Abstract
Recent investigation of one of the larger calcretes in the uppermost reaches of the Carey palaeochannel in the Yilgarn region of Western Australia revealed an unprecedented diversity of copepod crustaceans. Twenty-two different species and subspecies, from six copepod families, represent 70% of the previously recorded copepod α-diversity in the whole region, although the area investigated is less than 3% of its surface. The aims of this study were to explore the diversity of the harpacticoid genus Kinnecaris Jakobi, 1972 using both molecular and morphological methods, establish precise species boundaries, find their accurate area of distribution, reconstruct phylogenetic relationships, and explore colonisation pathways. To achieve this we sampled very intensively in the area, as well as in two neighbouring palaeochannels, analysing more than 700 samples from 230 different localities, half of which contained copepods. Seven species are described here as new, five of them from the Yeelirrie palaeochannel (K. esbe sp. nov., K. lined sp. nov., K. linel sp. nov., K. linesae sp. nov., and K. uranusi sp. nov.) and one each from two neighbouring palaeochannels (K. barrambie sp. nov. and K. lakewayi sp. nov.). Parastenocaris jane Karanovic, 2006 from the Pilbara region, along with a newly described third Australian parastenocaridid genus from the Yilgarn, were used as outgroups in our molecular analysis. The COI fragment was successfully PCR-amplified from 12 parastenocaridid specimens using a nested combination of primers. All analyses supported the presence of at least seven genetically divergent lineages, most supported with very high bootstrap values. Three genera formed three separate clades, and the average pairwise distances between Kinnecaris morpho-taxa were found to be very high (8.2-16.8 %), while the highest divergences within morpho-taxa were 0.3%. Some conflict between molecular phylogenies and morphological data was observed when it came to recognizing different groups of species. While morphology indicates that K. esbe, K. linel, and K. uranusi represent a group of very closely related species, supported by a number of synapomorphies, molecular analyses suggest that K. linel and K. uranusi are only remotely related. We argue in favor of morphological data, until more markers can be studied to try to resolve these differences. In Yeelirrie, morphological evidence would suggest a downstream colonisation history in the genus Kinnecaris, where the most plesiomorphic form (K. linesae) lives in the uppermost reaches of the palaeochannel, and the trend in the caudal rami elongation and denser somite ornamentation is obvious downstream the palaeochannel (K. uranusi, K. linel, and then K. esbe), with the only exception being K. lined, which probably represents an independent colonisation event. Parastenocarids are copepods of freshwater origin, and we argure that they can probably disperse downstream during periods of increased rainfall, evolving into separate species in isolated calcrete pockets during periods of increased aridity. Although some of the questions remained unanswered in this study, detailed morphological and molecular observations indicate that we are not dealing with one widely distributed and variable species in the Yilgarn region, but rather with a complex of short range endemics. Areas of distribution for different species range from 30 km to less than 5 km in diameter. Very strong seasonal dynamics in this subterranean community was observed, and this is a novel concept for these ecosystems globally. A key to nine Australian species of Kinnecaris is also included.
References
Adamowicz, S.J., Manu-Marque, S., Hebert, P.D.N. & Purvis, A. (2007) Molecular systematic and patterns of morphological evolution in the Centropagidae (Copepoda: Calanoida) of Argentina. Biological Journal of the Linnean Society, 90, 279–292.
Allford, A., Cooper, S.J.B., Humphreys, W.F. & Austin, A.D. (2008) Diversity and distribution of groundwater fauna in a limestone aquifer: does sampling alter the story? Invertebrate Systematics, 22, 127–138.
Beard, J.S. (1976) Vegetation survey of Western Australia, Murchison. University of Western Australia Press, Perth, 141 pp.
Beard, J.S. (1998) Position and development history of the central watershed of the Western Shield, Western Australia. Journal of the Royal Society of Western Australia, 81, 157–164.
Berera, R. & Cottarelli, V. (2003) Two new species of interstitial harpacticoids from southern Italy and proposal of a new genus Parastenocaris species-group. Italian Journal of Zoology, 70, 261–268.
Boxshall, G.A. & Halsey, S.H. (2004) An Introduction to Copepod Diversity. The Ray Society, London, 966 pp.
