Nematodes from terrestrial and freshwater habitats in the Arctic.

WE PRESENT AN UPDATED LIST OF TERRESTRIAL AND FRESHWATER NEMATODES FROM ALL REGIONS OF THE ARCTIC, FOR WHICH RECORDS OF PROPERLY IDENTIFIED NEMATODE SPECIES ARE AVAILABLE: Svalbard, Jan Mayen, Iceland, Greenland, Nunavut, Northwest territories, Alaska, Lena River estuary, Taymyr and Severnaya Zemlya and Novaya Zemlya. The list includes 391 species belonging to 146 genera, 54 families and 10 orders of the phylum Nematoda.

Introduction

Nematodes are one of the most numerous and abundant multicellular organisms on the planet in general and in the Arctic in particular. There are over 70 research papers published which include data on the fauna and distribution of nematodes in the region. Few faunistic overviews of the Arctic nematodes exist in the literature. The "Catalogue of free-living soil and fresh-water nematodes of Arctic and Subarctic" (Kuzmin and Gagarin 1990) includes a large number of subarctic nematodes and is normally inaccessible. Subsequent publication by Gagarin (2001b) focuses only on the fresh-water nematodes from the Russian Arctic and Subarctic, while Coulson and Refseth (2004) give a list of terrestrial and freshwater nematodes from Svalbard and Jan Mayen. The present paper aims to provide a comprehensive overview of all findings of nematodes in the entire Arctic region between the years of 1883 and 2013, with all species names adjusted according to the most recent classification and nomenclature.

Materials and methods

The species list has been compiled based on literature data and refers to the area of the Arctic (Fig. 1) as defined by the Circumpolar Arctic Vegetation Map (CAVM Team 2003) and Meltofte et al. (2013). The list below includes all species found in the Arctic, and specifies the regions where each species was found, as well as all relevant references. Current taxonomic status and systematic position of all species was updated according to the most recent views on nematode systematics. Section on nomenclature includes those synonymous names, and sometimes mistyped names, which were used in the original publications on the Arctic nematodes; it does not include any other existing synonyms or combinations. Asterisks indicate those new species names, which were originally proposed based on populations described from the Arctic.

Figure 1.

Known localities in the Arctic for which detailed information about nematode fauna (identified to species level) is available in the literature.

Taxa identified in the literature to the genus level only were not considered in the review for the following reasons: 1) It is impossible to make sure how many actual species were considered by the author – just one, or if s/he had treated multiple species together in the same unit. 2) Taxa identified only to the genus level have limited usefulness when analyzing distribution, endemism and other aspects of biogeography. Nematode systematics is continuously improving. Menzel (1920) described sp. from Spitzbergen (Svalbard). The context of the genus had changed considerably since 1920 and now includes members of at least families , , and , making it practically impossible to use Menzel's record in modern faunistic studies. Even genus-level identifications done more recently face similar problems. Species that are now considered to belong to the same genus may in future, when more data becomes available, be split into different, sometimes distantly related genera. It would thus be impossible to know in the future what author of the paper meant when he identified his material.

General classification and classification of the orders Chromadorida, and follows De Ley and Blaxter (2004) and Holovachov and Esquivel (2012). Classification of the order follows that of Smol et al. (2014) and Andrássy (2002); – Holovachov and Shoshin (2014); – Andrássy (2009); – Ahmad and Jairajpuri (2010); – Fonseca and Decraemer (2008); – Holovachov (2014); – Holovachov et al. (2009), Geraert (2008), Geraert (2011), Sudhaus (2011), Sudhaus and Fürst von Lieven (2003).

Checklists

Updated list of nematode species found in the Arctic

de Man, 1876

Notes

Taymyr and Severnaya Zemlya, Russia (Gagarin 1990, Gagarin 1993, Gagarin 1996, Gagarin 2001b, Gagarin 2001a, Kuzmin and Gagarin 1990).

Thorne, 1939

Svalbard (Loof 1971); Taymyr and Severnaya Zemlya, Russia (Kuzmin and Gagarin 1990).

Loof, 1971*

Svalbard (Loof 1971).

Thorne, 1939

Svalbard (Loof 1971).

de Man, 1880

Jan Mayen (Allgén 1953); Taymyr and Severnaya Zemlya, Russia (Gagarin 1990, Gagarin 2001b, Gagarin 2001a, Kuzmin and Gagarin 1990); Novaya Zemlya and Vaigach island, Russia (Steiner 1916b).

Thorne, 1939

Taymyr and Severnaya Zemlya, Russia (Kuzmin 1986, Kuzmin and Gagarin 1990).

Thorne, 1939

Taymyr and Severnaya Zemlya, Russia (Kuzmin and Gagarin 1990).

(Loof, 1964)

Loof, 1964

Taymyr and Severnaya Zemlya, Russia (Kuzmin 1986, Kuzmin and Gagarin 1990).

(de Man, 1921)

de Man, 1921

Jan Mayen (Allgén 1953); Taymyr and Severnaya Zemlya, Russia (Kuzmin and Gagarin 1990).

(de Man, 1876)

(de Man, 1876)

Svalbard (Loof 1971); Taymyr and Severnaya Zemlya, Russia (Kuzmin and Gagarin 1990).

Gagarin, 1991*

Taymyr and Severnaya Zemlya, Russia (Gagarin 1990, Gagarin 1991b, Gagarin 1993, Gagarin 2001b, Gagarin 2001a).

(Eliashvili, 1971)

Novaya Zemlya and Vaigach island, Russia (Gagarin 1997a, Gagarin 2001b).

(Thorne, 1939)

Thorne, 1939

Taymyr and Severnaya Zemlya, Russia (Kuzmin and Gagarin 1990).

de Man, 1880

Svalbard (Loof 1971); Nunavut, Canada (Mulvey 1969c); Taymyr and Severnaya Zemlya, Russia (Kuzmin 1976, Kuzmin and Gagarin 1990); Novaya Zemlya and Vaigach island, Russia (Gagarin 1997a, Gagarin 2001b).

(de Man, 1880)

(de Man, 1880)

Nunavut, Canada (Mulvey 1969c).

(de Man, 1880)

Taymyr and Severnaya Zemlya, Russia (Kuzmin and Gagarin 1990).

(W. Schneider, 1925)

Northwest territories, Canada (Ebsary 1982).

Gagarin, 1996*

Taymyr and Severnaya Zemlya, Russia (Gagarin 1996, Gagarin 2001b).

(W. Schneider, 1925)

Taymyr and Severnaya Zemlya, Russia (Gagarin 2001b); Novaya Zemlya and Vaigach island, Russia (Gagarin 1997a, Gagarin 2001b).

(Bastian, 1865)

Nunavut, Canada (Mulvey 1969c); Lena River estuary, Russia (Gagarin 2001b); Taymyr and Severnaya Zemlya, Russia (Gagarin 1990, Gagarin 1996, Gagarin 2001b, Gagarin 2001a, Gagarin and Kuzmin 1983, Kuzmin 1972, Kuzmin and Gagarin 1990); Novaya Zemlya and Vaigach island, Russia (Gagarin 1997a, Gagarin 1999a, Gagarin 2001b).

(Hofmänner in Hofmänner & Menzel, 1914)

Micoletzky 1925

Greenland (Ditlevsen 1927); Taymyr and Severnaya Zemlya, Russia (Gagarin 1996, Kuzmin and Gagarin 1990).

Gagarin, 1996*

Taymyr and Severnaya Zemlya, Russia (Gagarin 1996, Gagarin 2001b).

Gagarin, 1991*

Taymyr and Severnaya Zemlya, Russia (Gagarin 1990, Gagarin 1991a, Gagarin 1993, Gagarin 2001b, Gagarin 2001a).

Gagarin, 1991*

Taymyr and Severnaya Zemlya, Russia (Gagarin 1990, Gagarin 1991a, Gagarin 1993, Gagarin 2001b, Gagarin 2001a).

(Micoletzky, 1925)

Lena River estuary, Russia (Gagarin 2001b); Taymyr and Severnaya Zemlya, Russia (Gagarin 1990, Gagarin 1996, Gagarin 2001b, Gagarin 2001a, Kuzmin and Gagarin 1990); Novaya Zemlya and Vaigach island, Russia (Gagarin 1999a, Gagarin 2001b).

(Steiner, 1919)

Taymyr and Severnaya Zemlya, Russia (Gagarin 1996, Gagarin 2001b); Novaya Zemlya and Vaigach island, Russia (Gagarin 1997a, Gagarin 2001b).

Shoshin, 1988

sensu Gagarin, 1993, 2001;

Taymyr and Severnaya Zemlya, Russia (Gagarin 1990, Gagarin 1991a, Gagarin 1993, Gagarin 2001b, Gagarin 2001a, Gagarin 2009).

(Andrássy, 1953)

Taymyr and Severnaya Zemlya, Russia (Gagarin 1996, Gagarin 2001b).

(Altherr, 1963)

Taymyr and Severnaya Zemlya, Russia (Gagarin 2009, Kuzmin and Gagarin 1990).

