Article

Biogeography and phylogenetic relations within the Dinaric subgenus Monolistra (Microlistra) (Crustacea: Isopoda: Sphaeromatidae), with a description of two new species

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Abstract

A phylogenetic review of Monolistra (Microlistra), a freshwater cavernicolous subgenus of isopod crustaceans, distributed in the north-western part of the Dinaric karst, is presented. The distribution data and an identification key are provided for known taxa. Seven species are reviewed and two new species are described: Monolistra (Microlistra) fongi sp. nov. and Monolistra (Microlistra) jalzici sp. nov.Monolistra (Microlistra) pretneri spinulosa Sket is synonymysed with the nominate subspecies because of the morphological variability in the type subspecies and the genetic uniformity of the species. Two major, geographically vicariant and morphologically different clades have been identified by molecular analysis. Low genetic differentiation within the subgenus, as well as conspicuous dorsal sculpturing of animals, indicate their apparently recent colonization of the hypogean realm. These indications are confirmed by the distribution of Microlistra species within the current river systems, rather than palaeo-hydrographically defined basins, as is the case of other subterranean aquatic groups of crustaceans, including other members of the genus Monolistra.© 2010 The Linnean Society of London, Zoological Journal of the Linnean Society, 2010, 159, 1–21.

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... Metacyclops trisetosus Herbst, 1957 Thermocyclops dalmaticus Petkovski, 1956 Harpacticoida Morariopsis kieferi Petkovski, 1959 Ostracoda Podocopida Mixtacandona hvarensis (Danielopol, 1969) Pseudocandona sywulai Namiotko, Danielopol & Rađa, 2004 Sphaeromicola sphaeromidicola Hubault, 1938 Malacostraca Isopoda terrestria (kopneni jednakonožni rakovi) Aegonethes antilocapra Aegonethes cervinus (Verhoe , 1931) Alpioniscus balthasari (Frankenberger, 1937) Alpioniscus christiani Potočnik, 1983 Alpioniscus haasi (Verhoe , 1931) Alpioniscus kratochvili Alpioniscus magnus Alpioniscus trogirensis Buturović, 1955 Alpioniscus verhoe (Strouhal, 1938) Androniscus roseus histrianorum Androniscus stygius microcavernicolus Kesselyak, 1931 Androniscus wol Strouhal, 1939 Armadillidium dalmaticum Strouhal, 1939 Cyphodillidium absoloni (Strouhal, 1934) Cyphopleon kratochvili Oroniscus dalmaticus Strouhal, 1937 Oroniscus meledensis Strouhal, 1937 Oroniscus stentai (Arcangeli, 1926) Strouhaloniscellus biokovoensis Bedek & Taiti, 2009 Thaumatoniscellus speluncae I. Karaman, Bedek & Horvatović, 2009 Trichoniscus matulici metkovicensis Buturović, 1955 Troglocyphoniscus absoloni Strouhal, 1939 Typhlarmadillidium kratochvili Isopoda aquatica Merozoon vestigatum Sket, 2012 Monolistra berica hadzii Sket, 1959 Monolistra caeca meridionalis Deeleman-Reinhold, 1971 Monolistra fongi Prevorčnik, Verovnik, Zagmajster & Sket, 2010 Monolistra hercegovinensis atypica Sket, 1965 Monolistra hercegovinensis brevipes Sket, 1965 Monolistra jalzici Prevorčnik, Verovnik, Zagmajster & Sket, 2010 Monolistra pretneri Sket, 1964 Monolistra radjai Prevorčnik & Sket, 2007 Monolistra sketi Deeleman-Reinhold, 1971 Proasellus anophthalmus dalmatinus (S. Karaman, 1955) Proasellus anophthalmus rhausinus Proasellus coxalis lucifugus (Deeleman-Reinhold, 1965) Sphaeromides virei mediodalmatina Sket, 1964 Sphaeromides virei virei (Brian, 1923) Amphipoda Accubogammarus algor jalzici G. Karaman, 1988 Bogidiella sketi G. Karaman, 1989 Niphargus arbiter G. Karaman, 1984 Niphargus arcanus G. Karaman, 1988 Niphargus aulicus G. Niphargus buturovici S. Niphargus castellanus S. Karaman, 1960 Niphargus croaticus (Jurinac, 1887) Niphargus echion G. Karaman & Gottstein Matočec, 2006 Niphargus hebereri Schellenberg, 1933 Niphargus hvarensis S. Niphargus ilidzensis dalmatinus Schäferna, 1922 Niphargus jalzici G. Karaman, 1989 Niphargus krameri Schellenberg, 1935 Niphargus likanus S. Niphargus miljeticus Straškraba, 1959 Niphargus numerus G. Karaman & Sket, 1990 Niphargus pectencoronatae Sket & G. Karaman, 1990 Niphargus pretneri Sket, 1959 Niphargus redenseki Sket, 1959 Niphargus rejici jadranko Sket & G. Karaman, 1990 Niphargus rostratus Sket, 1971 Niphargus rucneri G. Karaman, 1962 Niphargus salonitanus S. Karaman, 1950 Niphargus steueri kolombatovici S. Karaman, 1950 Niphargus steueri liburnicus G. Karaman & Sket, 1989 Niphargus steueri steueri Schellenberg, 1935 Thermosbaenacea Tethysbaena halophila (S. ...
