Marine Bryozoa of Greece: an annotated checklist.

BACKGROUND: Until today, a complete checklist of Bryozoa of the Greek seas had never been published and species records were scattered in several taxonomic and ecological studies. The aim of this paper is to produce a first checklist of marine bryozoan species of Greece, in the framework of the Greek Taxon Information System (GTIS) initiative of the LifeWatchGreece Research Infrastructure (ESFRI), by reviewing the existing literature and following the recent trends in the taxonomy of this group. . NEW INFORMATION: The marine bryozoan fauna of Greece comprises 237 species, classified in 127 genera, 66 families, 3 orders, and 2 classes. The vast majority belongs to the class Gymnolaemata (177 Cheilostomatida and 21 Ctenostomatida), while the remaining 39 species are Stenolaemata (all Cyclostomatida). Among these species, 12 are considered endemic to the eastern Mediterranean, while another 12 species are non-indigenous.

Introduction

The Mediterranean bryozoan fauna represents about 9.6% of the known global diversity of (Rosso and Di Martino 2016). However, current knowledge on the bryozoan fauna of this basin is highly biased due to differences in research effort for different taxa, marine areas and habitat types. Recent efforts in the south-eastern Mediterranean Sea have significantly increased our knowledge from areas which were previously considered “data-poor” spots for bryozoans, revealing a considerable proportion of non-indigenous taxa (e.g. Harmelin et al. 2016, Sokolover et al. 2016). Furthermore, information on from several Mediterranean areas is fragmented in old publications and grey literature; the collation of updated checklists (e.g. Koçak and Aydın Önen 2014) by experts could contribute to the mapping of diversity for understudied taxa and areas (Bailly et al. 2016, this special collection).

Sporadic records of bryozoans from the Aegean Sea can be found in old taxonomic studies, including the descriptions of the cheilostome species , and from this marine region (Busk 1854), based on material collected from shallow water macroalgae by E. Forbes. The first studies focusing on of the eastern Mediterranean basin, the Greek seas included, took place in the late 1960s (Harmelin 1969), based on material that was collected with dredges during the oceanographic expedition of the French research vessel “Jean Charcot” (1968). A total of 101 species belonging to all the 3 living bryozoan orders, were found, including several new records for the Mediterranean fauna. Later, the taxonomic study by Hayward (1974) on the cheilostome of Chios Island (north-eastern Aegean) yielded 101 species, including rare records and new species. Sharing only 47 cheilostomes with Harmelin’s list mentioned above, this study increased the eastern Mediterranean bryozoan diversity by 54 species.

More recently, few taxonomic and ecological studies have focused on the bryozoan fauna of Greece (e.g. Castritsi-Catharios and Marcopoulou-Diacantoni 1983, Castritsi-Catharios and Kiortis 1984, Castritsi-Catharios and Kiortis 1985, Castritsi-Catharios et al. 1985a, Castritsi-Catharios et al. 1985b, Castritsi-Catharios et al. 1986a, Castritsi-Catharios et al. 1986b, Castritsi-Catharios and Ganias 1989, Ganias 1990), while a considerable number of records has been included in general ecological studies examining a variety of habitats (e.g. Simboura et al. 1995, Conides et al. 1999, Morri et al. 1999, Antoniadou and Chintiroglou 2005, Gerovasileiou et al. 2013, Gerovasileiou et al. 2015). A complete checklist of however had never been published from Greece. The aim of the present study is to give a first annotated checklist of from the Greek seas.

Materials and methods

The checklist presented in this paper was compiled in the framework of the Greek Taxon Information System (GTIS), an initiative of the LifeWatchGreece Research e-Infrastructure aiming to produce a complete inventory of the known biodiversity of Greece (Bailly et al. 2016, this special collection). As this is the first attempt to compile a checklist of bryozoan diversity of the Greek seas, we performed an exhaustive literature review of bryozoan records. All taxa were cross-checked for synonymies and dubious records against the World List of Bryozoa (recent and fossil), the World Register of Marine Species (Bock and Gordon 2016, WoRMS Editorial Board 2016), and the recent update of the Mediterranean bryozoan diversity by Rosso and Di Martino (2016). Taxa identified as cf., as well as those left in open nomenclature, and fossil records were not included in the list. The checklist is annotated with the literature references reporting the presence of each species from Greek waters, as well as with notes on the nomenclature of some taxa. Non-indigenous bryozoans are also marked (NIB). The checklist follows the alphabetical order.