Boxshall, G.A. & Jaume, D. (2000) Making waves: the repeated colonization of freshwater by copepod crustaceans. Advances in Ecological Research, 31, 61–79.
Bradford, T., Adams, M., Humphreys, W.F., Austin, A.D. & Cooper, S.J.B. (2010) DNA barcoding of stygofauna uncovers cryptic amphipod diversity in a calcrete aquifer in Western Australia’s arid zone. Molecular Ecology Resources, 10, 41–50.
Chappuis, P.A. (1936) Crustacea. III. Copepoda: Harpacticoida. Mission Scientifique de l'Omo. Mémoires du Muséum National d'Histoire Naturelle, Nouvelle Série, 3(29), 245–292.
Chappuis, P.A. (1937) Weitere subterrane Harpacticoiden aus Jugoslavien. Bulletin de la Société des Sciences de Cluj, Roumanie, 8, 503–532.
Chappuis, P.A. (1952) Copépodes harpacticoides psammiques de Madagascar. Mémoires de L’Institut Scientifique de Madagascar, Série A, 7, 145–160.
Chappuis, P.A. (1954) Recherches sur la faune interstitielle des sediments marins et d’eau douce a Madagascar. Mémoires de L’Institut Scientifique de Madagascar, Série A, 9, 45–73.
Chappuis, P.A. (1955) Harpacticoides psammiques du Lac Tanganika. Revue de Zoologie et de Botanique Africaines, 51, 68–80.
Cho, J-L., Humphreys, W.F. & Lee, S.-D. (2006a) Phylogenetic relationships within the genus Atopobathynella Schminke, 1973 (Bathynellacea, Parabathynellidae): with the description of six new species from Western Australia. Invertebrate Systematics, 20, 9–41.
Cho, J-L., Park, J-G. & Ranga Reddy, Y. (2006b) Brevisomabathynella gen. nov. with two new species from Western Australia (Bathynellacea, Syncarida): the first definitive evidence of predation in Parabathynellidae. Zootaxa, 1247, 25–42.
Clarke, J.D.A. (1994) Evolution of the Lefroy and Cowan palaeodrainage channels, Western Australia. Australian Journal of Earth Sciences, 41, 55–68.
Coineau, N. (2001) Syncarida. In: Juberthie C. & Decu, V. (Eds.), Encyclopedia Biospeologica. Societé de Biospéologica and Acdémie Roumanie, Moulis & Bucharest, pp. 863–876.
Cooper, S.J.B, Bradbury, J.H., Saint, K.M., Leys, R., Austin, A.D. & Humphreys, W.F. (2007) Subterranean archipelago in the Australian arid zone: mitochondrial DNA phylogeography of amphipods from central Western Australia. Molecular Ecology, 16, 1533–1544.
Cooper, S.J.B., Hinze, S., Leys, R., Watts, C.H.S. & Humphreys, W.F. (2002) Islands under the desert: molecular systematic and evolutionary origin of stygobitic water beetles (Coleoptera: Dytiscidae) from central Western Australia. Invertebrate Systematics, 16, 589–598.
Cooper, S.J.B., Saint, K.M., Taiti, S., Austin, A.D. & Humphreys, W.F. (2008) Subterranean archipelago: mitochondrial DNA phylogeography of stygobitic isopods (Oniscidea: Haloniscus) from the Yilgarn region of Western Australia. In: Austin, A.D., Cooper, S.J.B. & Humphreys, W.F. (Eds.) Subterranean Connections: Biology and evolution in troglobiont and groundwater ecosystems. Invertebrate Systematics, 22, 195–203.
Corgosinho, P.H.C. & Martínez Arbizu, P. (2005) Two new interstitial species of Remaneicaris Jakobi (Copepoda, Harpacticoida, Parastenocarididae) from the Ribeirão do Ouro River, Brazil, with a redefinition of the genus. Senckenbergiana Biologica, 85, 147–162.
Corgosinho, P.H.C., Martínez Arbizu, P. & dos Santos-Silva, E.N. (2007) Three new species of Remaneicaris Jakobi, 1972 (Copepoda, Harpacticoida, Parastenocarididae) from the Ribeirão do Ouro River, Minas Gerais, Brazil, with some remarks on the groundpattern of the Parastenocarididae. Zootaxa, 1437, 1–28.
