COXI based phylogenetic analysis of Caucasian clade of European Troglocaris (Crustacea: Decapoda: Atyidae) with the suggestion of a new taxonomic group structure

Keywords: phylogeny, COXI mtDNA, Xiphocaridinella, stygobiotic, stygobionts, shrimps, Caucasus


New genetic data on Caucasian troglobiotic shrimps collected from the territory of Russia, Abkhazia and Western Georgia are presented. Based on new genetic data on the marker region of subunit I of cytochrome oxidase of mitochondrial DNA (COXI mtDNA) of Caucasian species and other taxa of European cave shrimps (Troglocaris s.l.) from GenBank (NCBI) database, a new generic structure is presented. Based on a significant genetic divergence of COXI mtDNA subgenera of Troglocaris s.l., namely Troglocaris s.s., Xiphocaridinella and Spelaeocaris, should be considered as separate genera while Troglocaris (Troglocaridella) hercegovinensis (Babić, 1922) is suggested to be transferred within Dinaric genus Spelaeocaris as Spealeocaris hercegovinensis (Babić, 1922) comb. nov. Besides, Troglocaris bosnica shows a significant a genetic difference (at the level of the separate genus) from the remaining representatives of the genus Troglocaris s.s. Moreover, p-distances (COXI) of about 17% are supposed for generic separation within European Troglocaris-related atyid shrimps and 5% for separation of cryptic species within Caucasian Xiphocaridinella. A list of all known taxa of Troglocaris-related atyid shrimps of the European part and a discussion of the general distribution of troglocaridid atyid shrimps in the Balkans and the Caucasus are presented. A new version of divergence events between Dinaric–Caucasian Troglocaris lineages (genera) based on new genetic data is suggested.


Babic, K. (1922). Über die drei Atyiden aus Jugoslavien. Glasnik der Kroatischen Naturwissenschaftlichen Gesellschaft, Zagreb, 34, 300–306.

Birštein, Y. A. (1939). O peshernysh krevetkah Abhazii [About cave shrimps of Abkhazia]. Zoologichesky Zhurnal, 18, 960–974 (in Russian).

Birštein, Y. A. (1948). O nahozdenii pesherinoy krevetki Troglocaris v gruntovyh vodah Abkhazii b svazannye s etim voprosy [The occurrence of the cave shrimp Troglocaris in underground water of Mazesta and related problems]. Byulleten’ Moskovskogo Obshchestva Ispytatelei Prirody, Otdel Biologicheskii, 53, 3–10 (in Russian).

D’Udekemd’Acoz, C. (1999). Inventaireet distribution des crustacés décapodes de l’Atlantiquenord-oriental, de la Méditerranée et des ea ux continentale sadjacentes au nord de 25 N. Patrimoines Naturels, 40, 1–383.

De Grave, S., & Fransen, C. H. J. M. (2011). Carideorum сatalogus: The recent species of the dendrobranchiate, stenopodidean, procarididean and caridean shrimps (Crustacea: Decapoda). Zoologische Mededelingen, 85(9), 195–589.

Franjević, D., Kalafatić, M., Kerovec, M., & Gottstein, S. (2010). Phylogeny of cave-dwelling atyid shrimp Troglocaris in the Dinaric Karst based on sequences of three mitochondrial genes. Periodicum Biologorum, 112(2), 159–166.

Hebert, P. D. N., Cywinska, A., Ball, S. L., & De Waard, J. R. (2003). Biological identifications through DNA barcodes. Proceedings of the Royal Society of London B, 270, 313–322.

Jaume, D., & Bréhier, F. (2005). A new species of Typhlatya (Crustacea: Decapoda: Atyidae) from anchialine caves on the French Mediterranean coast. Zoological Journal of the Linnean Society, 144, 387–414.

Jugovic, J., Jalžić, B., Prevorčnik, S., & Sket, B. (2012). Cave shrimps Troglocaris s. str. (Dormitzer, 1853), taxonomic revision and description of new taxa after phylogenetic and morphometric studies. Zootaxa, 3421, 1–31.

Jugovic, J., Prevorčnik, S., Aljančič, G., & Sket, B. (2010). The atyid shrimp (Crustacea: Decapoda: Atyidae) rostrum: Phylogeny versus adaptation, taxonomy versus trophic ecology. Journal of Natural History, 44(41–42), 2509–2533.

Jugovic, J., Prevorčnik, S., Blejec, A., & Sket, B. (2011). Morphological differentiation in the cave shrimps Troglocaris (Crustacea: Decapoda: Atyidae) of the Dinaric karst – A consequence of geographical isolation or adaptation? Journal of Zoological Systematic and Evolutionary Researches, 49(3), 185–195.

Juzbaš’jan, S. M. (1940). O Shakuranskoy pesernoy krevetke [On a cave shrimp from Shakuran]. Trudy Biologiceskoj Stancii Narkomprosa Gruzinskoj SSR, 1, 73–86 (in Russian).

Knowlton, N., & Weigt, L. A. (1998). New dates and new rates for divergence across the Isthmus of Panama. Proceedings of the Royal Society of London. Series B, 265, 2257–2263.

Knowlton, N., Weight, L. A., Solórzano, L. A., Mills, D. K., & Bermingham, E. (1993). Divergence in proteins, mitochondrial DNA and reproductive compatibility across the Isthmus of Panama. Science, 260(5114), 1629–1632.