Checklists

Checklist of known to occur in the Greek seas

Allman, 1856

Busk, 1852

Busk, 1884

Busk, 1884

(Canu & Bassler, 1930)

Notes

Gautier 1962

(Heller, 1867)

Castritsi-Catharios et al. 1985a, Castritsi-Catharios et al. 1985b, Castritsi-Catharios et al. 1986a, Ganias 1990, Rosso and Novosel 2010, Chimenz Gusso et al. 2014, Gerovasileiou et al. 2015​

Busk, 1884

(Johnston, 1847)

Harmelin 1969, Hayward 1974, Castritsi-Catharios and Marcopoulou-Diacantoni 1983, Castritsi-Catharios and Kiortis 1984, Castritsi-Catharios et al. 1986a (as , probably a mispelling), Ganias 1990, Morri et al. 1999, Antoniadou and Chintiroglou 2005

Smitt, 1868

Lamouroux, 1812

(Linnaeus, 1758)

Harmelin 1969, Hayward 1974, Castritsi-Catharios and Kiortis 1984, Castritsi-Catharios and Kiortis 1985, Ganias 1990

Waters, 1906

Ganias 1990, Morri et al. 1999

(Couch, 1844)

Harmelin 1969, Hayward 1974, Castritsi-Catharios and Marcopoulou-Diacantoni 1983, Castritsi-Catharios and Kiortis 1984, Castritsi-Catharios and Kiortis 1985, Castritsi-Catharios et al. 1985a, Castritsi-Catharios et al. 1986a, Castritsi-Catharios et al. 1986b, Castritsi-Catharios and Ganias 1989, Ganias 1990, Morri et al. 1999

(Landsborough, 1852)

Harmelin 1969, Hayward 1974, Castritsi-Catharios and Kiortis 1984, Castritsi-Catharios and Kiortis 1985, Castritsi-Catharios et al. 1985a, Castritsi-Catharios et al. 1986a, Castritsi-Catharios and Ganias 1989, Ganias 1990, Morri et al. 1999

Vigneaux, 1949

Jullien, 1888

(Audouin, 1826)

Harmelin 1969, Ganias 1990

Canu & Bassler, 1927

Johnston, 1840

(Hincks, 1886)

​Harmelin 1969, Ganias 1990, Morri et al. 1999

(Heller, 1867)

Harmelin 1969, Hayward 1974, Castritsi-Catharios and Marcopoulou-Diacantoni 1983, Castritsi-Catharios and Kiortis 1984, Castritsi-Catharios et al. 1986a, Castritsi-Catharios and Ganias 1989, Ganias 1990, Simboura et al. 1995, Morri et al. 1999

(Busk, 1852)

Harmelin 1969, Hayward 1974, Castritsi-Catharios and Kiortis 1984, Castritsi-Catharios and Ganias 1989, Ganias 1990

Johnston, 1840

​Hayward 1974, Castritsi-Catharios and Ganias 1989, Ganias 1990

MacGillivray, 1895

Canu & Bassler, 1917

(Calvet, 1903)

Harmelin 1969, Hayward 1974, Ganias 1990, Morri et al. 1999

Fischer, 1807

(Pallas, 1766)

Ganias 1990, Antoniadou and Chintiroglou 2005, Chimenz Gusso et al. 2014

Canu & Bassler, 1917

(Manzoni, 1869)

In the absence of type material, the validity of has been questioned by Reverter-Gil et al. (2012). Two new species, Reverter-Gil, Souto & Fernandez-Pulpeiro, 2012 and Reverter-Gil, Souto, Novosel & Tilbrook, 2016 have been erected to allocate some records from the western Basin and the Adriatic, respectively. Nevertheless, records and material from the Aegean were not considered and hence they are provisionally still reported as (see also Rosso and Di Martino 2016).

(Busk, 1859)

Harmelin 1969, Hayward 1974, Morri et al. 1999; the status of these records needs to be checked (see Rosso and Di Martino 2016​).

(Calvet, 1927)

Synonym valid for

Hayward 1974

(Heller, 1867)

Harmelin 1969, Hayward 1974, Castritsi-Catharios and Ganias 1989, Ganias 1990, Simboura et al. 1995, Morri et al. 1999; Mediterranean colonies recorded as , and possibly belong to this species.