(Brakenhoff, 1914)

Lena River estuary, Russia (Gagarin 2001b); Taymyr and Severnaya Zemlya, Russia (Gagarin 1996, Gagarin 2001b); Novaya Zemlya and Vaigach island, Russia (Gagarin 1997a, Gagarin 1999a, Gagarin 2001b).

(Tsalolikhin, 1977)

Taymyr and Severnaya Zemlya, Russia (Gagarin 1996, Gagarin 2001b).

(Leidy, 1852)

Lena River estuary, Russia (Gagarin 2001b); Taymyr and Severnaya Zemlya, Russia (Gagarin 1996, Kuzmin and Gagarin 1990).

Micoletzky, 1922

Taymyr and Severnaya Zemlya, Russia (Gagarin 1990, Gagarin 1993, Gagarin 2001b, Gagarin 2001a).

Gagarin, 1991*

Taymyr and Severnaya Zemlya, Russia (Gagarin 1990, Gagarin 1991a, Gagarin 1993, Gagarin 2001b, Gagarin 2001a).

(Gagarin, 1991)

Gagarin, 1991*

Taymyr and Severnaya Zemlya, Russia (Gagarin 1990, Gagarin 1991a, Gagarin 1993, Gagarin 2001b, Gagarin 2001a).

Gagarin, 1999*

Lena River estuary, Russia (Gagarin 1999b, Gagarin 2001b).

(Gagarin, 1991)

Gagarin, 1991*

Taymyr and Severnaya Zemlya, Russia (Gagarin 1990, Gagarin 1991a, Gagarin 1993, Gagarin 2001b, Gagarin 2001a).

(Micoletzky, 1925)

(Micoletzky, 1925);

Lena River estuary, Russia (Gagarin 2001b); Taymyr and Severnaya Zemlya, Russia (Gagarin 1990, Gagarin 1993, Gagarin 1996, Gagarin 2001b, Gagarin 2001a, Kuzmin 1986, Kuzmin and Gagarin 1990).

Tsalolikhin, 2000*

sensu Ebsary, 1982

Northwest territories, Canada (Ebsary 1982).

(Schneider, 1923)

(Schneider, 1923)

Taymyr and Severnaya Zemlya, Russia (Kuzmin 1972, Kuzmin and Gagarin 1990).

(Ebsary, 1982)

Ebsary, 1982*

Northwest territories, Canada (Ebsary 1982); Taymyr and Severnaya Zemlya, Russia (Gagarin 1990, Gagarin 1993, Gagarin 2001b, Gagarin 2001a).

(Ebsary, 1982)

Lena River estuary, Russia (Gagarin 2001b).

de Man, 1880

Nunavut, Canada (Mulvey 1969c); Taymyr and Severnaya Zemlya, Russia (Kuzmin and Gagarin 1990); Novaya Zemlya and Vaigach island, Russia (Steiner 1916b).

Gagarin, 1997*

Novaya Zemlya and Vaigach island, Russia (Gagarin 1997a, Gagarin 2001b).

de Man, 1880

Taymyr and Severnaya Zemlya, Russia (Kuzmin 1986, Kuzmin and Gagarin 1990).

Bastian, 1865

Bütschli, 1873

Svalbard (Loof 1971); Greenland (Ditlevsen 1927); Taymyr and Severnaya Zemlya, Russia (Gagarin 1996, Kuzmin 1972, Kuzmin and Gagarin 1990); Novaya Zemlya and Vaigach island, Russia (Gagarin 1997a, Gagarin 1999a, Gagarin 2001b).

Brzeski & Winiszewska-Slipinska, 1993

Altherr in Altherr & Delamare Deboutteville, 1972

Svalbard (Janiec 1997); Taymyr and Severnaya Zemlya, Russia (Gagarin 1990, Gagarin 1993, Gagarin 1996, Gagarin 2001b, Gagarin 2001a).

Altherr in Altherr & Delamare Deboutteville, 1972

Lena River estuary, Russia (Gagarin 2001b); Taymyr and Severnaya Zemlya, Russia (Gagarin 1990, Gagarin 1993, Gagarin 1996, Gagarin 2001b, Gagarin 2001a).

Bütschli, 1873

Lena River estuary, Russia (Gagarin 2001b); Taymyr and Severnaya Zemlya, Russia (Kuzmin and Gagarin 1990).

de Man, 1880

Svalbard (Loof 1971); Jan Mayen (Allgén 1953); Greenland (Ditlevsen 1927); Taymyr and Severnaya Zemlya, Russia (Kuzmin and Gagarin 1990); Novaya Zemlya and Vaigach island, Russia (Steiner 1916b).

(Bütschli, 1873)

Svalbard (Loof 1971); Jan Mayen (Allgén 1953); Nunavut, Canada (Mulvey 1969c); Taymyr and Severnaya Zemlya, Russia (Kuzmin and Gagarin 1990).

Loof, 1971*

Svalbard (Loof 1971).

Loof, 1971*

Svalbard (Loof 1971).

Gagarin & Kuzmin, 1972*

Taymyr and Severnaya Zemlya, Russia (Gagarin 1993, Gagarin and Kuzmin 1972, Kuzmin 1976, Kuzmin and Gagarin 1990).

Hirschmann, 1952

Novaya Zemlya and Vaigach island, Russia (Gagarin 1997a, Gagarin 2001b).

de Man, 1876

Svalbard (Loof 1971); Nunavut, Canada (Mulvey 1969c); Taymyr and Severnaya Zemlya, Russia (Kuzmin and Gagarin 1990); Novaya Zemlya and Vaigach island, Russia (Steiner 1916b).

De Coninck, 1935

Taymyr and Severnaya Zemlya, Russia (Kuzmin and Gagarin 1990).

de Man, 1880

Svalbard (Loof 1971); Taymyr and Severnaya Zemlya, Russia (Kuzmin and Gagarin 1990).

de Man, 1880

Taymyr and Severnaya Zemlya, Russia (Kuzmin and Gagarin 1990).

Ivanova, 1958

Taymyr and Severnaya Zemlya, Russia (Kuzmin and Gagarin 1990).

Thorne, 1939

Taymyr and Severnaya Zemlya, Russia (Kuzmin and Gagarin 1990).

(Cobb, 1913)

Taymyr and Severnaya Zemlya, Russia (Kuzmin and Gagarin 1990).

(Andrássy, 1958)

Taymyr and Severnaya Zemlya, Russia (Kuzmin and Gagarin 1990).

(Cobb, 1920)

Taymyr and Severnaya Zemlya, Russia (Kuzmin and Gagarin 1990).

Allen, 1957

Alaska (Bernard 1992).

Bernard, 1992*

Alaska (Bernard 1992).

(de Man, 1880)

Taymyr and Severnaya Zemlya, Russia (Gagarin 1990, Gagarin 1996, Gagarin 2001b, Gagarin 2001a).

(Altherr, 1953)

Taymyr and Severnaya Zemlya, Russia (Kuzmin and Gagarin 1990).

(Thorne, 1930)

Taymyr and Severnaya Zemlya, Russia (Kuzmin and Gagarin 1990).

Gagarin, 1997*

Novaya Zemlya and Vaigach island, Russia (Gagarin 1997a, Gagarin 2001b).

Andrássy, 1986

Alaska (Andrássy 2003c).

(Bastian, 1865)

Bastian, 1865

Greenland (Allgén 1954).

Gagarin, 1996*

Taymyr and Severnaya Zemlya, Russia (Gagarin 2001b).

(Linstow, 1876)

(Linstow, 1876)

Taymyr and Severnaya Zemlya, Russia (Kuzmin and Gagarin 1990).

(Thorne & Swanger, 1936)

(Thorne & Swanger, 1936)

Taymyr and Severnaya Zemlya, Russia (Kuzmin 1986).

Altherr in Altherr & Delamare-Deboutteville, 1972

Northwest territories, Canada (Mulvey and Anderson 1979).

Dujardin, 1845

Greenland (Ditlevsen 1927); Northwest territories, Canada (Mulvey and Anderson 1979); Lena River estuary, Russia (Gagarin 2001b); Taymyr and Severnaya Zemlya, Russia (Gagarin 1990, Gagarin 1996, Gagarin 2001b, Gagarin 2001a, Kuzmin and Gagarin 1990); Novaya Zemlya and Vaigach island, Russia (Gagarin 1997a, Gagarin 2001b).

(Ditlevsen, 1911)

Taymyr and Severnaya Zemlya, Russia (Kuzmin and Gagarin 1990).

Thorne, 1975

Novaya Zemlya and Vaigach island, Russia (Gagarin 1997a, Gagarin 2001b).

(Micoletzly, 1927)

Taymyr and Severnaya Zemlya, Russia (Kuzmin and Gagarin 1990); Novaya Zemlya and Vaigach island, Russia (Gagarin 1997a, Gagarin 2001b).

(Bütschli, 1873)

Bütschli, 1873;

Svalbard (von Linstow 1900); Taymyr and Severnaya Zemlya, Russia (Kuzmin and Gagarin 1990); Novaya Zemlya and Vaigach island, Russia (Steiner 1916b).

(Cobb in Thorne & Swanger, 1936)

Taymyr and Severnaya Zemlya, Russia (Kuzmin and Gagarin 1990).

