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Larval Morphology and Phylogenetic Position of Micromalthus Debilis Leconte (Coleoptera: Micromalthidae)

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Larval Morphology and Phylogenetic Position of Micromalthus Debilis Leconte (Coleoptera: Micromalthidae) Systematic Entomology (2002) 27, 169±190 Larval morphology and phylogenetic position of Micromalthus debilis LeConte (Coleoptera: Micromalthidae) R. G. BEUTEL andT. HOÈ RNSCHEMEYER* Institut fuÈ r Spezielle Zoologie und Evolutionsbiologie, FSU, Jena, Germany and *Institut fuÈ r Zoologie und Anthropologie, Abteilung fuÈ r Morphologie und Systematik, UniversitaÈ tGoÈ ttingen, GoÈ ttingen, Germany Abstract. External and internal structures of the cerambycoid and triungulin larvae of Micromalthus debilis are described and compared to features found in larvae of other groups of Coleoptera. The morphological data are evaluated with respect to the systematic position of Micromalthidae. A cladistic analysis was carried out with fifty characters. Micromalthidae are not closely related to Lymexylidae (Polyphaga: Cucujiformia) but belong to Archostemata, which is con- firmed as a monophyletic unit. Micromalthidae are specialized in terms of morphology and life cycle and are characterized by a considerable number of larval autapomorphies. Their sister-group relationship with Cupedidae is supported by several apomorphic features, which are probably correlated with xylobiontic habits: head transverse and strongly rounded laterally, absence of stemmata, shortened antennae, presence of sternal asperities and presence of eversible lobes of segment IX. Cupedidae is monophyletic and Priamca is the sister group of the remaining genera of Cupedidae included in the analysis. A closer relationship between Tenomerga and Rhipsideigma is supported by several larval synapomorphies. The ancestral life style of larvae of Archostemata was probably xylobiontic. This is suggested by derived groundplan features of the suborder, which are also found in larvae of non-related, wood-associated families. Introduction South Africa and North America were pointed out by Patterson (1938). However, they were not considered as Micromalthus debilis LeConte is probably the most spec- sufficient evidence for description of a new species. tacular beetle species in terms of life cycle and morphology. The highly complex life cycle of M. debilis is character- It is the only known species of Micromalthidae and has ized by active, first-instar triungulins, different types of a natural range restricted to northeastern North America larviform reproductive stages, vivipary and parthenogenesis (Lawrence, 1991). However, it was apparently introduced (Barber, 1913a,b; Pringle, 1938; Lawrence, 1991). This with timber to different parts of the world, such as Burgen- reproductive system is unparalleled in Coleoptera and land (southeastern Austria) (JaÈ ch & Komarek, 2000), other groups of insects. British Columbia, New Mexico, Cuba, Brazil (Minas Gerais, The systematic position of Micromalthidae has been the SaÄ o Paulo), South Africa (Witwatersrand, Johannesburg), subject of a long controversy. The species was described by Hawaii (Oahu), Hong Kong and Gibraltar (Lawrence, LeConte (1878), who emphasized that the adults (Fig. 1) are 1991). In contrast to all other known micromalthid triungu- `feeble and ill developed'. He included it in Lymexylidae. lins, those found in Hong Kong possess a pair of stemmata This was supported by Hubbard (1878), who provided the (Marshall & Thornton, 1963) and may therefore belong first larval description and pointed out similarities in anten- to a separate species. Differences between adults from nae and mouthparts of micromalthid and lymexylid larvae. A close relationship with Lymexylidae was also sugges- ted by Jeannel & Paulian (1949), whereas affinities with Correspondence: Rolf G. Beutel, Institut fuÈ r Spezielle Zoologie Adephaga were discussed by van Emden (1932). Micro- und Evolutionsbiologie, FSU, 03641 Jena, Germany. E-mail: malthidae was placed with Cupedidae (Archostemata) based [email protected] on the wing venation and larval mouthparts by Forbes (1926) # 2002 The Royal Entomological Society 169 170 R. G. Beutel and T. HoÈrnschemeyer (Ross & Pothecary, 1970) are concise and largely or completely restricted to external structures. The main purpose of this study was to give a detailed account of the larval morphology of M. debilis. Cladistic analyses were carried out in order to clarify the archoste- matan affinities of Micromalthidae and to reconstruct the ancestral life style of this suborder. Features of archostema- tan larvae are compared to those of immature stages of other xylobiontic groups, and correlations of structural features to specialized wood-boring habits are examined. Materials and methods Material and morphology Larvae of M. debilis (one first-instar triungulin larva, one second-instar