Abstract
The primary chaetotaxy of the larval head capsule and head appendages of the family Hydrophilidae (Insecta: Coleoptera) is described and illustrated using the larva of Hydrobius fuscipes (Linnaeus, 1758) as a model, and compared with fifteen hydrophilid taxa representing all main taxonomic groups within the family; brief comparative notes with representatives of the families Helophoridae, Spercheidae, Hydrochidae and Histeridae are also provided. Primary chaetotaxic nomenclature is developed for the Hydrophilidae, allowing the use of chaetotaxic characters for phylogenetic studies as well as diagnostic purposes. The study of representatives of the families Helophoridae, Hydrochidae and Spercheidae suggests that this nomenclature can also be effectively applied to other hydrophiloid families. Chaetotaxic nomenclature systems used in larvae of other groups of Coleoptera are briefly reviewed.
References
Ahn, K.J. & Ashe, J.S. (1996) Phylogeny of the intertidial aleocharine tribe Liparocephalini (Coleoptera: Staphylinidae). Systematic Entomology, 21, 99–114.
Alarie, Y. (1991) Primary setae and pores on the cephalic capsule and head appendages of larval Hydroporinae (Coleoptera: Dytiscidae: Hydroporinae). Canadian Journal of Zoology, 69, 2255–2265.
Alarie, Y. (1995) Primary setae and pores on the legs, the last abdominal segment, and the urogomphi of larvae of Nearctic Colymbetinae (Coleoptera: Adephaga: Dytiscidae) with an analysis of their phylogenetic relationships. Canadian Entomologist., 127, 913–943.
Alarie, Y. (1998) Phylogenetic relationships of Nearctic Colymbetinae (Coleoptera: Adephaga: Dytiscidae) based on chaetotaxic and porotaxic analysis of head capsule and appendages of larvae. Canadian Entomologist, 130, 803–824.
Alarie, Y., Archangelsky, M., Nilsson, A. N. & Watts, C.H.S. (2002) Larval morphology of genus Lancetes (Coleoptera: Adephaga: Dytiscidae): the hypothesis of sister-group relationships with the subfamily Dytiscinae revisited. Canadian Entomologist, 134, 467–501.
Alarie, Y., Beutel R.G. & Watts, C.H.S. (2004) Larval morphology of three species of Hygrobiidae (Coleoptera: Adephaga: Dytiscoidea) with phylogenetic considerations. European Journal of Entomology, 101, 293–311.
Alarie, Y. & Bilton, D.T. (2005) Larval morphology of Aspidytidae (Coleoptera: Adephaga) and its phylogenetic implications. Annals of the Entomological Society of America, 98, 417–430.
Alarie, Y., Nilsson, A.N. & Hendrich, L. (1999) Larval morphology of the Palaearctic genera Deronectes Sharp and Scarodytes Gozis (Coleoptera: Dytiscidae: Hydroporinae), with implications for the phylogeny of the Deronectes-group of genera. Entomologica Scandinavica, 30, 173–195.
Alarie, Y., Nilsson, A.N., Hendrich, L., Watts, C.H.S. & Balke, M. (2000) Larval morphology of four genera of Laccophilinae (Coleoptera: Adephaga: Dytiscidae) with an analysis of their phylogenetic relationships. Insect Systematics & Evolution, 31, 121–164.
Alekseev, M.A., Sinitsina, E.E. & Chaika, S. Yu. (2006) Sensory organs of the antennae and mouthparts of beetle larvae (Coleoptera). Entomological Review, 86, 638–648. (Russian original: Entomologicheskoe Obozrenie, 85 (2006), 508–518).
Angus, R.B. (1970) A revision of the beetles of the genus Helophorus F. (Coleoptera: Hydrophilidae) subgenera Orphelophorus d’Orchymont, Gephelophorus Sharp and Meghelophorus Kuwert. Acta Zoologica Fennica, 129, 1–62.