Loof, 1969

Taymyr and Severnaya Zemlya, Russia (Gagarin 1990, Gagarin 2001b, Gagarin 2001a).

(Tulaganov, 1949)

Taymyr and Severnaya Zemlya, Russia (Gagarin 1990, Gagarin 2001b, Gagarin 2001a).

(de Man, 1884)

(de Man, 1884)

Greenland (Ditlevsen 1927); Alaska (Andrássy 2003c); Lena River estuary, Russia (Gagarin 2001b); Taymyr and Severnaya Zemlya, Russia (Kuzmin 1972, Kuzmin and Gagarin 1990).

Eveleigh, 1982*

Northwest territories, Canada (Eveleigh 1982).

(Cobb in Thorne, 1939)

Taymyr and Severnaya Zemlya, Russia (Kuzmin and Gagarin 1990).

Mulvey & Anderson, 1979*

Northwest territories, Canada (Mulvey and Anderson 1979).

(Meyl, 1955)

Taymyr and Severnaya Zemlya, Russia (Kuzmin and Gagarin 1990).

(Cobb in Thorne & Swanger, 1936)

Taymyr and Severnaya Zemlya, Russia (Kuzmin and Gagarin 1990).

(Andrássy, 1959)

Taymyr and Severnaya Zemlya, Russia (Kuzmin and Gagarin 1990).

(Andrássy, 1960)

Andrássy, 1960;

Lena River estuary, Russia (Gagarin 1999a, Gagarin 1999b, Gagarin 2001b); Taymyr and Severnaya Zemlya, Russia (Gagarin 1990, Gagarin 1996, Gagarin 1999a, Gagarin 2001a, Gagarin 2001b); Novaya Zemlya and Vaigach island, Russia (Gagarin 1997a, Gagarin 1999a, Gagarin 2001b).

(Thorne, 1929)

Taymyr and Severnaya Zemlya, Russia (Kuzmin and Gagarin 1990); Novaya Zemlya and Vaigach island, Russia (Gagarin 1997a, Gagarin 1999a, Gagarin 2001b).

(de Man, 1880)

de Man, 1880;

Jan Mayen (Allgén 1953); Greenland (Allgén 1954, Ditlevsen 1927); Taymyr and Severnaya Zemlya, Russia (Kuzmin 1972, Kuzmin and Gagarin 1990); Novaya Zemlya and Vaigach island, Russia (Steiner 1916b).

(Thorne & Swanger, 1936)

Taymyr and Severnaya Zemlya, Russia (Kuzmin 1986, Kuzmin and Gagarin 1990).

(Andrássy, 1958)

Taymyr and Severnaya Zemlya, Russia (Kuzmin and Gagarin 1990).

(Bastian, 1865)

Bastian, 1865

Svalbard (Klekowski and Opalinski 1986, Klekowski and Opalinski 1990, Klekowski and Opalinski 1992, Opalinski and Klekowski 1992); Jan Mayen (Allgén 1953); Greenland (Ditlevsen 1927); Alaska (Andrássy 2003c); Taymyr and Severnaya Zemlya, Russia (Gagarin 1996, Gagarin and Kuzmin 1983, Kuzmin 1972, Kuzmin 1976, Kuzmin and Gagarin 1990); Novaya Zemlya and Vaigach island, Russia (Steiner 1916b).

(de Man, 1880)

de Man, 1880;

Jan Mayen (Allgén 1953); Taymyr and Severnaya Zemlya, Russia (Gagarin 1990, Gagarin 2001b, Gagarin 2001a, Kuzmin and Gagarin 1990).

(Bastian, 1865)

de Man, 1876

Jan Mayen (Allgén 1953).

(Bütschli, 1873)

Micoletzky, 1922

Greenland (Allgén 1954).

Loof, 1971*

Svalbard (Loof 1971).

Loof, 1971*

Svalbard (Loof 1971).

Loof, 1971*

Svalbard (Loof 1971).

(de Man, 1880)

de Man, 1880;

Svalbard (Loof 1971); Jan Mayen (Allgén 1953); Greenland (Allgén 1954); Taymyr and Severnaya Zemlya, Russia (Kuzmin 1986, Kuzmin and Gagarin 1990); Novaya Zemlya and Vaigach island, Russia (Steiner 1916b).

(Thorne & Swanger, 1936)

(Thorne & Swanger, 1936)

Taymyr and Severnaya Zemlya, Russia (Kuzmin 1972, Kuzmin and Gagarin 1990).

(de Man, 1918)

Taymyr and Severnaya Zemlya, Russia (Gagarin 1996, Kuzmin and Gagarin 1990).

(Thorne & Swanger, 1936)

(Thorne & Swanger, 1936)

Taymyr and Severnaya Zemlya, Russia (Kuzmin 1972, Kuzmin and Gagarin 1990).

(de Man, 1880)

de Man, 1880;

Svalbard (Loof 1971); Jan Mayen (Allgén 1953); Greenland (Ditlevsen 1927); Alaska (Andrássy 2003c); Taymyr and Severnaya Zemlya, Russia (Kuzmin and Gagarin 1990).

(Skwarra, 1921)

Taymyr and Severnaya Zemlya, Russia (Kuzmin and Gagarin 1990).

Gagarin, 1991*

Taymyr and Severnaya Zemlya, Russia (Gagarin 1990, Gagarin 1991b, Gagarin 2001b, Gagarin 2001a).

(de Man, 1880)

(de Man, 1880)

Svalbard (Loof 1971); Taymyr and Severnaya Zemlya, Russia (Kuzmin and Gagarin 1990).

(Loof, 1961)

Loof, 1961

Svalbard (Loof 1971).

(de Man, 1885)

(de Man, 1885);

Taymyr and Severnaya Zemlya, Russia (Kuzmin 1972, Kuzmin and Gagarin 1990).

(de Man, 1906)

(de Man, 1906)

Taymyr and Severnaya Zemlya, Russia (Kuzmin and Gagarin 1990).

(de Man, 1880)

de Man, 1880

Jan Mayen (Allgén 1953).

(de Man, 1880)

de Man, 1880

Jan Mayen (Allgén 1953).

(de Man, 1880)

Taymyr and Severnaya Zemlya, Russia (Kuzmin and Gagarin 1990).

Thorne, 1939

Taymyr and Severnaya Zemlya, Russia (Kuzmin and Gagarin 1990).

Altherr in Altherr & Delamare-Deboutteville, 1972

Taymyr and Severnaya Zemlya, Russia (Gagarin 1990, Gagarin 2001b, Gagarin 2001a).

Andrássy, 1978

Lena River estuary, Russia (Gagarin 2001b); Novaya Zemlya and Vaigach island, Russia (Gagarin 1997a).

Altherr, 1968

Alaska (Andrássy 2003c).

(Cobb, 1906)

Taymyr and Severnaya Zemlya, Russia (Kuzmin and Gagarin 1990).

(Ditlevsen, 1928)

(Ditlevsen, 1928)

Taymyr and Severnaya Zemlya, Russia (Gagarin 2001b, Kuzmin 1972, Kuzmin and Gagarin 1990); Novaya Zemlya and Vaigach island, Russia (Gagarin 2001b).

(Thorne & Swanger, 1936)

(Thorne & Swanger, 1936)

Taymyr and Severnaya Zemlya, Russia (Kuzmin 1972, Kuzmin and Gagarin 1990).

(Bastian, 1865)

Bastian, 1865;

Jan Mayen (Allgén 1953, Allgén 1954); Greenland (Allgén 1954, Ditlevsen 1927); Taymyr and Severnaya Zemlya, Russia (Gagarin 1990, Gagarin 2001b, Gagarin 2001a, Kuzmin and Gagarin 1990).

(Bastian, 1865)

(Bastian, 1865)

Taymyr and Severnaya Zemlya, Russia (Kuzmin and Gagarin 1990).

(Micoletzky, 1922)

Taymyr and Severnaya Zemlya, Russia (Kuzmin and Gagarin 1990).

(Bastian, 1865)

Bastian, 1865

Jan Mayen (Allgén 1953); Greenland (Allgén 1954).

(Ditlevsen, 1911)

Taymyr and Severnaya Zemlya, Russia (Kuzmin and Gagarin 1990).

Andrássy, 2003*

Alaska (Andrássy 2003c).

(de Man, 1880)

Taymyr and Severnaya Zemlya, Russia (Kuzmin and Gagarin 1990).

(de Man, 1880)

de Man, 1880

Jan Mayen (Allgén 1953).

(Altherr, 1963)

(Altherr, 1963)

Svalbard (Klekowski and Opalinski 1986, Klekowski and Opalinski 1990, Loof 1971).

(Ditlevsen, 1927)

Ditlevsen, 1927*

Svalbard (Loof 1971); Greenland (Ditlevsen 1927).

(Ditlevsen, 1927)

Ditlevsen 1927*

Greenland (Ditlevsen 1927); Taymyr and Severnaya Zemlya, Russia (Elshishka et al. 2012).

(Steiner, 1914)

Svalbard (Loof 1971).

(de Man, 1880)

de Man, 1880

Svalbard (Loof 1971); Jan Mayen (Allgén 1953); Lena River estuary, Russia (Gagarin 2001b); Taymyr and Severnaya Zemlya, Russia (Kuzmin and Gagarin 1990); Novaya Zemlya and Vaigach island, Russia (Gagarin 1997a, Gagarin 2001b, Steiner 1916a).