Corgosinho, P.H.C., Martínez Arbizu, P. & dos Santos-Silva, E.N. (2010) Revision of Brasilibathynellocaris Jakobi, 1972 (Copepoda: Harpacticoida: Parastenocarididae) with redefinition of the genus. Zoological Journal of the Linnean Society, 159, 527–566.
Cottarelli, V. & Bruno, M.C. (1994) Parastenocaris lyncaea n. sp. and Stygoelaphoidella africana n. sp. from interstitial continental waters of Sierra Leone. Ricerche Biologiche in Sierra Leone, parte IV, Accademia Nazionale dei Lincei, 267, 97–109.
Cottarelli, V. & Bruno, M.C. (1995) First record of Parastenocarididae (Crustacea, Copepoda, Harpacticoida) from subterranean waters of Ethiopia and the description of three new species. Journal of African Zoology, 109, 467–479.
Cottarelli, V., Bruno, M.C. & Berera, R. (2006) A new species of Parastenocaris from Mindoro Island, Philippines: Parastenocaris distincta sp. nov. (Crustacea: Copepoda: Harpacticoida: Parastenocarididae). Zootaxa, 1368, 57–68.
Cottarelli, V., Bruno, M.C. & Berera, R. (2007) Interstitial harpacticoids from groundwater in Tuscany (Central Italy): Parastenocaris reidae sp. nov., Nitocrella ensifera sp. nov., and notes on the morphology of Parastenocaris cf. glacialis Noodt (Crustacea: Copepoda). Italian Journal of Zoology, 74, 83–99.
Cottarelli, V., Bruno, M.C. & Berera, R. (2008) Two new species of Parastenocaris (Copepoda, Harpacticoida, Parastenocarididae) from groundwater of Sardinia and Sicily. Crustaceana, 81, 537–555.
Cottarelli, V., Bruno, M.C. & Berera, R. (2010) First record of Parastenocarididae from Thailand and description of a new genus (Copepoda: Harpacticoida). Journal of Crustacean Biology, 30, 478–494.
Culver, D.C., Kane, T. & Fong, D.W. (1995) Adaptation and Natural Selection in Caves: The Evolution of Gammarus minus. Harvard University Press, Cambridge, 223 pp.
Culver, D. & Pipan, T. (2009) The biology of caves and other subterranean habitats. Oxford University Press, Oxford, 256 pp.
Dahms, H.U. (1988) Development of functional adaptation to clasping behaviour in harpacticoid copepods (Copepoda, Harpacticoida). Hydrobiologia, 167/168, 505–513.
Dussart, B.H. (1979) Algunos copepodos de America del Sur. Publicación Ocasional, Museo Nacional de Historia Natural, Santiago, 30, 1–13.
Dussart, B. & Defaye, D. (1990) Répertoire mondial des crustacés copépodes des eaux intérieures, III. Harpacticoïdes. Crustaceana supplement, 16, 1–384.
Felsenstein, J. (1985) Confidence limits on phylogenies: an approach using the bootstrap. Evolution, 39, 783–791.
Finston, T.L., Johnson, M.S., Humphreys, W.F., Eberhard, S. & Halse, S. (2007) Cryptic speciation in two widespread subterranean amphipod genera reflects historical drainage patterns in an ancient landscape. Molecular Ecology, 16, 355–365.
Folmer, O., Black, M., Hoeh, W., Lutz, R. & Vrijenoek, R. (1994) DNA primers for amplification of mitochondrial cytochrome c oxidase subunit 1 from diverse metazoan invertebrates. Molecular Marine Biology and Biotechnology, 3, 294–299.
Frakes, L.A. (1999) Evolution of Australian environments. Flora of Australia (2nd edition), 1, pp. 163–203.
Fryer, G. (1956) New species of cyclopoid and harpacticoid copepods from sandy beaches of Lake Nyasa. Annals and Magazine of Natural History, Serie 12, 9, 225–249.