Archangelsky, M. (1997) Studies on the biology, ecology, & systematics of the immature stages of New World Hydrophiloidea (Coleoptera: Staphyliniformia). Bulletin of the Ohio Biological Survey (New Series), 12(1), ix + 207.
Archangelsky, M. (1998) Phylogeny of Hydrophiloidea (Coleoptera: Staphyliniformia) using characters from adult and preimaginal stages. Systematic Entomology, 23, 9–24.
Archangelsky, M. (1999) Adaptations of immature stages of Sphaeridiinae (Staphyliniformia, Hydrophiloidea: Hydrophilidae) and state of knowledge of preimaginal Hydrophilidae. Coleopterist’s Bulletin, 53, 64–79.
Archangelsky, M (2004). Higher-level phylogeny of Hydrophilinae (Coleoptera: Hydrophilidae) based on larval, pupal and adult characters. Systematic Entomology, 29, 188–214.
Archangelsky, M., Beutel, R.G. & Komarek, A. (2005) Hydrophiloidea. 10.1. Hydrophilidae. In: Beutel, R.G. & Leschen, R.A.B. (Eds.), Coleoptera, Volume 1: Morphology and Systematics (Archostemata, Adephaga, Myxophaga, Polyphaga partim). Handbook of Zoology, Vol. IV: Arthropoda: Insecta). Walter De Gruyter, Berlin, pp. 157–183.
Ashe, J.S. (1985) Structural features and phylogenetic relationships among larvae of genera of gyrophaeninae staphylinids (Coleoptera: Staphylinidae: Aleocharinae). Fieldiana Zoology (N.S.), 30, 1–60.
Ashe, J.S. & Watrous, L.E. (1984) Larval chaetotaxy of Aleocharinae (Staphylinidae) based on a description of Atheta coriaria Kraatz. Coleopterist’s Bulletin, 38, 165–179.
Balduf, W. V. (1935) The bionomics of entomophagous Coleoptera. John S. Swift Co., St. Louis.
Berge Henegouwen, A.L. van (1988) Hydrochus megaphallus, a new and widespread European water beetle described from the Netherlands (Coleoptera, Hydrophilidae). Balfour-Browne Club Newsletter, 42, 18–21.
Bernhard, D., Schmidt, C., Korte, A., Fritzsch, G. & Beutel, R.G. (2006) From terrestrial to aquatic habitats and back again – molecular insights into the evolution and phylogeny of Hydrophiloidea (Coleoptera) using multigene analyses. Zoologica Scripta, 35, 597–606.
Beutel, R.G. (1994) Phylogenetic analysis of Hydrophiloidea based on characters of the head of adults and larvae (Coleoptera: Staphyliniformia). Koleopterologische Rundschau, 64, 103–131.
Beutel, R.G. (1999) Morphology and evolution of the larval head of Hydrophiloidea and Histeroidea (Coleoptera: Staphyliniformia). Tijdschrift voor Entomologie, 142, 9–30.
Beutel, R.G. & Leschen, R.A.B. (2005) Phylogenetic analysis of Staphyliniformia (Coleoptera) based on characters of larvae and adults. Systematic Entomology, 30, 510–548.
Bousquet, Y. & Goulet, H. (1984) Notation of primary setae and pores on larvae of Carabidae (Coleoptera: Adephaga). Canadian Journal of Zoology, 62, 573–588.
Bousquet, Y. & Tchang, J.P. (1992) Anisodactyline larvae (Coleoptera: Carabidae: Harpalini): descriptions of genus-group taxa of Eastern Canada and phylogenetic remarks. Canadian Entomologist., 124, 751–783.
Böving, A.G. & Henriksen, K.L. (1938) The developmental stages of the Danish Hydrophilidae (Ins., Coleoptera). Videnskabelige Meddedelerser dansk naturhistorisk Forening i Kjobenhaven, 102, 27–162.
Caterino, M. & Tishechkin, A.K. (2006) DNA identification and morphological description of the first confirmed larvae of Hetaeriinae (Coleoptera: Histeridae). Systematic Entomology, 31, 405–418.