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... typ.: Sorgenti Sardos e Moschenizze Specie da ritenersi al momento endemica del carso triestino ed isontino, dove è piuttosto rara. Recenti analisi molecolari (Prevorčnik et al., 2010) dimostrano come le specie attribuite al sottogenere Microlistra costituiscano un raggruppamento monofiletico, sebbene l'insieme dei sottogeneri di Monolistra costituisca dei raggruppamenti artificiali. Dalle analisi genetiche risulta che Monolistra schottlaenderi è affine a M. spinosa (Racovitza, 1929) e M. spinosissima (Racovitza, 1929 della Slovenia occidentale, con le quali costituisce un clade. ...
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... Однако представитель другого рода перитрих -Lagenophrys monolistrae Stammer, 1935, был описан Г. Стаммером (Stammer, 1935) на стигобионтных (Prevorčnik et al., 2010) Hadzi, 1940(Hadzi, 1940. ...
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The ecological and evolutionary processes leading to isolation and adaptation of cave animals compared to their surface ancestors are not yet unequivocally understood. In this study the genetic relations of four cave and three surface population of the freshwater crustacean Asellus aquaticus in the Karst region of SW Slovenia and NE Italy were assessed using RAPDs as genetic markers. The results suggest that specialized populations from two caves invaded their subterranean habitat independently, and that their morphological similarity is a result of convergent evolution. Another, less specialized cave population seems to originate from a later colonization of a cave system already inhabited by a more specialized population, but the two populations do not interbreed. This series of temporally and spatially independent invasions has generated a diversity hotspot of non-interbreeding populations of a ubiquitous freshwater crustacean, which is uniform over most of its range. Genetic variability estimated by the percentage of polymorphic RAPD fragments was similar (86-91%) in most cave and surface populations. Substantially lower values (as low as 49%) were found in two cave populations affected by heavy pollution. Two a priori groupings of populations, traditional subspecies and hydrologically connected groups, were rejected as not significant by means of nested analysis of molecular variance (AMOVA). On the other hand, groupings revealed by UPGMA clustering displayed a significant component of among-group variance. An analysis of gene flow between populations using estimated migration rates confirmed these findings.
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Following the first systematic part, in this paper the author describes the biological observations made on these Isopod Crustaceans of underground waters. The different sexual characters of the particular groups and the related differences in the behaviour before copulation are described. During the embryonic and larval development small differences between sub-genus are reported. To the differences in Caecospaeroma (according to Daum) the first and second “mancastadium” and another “postmanca stadium” with pereiopods VII not wholly formed have to be added. The author describes the growth of the different parts of the corps and the extremities, comprising the sexual characters.
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Bilistra gen. nov., B. millari n. sp., type species, B. mollicopulans n. sp. and B. cavernicola n. sp. are described from karst areas in the northwest of South Island, New Zealand. Bilistra millari n. sp. occurs mainly in surface waters, while B. mollicopulans and B. cavernicola are stygobiotic and troglomorphic, occurring in cave waters. The genus is related to Benthosphaeroma Bruce, 1994, Neosphaeroma Baker, 1926, and Thermosphaeroma Cole & Bane, 1978, but each of these genera exhibits its own apomorphies, mainly in brood-pouch structure and morphology of pleopods IV-V. German Bilistra gen. nov., B. millari n. sp. als Typusart, B. mollicopulans n. sp. und B. cavernicola n. sp. aus Karstgebieten im Nordwesten der Südinsel von Neuseeland werden beschrieben. Bilistra millari n. sp., komt vor allem in oberirdischen Gewässern vor, während B. mollicopulans und B. cavernicola Stygobionten sind, die in Höhlengewässern vorkommen und Troglomorphien aufweisen. Die Gattung ist mit Benthosphaeroma Bruce, 1994, Neosphaeroma Baker, 1926 und Thermosphaeroma Cole & Bane, 1978, verwandt. Jede dieser Gattungen weist jedoch ihre eigenen Apomorphien auf, hauptsächlich in der Struktur des Marsupium und Morphologie der Pleopoden IV-V.