Galassi, D.M.P. & De Laurentiis, P. (2004) Towards a revision of the genus Parastenocaris Kessler, 1913: establishment of Simplicaris gen. nov. from groundwaters in central Italy and review of the P. brevipes-group (Copepoda, Harpacticoida, Parastenocarididae). Zoological Journal of the Linnean Society, 140, 417–436.
Gibert, J., Danielopol, D.L. & Stanford, J.A. (1994) Groundwater Ecology. Academic Press, London, 571 pp.
Giribet , G. & Edgecombe, G.D. (2006) The importance of looking at small-scale patterns when inferring Gondwanan biogeography: a case study of the centipede Paralamyctes (Chilopoda, Lithobiomorpha, Henicopidae). Biological Journal of the Linnean Society, 89, 65–78.
Glatzel, T. (1996) Mating behaviour of the groundwater copepod Parastenocaris phyllura Kiefer, 1938 (Copepoda: Harpacticoida). Contributions to Zoology, 66, 103–108.
Guzik, M., Abrams, K.M., Cooper, S.J.B., Humphreys, W.F., Cho, J.-L. & Austin, A.D. (2008) Phylogeography of the ancient Parabathynellidae (Crustacea: Bathynellacea) from the Yilgarn region of Western Australia. In: Austin, A.D., Cooper, S.J.B. & Humphreys, W.F. (Eds.) Subterranean Connections: Biology and evolution in troglobiont and groundwater ecosystems. Invertebrate Systematics, 22, 205–216.
Guzik, M.T., Austin, A.D., Cooper, S.J.B., Harvey, M.S., Humphreys, W.F., Bradford, T., Eberhard, S.M., King, R.A., Leys, R., Muirhead, K.A. & Tomlinson, M. (2011) Is the Australian subterranean fauna uniquely diverse? Invertebrate Systematics (in press 10/1/2011).
Harvey, M.S. (2002) Short-range endemism among the Australian fauna: some examples from non-marine environments. Invertebrate Systematics, 16, 555–570.
Hasegawa, M., Kishino, H. & Yano, T. (1985) Dating of the human-ape splitting by a molecular clock of mitochondrial DNA. Journal of Molecular Evolution, 22, 160–174.
Hebert, P.D.N., Cywinska, A., Ball, S.L. & deWaard, J.R. (2003) Biological identifications through DNA barcodes. Proceedings of the Royal Society of London, 270, 313–321.
Holmgren, M., Stapp, P., Dickman, C.R., Gracia, C., Graham, S., Gutiérrez, J.R., Hice, C., Jaksic, F., Kelt, D.A., Letnic, M., Lima, M., López, B.C., Meserve, P.L., Milstead, W.B., Polis, G.A., Previtali, M.A., Richter, M., Sabate, S. & Squeo, F.A. (2006) Extreme climatic events shape arid and semiarid ecosystems. Frontiers in Ecology and the Environment, 4, 87–95.
Humphreys, W.F. (2000) Background and glossary. In: Wilkens, H., Culver, D.C. & Humphreys, W.F. (Eds.), Ecosystems of the World, 30, Subterranean ecosystems, Elsevier, Amsterdam, pp. 3–14.
Humphreys, W.F. (2001) Groundwater calcrete aquifers in the Australian arid zone: the context to an unfolding plethora of stygal biodiversity. Records of the Western Australian Museum, Supplement 64, 63–83.
Humphreys, W.F. (2006) Aquifers: the ultimate groundwater-dependent ecosystems. Australian Journal of Botany, 54, 115–132.
Humphreys, W.F. (2008) Rising from Down Under: developments in subterranean biodiversity in Australia from a groundwater perspective. In: Austin, A.D., Cooper, S.J.B. & Humphreys, W.F. (Eds.) Subterranean Connections: Biology and evolution in troglobiont and groundwater ecosystems. Invertebrate Systematics, 22, 85–101.
Humphreys, W.F., Watts, C.H.S., Cooper, S.J.B., Leijs, R. & Bradbury, J.H. (2009) Groundwater estuaries of saltlakes: buried pools of endemic biodiversity on the Western Plateau, Australia. Hydrobiologia, 626, 79–95.
Huys, R. (2009) Unresolved cases of type fixation, synonymy and homonymy in harpacticoid copepod nomenclature (Crustacea: Copepoda). Zootaxa, 2183, 1–99.