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 delimitation. Molecular Phylogenetics and Evolution, 40(2), 435–447.

Lushai, G., Smith, D. A. S., Goulson, D., Allen, J. A., & Maclean, N. (2003). Mitochondrial DNA clocks and the phylogeny of Danaus butterflies. Insect Science and its Application, 2, 309–315.

Marin, I. (2017). Troglocaris (Xiphocaridinella) kumistavi sp. nov., a new species of stygobiotic atyid shrimp (Crustacea: Decapoda: Atyidae) from Kumistavi Cave, Imereti, Western Georgia, Caucasus. Zootaxa, 4311(4), 576–588.

Marin, I. N. (2015). Atlas of decapod crustaceans of Russia. KMK Scientific Press, Moscow.

Marin, I. N., & Sinelnikov, S. Y. (2017). Preliminary data on larval development of Caucasian cave-dwelling shrimp Troglocaris (Xiphocaridinella) kumistavi Marin, 2017 (Crustacea: Decapoda: Atyidae). Arthropoda Selecta, 26(4), 297–302.

Marin, I. N., & Turbanov, I. S. (2015). Peshernye krevetki roda Troglocaris Dormitzer, 1853 (Crustacea: Decapoda: Atyidae): Raznoobrazie, ecologiya, proiskhozdenie [Cave shrimps of the genus Troglocaris Dormitzer, 1853 (Crustacea: Decapoda: Atyidae): Diversity, ecology, origin]. In: Turbanov, I. S., Marin, I. N., & Gongalsky, K. B. (Eds.). Abstracts of the Youth Conference "Biospeleology of the Caucasus and other regions of Russia" (IPEE RAS, Moscow, December, 3–4, 2015). Kostroma Printing House, Kostroma. pp. 36–41.

Marin, I., & Sokolova, A. (2014). Redescription of the stygobiotic shrimp Troglocaris (Xiphocaridinella) jusbaschjani Birštein, 1948 (Decapoda: Caridea: Atyidae) from Agura River, Sochi, Russia, with remarks on other representatives of the genus from Caucasus. Zootaxa, 3754(3), 277–298.

Matjasic, J. (1956). Ein neuer Höhlen decapods e aus Jugoslawien. Zoologischer Anzeiger, 157, 65–68.

Sadowsky, A. A. (1930). Xiphocaridinella kutaissiana nov. gen. et sp. (fam. Atyidae) iz podzemnoi peshchery pod Kutaisom [Xiphocaridinella kutaissiana nov. gen. et sp. (fam. Atyidae) from subterranean cave near Kutaisi]. Zakavkazskij Kraevedstenny Sbornik Naučnoissledovatel’nogo Kraevedstvenogo Kabineta Universiteta Tiflis, 1, 93–104 (in Russian).

Sanz, S., & Platvoet, D. (1995). New perspectives on the evolution of the genus Typhlatya (Crustacea, Decapoda): First record of a cavernicolous atyid in the Iberian Peninsula, Typhlatya miravetensis n. sp. Contributions to Zoology, 65, 79–99.

Sendra, A., & Reboleira, A. S. P. S. (2012). The world’s deepest subterranean community – Krubera-Voronja Cave (Western Caucasus). International Journal of Speleology, 42(2), 221–230.

Sites, J. W., & Marshall, J. C. (2004). Operational criteria for delimiting species. Annual Review of Ecology, Evolution, and Systematics, 35, 199–227.

Sket, B. (2003). Distribution of Proteus (Amphibia: Urodela: Proteidae) and its possible explanation. Journal of Biogeography, 24(3), 263–280.

Sket, B., & Zakšek, V. (2009). European cave shrimp species (Decapoda: Caridea: Atyidae), redefined after a phylogenetic study; redefinition of some taxa, a new genus and four new Troglocaris species. Zoological Journal of the Linnean Society, 155(4), 786–818.

Turbanov, I. S., & Marin, I. N. (2015). Typhlogammaridae (Crustacea: Amphipoda) – Novoe semeistvo stigobontnyh amphipod dla fauny Rossii is podzemnyh vod Krasnodaskogo kraya [Typhlogammaridae (Crustacea: Amphipoda) – A new family of stygobiotic amphipods for the Russian fauna in groundwater of the Krasnodar Territory]. In: Turbanov, I. S., Marin, I. N., & Gongalsky, K. B. (Eds.). Abstracts of the Youth Conference "Biospeleology of the Caucasus and other regions of Russia" (IPEE RAS, Moscow, December, 3–4, 2015). Kostroma Printing House, Kostroma. pp. 87–89.

Turbanov, I. S., Palatov, D. M., & Golovatch, S. I. (2016). The state of the art of biospeleology in Russia and other countries of the former Soviet Union: A review of the cave (endogean) invertebrate fauna. 1. Introduction. Crustacea. Entomological Review, 96, 926–963.

Zakšek, V., Sket, B., & Trontelj, P. (2006). Phylogeny of the cave shrimp Troglocaris: Evidence of a young connection between Balkans and Caucasus. Molecular Phylogenetics and Evolution, 42, 223–235.

Zakšek, V., Sket, B., Gottstein, S., Franjevic, D., & Trontelj, P. (2009). The limits of cryptic diversity in groundwater: Phylogeography of the cave shrimp Troglocaris anophthalmus (Crustacea: Decapoda: Atyidae). Molecular Ecology, 18, 931–946.