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Approximately 250 localities of the nominal species Proteus anguinus Laurenti 1768 have been evaluated and listed. The species is limited to the Dinaric Karat; it ranges from the Tsonzo-Soca River in southeastern Venezia Giulia, Italy, through the southern half of Slovenia, southern Croatia, and parts of Bosnia and Hercegovina, to the Trebisnica River in eastern Hercegovina. In some regions, populations have been extinguished or endangered by pollution or human-induced hydrographical changes. The distribution of Proteus is compared with those of some cave Crustacea: Troglocaris (Crustacea: Decapoda), Monolistra, and Titanethes (Crustacea: Isopoda). The similarity of distribution patterns within this ecologically diverse assemblage supports their paleogeographic rather than ecological foundation. The paleogeographical and paleoclimatological data, in combination with the physiological requirements of Proteus, strongly suggest that these animals invaded the caves, at least in the NW parts of their ranges, only after the last glaciations, within the last 10,000 years. It is suggested that the high heterozygosity of populations can best be explained by fusion of some locally restricted immigration waves. The close morphological similarity of nearly all populations of Proteus is probably due to the convergent evolution of previously differentiated populations (or even species) after their withdrawal underground.
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A selection of the most representative distribution patterns of stygobiont animals is presented. The range of a genus may extend beyond the Dinaride karst, while some species exhibit a holo-Dinaric distribution. The mero-Dinaric distribution is organised in two vicarious centres (in the NW and SE) as well as an epilittoral and a paralittoral belt. The paralittoral distribution is a result of paleogeographic rather than present-day ecological conditions. Important differences between the interstitial faunas of Slovenia (and NE Italy) and Macedonia are probably the result of their different sources. Some other distribution patterns are discussed or summarized from the existing literature. The distribution patterns of this stygobiont fauna are extremely diverse, and are influenced by the geological complexity of the territory as well as by the richness of the fauna concerned. New distribution data for some Copepoda, Thermosbaenacea, and Amphipoda are appended.Predstavljen je izbor najznailnejih vzorcev razirjenosti stigobiontskih ivali. Areal rodu lahko vkljuuje ve kot le Dinarski kras, holodinarsko razirjenost pa e kaejo posamezne vrste. Merodinarska razirjenost izhaja iz dveh centrov (na SZ in na JV) ali pa je epilitoralna ali paralitoralna. Paralitoralna razirjenost je vezana na paleogeografske in ne na dananje ekoloke razmere. Pomembne razlike med intersticialnima favnama Slovenije (in SV Italije) in Makedonije so verjetno posledica njunega razlinega izvora. Obravnavani ali pa povzeti iz obstojeih razprav so e nekateri vzorci razirjenosti. Vzorci razirjenosti tukajnje stigobiontske favne so izredno raznoliki, kar je posledica geoloke pestrosti ozemlja, kot tudi izrednega bogastva favne. Dodani so novi podatki o razirjenosti nekaterih kopepodov, termosbenacejev in amfipodov.
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We describe MUSCLE, a new computer program for creating multiple alignments of protein sequences. Elements of the algorithm include fast distance estimation using kmer counting, progressive alignment using a new profile function we call the log‐expectation score, and refinement using tree‐dependent restricted partitioning. The speed and accuracy of MUSCLE are compared with T‐Coffee, MAFFT and CLUSTALW on four test sets of reference alignments: BAliBASE, SABmark, SMART and a new benchmark, PREFAB. MUSCLE achieves the highest, or joint highest, rank in accuracy on each of these sets. Without refinement, MUSCLE achieves average accuracy statistically indistinguishable from T‐Coffee and MAFFT, and is the fastest of the tested methods for large numbers of sequences, aligning 5000 sequences of average length 350 in 7 min on a current desktop computer. The MUSCLE program, source code and PREFAB test data are freely available at http://www.drive5. com/muscle.
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