Abstract
The larvae of 13 species (six epigaeic and seven hypogaeic) of the Australian endemic genus Paroster Sharp, 1882 are described with an emphasis on chaetotaxy of the head capsule, head appendages, legs, last abdominal segment and urogomphi. A cladistic analysis of 41 Hydroporinae species and 63 informative characters supports a monophyletic origin of members of Paroster, which share a labial palpus comprised of three palpomeres and the absence of the primary seta FE7. Contrary to their epigaeic counterparts, larvae of hypogaeic Paroster species have turned out to be very divergent morphologically. In addition to the common characteristics associated with hypogaeic living (i.e., absence of eye, reduced pigmentation, and thin or soft exoskeleton), larvae of these species have undergone a relative elongation/ enlargement of the head capsule and a more elongate and narrower mandible and have developed a variable number of secondary temporal spines. Compared to the other stygobitic species studied, larvae of Paroster hinzeae (Watts & Humphreys, 2001), P. macrosturtensis (Watts & Humphreys, 2006), P. stegastos (Watts & Humphreys, 2003) and P. darlotensis (Watts & Humphreys, 2003) evolved a disproportionately large head capsule relative to body size. Larvae of P. wedgeensis (Watts & Humphreys, 2003) and P. mesosturtensis (Watts & Humphreys, 2006) are deemed to have deviated the least from the hypothetical epigaeic Paroster groundplan.References
Alarie, Y. (1991) Primary setae and pores on the cephalic capsule and head appendages of larval Hydroporinae (Coleoptera: Dytiscidae). Canadian Journal of Zoology, 69, 2255–2265.
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. & Harper, P.-P. (1990) Primary setae and pores on the last abdominal segment and the urogomphi of larval Hydroporinae (Coleoptera: Adephaga: Dytiscidae), with notes on other dytiscid larvae. Canadian Journal of Zoology, 68, 368–374.
Alarie, Y., Harper, P.-P. & Maire, A. (1990) Primary setae and pores on legs of larvae of Nearctic Hydroporinae (Coleoptera: Dytiscidae). Quaestiones Entomologicae, 26, 199–210.
Alarie, Y. & Michat, M.C. (2007a) Phylogenetic analysis of Hydroporinae (Coleoptera: Dytiscidae) based on larval morphology, with description of first-instar of Laccornellus lugubris. Annals of the Entomological Society of America, 100, 655–665.
Alarie, Y. & Michat, M.C. (2007b) Primary setae and pores on the maxilla of larvae of the subfamily Hydroporinae (Coleoptera: Adephaga: Dytiscidae): ground plan pattern reconsidered. The Coleopterists Bulletin, 61, 310–317.
Alarie, Y. & Wewalka, G. (2001) Description of the mature larva of Glareadessus stocki Wewalka and Biström (Coleoptera: Dytiscidae), a stygobiontic Bidessini from the Persian Gulf region. The Coleopterists Bulletin, 55, 144–151.
Bertrand, H. (1972) Larves et nymphes des Coléoptères aquatiques du globe. F. Paillart, France. 804 pp.
Clark, H. (1862) Catalogue of the Dytiscidae and Gyrinidae of Australasia, with descriptions of new species. The Journal of Entomology. Descriptive and Geographical, 1, 399–421.
Farris, J.S. (1989) The retention index and the rescaled consistency index. Cladistics, 5, 417–419.
Hendrich, L. & Fery, H. (2008) Paroster baylyi sp. n., P. ursulae sp. n. (Coleoptera: Dytiscidae: Hydroporinae) and the water beetle diversity of pan-gnammas on isolated granite outcrops in the mallee of south-western Australia. Zootaxa, 1704, 27–41.
Kluge, A.G. & Farris, J.S. (1969) Quantitative phyletics and the evolution of anurans. Systematic Zoology, 30, 1–32.
Larson, D.J. & Labonte J.R. (1994) Stygoporus oregonensis, a new genus and species of hypogaeic water beetle (Coleoptera: Dytisicdae: Hydroporini) from the United States. The Coleopterists Bulletin, 48, 371–379.
Lawrence, J.F. (1991) Order Coleoptera. In: Sther, F.W. (ed.), Immature Insects. Volume 2. Kendall/Hunt Publishing Company, Dubuque (IA), pp. 144–658.
Leys, R. & Watts, C.H.S. (2008) Systematics and evolution of the Australian hypogaeic hydroporine diving beetles (Dytiscidae: Coleoptera). Invertebrate Systematics, 22, 1–9.
Macleod, N. & Forey, P.L. (2002) Introduction: morphology, shape and phylogenetics. In: Macleod, N. & Forey, P.L. (eds), Morphology, shape and phylogeny. Taylor & Francis Inc, New York, pp. 1–7.
Maddison, W.P. & Maddison, D.R. (2000) MacClade. Analysis of Phylogeny and /Character Evolution. Sinauer Associates, Inc. Sunderland, Massachusetts.