Chapman, R.F. (1998) The Insects. Structure and Function. 4th edition. Cambridge University Press, Cambridge.
Charpentier, T. de (1825) Horae Entomologicae. A. Gosohorsky, Wratislav.
Delgado, J.A. & Archangelsky, M. (2005) Description of the larval stages of Gymnochthebius jensenhaarupi and phylogenetic analysis of the relationships with other species of the subfamily Ochthebiinae (Coleoptera: Hydraenidae). European Journal of Entomology, 102, 231–240.
Delgado. J.A. & Soler, A.G. (1997a) Morphology and chaetotaxy of larval Hydraenidae (Coleoptera) I: genus Limnebius Leach, 1815 based on a description of Limnebius cordobanus d’Orchymont. Aquatic Insects, 19, 37–49.
Delgado, J.A. & Soler, A.G. (1997b) Morphology and chaetotaxy of larval Hydraenidae (Coleoptera) II: the subgenus Ochthebius s.str. Leach. Bulletin de l’Institut Royal des Sciences Naturelles de Belgique, Entomologie, 67, 45–55.
Fabricius, J.C. (1775) Systema Entomologiae. Libraria Korte, Fensburg & Lipsia.
Fabricius, J.C. (1792) Entomologica Systematica, Vol. 1, Pars 1. C. G. Proft, Hafnia.
Fabricius, J.C. (1801) Systema Eleutheatorum, Vol. 1. Bibliopoli Academici Novi, Kilia.
Fikáček, M. (2006) Primary chaetotaxy of the larval head of the hydrophiloid beetles (Coleoptera : Hydrophiloidea). M.Sc. thesis, Department of Zoology, Faculty of Science, Charles University in Prague, Praha.
Frania, H. (1986) Larvae of Eustilicus Sharp, Rugilus Leach, Deroderus Sharp, Stilocharis Sharp, and Medon Stephens (Coleoptera: Staphylinidae: Paederinae: Paederini), and their phylogenetic significance. Canadian Journal of Zoology, 64, 2543–2557.
Germar, E.F. (1824) Insectorum species novae aut minus cognitae, descriptionibus illustratae. Vol. 1. Coleoptera. J. C. Hendel et fil, Hala.
Grebennikov, V.V. (2004) Review of larval morphology of beetles of the suborder Archostemata (Insecta: Coleoptera), including first-instar chaetotaxy. European Journal of Entomology, 102, 273–292.
Grebennikov, V.V. & Beutel, R.G. (2002) Morphology of the minute larva of Ptinella tenella, with special reference to effects of miniaturisation and the systematic position of Ptiliidae (Coleoptera: Staphylinoidea). Arthropod Structure & Development, 31, 157–172.
Gredler, V.M. (1863) Die Käfer von Tirol nach ihrer horizontalen vaertikalen Verbreitung verzeichnet. I. Hälfte: Cicindelidae-Dascillidae. J. Eberle’sche Buchdruckerei, Bozen.
Hanley, R.S. & Goodrich, M.A. (1994) Natural history, development and immature stages of Oxyporus stygicus Say (Coleoptera: Staphylinidae: Oxyporinae). Coleopterist’s Bulletin, 48, 213–225.
Hansen, M. (1991) The hydrophiloid beetles: Phylogeny, classification and a revision of the genera (Coleoptera, Hydrophiloidea). Biologiske Skrifter, 40, 1–367.
Hansen, M. (1999). Hydrophiloidea (Coleoptera) World Catalogue of Insects, 2, 1–416.
Hansen, M. & Richardson, B.A. (1998) A new species of Omicrus Sharp (Coleoptera: Hydrophilidae) from Puerto Rico and its larva, the first known larva of Omicrini. Systematic Entomology, 23: 1–8.
Herbst, J.F.W. (1797) Natursystem aller bekannten in- und ausländischen Insekten, Käfer 7. Pauli, Berlin.
Hoffman. K.V. & Brushwein, J.R. (1992) Descriptions of the larvae and pupae of some North American Mantispinae (Neuroptera: Mantispidae) and development of a system of larval chaetotaxy for Neuroptera. Transactions of the American Entomological Society, 118, 159–196.