Huys, R. & Boxshall, G.A. (1991) Copepod Evolution. The Ray Society, London, 468 pp.
ICZN (1999) Fourth edition. The International Trust for Zoological Nomenclature, London.
Jakobi, H. (1969) Forficatocaris noodti n. gen., n. sp. (Copepoda, Harpacticoidae) aus brasilianischem Limnopsammal. Crustaceana, 17, 231–238.
Jakobi, H. (1972) Trends (Enp.P4 Mänchen) innerhalb der Parastenocarididen (Copepoda Harpacticoidea). Crustaceana, 22, 127–146.
Johnson, S.L., Commander, D.P. & O’Boy, C.A. (1999) Groundwater resources of the Northern Goldfields, Western Australia. Water and Rivers Commission, Hydrogeological Record Series, Report HG 2, 57pp.
Karanovic, I. (2007) Candoninae ostracods from the Pilbara region in Western Australia. Crustaceana Monographs, 7, Brill, Leiden, 432 pp.
Karanovic, T. (2004) Subterranean Copepoda from arid Western Australia. Crustaceana Monographs, 3, Brill, Leiden, 366 pp.
Karanovic, T. (2005) Two new subterranean Parastenocarididae (Crustacea, Copepoda, Harpacticoida) from Western Australia. Records of the Western Australian Museum, 22, 353–374.
Karanovic, T. (2006) Subterranean Copepods (Crustacea, Copepoda) from the Pilbara Region in Western Australia. Records of the Western Australian Museum, Supplement, 70, 1–239.
Karanovic, T. (2008) Marine Interstitial Poecilostomatoida and Cyclopoida (Copepoda) of Australia. Crustaceana Monographs, 9, Brill, Leiden, 331 pp.
Karanovic, T. (2010) First record of the harpacticoid genus Nitocrellopsis (Copepoda, Ameiridae) in Australia, with descriptions of three new species. International Journal of Limnology, 46, 249–280.
Karanovic, T. & Bobic, M. (1998) Two new species of Copepoda Harpacticoida from east Serbia (Balkan Peninsula): Parastenocaris serbica sp. n. and Bryocamptus (R.) borus sp.n. Crustaceana, 71, 171–184.
Karanovic, T. & Cooper, S.J.B. (in press) Third genus of parastenocaridid copepods from Australia supported by molecular evidence (Harpacticoida: Parastenocarididae). Crustaceana Monographs, 16.
Karanovic, T. & Cooper, S.J.B. (submitted) Explosive radiation of the genus Schizopera Sars (Copepoda: Harpacticoida) in a small subterranean island in Western Australia; Unraveling the cases of cryptic speciation, size differentiation, and multiple invasions. Organisms Diversity & Evolution.
Karanovic, T. & Hancock, P. (2009) On the diagnostic characters of the genus Stygonitocrella (Copepoda, Harpacticoida), with descriptions of seven new species from Australian subterranean waters. Zootaxa, 2324, 1–85.
Karanovic, T., Eberhard, S.M. & Murdoch, A. (2011) A cladistic analysis and taxonomic revision of Australian Metacyclops and Goniocyclops, with description of four new species and three new genera (Copepoda, Cyclopoida). Crustaceana, 84, 1–67.
Kumar, S., Dudley, J., Nei, M. & Tamura, K. (2008) MEGA: A biologist-centric software for evolutionary analysis of DNA and protein sequences. Briefings in Bioinformatics, 9, 299–306.
Kunz, H. (1938). Harpacticoiden vom Sandstrand der Kurischen Nehrung. Kieler Meeresforschung, 3, 148–157.
Lang, K. (1948) Monographie der Harpacticiden,1-2. Nordiska Bokhandeln, Lund, 1682 pp.
Langecker, T.G. (2000) The effect of continuous darkness on cave ecology and cavernicolous evolution. In: Wilkens, H., Culver, D.C. & Humphreys, W.F. (Eds.), Ecosystems of the World, 30, Subterranean ecosystems, Elsevier, Amsterdam, pp. 135–157.