Elshishka et al., 2012*

Taymyr and Severnaya Zemlya, Russia (Elshishka et al. 2012).

Baqri & Jairajpuri, 1974

Taymyr and Severnaya Zemlya, Russia (Kuzmin 1986, Kuzmin and Gagarin 1990).

Loof, 1971*

Svalbard (Loof 1971); Taymyr and Severnaya Zemlya, Russia (Kuzmin 1986, Kuzmin and Gagarin 1990).

Thorne, 1939

Taymyr and Severnaya Zemlya, (Kuzmin and Gagarin 1990).

(Thorne, 1939)

Thorne, 1939

Nunavut, Canada (Elshishka et al. 2013); Taymyr and Severnaya Zemlya, Russia (Elshishka et al. 2013, Kuzmin and Gagarin 1990).

(Ditlevsen, 1927)

Ditlevsen, 1927*;

Svalbard (Loof 1971); Greenland (Ditlevsen 1927).

Altherr, 1952

Taymyr and Severnaya Zemlya, Russia (Elshishka et al. 2013).

(Thorne & Swanger, 1936)

Taymyr and Severnaya Zemlya, Russia (Kuzmin and Gagarin 1990).

(Brzeski, 1962)

Brzeski, 1962*

Svalbard (Brzeski 1962, Loof 1971).

Andrássy, 2003*

Alaska (Andrássy 2003c).

Ebsary, 1984*

Nunavut, Canada (Ebsary 1984).

Loof, 1971*

Svalbard (Loof 1971); Alaska (Andrássy 2003c).

Thorne, 1939

Taymyr and Severnaya Zemlya, Russia (Kuzmin and Gagarin 1990).

(Thorne & Swanger, 1936)

Taymyr and Severnaya Zemlya, Russia (Kuzmin and Gagarin 1990).

Heyns, 1969

Taymyr and Severnaya Zemlya, Russia (Kuzmin and Gagarin 1990).

Loof & Jairajpuri, 1968

Taymyr and Severnaya Zemlya, Russia (Kuzmin and Gagarin 1990).

(Bütschli, 1873)

Taymyr and Severnaya Zemlya, Russia (Kuzmin and Gagarin 1990).

Thorne, 1939

Svalbard (Loof 1971); Taymyr and Severnaya Zemlya, Russia (Kuzmin and Gagarin 1990).

Steiner, 1914

Taymyr and Severnaya Zemlya, Russia (Kuzmin and Gagarin 1990).

Thorne, 1939

Svalbard (Loof 1971); Taymyr and Severnaya Zemlya, Russia (Kuzmin 1972, Kuzmin 1976, Kuzmin and Gagarin 1990).

Baqri & Jairajpuri, 1969

Taymyr and Severnaya Zemlya, Russia (Kuzmin 1972, Kuzmin and Gagarin 1990).

Thorne, 1939

Taymyr and Severnaya Zemlya, Russia (Kuzmin 1972, Kuzmin and Gagarin 1990).

(Tsalolikhin, 1974)

Lena River estuary, Russia (Gagarin 2001b); Taymyr and Severnaya Zemlya, Russia (Gagarin 1990, Gagarin 1993, Gagarin 2001b, Gagarin 2001a); Novaya Zemlya and Vaigach island, Russia (Gagarin 1997a, Gagarin 2001b).

Gagarin, 1984

Taymyr and Severnaya Zemlya, Russia (Gagarin 1996, Gagarin 2001b).

Coetzee, 1968

Taymyr and Severnaya Zemlya, Russia (Gagarin 2001b).

Schneider, 1925

Northwest territories, Canada (Mulvey 1978).

Skwarra, 1921

Nunavut, Canada (Mulvey 1967b, Mulvey 1978).

Mulvey, 1978*

Northwest territories, Canada (Mulvey 1978).

Gagarin, 1991*

Taymyr and Severnaya Zemlya, Russia (Gagarin 1990, Gagarin 1991b, Gagarin 1993, Gagarin 2001a, Gagarin 2001b).

Bastian, 1865

Northwest territories, Canada (Mulvey 1978); Taymyr and Severnaya Zemlya, Russia (Gagarin 1990, Gagarin 1996, Gagarin 2001b, Gagarin 2001a, Kuzmin and Gagarin 1990); Novaya Zemlya and Vaigach island, Russia (Gagarin 1997a, Gagarin 2001b).

Mulvey, 1978*

Northwest territories, Canada (Mulvey 1978).

(Thorne, 1929)

Nunavut, Canada (Mulvey 1967a, Mulvey 1978); Taymyr and Severnaya Zemlya, Russia (Kuzmin and Gagarin 1990).

Gagarin, 2001*

Novaya Zemlya and Vaigach island, Russia (Gagarin 2001c, Gagarin 2001b).

(Dujardin, 1845)

Svalbard (Klekowski and Opalinski 1986, Klekowski and Opalinski 1990); Nunavut, Canada (Mulvey 1967a); Alaska (Andrássy 2003a); Taymyr and Severnaya Zemlya, Russia (Kuzmin and Gagarin 1990); Novaya Zemlya and Vaigach island, Russia (Gagarin 1997a, Gagarin 2001b).

Cobb, 1917

Nunavut, Canada (Mulvey 1967a).

Gagarin, 2000*

Lena River estuary, Russia (Gagarin 2000, Gagarin 2001b).

(Ditlevsen, 1911)

Ditlevsen, 1911

Greenland (Ditlevsen 1927).

(Mulvey, 1967)

Taymyr and Severnaya Zemlya, Russia (Kuzmin and Gagarin 1990).

(Bastina, 1865)

Bastian, 1865

Jan Mayen (Allgén 1953); Greenland (Allgén 1954, Ditlevsen 1927); Taymyr and Severnaya Zemlya, Russia (Kuzmin and Gagarin 1990); Novaya Zemlya and Vaigach island, Russia (Steiner 1916b).

Mulvey, 1978*

Nunavut, Canada (Mulvey 1978).

(de Man, 1904)

Northwest territories, Canada (Mulvey 1978).

(de Man, 1880)

Alaska (Andrássy 2003a).

(Menzel, 1913)

(Menzel, 1913)

Taymyr and Severnaya Zemlya, Russia (Kuzmin 1972, Kuzmin and Gagarin 1990).

(Bütschli, 1873)

Bütschli, 1873

Greenland (Ditlevsen 1927); Alaska (Andrássy 2003a); Taymyr and Severnaya Zemlya, Russia (Kuzmin and Gagarin 1990); Novaya Zemlya and Vaigach island, Russia (Steiner 1916b).

Gagarin, 1993

Novaya Zemlya and Vaigach island, Russia (Gagarin 1997a, Gagarin 2001b).

Cobb, 1917

Taymyr and Severnaya Zemlya, Russia (Gagarin 2001b).

(Cobb in Cobb, 1915)

Northwest territories, Canada (Mulvey 1978).

(Cobb, 1917)

Taymyr and Severnaya Zemlya, Russia (Kuzmin and Gagarin 1990).

Andrássy, 2003*

Alaska (Andrássy 2003a).

Gagarin, 1999*

Gagarin, 1999 (lapsus)

Novaya Zemlya and Vaigach island, Russia (Gagarin 1999a, Gagarin 2001b).

(de Man, 1876)

Lena River estuary, Russia (Gagarin 2001b); Taymyr and Severnaya Zemlya, Russia (Gagarin 1996); Novaya Zemlya and Vaigach island, Russia (Gagarin 1997a, Gagarin 2001b).

(Linstow, 1876)

Novaya Zemlya and Vaigach island, Russia (Gagarin 1997a, Gagarin 2001b).

(Stefanski, 1915)

Steiner, 1916*;

Taymyr and Severnaya Zemlya, Russia (Kuzmin 1976, Kuzmin and Gagarin 1990); Novaya Zemlya and Vaigach island, Russia (Steiner 1916b).

(de Man, 1880)

Nunavut, Canada (Mulvey 1969c); Taymyr and Severnaya Zemlya, Russia (Kuzmin 1972, Kuzmin and Gagarin 1990).

Altherr, 1952

Svalbard (Loof 1971).

(de Man, 1876)

Svalbard (Janiec 1997, Loof 1971).

(de Man, 1880)

Steiner, 1916*

Greenland (Ditlevsen 1927); Nunavut, Canada (Mulvey 1969c); Lena River estuary, Russia (Gagarin 2001b); Taymyr and Severnaya Zemlya, Russia (Kuzmin and Gagarin 1990); Novaya Zemlya and Vaigach island, Russia (Gagarin 1997a, Gagarin 2001b, Steiner 1916a).

de Man, 1880

Steiner, 1916*

Nunavut, Canada (Mulvey 1969c); Lena River estuary, Russia (Gagarin 2001b); Taymyr and Severnaya Zemlya, Russia (Gagarin 1990, Gagarin 1996, Gagarin 2001b, Gagarin 2001a, Kuzmin 1972, Kuzmin and Gagarin 1990); Novaya Zemlya and Vaigach island, Russia (Gagarin 1997a, Gagarin 2001b, Steiner 1916a).

de Man, 1880

Taymyr and Severnaya Zemlya, Russia (Kuzmin and Gagarin 1990).