Kilian, A. (1998) Morphology and phylogeny of the larval stages of the tribe Agathidini (Coleoptera: Leiodidae: Leiodinae). Annales Zoologici (Warszawa), 48, 125–220.
Kilian, A. (2001) A description of the larva of Agathidium kumaonicum Angelini et De Marzo, 1985 (Coleoptera Leiodidae). Genus, 12, 563–575.
Kilian, A. (2007) Comparative morphology of larval Camiarinae (Coleoptera: Leiodidae). Part I. Genus Paragyrtodes Szymczakowski 1966. Zootaxa, 1640, 1–39.
Kiselyova, T. & McHugh, J. V. (2006) A phylogenetic study of Dermestidae (Coleoptera) based on larval morphology. Systematic Entomology, 31, 469–507.
Knisch, A. (1925) Coleoptera aus dem tropischen Amerika. Hydrophilidae (Palpicornia). Bolletino dei Musei di Zoologia e di Anatomia comparata della R. Universita di Torino (N. S.), 30 (1924), 1–12.
Kovarik, P.W. & Passoa, S. (1993) Chaetotaxy of larval Histeridae (Coleoptera: Hydrophiloidea) based on a description of Onthophilus nodatus LeConte. Annals of the Entomological Society of America, 86, 560–576.
Leach, W.E. (1815) Entomology. In: Brewster, D. (Ed.), Edinburgh Encyclopaedia, Vol. 9. Balfour, Edinburgh, pp. 57–172.
LeConte, J.E. (1845) A monography of the North American Histeroides. Boston Journal of Natural History, 5 (1845–1847), 32–86.
Lee, C.-F., Satô, M., Shepard, W.D. & Jäch, M.A. (2007) Phylogeny of Psephenidae (Coleoptera: Byrrhoidea) based on larval, pupal and adult characters. Systematic Entomology, 32, 502–538.
Linnaeus, C. (1758) Systema Naturae, Vol. 1 (10. edition). Laur. Salvi, Holmia.
Makarov, K.V. (1996) Patterns of chaetome modifications in ground-beetle larvae (Coleoptera: Carabidae). Acta Societatis Zoologicae Bohemicae, 60, 391–413.
Marvaldi, E. (1999) Morfología larval en Curculionidae (Insecta: Coleoptera). Acta Zoologíca Lilloana, 45, 7–24.
Marvaldi, E. (2003) Key to larvae of the South American subfamilies of weevils (Coleoptera, Curculinoidea). Revista Chilena de Historia Natural, 76, 603–612.
May, B.M. (1964) Immature stages of Curculionidae. 1. Some genera in the tribe Araucariini (Cossoninae). New Zealand Journal of Science, 10, 644–660.
May, B.M. (1994) An introduction to the immature stages of Australian Curculionoidea. In: Zimmerman, E. C. (Ed.), Australian Weevils (Coleoptera: Curculionoidea), Vol. 2. CSIRO, East Melbourne, pp. 365–728.
Mazur, S. (1972) Systematic and synonymic notes upon certain species of Histeridae (Coleoptera). Polske Pismo Entomologiczne, 42, 137–143.
Micó, E., Morón, M.Á., Šípek, P. & Galante, E. (2008) Larval morphology enhances phylogenetic reconstruction in Cetoniidae (Coleoptera: Scarabaeoidea) and allows the interpretation of the evolution of larval feeding habits. Systematic Entomology, 33, 128–144.
Miller, D.C. (1963) The biology of Hydrophilidae. The Biologist, 45, 33–38.
Moulins, M. (1959) Contribution à la connaissance de quelques types larvaires d’Hydrophilidae (Coléoptères). Travaux du Laboratoire de Zoologie et de la Station Aquicole Grimaldi de la Faculté des Sciences de Dijon, 30, 1–46 + i–v + 42 Figs.