Lefébure, T., Douady, C.J., Gouy, M. & Gibert, J. (2006) Relationship between morphological taxonomy and molecular divergence within Crustacea: Proposal of a molecular threshold to help species delimination. Molecular Phylogenetics and Evolution, 40, 435–447.
Lescher-Moutoué, F. (1973) Sur la biologie et l’écologie des copepods cyclopides hypogés (Crustacés). Annales de Spéléologie, 28, 429–674.
Leys, R. & Watts, C.H. (2008) Systematics and evolution of the Australian subterranean hydroporine diving beetles (Dytiscidae), with notes on Carabhydrus. In: Austin, A.D., Cooper, S.J.B. & Humphreys, W.F. (Eds.) Subterranean Connections: Biology and evolution in troglobiont and groundwater ecosystems. Invertebrate Systematics, 22, 217–225.
Leys, R., Watts, C.H.S., Cooper, S.J.B. & Humphreys, W.F. (2003) Evolution of subterranean diving beetles (Coleoptera: Dytiscidae: Hydroporini, Bidessini) in the arid zone of Australia. Evolution, 57, 2819–2834.
Martínez Arbizu, P. & Moura, G. (1994) The phylogenetic position of the Cylindropsyllinae Sars (Copepoda, Harpacticoida) and the systematic status of the Leptopontiinae Lang. Zoologische Beiträge, 35, 55–77.
Noodt, W. (1962) Limnisch-subterrane Copepoden der Gattung Parastenocaris Kessler aus Mittelamerika. In: Titschack, E. & Koepcke, H.-W. (Eds.). Beiträge zur neotropischen Fauna 2, VEB Gustav Fischer Verlag, Jena, pp 223–248.
Noodt, W. (1963) Subterrane Crustaceen der zentralen Neotropis. Zoologischer Anzeiger, 171, 114–147.
Noodt, W. (1972) Drei neue Parastenocaris aus Kolumbien (Crustacea Copepoda), 1. Mitteilung über kolumbianische Grundwasser-Crustaceen. Studies on Neotropical Fauna, 7, 101–112.
Pipan, T. & Brancelj, A. (2003) Fauna of epikarst - Copepoda (Crustacea) in percolation water of caves in Slovenia. Annales Series Historia Naturalis, Koper, 13, 223–228.
Pipan, T. & Brancelj, A. (2004) Distribution patterns of copepods (Crustacea: Copepoda) in percolation water of the Postojnska Jama cave system (Slovenia). Zoological Studies, 43, 206–210.
Playford, P. (2003) The Permo-Carboniferous glaciations of Gondwana: its impact on Western Australia. Western Wildlife, 7, 1–5.
Por, F.D. & Hadel, V.F. (1986) Two new species of Attheyella (Copepoda: Harpacticoida: Canthocamptidae) from bromeliads of the Serra da Juréia (São Paulo, Brazil). Journal of Crustacean Biology, 6, 777–788.
Ranga Reddy, Y. (2001) Discovery of Parastenocarididae (Copepoda, Harpacticoida) in India, with the description of three new species of Parastenocaris Kessler, 1913, from the River Krishna at Vijayawada. Crustaceana, 74, 705–733.
Ranga Reddy, Y. & Defaye, D. (2007) Parastenocarididae (Crustacea, Copepoda, Harpacticoida) of India: description of Parastenocaris mahanadi n. sp., and redescription of P. curvispinus Enckell, 1970 from hyporheic habitats. Zootaxa, 1580, 1–26.
Ranga Reddy, Y. & Defaye, D. (2009) Two new Parastenocarididae (Copepoda, Harpacticoida) from India: Parastenocaris muvattupuzha n. sp. from a river and P. kotumsarenis n. sp. from a cave. Zootaxa, 2077, 31–55.
Ranga Reddy, Y. & Schminke, H.K. (2009) Discovery of the genus Kinnecaris Jakobi, 1972 (Copepoda, Harpacticoida, Parastenocarididae) in southeastern India, with description of a new species. Crustaceana, 82, 311–326.
Reid, J.W. (1994) Murunducaris juneae, new genus, new species (Copepoda: Harpacticoida: Parastenocarididae) from wet campo in central Brazil. Journal of Crustacean Biology, 14, 771–781.