(Mulvey, 1969)

Mulvey 1969*

Nunavut, Canada (Mulvey 1969c); Novaya Zemlya and Vaigach island, Russia (Gagarin 1997a, Gagarin 2001b).

(Micoletzky, 1913)

Lena River estuary, Russia (Gagarin 2001b); Taymyr and Severnaya Zemlya, Russia (Gagarin 1996, Kuzmin 1976, Kuzmin and Gagarin 1990); Novaya Zemlya and Vaigach island, Russia (Gagarin 1997a, Gagarin 2001b).

Mulvey, 1969*

Nunavut, Canada (Mulvey 1969c).

(Bütschli, 1873)

Taymyr and Severnaya Zemlya, Russia (Gagarin 1996, Gagarin and Kuzmin 1983, Kuzmin and Gagarin 1990).

Gagarin, 1993*

Taymyr and Severnaya Zemlya, Russia (Gagarin 1990, Gagarin 1993, Gagarin 2001b, Gagarin 2001a).

Gagarin, 2000*

Lena River estuary, Russia (Gagarin 2000, Gagarin 2001b).

(Ditlevsen, 1918)

Novaya Zemlya and Vaigach island, Russia (Gagarin 1999a, Gagarin 2001b).

(de Man, 1880)

de Man, 1880

Jan Mayen (Allgén 1953, Tchesunov and Pletnikova 1986); Taymyr and Severnaya Zemlya, Russia (Kuzmin 1972, Kuzmin 1976, Kuzmin and Gagarin 1990).

Schuurmans Stekhoven, 1935

Novaya Zemlya and Vaigach island, Russia (Gagarin 1997b, Gagarin 2001a).

Andrássy, 1981

Novaya Zemlya and Vaigach island, Russia (Gagarin 1997b, Gagarin 2001a).

(Bastian, 1865)

Bastian, 1865

Svalbard (Loof 1971); Nunavut, Canada (Mulvey 1969c); Taymyr and Severnaya Zemlya, Russia (Gagarin 1996); Novaya Zemlya and Vaigach island, Russia (Gagarin 1997b, Gagarin 1997a, Gagarin 1999a, Gagarin 2001b).

(Bastian, 1865)

Bastian, 1865 (lapsus);

Svalbard (Janiec 1997, Loof 1971); Lena River estuary, Russia (Gagarin 2001b); Taymyr and Severnaya Zemlya, Russia (Kuzmin 1972, Kuzmin and Gagarin 1990); Novaya Zemlya and Vaigach island, Russia (Gagarin 1997a, Gagarin 2001b, Steiner 1916a).

Gagarin, 1997*

Novaya Zemlya and Vaigach island, Russia (Gagarin 1997b, Gagarin 1997a, Gagarin 1999a, Gagarin 2001b).

Gagarin, 1996

Taymyr and Severnaya Zemlya, Russia (Gagarin 2001b).

(Daday, 1899)

Taymyr and Severnaya Zemlya, Russia (Gagarin 1996, Gagarin 2001b).

(Daday, 1896)

Daday, 1896 (lapsus)

Greenland (Ditlevsen 1927).

(Bütschli, 1873)

Taymyr and Severnaya Zemlya, Russia (Gagarin 2001b, Kuzmin and Gagarin 1990).

(de Man, 1880)

de Man, 1880

Taymyr and Severnaya Zemlya, Russia (Kuzmin 1972, Kuzmin and Gagarin 1990).

Gagarin, 1991

Novaya Zemlya and Vaigach island, Russia (Gagarin 1999a, Gagarin 2001b).

(de Man, 1880)

de Man, 1880;

Svalbard (Janiec 1997, Loof 1971); Jan Mayen (Allgén 1953); Nunavut, Canada (Mulvey 1969c); Lena River estuary, Russia (Gagarin 2001b); Taymyr and Severnaya Zemlya, Russia (Gagarin 1996, Kuzmin and Gagarin 1990); Novaya Zemlya and Vaigach island, Russia (Gagarin 1997a, Gagarin 1999a, Gagarin 2001b, Steiner 1916a).

(Meyl, 1953)

Novaya Zemlya and Vaigach island, Russia (Gagarin 1997a, Gagarin 2001b).

(Meyl, 1954)

Meyl, 1954

Nunavut, Canada (Mulvey 1969c).

(Butschli, 1873)

Butschli, 1873 (lapsus)

Svalbard (Loof 1971); Taymyr and Severnaya Zemlya, Russia (Kuzmin and Gagarin 1990); Novaya Zemlya and Vaigach island, Russia (Steiner 1916b).

(Bastian, 1865)

Bastian, 1865

Jan Mayen (Allgén 1953); Novaya Zemlya and Vaigach island, Russia (Gagarin 1999a, Gagarin 2001b).

Gagarin, 1997*

Novaya Zemlya and Vaigach island, Russia (Gagarin 1997b, Gagarin 1997a, Gagarin 2001c, Gagarin 2001b).

Bastian, 1865

Svalbard (Janiec 1997, Loof 1971); Taymyr and Severnaya Zemlya, Russia (Gagarin and Kuzmin 1983, Kuzmin and Gagarin 1990).

(de Man, 1880)

de Man, 1880

Jan Mayen (Allgén 1953).

(Gagarin, 1997)

Gagarin, 1997*;

Novaya Zemlya and Vaigach island, Russia (Gagarin 1997b, Gagarin 1997a, Gagarin 2001b).

(Gagarin, 2000)

Gagarin, 2000

Novaya Zemlya and Vaigach island, Russia (Gagarin 2001b).

Andrássy, 1959

Taymyr and Severnaya Zemlya, Russia (Kuzmin 1972, Kuzmin and Gagarin 1990).

de Man, 1880

Taymyr and Severnaya Zemlya, Russia (Kuzmin and Gagarin 1990).

de Man, 1880

Svalbard (Loof 1971); Nunavut, Canada (Mulvey 1969c); Lena River estuary, Russia (Gagarin 2001b); Taymyr and Severnaya Zemlya, Russia (Kuzmin 1972, Kuzmin and Gagarin 1990).

de Man, 1880

Greenland (Ditlevsen 1927).

(de Man, 1921)

Taymyr and Severnaya Zemlya, Russia (Kuzmin and Gagarin 1990).

(Ditlevsen, 1928)

(Hirschmann, 1952)

Svalbard (Janiec 1997); Alaska (Andrássy 2003b); Lena River estuary, Russia (Gagarin 2001b); Taymyr and Severnaya Zemlya, Russia (Gagarin 1990, Gagarin 1993, Gagarin 2001b, Gagarin 2001a, Kuzmin and Gagarin 1990).

(Bastian, 1865)

Bastian, 1865

Svalbard (Loof 1971); Jan Mayen (Allgén 1953); Greenland (Ditlevsen 1927); Taymyr and Severnaya Zemlya, Russia (Gagarin 1990, Gagarin 1996, Gagarin 2001b, Gagarin 2001a, Kuzmin and Gagarin 1990); Novaya Zemlya and Vaigach island, Russia (Gagarin 1997a, Gagarin 1999a, Gagarin 2001b, Steiner 1916a).

Allen and Noffsinger, 1968

Svalbard (Klekowski and Opalinski 1986, Klekowski and Opalinski 1990, Klekowski and Opalinski 1992, Loof 1971).

Andrássy, 2003*

Alaska (Andrássy 2003b).

(Sanwal, 1968)

Sanwal, 1968*

Nunavut, Canada (Sanwal 1968, Zell 1993).

Andrássy, 2003*

Alaska (Andrássy 2003b).

Bastian, 1865

Svalbard (Loof 1971); Alaska (Andrássy 2003b); Taymyr and Severnaya Zemlya, Russia (Gagarin 2001b, Kuzmin and Gagarin 1990); Novaya Zemlya and Vaigach island, Russia (Gagarin 1997a, Gagarin 2001b).

Ebsary, 1985

Svalbard (Zell 1993).

Andrássy, 1984

Svalbard (Janiec 1997).

Butschli, 1873

Svalbard (Loof 1971).

Bütschli, 1873

(Bütschli, 1873)

Svalbard (Loof 1971); Greenland (Zell 1993); Alaska (Andrássy 2003b); Taymyr and Severnaya Zemlya, Russia (Kuzmin and Gagarin 1990).

Zullini, 1978

sensu Zell, 1993

Nunavut, Canada (Ebsary 1985, Zell 1993).

Bastian, 1865

Svalbard (Klekowski and Opalinski 1986, Klekowski and Opalinski 1990, Klekowski and Opalinski 1992, Menzel 1920); Jan Mayen (Allgén 1953); Greenland (Ditlevsen 1927); Alaska (Andrássy 2003b); Lena River estuary, Russia (Gagarin 2001b); Taymyr and Severnaya Zemlya, Russia (Gagarin 1990, Gagarin 1996, Gagarin 2001b, Gagarin 2001a, Kuzmin and Gagarin 1990); Novaya Zemlya and Vaigach island, Russia (Gagarin 1997a, Gagarin 1999a, Gagarin 2001b).