Müller, O. F. (1776) Zoologicae Danicae Prodromus, seu Animalium, Daniae et Norvegiae Indigenarum. Hallger, Havnia.
Nichols, S.W., Schuh, R.T. & Torre-Bueno, J.R. de la (1989) The Torre-Bueno Glossary of Entomology. American Museum of Natural History, New York.
Ratajczak, A. (1995) Redescription of the third larval instar of Anisotoma humeralis (F.) (Coleoptera: Leiodidae). Genus, 6, 277–288.
Ratajczak, A. (1996) Description of third larval stage of Anisotoma orbicularis (Herbst) and redescription of the third larval stage of A. castanea (Herbst) (Coleoptera: Leiodidae). Genus, 7, 351–378.
Remane A. (1956) Die Grundlagen des natürlichen Systems, der vergleichenden Anatomie und Phylogenetik. Akademische Verlagsgesselschaft, Leipzig.
Richmond, E.A. (1920) Studies on the biology of the aquatic Hydrophilidae. Bulletin of the American Museum of Natural History, 42, 1–93 + 16 pls.
Say, T. (1825) Descriptions of new species of Coleopterous Insects inhabiting the United States. Journal of the Academy of Natural Sciences of Philadelphia, 5, 160–204.
Schaller, J.G. (1783) Neue Insekten. Abhandlungen der Hallischen naturforschenden Gesellschaft, 1, 217–332.
Short, A.E.Z. & Hebauer, F. (2006) World Catalogue of Hydrophiloidea – additions and corrections, 1 (1999–2005) (Coleoptera). Koleopterologische Rundschau, 76, 315–359.
Skuhrovec, J. (2006) Identification of instars of Hypera postica using chaetotaxy. Journal of Economic Entomology., 99, 2216–2218.
Skuhrovec, J. (2007) Descriptions of larvae of the tribe Hyperini (Coleoptera: Curculionidae): III. Mature larvae of the genus Donus. Zootaxa, 1606, 1–28.
Solodovnikov, A.Y. (2007) Larval chaetotaxy of Coleoptera (Insecta) as a tool for evolutionary research and systematics: less confusion, more clarity. Journal of Zoological Systematics and Evolutionary Research, 45, 120–127.
Solodovnikov, A.Y. & Newton, A.F. (2005) Phylogenetic placement of Arrowinini trib.n. within the subfamily Staphylininae (Coleoptera: Staphylinidae), with revision of the relict South African genus Arrowinus and description of its larva. Systematic Entomology, 30, 398–441.
Spangler, P.J. (1991) Hydrophilidae. In: Stehr, F. E. (Ed.), Immature Insects, Volume 2. Kendall/Hunt Publishing Co. Dubuque, Iowa, pp. 355–359.
Stephens, J.F. (1829) Illustrations of British Entomology. Mandibulata, Vol. 2. Baldwin and Cradock, London.
Thayer, M.K., Ashe, J.S. & Hanley, R.S. (2004) Discovery of the remarkable larvae of Hopladriini (Coleoptera: Staphylinidae: Aleocharinae). Annals of the Entomological Society of America, 97, 624–634.
Thomson, C.G. (1859) Scandinaviens Coleoptera. Vol. 1. Berlingska Boktryckeriet, Lund.
Valladares, L.F. (1994) Description de la larva de Laccobius canariensis Orchymont, 1940 (Coleoptera: Hydrophilidae). Nouvelle Revue d’Entomologie (N.S.), 11, 117–121.
Wheeler, Q.D. (1990) Morphology and ontogeny of postembryonic larval Agathidium and Anisotoma (Coleoptera: Leiodidae). American Museum Novitates, 2986, 1–46.
Wilson, C.B. (1923a) Life history of the scavenger water beetle Hydrous (Hydrophilus) triangularis and its economic relation to fish breeding. Bulletin of the Bureau of Fisheries, 39, 9–38.
Wilson, C.B. (1923b) Water beetles in relation to pond fish culture, with life histories of those found in fishponds at Fairport, Iowa. Bulletin of the Bureau of Fisheries, 39, 231–345.