cerambycoid larva, seven third-instar ceram- bycoid larvae, two damaged fourth-instar cerambycoid larvae) were collected from wood in the laboratory and fixed in Bouin's solution. The body segments of the larvae were less well preserved than the head. The triungulin and two third-instar cerambycoid larvae were imbedded in Histo- resin medium, cut into 3±5 mm sections with a rotation micro- tome (Microm GmbH, Walldorf, Germany) and stained with methylene-blue and acid fuchsin. Drawings were made using an ocular grid or a camera lucida (cross- Fig. 1. Micromalthus debilis: adult habitus. Specimen from Austria (Burgenland), drawn after photograph, no scale available. Length sections). For scanning electron microscopy, two third-instar of specimen approximately 2.5 mm. cerambycoid larvae were cleaned with ultrasonic sound, critical-point dried and coated with gold. The concept of the order and families follows Lawrence and BoÈ ving & Craighead (1931), respectively. This view & Newton (1995). von Ke ler's (1963) nomenclature for was adopted by Crowson (1955). Machatschke (1962) and muscles is used in the text, and the corresponding numbers Hennig (1969) rejected this placement, and the family are used in the illustrations. The following material was was tentatively transferred to Cantharoidea by Arnett (1968). examined. Archostematan affinities were also clearly rejected by Megaloptera, Sialidae. Larvae: Sialis sp. (fixed in Klausnitzer (1975): `die Archostemata im Sinne Crowson's FAE formol-ethanol-acetic acid; diss. dissection). Adults: sind zweifellos paraphyletisch'. The concept of Micro- Sialis lutaria (Linnaeus) (FAE; diss.). malthidae as an archostematan family was again strongly Archostemata, Micromalthidae. Larvae: M. debilis supported in several systematic studies (e.g. Lawrence, (Bouin's solution, micr. microtome sections, SEM) 1982; Lawrence & Newton, 1982, 1995; Kukalova -Peck & (from decaying wood, collected in south-central Wisconsin Lawrence, 1993). Nevertheless, Barlet (1996) decidedly by D. K. Young; material deposited in collection of R.G.B). advocated a position close to Lymexylidae (Polyphaga: Adults: M. debilis (dried, SEM). Cucujiformia) based on an examination of thoracic features Cupedidae. Larvae: Rhipsideigma raffrayi Neboiss of adults. (Pampel's fluid, ethanol), Tenomerga cinereus (Westwood) If Micromalthus belongs to the oldest suborder of (ethanol; examined at the Australian National Insect Coleoptera (Lawrence, 1999), a profound knowledge of Collection, Canberra ANIC), Distocupes varians (Lea) the morphology of its immature stages is important for (ethanol, micr.). Adults: Priacma serrata LeConte (diss., understanding the evolution of beetles. Nevertheless, little micr., SEM), Prolixocupes latreillei (Solier) (dried), Disto- information on micromalthid, cupedid and ommatid larvae cupes varians (dried, SEM). is presently available. Larvae of species of Tetraphalerinae, Ommatidae. Larvae: Omma sp. (ethanol; at ANIC). Adults: Crowsoniellidae and several genera of Cupedidae are Omma stanleyi Newman (diss.; SEM), Tetraphalerus unknown. The descriptions of larvae of M. debilis (Hubbard, sp. (dried). 1878; Pringle, 1938; Costa et al., 1988), Omma sp. (one known Adephaga, Trachypachidae. Larvae and adults: specimen; Lawrence, 1999), Tenomerga cinereus (Say) Trachypachus holmbergi Mannerheim (FAE, diss., micr., (BoÈ ving, 1929; BoÈ ving & Craighead, 1931), T. mucida SEM). (Chevrolat) (Fukuda, 1938), Distocupes varians (Lea) (Neboiss, Myxophaga, Torridincolidae. Larvae and adults: 1968) and first-instar larvae of Priacma serrata (LeConte) Satonius kurosawai Satoà (ethanol, micr., SEM). # 2002 The Royal Entomological Society, Systematic Entomology, 27, 169±190 Larval morphology and phylogenetic position of Micromalthus 171 Polyphaga, Lymexylidae. Larvae and adults: Hylecoetus was conducted using MacClade version 3 (Maddison & sp. (FAE, micr.). Adults: Hylecoetus sp. (Bouin's solution, Maddison, 1992). Sialis sp. (larva), Sialis lutaria (adults) diss.), Lymexylon navale (Linnaeus) (Bouin's solution, (Megaloptera), Trachypachus holmbergi (Adephaga), diss.). Satonius kurosawai (Myxophaga), Hylecoetus sp. and Coccinellidae. Larvae and adults: Coccinella Coccinella septempunctata (Polyphaga) were used as outgroup septempunctata Linnaeus (FAE, micr.). taxa. The outgroup taxa were treated as all other groups Melandryidae. Larvae: Orchesia sp. (FAE, micr., SEM). in the analysis (simultaneous analysis; Nixon & Carpenter, 1993). All characters were weighted equally and not ordered. Cladistic analysis Morphological observations Most parsimonious cladograms were generated with PAUP version 3.1 (Swofford, 1991). Branch and bound Cerambycoid larvae are described in detail from specimens search was applied (thirteen taxa and fifty characters; prepared for scanning electron microscopy (SEM), micro- Appendix 1: data matrix). Analysis
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