Abstract
Two new species of the genus Plutomurus, P. revazi sp. nov. from Prometheus and Satsurblia caves and P. eristoi sp. nov. from Satevzia Cave are described, illustrated and differentiated from other morphologically closely related species. A high variability in the number of teeth in the claw, unguiculus and mucro of P. revazi sp. nov. demonstrate that these characters are not useful for species diagnosis. However, dorsal chaetotaxy was shown to be stable character for this purpose. Analysis of DNA sequences for the COI and 28S genes is congruent with species-level groups delimited by chaetotaxy, and provide additional support for chaetotaxy as the most reliable morphological character system to distinguish species in Plutomurus. A key to species of the genus Plutomurus found in Georgia is provided, which for the first time includes characters of the macrochaetotaxy.
References
Anslan, S. & Tedersoo, L. (2015) Performance of cytochrome oxidase I (COI), ribosomal DNA large subunit (LSU) and internal transcribed spacer 2 (ITS2) in DNA barcoding of Collembola. European Journal of Soil Biology, 69, 1–7.
http://dx.doi.org/10.1016/j.ejsobi.2015.04.001
Barjadze, Sh. & Djanashvili, R. (2008) Checklist of the springtails (Collembola) of Georgia. Caucasian Entomological Bulletin, 4, 187–193.
Barjadze, Sh. & Djanashvili, R. (2014) Redescription of Plutomurus kelasuricus Martynova, 1969 (Collembola: Tomoceridae) from Georgian Caves. Zoology in the Middle East, 60, 251–254.
http://dx.doi.org/10.1080/09397140.2014.939812
Barjadze, Sh., Murvanidze, M., Arabuli, T., Mumladze, L., Pkhakadze, V., Djanashvili, R. & Salakaia, M. (2015) Annotated List of Invertebrates of the Georgian Karst Caves. Georgian Academic Book, Tbilisi, 120 pp.
Börner, C. (1901) Über einige theilweise neue Collembolen aus den Höhlen der Gegend von Letmathe in Westfolen. Zoologische Anzeiger, 24, 333–345.
Carapelli, A., Frati, F., Fanciulli, P.P. & Romano, D. (1995) Genetic differentiation of six sympatric species of Isotomurus (Collembola, Isotomidae); is there any difference in their microhabitat preference? European Journal of Soil Biology, 31, 87–99.
Christiansen, K. (1964) A revision of the nearctic members of the genus Tomocerus (Collembola Entomobryidae). Revue d' Écologie et de Biologie du Sol, 1, 639–678.
Christiansen, K. (1980) A new nearctic species of the genus Tomocerus (Collembola: Entomobryidae). Proceedings of the Iowa Academy of Sciences, 87, 121–123.
Christiansen, K. & Bellinger, P. (1998) The Collembola of North America, North of the Rio Grande. A taxonomic analysis. Part 3. Grinnell College, Iowa. pp. 877–1173.
Cicconardi, F., Nardi, F., Emerson, B.C., Frati, F. & Fanciulli, P.P. (2010) Deep phylogeographic divisions and long-term persistence of forest invertebrates (Hexapoda: Collembola) in the North-Western Mediterranean basin. Molecular Ecology, 19, 386–400.
http://dx.doi.org/10.1111/j.1365-294X.2009.04457.x
Cicconardi, F., Fanciuli, P.P. & Emerson, B.C. (2013) Collembola, the biological species concept and the underestimation of global species richness. Molecular Ecology, 22, 5382–5396.
http://dx.doi.org/10.1111/mec.12472
D'Haese, C. (2002) Where the first springtails semi-aquatic? A phylogenetic approach by means of 28S rDNA and optimization alignment. Proceedings of the Royal Society, London, B, 269, 1143–1151.
Djanashvili, R. & Barjadze, S. (2011) A new species of the genus Plutomurus Yosii, 1956 (Collembola, Tomoceridae) from Georgian caves. Journal of Cave and Karst Studies, 73, 28–30.
http://dx.doi.org/10.4311/jcks2010lsc0147
Felderhoeff, K., Bernard, E.C. & Moulton, J.K. (2010) Survey of Pogonognathellus Börner (Collembola: Tomoceridae) in the southern Appalachians based on morphology and molecular data. Annals of the Entomological Society of America, 103, 472–491.
http://dx.doi.org/10.1603/AN09105
Folsom, J.W. (1913) North American Springtails of the sub-family Tomocerinae. Proceedings of the U.S. National Museum, 46, 451–472.
Frati, F., Dell'Ampio, E., Casasanta, S., Carapelli, A. & Fanciulli, P.P. (2000) Large amounts of genetic divergence among species of the genus Orchesella (Insecta, Collembola) and the relationships of two new species. Molecular Phylogenetics and Evolution, 17, 456–461.
http://dx.doi.org/10.1006/mpev.2000.0854
Gisin, H. (1963) Collemboles d’Europe. V. Revue Suisse de Zoologie, 70, 77–101.
Gisin, H. (1964) Collemboles d’Europe. VII. Revue Suisse de Zoologie, 71, 649–678.
Gisin, H. (1967) Espèces nouvelles et lignées évolutives de Pseudosinella endogés. Memórias e Estudos do Museu Zoológico da Universidade de Coimbra, 301, 5–25.