Bütschli, 1873

Svalbard (Menzel 1920, Zell 1993); Iceland (Zell 1993); Greenland (Allgén 1954).

Maggenti, 1961

(Maggenti, 1961)

Svalbard (Loof 1971, Zell 1993); Taymyr and Severnaya Zemlya, Russia (Kuzmin 1976, Kuzmin and Gagarin 1990).

Maggenti, 1961

Taymyr and Severnaya Zemlya, Russia (Kuzmin and Gagarin 1990).

de Man, 1880

Novikova & Gagarin, 1971

Svalbard (Loof 1971, Zell 1993); Jan Mayen (Allgén 1953); Taymyr and Severnaya Zemlya, Russia (Kuzmin 1986, Kuzmin and Gagarin 1990).

Andrássy, 1958

Svalbard (Loof 1971, Zell 1993); Alaska (Andrássy 2003b); Taymyr and Severnaya Zemlya, Russia (Kuzmin and Gagarin 1990).

Cobb, 1893

Nunavut, Canada (Zell 1993).

Butschli, 1873

Svalbard (Loof 1971, Zell 1993); Jan Mayen (Allgén 1953); Iceland (Zell 1993); Taymyr and Severnaya Zemlya, Russia (Kuzmin and Gagarin 1990); Novaya Zemlya and Vaigach island, Russia (Steiner 1916b).

Kuzmin, 1979

Svalbard (Zell 1993).

Yeates, 1970

Svalbard (Zell 1993).

Andrássy, 1952

Svalbard (Janiec 1997).

de Man, 1880

Lena River estuary, Russia (Gagarin 2001b); Taymyr and Severnaya Zemlya, Russia (Gagarin 1996, Gagarin 2001b); Novaya Zemlya and Vaigach island, Russia (Gagarin 1997a, Gagarin 1999a, Gagarin 2001b).

Bastian, 1865

Bastian, 1865

Svalbard (Janiec 1997, Klekowski and Opalinski 1986, Klekowski and Opalinski 1990, Klekowski and Opalinski 1992, Loof 1971, Opalinski and Klekowski 1992); Jan Mayen (Allgén 1953); Iceland (Zell 1993); Alaska (Andrássy 2003b); Taymyr and Severnaya Zemlya, Russia (Gagarin 1990, Gagarin 1996, Gagarin 2001b, Gagarin 2001a, Gagarin and Kuzmin 1983, Kuzmin 1976, Kuzmin and Gagarin 1990); Novaya Zemlya and Vaigach island, Russia (Gagarin 1997a, Gagarin 2001b, Steiner 1916a).

Bastian, 1865

Svalbard (Loof 1971, Zell 1993); Iceland (Zell 1993); Taymyr and Severnaya Zemlya, Russia (Kuzmin and Gagarin 1990); Novaya Zemlya and Vaigach island, Russia (Gagarin 1997a, Gagarin 2001b).

de Man, 1880

de Man, 1880

Svalbard (Loof 1971, Zell 1993); Jan Mayen (Allgén 1953, Steiner 1916a); Greenland (Allgén 1954); Lena River estuary, Russia (Gagarin 2001b); Taymyr and Severnaya Zemlya, Russia (Gagarin and Kuzmin 1983, Kuzmin and Gagarin 1990); Novaya Zemlya and Vaigach island, Russia (Gagarin 1997a, Gagarin 2001b, Steiner 1916a).

Zell, 1993

Iceland (Zell 1993).

Bastian, 1865

Lena River estuary, Russia (Gagarin 2001b); Taymyr and Severnaya Zemlya, Russia (Gagarin 1996); Novaya Zemlya and Vaigach island, Russia (Gagarin 1997a, Gagarin 1999a, Gagarin 2001b).

Maggenti, 1961

Iceland (Zell 1993); Taymyr and Severnaya Zemlya, Russia (Kuzmin 1976).

Bastian, 1865

Svalbard (Zell 1993).

(Bütschli, 1873)

Bütschli, 1873

Jan Mayen (Steiner 1916a); Nunavut, Canada (Anderson 1966); Taymyr and Severnaya Zemlya, Russia (Kuzmin and Gagarin 1990).

Loof, 1971*

Svalbard (Boström 1987a, Holovachov et al. 2003, Loof 1971).

Anderson, 1966*

Nunavut, Canada (Anderson 1966).

(de Man, 1880)

Taymyr and Severnaya Zemlya, Russia (Kuzmin and Gagarin 1990); Novaya Zemlya and Vaigach island, Russia (Gagarin 1997a, Gagarin 2001b).

(de Man, 1880)

de Man, 1880;

Svalbard (Boström 1989, Loof 1971); Jan Mayen (Allgén 1953); Taymyr and Severnaya Zemlya, Russia (Kuzmin 1972, Kuzmin 1976, Kuzmin and Gagarin 1990); Novaya Zemlya and Vaigach island, Russia (Gagarin 1997a, Gagarin 2001b, Steiner 1916a).

(de Man, 1880)

Taymyr and Severnaya Zemlya, Russia (Kuzmin 1972, Kuzmin and Gagarin 1990).

de Man, 1880

Taymyr and Severnaya Zemlya, Russia (Kuzmin 1986, Kuzmin and Gagarin 1990).

Andrássy, 1958

Anderson, 1969*

Svalbard (Boström 1989, Loof 1971); Nunavut, Canada (Anderson 1969); Taymyr and Severnaya Zemlya, Russia (Kuzmin and Gagarin 1990); Novaya Zemlya and Vaigach island, Russia (Gagarin 1997a, Gagarin 2001b).

Andrássy, 1968

Anderson, 1969*

Nunavut, Canada (Anderson 1969).

Anderson, 1969*

Svalbard (Boström 1989, Loof 1971); Nunavut, Canada (Anderson 1969); Taymyr and Severnaya Zemlya, Russia (Kuzmin and Gagarin 1990).

Andrássy, 1958

Taymyr and Severnaya Zemlya, Russia (Kuzmin and Gagarin 1990).

(Bütschli, 1873)

Svalbard (Boström 1989); Jan Mayen (Allgén 1953); Taymyr and Severnaya Zemlya, Russia (Kuzmin 1972, Kuzmin 1976, Kuzmin and Gagarin 1990); Novaya Zemlya and Vaigach island, Russia (Steiner 1916b).

(Bütschli, 1873)

Cobb, 1924

Taymyr and Severnaya Zemlya, Russia (Kuzmin 1972, Kuzmin and Gagarin 1990).

Loof, 1971*

Svalbard (Loof 1971).

(Schneider, 1866)

Svalbard (Loof 1971); Taymyr and Severnaya Zemlya, Russia (Kuzmin 1976, Kuzmin and Gagarin 1990).

de Man, 1876

species inquirenda

Jan Mayen (Allgén 1953).

Bastian, 1865

Svalbard (Loof 1971); Jan Mayen (Allgén 1953); Taymyr and Severnaya Zemlya, Russia (Kuzmin and Gagarin 1990).

(de Man, 1884)

de Man, 1884

Svalbard (Menzel 1920); Jan Mayen (Steiner 1916a); Taymyr and Severnaya Zemlya, Russia (Kuzmin and Gagarin 1990).

Loof, 1971*

Svalbard (Loof 1971).

(de Man, 1880)

de Man, 1880

Svalbard (Loof 1971).

(Thorne, 1925)

Svalbard (Boström 1987a, Loof 1971).

Loof, 1971*

Svalbard (Boström 1987a, Loof 1971).

(Kozlovska & Roguska-Wasilewska, 1963)

Taymyr and Severnaya Zemlya, Russia (Kuzmin and Gagarin 1990).

(de Man, 1876)

de Man, 1876

Svalbard (Loof 1971); Jan Mayen (Allgén 1953).

De Coninck, 1943

Taymyr and Severnaya Zemlya, Russia (Kuzmin 1972, Kuzmin and Gagarin 1990).

(Bastian, 1865)

Taymyr and Severnaya Zemlya, Russia (Kuzmin 1972, Kuzmin and Gagarin 1990).

(de Man, 1880)

de Man, 1880;

Svalbard (Loof 1971); Jan Mayen (Allgén 1953); Taymyr and Severnaya Zemlya, Russia (Gagarin 2001b, Kuzmin 1972, Kuzmin and Gagarin 1990).

(Maupas, 1900)

Taymyr and Severnaya Zemlya, Russia (Kuzmin 1972, Kuzmin and Gagarin 1990).

Sumenkova in Sumenkova & Razjivin, 1968

Loof, 1971*

Svalbard (Boström 1987a, Loof 1971).

(Thorne, 1925)

Taymyr and Severnaya Zemlya, Russia (Kuzmin and Gagarin 1990).

Steiner, 1940

Taymyr and Severnaya Zemlya, Russia (Kuzmin and Gagarin 1990).

(Steiner, 1914)

(Steiner, 1914);

Taymyr and Severnaya Zemlya, Russia (Kuzmin 1972, Kuzmin and Gagarin 1990).

(Thorne, 1925)

Loof, 1971*

Svalbard (Loof 1971).