Michat, M.C., Alarie, Y., Torres, P.L.M. & Megna, Y.S. (2007) Larval morphology of the diving beetle Celina and the phylogeny of ancestral hydroporines (Coleoptera: Dytiscidae: Hydroporinae). Invertebrate Systematics, 21, 239–254.
Michat, M.C. & Torres, P.L.M. (2005) Larval morphology of Macrovatellus haagi (Wehncke) and phylogeny of Hydroporinae (Coleoptera: Dytiscidae). Insect Systematics & Evolution, 36, 199–217.
Nilsson, A.N. (2001) World Catalogue of Insects. Volume 3, Dytiscidae (Coleoptera). Apollo Books, Stenstrup, 395 pp.
Nilsson, A. N. (2003) World Catalogue of Dytiscidae - corrections and additions, 1 (Coleoptera: Dytiscidae). Koleopterologische Rundschau, 73, 65–74.
Nilsson, A. N. (2004) World Catalogue of Dytiscidae - corrections and additions, 2 (Coleoptera: Dytiscidae). Koleopterologische Rundschau, 74, 157–174.
Nilsson, A. N. & Fery, H. (2006) World Catalogue of Dytiscidae - corrections and additions, 3 (Coleoptera: Dytiscidae). Koleopterologische Rundschau, 76, 55–74.
Régimbart, M. (1908) Dytiscidae, Gyrinidae et Hydrophilidae. In: Michaelsen, W. & Hartmeyer, R. (eds), Die Fauna Südwest-Australiens. Ergebnisse der hamburger südwest-australischen Forschungsreise 1905, 1(8), 309–316.
Ribera, I., Vogler, A.P & Balke, M. (2008) Phylogeny and diversification of diving beetles (Coleoptera: Dytiscidae). Cladistics, 24, 563–590.
Sharp, D. (1882) On aquatic carnivorous Coleoptera or Dytiscidae. Scientific Transactions of the Royal Dublin Society, 2(2), 179–1003.
Smrz, J. (1983) Morhological adaptations of sensory organs and wings in hypogaeic water beetles (Coleoptera:, Dytiscidae). Acta Entomologica Bohemoslovaca, 80, 246–255.
Swofford, D.L. (2002) PAUP*: Phylogenetic Analysis Using Parsimony (* and Other Methods). Version 4.0b10. Sinauer Associates, Sunderland.
Watts, C.H.S. (1963) The larvae of Australian Dytiscidae. Transactions of the Royal Society of South Australia, 87, 23–40.
Watts, C.H.S. (1978) A revision of the Australian Dytiscidae (Coleoptera). Australian Journal of Zoology, Supplementary Series No 57, 1–166.
Watts, C.H.S. & Humphreys, W.F. (2001) A new genus and six new species of Dytiscidae (Coleoptera) from underground waters in the Yilgarn palaeodrainage system of Western Australia. Records of the South Australian Museum, 34, 99–114.
Watts, C.H.S. & Humphreys, W.F. (2003) Twenty-five new Dytiscidae (Coleoptera) of the genera Tjirtudessus Watts & Humphreys, Nirripirti Watts & Humphreys and Bidessodes Régimbart from underground waters in Australia. Records of the South Australian Museum, 36, 135–187.
Watts, C.H.S. & Humphreys, W.F. (2004) Thirteen new Dytiscidae (Coleoptera) of the genera Boongurrus Larson, Tjirtudessus Watts & Humphreys and Nirripirti Watts & Humphreys, from underground waters in Australia. Transactions of the Royal Society of South Australia, 128, 99–129.
Watts, C.H.S. & Humphreys, W.F. (2006) Twenty-six new Dytiscidae (Coleoptera) of the genera Limbodessus Guignot and Nirripirti Watts and Humphreys, from underground waters in Australia. Transactions of the Royal Society of South Australia, 130, 123–185.
Watts, C.H.S. & Leys, R. (2008). Review of the epigaeic species of Paroster Sharp, 1882, with descriptions of three new species, and phylogeny based on DNA sequence data of two mitochondrial genes (Coleoptera: Dytiscidae: Hydroporinae). Koleopterologische Rundschau, 78, 9–38.
Watts, C.H.S. & Humphreys, W.F. (2009) Fourteen new Dytiscidae (Coleoptera) of the genera Limbodessus Guignot, Paroster Sharp, and Exocelina Broun from undeground waters in Australia. Transactions of the Royal Society of South Australia, 133, 62–107.
West-Eberhard, M.J. (2003) Developmental plasticity and evolution. Oxford University Press, New York, 794 pp.
Wolfe, G.W. (1985) A phylogenetic analysis of plesiotypic hydroporine lineages with an emphasis on Laccornis Des Gozis (Coleoptera: Dytiscidae). Proceedings of the Academy of Natural Sciences of Philadelphia, 137, 132–155.