Greenslade, P. & Jordana, R. (2014) Description and conservation status of a new species of Australotomurus (Collembola: Entomobryidae: Orchesellinae) from urban Perth remnant bushland. Zootaxa, 3872 (5), 561–76.
http://dx.doi.org/10.11646/zootaxa.3872.5.7
Hogg, I.D. & Hebert P.D.N. (2004) Biological identification of springtails (Hexapoda: Collembola) from the Canadian Arctic, using mitochondrial DNA barcodes. Canadian Journal of Zoology, 82, 749–754.
http://dx.doi.org/10.1139/Z04-041
Jordana, R. & Baquero, E. (2005) A proposal of characters for taxonomic identification of Entomo-brya species (Collembola, Entomobryomorpha), with description of a new species. Abhan-dlungen und Berichte des Naturkundemuseums Görlitz, 76, 117–134.
Jordana, R., Baquero, E., Reboleira, S. & Sendra, R. (2012) Reviews of the genera Schaefferia Absolon, 1900, Deuteraphorura Absolon, 1901, Plutomurus Yosii, 1956 and the Anurida Laboulbène, 1865 species group without eyes, with the description of four new species of cave springtails (Collembola) from Krubera-Voronya cave, Arabika Massif, Abkhazia. Terrestrial Arthropod Reviews, 5, 35–85.
http://dx.doi.org/10.1163/187498312X622430
Joy, D.A. & Conn, J.E. (2001) Molecular and morphological phylogenetic analysis of an insular radiation in Pacific black flies (Simulium). Systematic Biology, 50, 18–38.
Katz, A., Giordano, R. & Soto-Adames, F.N. (2015) Taxonomic review and phylogenetic analysis of fifteen North American Entomobrya (Collembola, Entomobryidae), including four new species. Zookeys, 525, 1–75.
http://dx.doi.org/10.3897/zookeys.525.6020
Kniss, V. & Thibaud, J. (1999) Genus Plutomurus in Russia and Georgia (Collembola, Tomoceridae). Revue française d’Entomologie, New Series, 21, 57–64 [in French]
Liponava, K. (1990) Speleological news on the Tskaltubo limestone massif. Proceedings of the Georgian SSR geographical society, 27, 8–13 [in Georgian]
Martynova, E. (1969) Springtails of the family Tomoceridae (Collembola) from the fauna of the USSR. Entomological Review, 43, 299–314. [in Russian]
Maynard, E.A. (1951) A Monograph of the Collembola or Springtail Insects of New York State. Comstock Publishing Company Inc., Ithaca, New York, 339 pp.
Porco, D., Bedos, A., Greenslade, P., Janion, C., Skazynski, D., Stevens, M.I., Jansen Van Vuuren, B. & Deharveng, L. (2012) Challenging species delimitation in Collembola: cryptic diversity among common springtails unvailed by DNA barcoding. Invertebrate Systematics, 26, 470–477. Avaliable from: http://www.publish.csiro.au/?paper=IS12026 (Accessed 14 Jun. 2016)
Simon, C., Frati, F., Beckenbach, A., Crespi, B., Liu, H. & Flook, P. (1994) Evolution, weighting, and phylogenetic utility of mitochondrial gene sequences and a compilation of conserved polymerase chain reaction primers. Annals of the Entomological Society of America, 87, 651–700.
Skorikov, A. (1900) Zoologische Ergebnisse der russischen Expedition nach Spitzbergen im Jahre 1899. Collembola. Annuaire du Musee Zoologique de l'Academie d. Sciences de St. Petersbourg, 5, 190–209.
Soto-Adames, F.N. (2002) Molecular Phylogeny of the Puerto Rican Lepidocyrtus and Pseudosinella (Hexapoda: Collembola: Entomobryidae), a validation of Yoshii's "colour pattern species". Molecular Phylogenetic and Evolution, 25, 27–42.
http://dx.doi.org/10.1016/S1055-7903(02)00250-6
Stomp, N. & Weiner, W.M. (1994) Redescription of Plutomurus unidentatus (Börner, 1901) comb. nov. (Collembola: Tomoceridae). Bulletin de la Société des naturalistes luxembourgeois, 95, 359–465.
Tatashidze, Z.K., Tsikarishvili, K.D. & Jishkariani, J.M. (2009a) The Cadastre of the Karst Caves of Georgia. Petiti Publishing House, Tbilisi, 670 pp. [in Georgian]
Tatashidze, Z., Tsikarishvili, K., Jishkariani, J., Jamrishvili, A., Geladze, G. & Lominadze, G. (2009b) Tskaltubo Cave System. Petiti Publishing House, Tbilisi, 72 pp. [in Georgian]
Yosii, R. (1939) Two new species of Tomocerid Collembola from limestone caves of Japan. Annotationes Zoologicae Japonensis, Tokyo, 18, 177–179.
Yosii, R. (1956) Monographie zur Höhlencollembolen Japans. Contribution from the Biological Laboratory Kyoto University, 3, 1–109.
Yosii, R. (1967) Studies on the Collembolan family Tomoceridae, with special reference to Japanese forms. Contributions from the Biological Laboratory Kyoto University, 20, 1–54.
Yoshii, R. (1991) A new species of Tomocerid Collembola from the Cave of Pref. Iwate. Annales of the Speleological Institute of Japan (Iwaizumi), 9, 1–2.
Zhang, F., You, D., Luo, Y. Ho, S.Y.W., Wang, B. & Zhu, C. (2014) Cryptic diversity, diversification and vicariance in two species complexes of Tomocerus (Collembola, Tomoceridae) from China. Zoologica Scripta, 43, 393–404.
http://dx.doi.org/10.1111/zsc.12056