Reid, J.W. (1995) Redescription of Parastenocaris brevipes Kessler and description of a new species of Parastenocaris (Copepoda: Harpacticoida: Parastenocarididae) from the U.S.A. Canadian Journal of Zoology, 73, 173–187.
Sakaguchi, S.O. & Ueda, H. (2010) A new species of Pseudodiaptomus (Copepoda: Calanoida) from Japan, with notes on closely related P. inopinus Bruckhardt, 1913 from Kyushu Island. Zootaxa, 2623, 52–68.
Schminke, H.K. (1976) Systematische Untersuchungen an Grundwasserkrebsen eine Bestandsaufnahme (mit der Beschreibung zweier neuer Gattungen der Familie Parabathynellidae, Bathynellacea). International Journal of Speleology, 8, 195–216.
Schminke, H.K. (1981) Perspectives in the study of the zoogeography of interstitial Crustacea: Bathynellacea (Syncarida) and Parastenocarididae (Copepoda). International Journal of Speleology, 11, 83–89.
Schminke, H.K. (1991) Sexual dimorphism in furcal rami of Parastenocarididae (Copepoda: Harpacticoida). Bulletin of the Plankton Society of Japan, Special Volume, 1991, 573–584.
Schminke, H.K. (2008) First record of groundwater fauna from Papua New Guinea: Kinnecaris Jakobi, 1971 redefined (Copepoda, Harpacticoida, Parastenocarididae), and description of a new species. Crustaceana, 81, 1241–1253.
Schminke, H.K. (2009) Monodicaris gen. n. (Copepoda, Harpacticoida, Parastenocarididae) from West Africa. Crustaceana, 82, 367–378.
Schminke, H.K. (2010) High-level phylogenetic relationships within Parastenocarididae (Copepoda, Harpacticoida). Crustaceana, 83, 343–367.
Schram, F.R. (2008) Does biogeography have a future in a globalized world with globalized faunas? Contributions to Zoology, 77, 127–133.
Stamatakis, A., Hoover, P. & Rougemont, J. (2008) A Rapid Bootstrap Algorithm for the RAxML Web-Servers. Systematic Biology, 75, 758–771.
Stock, J.K. & von Vaupel Klein, J.C. (1996) Mounting media revisited: the suitability of Reyne’s fluid for small crustaceans. Crustaceana, 69, 749–798.
Swofford, D.L. (2002) PAUP*. Phylogenetic Analysis Using Parsimony (* and Other Methods), Version 4.0b10. (Sinauer Associates, Sunderland, Massachusetts).
Tamura, K., Dudley, J., Nei, M. & Kumar, S. (2007) MEGA4: Molecular Evolutionary Genetics Analysis (MEGA) software version 4.0. Molecular Biology and Evolution, 24, 1596–1599.
Tamura, K., Peterson, D., Peterson, N., Stecher, G., Nei, M. & Kumar, S. (in press) MEGA5: Molecular Evolutionary Genetics Analysis using Maximum Likelihood, Evolutionary Distance, and Maximum Parsimony Methods. Molecular Biology and Evolution.
Tang, D. & Knott, B. (2009) Freshwater cyclopoids and harpacticoids (Crustacea: Copepoda) from the Gnangara Mound region in Western Australia. Zootaxa, 2029, 1–70.
Timms, B.V. (1992) Lake Geomorphology. Gleneegles Publishing, Glen Osmond, South Australia, 180 pp.
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. Transaction of the Royal Society of South Australia, 130, 123–185.
Wells, J.B.J. (1964) Six new species of Parastenocaris (Crustacea, Copepoda) from Southern Rhodesia. Annals and Magazine of Natural History, Series 13, 7, 193–204.
Wells, J.B.J. (2007) An annotated checklist and keys to the species of Copepoda Harpacticoida. Zootaxa, 1568, 1–872.
Weston P.H. & Crisp, M.D. (1994) Cladistic biogeography of waraths (Proteaceae: Embothrieae) and their allies across the Pacific. Australian Systematic Botany, 7, 225–249.
Wilson, G.D.F. (2008) Gondwanan groundwater: subterranean connections of Australian phreatoicidean isopods to India and New Zealand. Invertebrate Systematics, 22, 301–310.