(Hincks, 1862)

Morri et al. 1999​

(Hassall, 1841)

Gautier 1962, Harmelin 1969, Hayward 1974, Castritsi-Catharios et al. 1985a, Ganias 1990

(Hincks, 1880)

Gautier 1962, Harmelin 1969, Hayward 1974, Morri et al. 1999

Vigneaux, 1949

Cossmann, 1906

(Busk, 1861)

Harmelin 1969​

Bishop & Hayward, 1989

Madurell, Zabala, Domínguez-Carrió & Gili, 2013

Harmelin 1969, Castritsi-Catharios et al. 1985a (as ), Castritsi-Catharios et al. 1986a. These records have been provisionally reported as following Madurell et al. (2013) and Rosso and Di Martino (2016).

(Ellis & Solander, 1786)

Ganias 1990​

Gray, 1848

(Busk, 1854)

Harmelin 1969​

(Norman, 1868)

Harmelin 1969​

Gray, 1848

Levinsen, 1909

(Linnaeus, 1758)

Antoniadou and Chintiroglou 2005

Oken, 1815

(Norman, 1868)

Hayward 1974

(Linnaeus, 1758)

Distribution

NIB

Castritsi-Catharios and Kiortis 1984, Castritsi-Catharios and Kiortis 1985, Castritsi-Catharios and Ganias 1989

Gray, 1848

(Linnaeus, 1758)

Ganias 1990

(Ryland, 1960)

NIB

Morri et al. 1999

(Hincks, 1886)

Castritsi-Catharios and Kiortis 1984

(Hincks, 1886)

Castritsi-Catharios and Marcopoulou-Diacantoni 1983

(Ryland, 1960)

Castritsi-Catharios and Kiortis 1984, Castritsi-Catharios and Kiortis 1985, Ganias 1990, Antoniadou and Chintiroglou 2005

(Alder, 1857)

​Castritsi-Catharios and Kiortis 1984, Castritsi-Catharios and Kiortis 1985

Gray, 1848

(Hincks, 1886)

Castritsi-Catharios and Marcopoulou-Diacantoni 1983

(Pallas, 1766)

NIB

Hayward 1974

(Lamarck, 1816)

NIB

Harmelin 1969, Ganias 1990

Norman, 1903

Norman, 1903

(Hincks, 1860)

Hayward 1974

Canu & Bassler, 1927

(Waters, 1898)

Hayward 1974

Gray, 1848

(Audouin, 1826)

Hayward 1974, Ganias 1990

(Linnaeus, 1767)

Harmelin 1969, Ganias 1990

Harmer, 1926

(Hincks, 1880)

Hayward 1974, Castritsi-Catharios et al. 1986b, Castritsi-Catharios and Ganias 1989, Ganias 1990

(Hincks, 1880)

Hayward 1974, Castritsi-Catharios et al. 1986a, Castritsi-Catharios and Ganias 1989, Ganias 1990

Gordon, 1984

(Waters, 1898)

Hayward 1974, Ganias 1990, Morri et al. 1999

Canu, 1900

(Hincks, 1880)

Gautier 1962

(Hincks, 1860)

Ganias 1990

Norman, 1903

Fernandez Pulpeiro & Reverter Gil, 1993

​Ganias 1990

Osburn, 1940

(Hincks, 1862)

Harmelin 1969, Hayward 1974, Ganias 1990

Brown, 1948

Chimenz, Nicoletti & Lippi Boncambi, 1997

Morri et al. 1999

d'Orbigny, 1851

Lamouroux, 1816

(Audouin, 1826)

Harmelin 1969, Hayward 1974, Ganias 1990, Morri et al. 1999, Simboura et al. 1995

Vieira, Spencer Jones & Winston, 2013

(Audouin, 1826)

Hayward 1974, Ganias 1990

(Gautier, 1962)

Harmelin 1969, Hayward 1974

Vieira, Spencer Jones, Winston, Migotto & Marques, 2014

(Busk, 1860)

Castritsi-Catharios et al. 1986a, Castritsi-Catharios et al. 1986b, Castritsi-Catharios and Ganias 1989, Ganias 1990

Van Beneden, 1845

Harmelin, 1969

Harmelin 1969, Ganias 1990

(Audouin, 1826)

Harmelin 1969, Ganias 1990, Morri et al. 1999

Waters, 1897

Harmelin 1969, Ganias 1990

(Lamouroux, 1816)

Castritsi-Catharios and Kiortis 1984, Castritsi-Catharios et al. 1986a

Busk, 1852

Gautier 1962, Harmelin 1969, Hayward 1974, Castritsi-Catharios and Kiortis 1984, Castritsi-Catharios and Kiortis 1985, Castritsi-Catharios et al. 1986a, Ganias 1990, Simboura et al. 1995, Morri et al. 1999

(Linnaeus, 1758)

NIB

Hayward 1974, