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Tve334 Beutel.Qxp R. G. BEUTEL Institut für Spezielle Zoologie und Evolutionsbiologie, Jena MORPHOLOGY AND EVOLUTION OF THE LARVAL HEAD OF HYDROPHILOIDEA AND HISTEROIDEA (COLEOPTERA: STAPHYLINIFORMIA) R. G. Beutel, 1999. Morphology and evolution of the larval head of Hydrophiloidea and Histeroidea (Coleoptera: Staphyliniformia). – Tijdschrift voor Entomologie 142: 9-30, figs. 1- 29, table 1. [ISSN 0040-7496]. Published 22 September 1999. Internal and external features of the larval head of Spercheus emarginatus, Hydrochus sp., Helophorus sp., and of Hister sp. are described in detail. Character transformations of these struc- tures apparently play an important role in the evolution of Hydrophiloidea and Histeroidea. A basal position of Spercheidae within this lineage of Staphyliniformia is suggested by a considerable number of plesiomorphic features: head subprognathous, adnasalia and nasale absent, gula broad and short, posterior tentorial arms arise close to hind margin of head capsule, tentorial bridge fair- ly broad and straight, maxillary groove deep, with well developed articulating membrane, uniden- tate mandibular retinaculum, cardo undivided, moveability between cardo and stipes fully re- tained, stipes not tube-like, maxillary palp inserted laterally, lacinia strongly developed and hook-like, prepharynx short, anatomical mouth dilatable, brain located within head capsule, glands present in labial region. Some of these character states are also found in larvae of Hydrochus but not in other larvae of Hydrophiloidea and Histeroidea. A cladistic analysis, which is exclu- sively based on larval characters, results in the following branching pattern (strict consensus tree): Scarabaeidae + (Agyrtidae + Leiodidae + Hydraenidae + (Spercheidae + (Hydrochidae + (Histeri- dae + Synteliidae + Sphaeritidae) + (Helophoridae + ((Georissidae + Epimetopidae) + Hydrophil- idae)))))). A monophyletic unit which comprises Histeroidea and subgroups of Hydrophiloidea is in contrast to other phylogenetic hypotheses. It cannot be fully excluded that derived character states which are related with predacious habits have independently evolved in both superfamilies. R. G. Beutel, Institut für Spezielle Zoologie und Evolutionsbiologie, FSU Jena, D-07743 Jena, Germany. E-mail: [email protected] Key words. – Morphology; larval head; Hydrophiloidea; Histeroidea; phylogeny. Hydrophiloidea is a superfamily of the polyphagan 1997b). Hydrophiloid larvae hatch from a silk cocoon series Staphyliniformia (Lawrence & Newton 1995, whereas eggs are deposited uncovered by histeroid fe- Hansen 1997a) and comprises about 2400 described males. The larvae of both groups are carnivorous and de- species (Hansen 1991). Adults are found in aquatic, velop fast. However, histeroid larvae seem to be special- semiaquatic, or terrestrial habitats, and some repre- ized on maggots (Newton 1991), a food preference which sentatives of the subfamily Sphaeridiinae have spe- is also found in some sphaeridiines but not in other groups cialised on life in dung or decaying matters (Hansen of Hydrophiloidea. 1997b). In most cases, larvae are found in the same The systematic concept of the superfamily Hy- environment as the adults. However the feeding drophiloidea, the interrelationships of subgroups, and habits differ considerably. Adults rely on various sorts the affinities with Histeroidea were discussed contro- of plant material, mainly decaying tissue (Hansen versially in older and more recent studies (Böving & 1997b), whereas larvae of most subgroups are carniv- Craighead 1931, Crowson 1955, Lawrence & New- orous (Böving & Henriksen 1938, Bertrand 1972, ton 1982, 1995, Hansen 1991, 1997a, Beutel 1994, Hansen 1997b). Archangelsky 1998). The phylogenetic analyses pre- A sister group relationship between Histeroidea and sented by Hansen (1991, 1997a) are based on many Hydrophiloidea was postulated by Hansen (1997a). external features of adults and some external charac- However, the life styles of both groups are clearly differ- ters of larvae. External features of immature stages ent. All stages of Histeroidea are terrestrial and the adults were also described and analysed by Archangelsky are predominantly or exclusively predacious (Hansen (1997, 1998). Internal features of larvae (and adults) 9 Downloaded from Brill.com09/28/2021 06:03:57PM via free access T E, 142, 1999 are treated in very few studies and not in great detail Histeridae (e.g. Quennedey 1965, Moulins 1959). They were Hister sp. (?) (identified with Klausnitzer 1978, not considered in cladistic analyses so far. Important several genera are missing in the key) structural and functional transformations of the larval Hololepta aequalis (Say, 1825) head have apparently taken place in the evolution of Scarabaeidae