fossorial Lawrence, J. F. (1988): Rhinorhipidae, a new beetle fam- habits. The ovipositor is relatively unspecialized, ily from Australia, with comments on the phylogeny so it is unlikely that the eggs are embedded in plant of the Elateriformia. Invertebrate 2: 1–53. tissue or placed deep in soil. One female laid several Lawrence, J. F. & Newton, A. F., Jr. (1982): Evolution eggs in the laboratory, but none of them hatched. and classifi cation of beetles. – Annual Review of Ecol- ogy and Systematics 13: 261–290. Morphology, Adults (Figs. 4.1.1–3). Length 5–8.5 Lawrence, J. F., Nikitsky, N. B., Kirejtshuk, A. G. (1995): mm. Body about 3 times as long as wide; slightly Phylogenetic position of Decliniidae (Coleoptera: ) and comments on the classifi cation fl attened above but moderately convex below. of Elateriformia (sensu lato), pp. 375–410. In Heavily sclerotized and clothed with relatively J. Pakaluk and S. A. S´ lipin´ ski (eds.), Biology, stout and somewhat fl attened, decumbent hairs. Phylogeny, and Classifi cation of Coleoptera: Papers Head longer than wide, strongly declined, Celebrating the 80th Birthday of Roy A. Crowson. abruptly constricted immediately behind eyes, Muzeum i Instytut Zoologii Polska Akademia so that no temples are present. With very short, Nauk, Warsaw, Poland. median occipital endocarina but no transverse Mjöberg, E. (1925): The mystery of the so called “trilo- line. Eyes moderately large, protuberant, more or bite larvae” or “Perty’s larvae” defi nitely solved. less circular, fi nely faceted, without interfacetal Psyche 32: 119–157. setae; ommatidium of exocone type with thick