(de Man, 1880)

(Thorne, 1925);

Svalbard (Loof 1971); Taymyr and Severnaya Zemlya, Russia (Kuzmin 1972, Kuzmin and Gagarin 1990).

Boström, 1987*

Svalbard (Boström 1987b).

von Linstow, 1877

Svalbard (Loof 1971); Taymyr and Severnaya Zemlya, Russia (Kuzmin and Gagarin 1990).

Sanwal, 1965*

Svalbard (Loof 1971); Nunavut, Canada (Sanwal 1965); Lena River estuary, Russia (Gagarin 2001b).

Franklin, 1952

Lena River estuary, Russia (Gagarin 2001b).

Haque, 1968

Taymyr and Severnaya Zemlya, Russia (Kuzmin 1972, Kuzmin and Gagarin 1990).

(Steiner, 1914)

Steiner, 1914

Jan Mayen (Steiner 1916a).

Sanwal, 1965*

Nunavut, Canada (Sanwal 1965); Taymyr and Severnaya Zemlya, Russia (Gagarin 2001b).

(Bastian, 1865)

de Man, 1876

Jan Mayen (Steiner 1916a); Taymyr and Severnaya Zemlya, Russia (Gagarin 1996, Kuzmin and Gagarin 1990); Novaya Zemlya and Vaigach island, Russia (Steiner 1916b).

Gagarin, 1997*

Novaya Zemlya and Vaigach island, Russia (Gagarin 1997c, Gagarin 1997a, Gagarin 2001b).

Franklin, 1957

Taymyr and Severnaya Zemlya, Russia (Kuzmin 1986, Kuzmin and Gagarin 1990).

(Cobb, 1926)

Taymyr and Severnaya Zemlya, Russia (Kuzmin 1972, Kuzmin and Gagarin 1990).

(de Man, 1884)

Greenland (Brzeski 1999); Taymyr and Severnaya Zemlya, Russia (Kuzmin and Gagarin 1990); Novaya Zemlya and Vaigach island, Russia (Gagarin 1997a, Gagarin 2001b).

(de Man, 1921)

de Man, 1921

Svalbard (Loof 1971); Greenland (Brzeski 1999); Taymyr and Severnaya Zemlya, Russia (Kuzmin and Gagarin 1990); Novaya Zemlya and Vaigach island, Russia (Gagarin 1997a, Gagarin 2001b).

Loof, 1971*

Svalbard (Loof 1971).

(de Man, 1880)

de Man, 1880;

Svalbard (Loof 1971); Jan Mayen (Allgén 1953); Taymyr and Severnaya Zemlya, Russia (Kuzmin 1986, Kuzmin and Gagarin 1990); Novaya Zemlya and Vaigach island, Russia (Steiner 1916b).

(Wu, 1969)

Wu, 1969*

Nunavut, Canada (Wu 1969b).

(Thorne & Malek, 1968)

Lena River estuary, Russia (Gagarin 2001b).

(Andrássy, 1954)

Andrássy, 1954;

Taymyr and Severnaya Zemlya, Russia (Kuzmin 1986, Kuzmin and Gagarin 1990).

(Brzeski, 1963)

Brzeski, 1963

Taymyr and Severnaya Zemlya, Russia (Gagarin and Kuzmin 1983, Kuzmin and Gagarin 1990).

(Szczygiel, 1970)

Greenland (Brzeski 1999).

(Bütschli, 1873)

species inquirenda;

Taymyr and Severnaya Zemlya, Russia (Kuzmin 1972, Kuzmin 1976, Kuzmin and Gagarin 1990); Novaya Zemlya and Vaigach island, Russia (Steiner 1916b).

(Thorne & Malek, 1968)

Greenland (Brzeski 1999).

(Wu, 1969)

Wu, 1969*;

Nunavut, Canada (Wu 1969b, Wu 1969a).

(Cobb, 1893)

species inquirenda

Taymyr and Severnaya Zemlya, Russia (Kuzmin and Gagarin 1990).

(Andrássy, 1958)

Greenland (Brzeski 1999).

Andrássy, 1954

Wu, 1969*

Greenland (Brzeski 1999); Nunavut, Canada (Wu 1969b).

(Szczygiel, 1969)

Greenland (Brzeski 1999).

(Andrássy, 1954)

Andrássy, 1954

Svalbard (Loof 1971); Taymyr and Severnaya Zemlya, Russia (Kuzmin and Gagarin 1990).

Andrássy, 1968

Lena River estuary, Russia (Gagarin 2001b); Taymyr and Severnaya Zemlya, Russia (Kuzmin and Gagarin 1990).

(Steiner, 1914)

Steiner, 1914

Svalbard (Loof 1971); Novaya Zemlya and Vaigach island, Russia (Gagarin 1997a, Gagarin 2001b).

(Massey, 1969)

Wu, 1970*

Nunavut, Canada (Wu 1970).

Bastian, 1865

(Bastian, 1865);

Svalbard (Loof 1971); Jan Mayen (Allgén 1953); Greenland (Brzeski 1999, Ditlevsen 1927); Lena River estuary, Russia (Gagarin 2001b); Taymyr and Severnaya Zemlya, Russia (Gagarin and Kuzmin 1983, Kuzmin 1972, Kuzmin 1976, Kuzmin and Gagarin 1990); Novaya Zemlya and Vaigach island, Russia (Gagarin 1997a, Gagarin 2001b, Steiner 1916a).

Wu, 1969*

Nunavut, Canada (Wu 1969b).

Wu, 1969*

Nunavut, Canada (Wu 1969b).

(Anderson & Ebsary, 1982)

Anderson & Ebsary, 1982*

Nunavut, Canada (Anderson and Ebsary 1982).

(Mulvey, 1969)

Mulvey, 1969*

Svalbard (Loof 1971); Nunavut, Canada (Mulvey 1969a); Taymyr and Severnaya Zemlya, Russia (Kuzmin 1986, Kuzmin and Gagarin 1990).

(Allen, 1955)

(Allen, 1955)

Taymyr and Severnaya Zemlya, Russia (Kuzmin and Gagarin 1990).

(Anderson & Ebsary, 1982)

Anderson & Ebsary, 1982*

Nunavut, Canada (Anderson and Ebsary 1982).

(Thorne, 1949)

Thorne, 1949

Taymyr and Severnaya Zemlya, Russia (Kuzmin 1986, Kuzmin and Gagarin 1990).

(Siddiqi, 1979)

sensu Loof, 1971;

Svalbard (Loof 1971, Siddiqi 1979).

(Geraert, 1966)

Geraert, 1966

Svalbard (Loof 1971).

(Anderson & Ebsary, 1982)

Anderson & Ebsary, 1982*

Nunavut, Canada (Anderson and Ebsary 1982).

Powers, Baldwin & Bell, 1983*

Alaska (Powers et al. 1983).

(Allen, 1955)

Allen, 1955

Taymyr and Severnaya Zemlya, Russia (Kuzmin 1972, Kuzmin and Gagarin 1990).

(Allen, 1955)

Allen, 1955

Svalbard (Loof 1971); Greenland (Brzeski 1999); Nunavut, Canada (Mulvey 1969a); Taymyr and Severnaya Zemlya, Russia (Kuzmin and Gagarin 1990).

(Allen, 1955)

Taymyr and Severnaya Zemlya, Russia (Kuzmin and Gagarin 1990).

(Allen, 1955)

Taymyr and Severnaya Zemlya, Russia (Kuzmin and Gagarin 1990).

(Mulvey, 1969)

Mulvey, 1969*

Nunavut, Canada (Mulvey 1969a).

(Bütschli, 1873)

(Bütschli, 1873)

Taymyr and Severnaya Zemlya, Russia (Kuzmin 1972, Kuzmin 1976, Kuzmin and Gagarin 1990).

Allen, 1955

(Allen, 1955);

Greenland (Brzeski 1999); Taymyr and Severnaya Zemlya, Russia (Kuzmin 1986, Kuzmin and Gagarin 1990).

Allen, 1955

Svalbard (Loof 1971).

Allen, 1955

(Allen, 1955)

Taymyr and Severnaya Zemlya, Russia (Kuzmin and Gagarin 1990).

(Hopper, 1959)

(Hopper, 1959)

Taymyr and Severnaya Zemlya, Russia (Kuzmin and Gagarin 1990).

Perry in Perry, Darling & Thorne, 1959

Greenland (Brzeski 1999).

Loof, 1971*

Svalbard (Loof 1971); Greenland (Brzeski 1999).

Yuen, 1964

Taymyr and Severnaya Zemlya, Russia (Kuzmin 1972, Kuzmin and Gagarin 1990).

Winslow, 1958

Svalbard (Loof 1971).

(Thorne, 1935)

(Thorne, 1935)

Svalbard (Loof 1971).

(de Man, 1880)

(de Man, 1880)

Jan Mayen (Allgén 1953).

(de Man, 1880)

Cobb (lapsus)

Svalbard (von Linstow 1900).

Gagarin, 2000*

Novaya Zemlya and Vaigach island, Russia (Gagarin 1997a, Gagarin 2000, Gagarin 2001b).

Cobb in Taylor, 1936

Wu, 1965*

Svalbard (Loof 1971); Nunavut, Canada (Wu 1965).

(de Man, 1880)

species inquirenda;

Jan Mayen (Allgén 1953).

Jenkins, 1956

Greenland (Brzeski 1999).

(Steinbuch, 1799)

Nunavut, Canada (Mulvey 1963).

(Bütschli, 1873)

(Bütschli, 1873)

Taymyr and Severnaya Zemlya, Russia (Gagarin 2001b).

Brzeski, 1991

Greenland (Brzeski 1999).

Anderson & Mulvey, 1980*

Nunavut, Canada (Anderson and Mulvey 1980).

Heyns, 1964

Greenland (Brzeski 1999).

(de Man, 1880)

Taymyr and Severnaya Zemlya, Russia (Kuzmin 1972, Kuzmin and Gagarin 1990).

Thorne, 1941

Nunavut, Canada (Mulvey 1969b); Taymyr and Severnaya Zemlya, Russia (Kuzmin and Gagarin 1990).

Khan, 1965

Nunavut, Canada (Mulvey 1969b).

Mulvey, 1969*

Nunavut, Canada (Mulvey 1969b).

Andrássy, 1960

Nunavut, Canada (Mulvey 1969b).

(Cobb, 1922)

Nunavut, Canada (Das 1964).

Mulvey, 1969*

Nunavut, Canada (Mulvey 1969b).

Das, 1964*

Nunavut, Canada (Das 1964).

(Thorne & Malek, 1968)

Thorne & Malek, 1968

Greenland (Brzeski 1999); Nunavut, Canada (Mulvey 1969b).

Steiner, 1914

Novaya Zemlya and Vaigach island, Russia (Steiner 1916b).

Richters, 1905

Svalbard (Loof 1971).

(Leydig, 1854)

Lena River estuary, Russia (Gagarin 2001b).

(Micoletzky, 1922)

Taymyr and Severnaya Zemlya, Russia (Gagarin 1990, Gagarin 2001b, Gagarin 2001a, Gagarin 2008).

(Bastian, 1865)

Taymyr and Severnaya Zemlya, Russia (Kuzmin and Gagarin 1990).

Gagarin, 1995*

Lena River estuary, Russia (Gagarin 1995, Gagarin 2001b); Novaya Zemlya and Vaigach island, Russia (Gagarin 1997a).

(Gagarin, 1983)

(Gagarin, 1983)

Lena River estuary, Russia (Gagarin 2001b); Taymyr and Severnaya Zemlya, Russia (Gagarin 1990, Gagarin 2001b, Gagarin 2001a).

(Gagarin, 1983)

(Gagarin, 1983)

Taymyr and Severnaya Zemlya, Russia (Gagarin 2001b).

Gagarin, 1993

Taymyr and Severnaya Zemlya, Russia (Gagarin 1996, Gagarin 2001b).

(de Man, 1876)

de Man, 1876

Jan Mayen (Allgén 1953).

(Schneider, 1866)

(de Man, 1880)

Jan Mayen (Allgén 1953).

Dujardin, 1844

Bütschli, 1873

Jan Mayen (Allgén 1953).

Aurivillius, 1883*

Inquirenda et incertae sedis. Svalbard (Aurivillius 1883).

Allgén, 1953*

Inquirenda et incertae sedis. Jan Mayen (Allgén 1953).

Allgén, 1953*

Inquirenda et incertae sedis. Jan Mayen (Allgén 1953).

Allgén, 1953*

Inquirenda et incertae sedis. Jan Mayen (Allgén 1953).

Gagarin, 1993*

Inquirenda et incertae sedis. Taymyr and Severnaya Zemlya, Russia (Gagarin 1990, Gagarin 1993, Gagarin 2001b).

Rahm, 1924

Inquirenda et incertae sedis. Taymyr and Severnaya Zemlya, Russia (Kuzmin 1972, Kuzmin and Gagarin 1990).

Gagarin, 1997

Nomen nudum. Novaya Zemlia and Vaigach island, Russia (Gagarin 1997a).

Discussion

Despite of more than 130 years of research, our knowledge on the diversity and distribution of Arctic nematodes is still very incomplete and limited to only few relatively small areas of the region. The first studies of Arctic nematodes date back to 1883, when the first nematode was described from Spitzbergen (Aurivillius 1883). The taxonomic position of the species Aurivillius, 1883 remains uncertain and its type material can not be located for re-study. After Aurivillius, the nematode fauna of Svalbard archipelago was studied by von Linstow (1900), Menzel (1920), Boström (1987a), Boström (1987b), Boström (1989), Zell (1993) and to a lesser extent by Brzeski (1962), Siddiqi (1979), Janiec (1997), Klekowski and Opalinski (1986), Klekowski and Opalinski (1990), Klekowski and Opalinski (1992), Opalinski and Klekowski (1992), Holovachov et al. (2003) adding up to the list of 101 species. Most of the sampling on Svalbard was concentrated around the west part of the Spitsbergen island and a large part of the archipelago remains unstudied. From other territories belonging to the Kingdom of Norway, 43 species of nematodes were found on the small island of Jan Mayen (Allgén 1953, Steiner 1916a), while no information is available for the northernmost part of the Finnmark.

Most of the nematological research in the Iceland was focused in southern and western part of the island (De Coninck 1940, De Coninck 1943, De Coninck 1944), which is considered as Subarctic in the current publication. Sampling sites from the northern part of the country were also outside the boundary considered in the present paper (Kreis 1963, Starmühlner 1969). A small part of the Arctic of the Iceland has not been studied to any considerable extent – only six species from the genus are listed from the areas near Húsavík and Tjörnes by Zell (1993). Terrestrial nematode fauna of Greenland, on the other hand, received considerable attention from Ditlevsen (1927), Allgén (1954), Zell (1993), Brzeski (1999), who altogether reported 43 species of nematodes collected in several distant localities along the eastern (Little Pendulum Island and Angmagssalik) and western (Disko island, Kangerlussuaq, Kapisillit, Sukkertoppen and Upernavik) sides of the island.

Nematode fauna of the Arctic regions of Canada is studied comparatively better. The research was focused in several localities in the Nunavut province including Bathhurst, Ellef Ringnes, Ellesmere and Somerset islands (Anderson 1966, Anderson 1969, Anderson and Ebsary 1982, Anderson and Mulvey 1980, Das 1964, Ebsary 1984, Ebsary 1985, Elshishka et al. 2013, Mulvey 1963, Mulvey 1967b, Mulvey 1967a, Mulvey 1969c, Mulvey 1969a, Mulvey 1969b, Mulvey 1978, Sanwal 1965, Sanwal 1968, Wu 1965, Wu 1969b, Wu 1969a, Wu 1970, Zell 1993), and in the small area of the Mackenzie River delta and Tuktoyaktok in the Northwest territories (Ebsary 1982, Eveleigh 1982, Mulvey 1978, Mulvey and Anderson 1979). Respectively, 55 and 13 species were described from these regions between 1963 and 1993. Much less is known about the nematodes from the Arctic regions of Alaska – only 22 species were recorded there by Andrássy (2003a), Andrássy (2003b), Andrássy (2003c), Bernard (1992), Powers et al. (1983). Studied localities include Prudhoie Bay and neighbouring North Slope and Brooks Range, as well as localities along Kobuk River near Noorvik and Unalakleet River near Unalakleet.

Fauna of the Russian part of the Arctic lists the largest number of nematode species, including many typical freshwater nematodes, but known from only three distinct geographic regions: Novaya Zemlya archipelago together with Vaygach island, Taymyr peninsula together with Severnaya Zemlya, and the Lena River estuary. So far, only 37 species of freshwater nematodes are know from the Lena River estuary (Gagarin 1995, Gagarin 1999b, Gagarin 2000, Gagarin 2001b). The list of nematodes from the Novaya Zemlya archipelago and Vaygach island includes 81 nematode species, both terrestrial (Steiner 1916b) and freshwater (Gagarin 1997c, Gagarin 1997b, Gagarin 1997a, Gagarin 1999a, Gagarin 2000, Gagarin 2001c, Gagarin 2001b). Arctic areas of the Taymyr peninsula and Severnaya Zemlya are the most intensively studied part of the Arctic with localities including the Taymyr Lake, Enisei River, Putorana Plateau, Cape Chelyuskin – 229 species of freshwater and terrestrial nematodes were reported from different sampling sites of this region over the past 40 years (Elshishka et al. 2012, Elshishka et al. 2013, Gagarin 1990, Gagarin 1991a, Gagarin 1991b, Gagarin 1993, Gagarin 1996, Gagarin 1999a, Gagarin 2001b, Gagarin 2001a, Gagarin 2008, Gagarin 2009, Gagarin and Kuzmin 1972, Gagarin and Kuzmin 1983, Kuzmin 1972, Kuzmin 1976, Kuzmin 1986, Kuzmin and Gagarin 1990).

In summary, 391 species, belonging to 146 genera, 54 families, and 10 orders of the phylum are known from the entire Arctic (including species of dubious taxonomic status, but exclusing single nomen nudum). Even taking into consideration relatively low variety of habitats in this region, comparing to other areas of the world, as well as the rather harsh environmental conditions, this list is considered to be very incomplete and preliminary.