Hydrophiloidea (Beutel 1994). Therefore, this con- Liocola sp. tribution is aiming at an improved knowledge of the anatomy and morphology of larvae in order to pro- Specimens of Spercheus emarginatus, Helophorus, and vide a broader basis for phylogenetic analyses. Hololepta were embedded in Historesin, cut at 5 ␮m, The cladistic analysis presented in this study is main- and stained with methylene-blue and acid fuchsine. ly aiming at a more reliable character state polarity as- Available specimens of larvae of Hydrochus were mount- sessment. The results have to be considered as prelimi- ed on microscope slides. Larvae of other taxa were dis- nary for several reasons. Only larval features, mainly of sected. Drawings were made using an ocular grid. the head are included. Different higher ranking taxa are Von Kéler’s muscular nomenclature is used in the only represented by the larvae of one or a few species ex- text and the corresponding number are used in the il- amined (e.g. Hydrophilidae, Histeridae) and many data lustrations (Von Kéler 1963). concerning internal features are missing. This is due to The phylogenetic analysis was carried out using the the general lack of anatomical studies and the restricted cladistic computer programs PAUP version 3.1 (Swof- availability of larvae, especially of material suitable for ford 1991) and MacClade version 3 (Maddison & histological work. Detailed study of internal structures Maddison 1992). of larvae of Hydrochidae, Georissidae, Epimetopidae, Synteliidae, and Sphaeritidae should be considered as MORPHOLOGICAL RESULTS an important future project. Spercheus emarginatus MATERIAL AND TECHNIQUES Head capsule, external features (figs. 1, 7) Species examined Head slightly inclined, subprognathous, moderately Hydraenidae: compressed dorsoventrally, rounded laterally. Sclero- Davidraena sp. tized parts brownish to testaceous. Setae unusually thin, Hydraena sp. distribution irregular (fig. 1). Dorsal side of head capsule Ochthebius sp. covered with minute spines. Five stemmata present. An- Agyrtidae teriormost stemma large, located on a prominent, an- Necrophilus hydrophiloides terolateral elevation. Posterodorsal stemma adjacent Guérin-Menneville, 1835 with an unpigmented, semitransparent area. Labrum Silphidae fused to clypeus, posterior margin defined by indistinct- Silpha obscura Linnaeus, 1758 ly impressed transverse line. Clypeofrontal suture com- Spercheidae pletely absent. Anterior tentorial grooves not apparent. Spercheus emarginatus (Schaller, 1783) Frontal suture very indistinct in 3rd instar larvae, oblit- Hydrochidae erated posteriorly. Conspicuous transverse sulcus pre- Hydrochus elongatus (Schaller, 1783) sent in the posterodorsal area of the head capsule. Coro- H. megaphallus Berge Henegouwen, 1988 nal suture absent. Maxillae inserted in deep maxillary H. ignicollis Motschulsky, 1860 grooves, with well developed articulating membrane (mounted on slides) (fig. 7). Distinct ridge present lateral to the maxillary Helophoridae grooves. Gula broad and sclerotized. Posterior tentorial Helophorus sp. grooves widely separated, fairly long and fissure-shaped, H. aequalis Thomson, 1868 scarcely discernible externally. H. redtenbacheri Kuwert, 1885 (3rd instar) Georissidae Internal skeletal structures (figs. 13, 17, 21) Georissus crenulatus Rossi, 1794 Posterior tentorial arms arise immediately close to Sphaeridiinae hind margin of head capsule, connected with the Sphaeridium sp. postoccipital ridge posterolaterally. Tentorial bridge Hydrophilinae straight and fairly broad (figs. 17, 21). Anterior arms Berosus sp. moderately strong. Dorsal arms normally developed, Hydrochara caraboides (Linnaeus, 1758) attached to dorsal wall of head capsule close to Hydrophilus piceus (Linnaeus, 1758) frontal suture. 10 Downloaded from Brill.com09/28/2021 06:03:57PM via free access BEUTEL: Larval head of Hydrophiloidea and Histeroidea Labrum (figs. 1, 13, 21) almost vertical bundles, O: posterior tentorial arm, Fused with clypeus, rounded anteriorly, without below M 17, I: mesally and ventromesally on pro- nasal or adnasal lobes. Lateral margin with several ximal part of stipes; M 19: M. craniolacinialis, O: ventrally directed, triangular, sclerotized processes posterolaterally from head capsule, I: base of lacinia; (fig. 21). M 20: M. stipitolacinialis: absent; M 22: M. stipi- Musculature: absent topalpalis externus, O: mesally from stipes, lateral to insertion of M. craniolacinialis, I: ventrally on base of Antenna (figs. 1, 13) palpifer; M 23: M. stipitopalpalis internus, O: mesal- 3-segmented, anteriorly directed, fairly slender. In- ly from stipes, close to origin of M 22, I: anterodor- serted on a conspicuous prominence resembling a basal sally on base of palpifer. antennomere. Antennomere
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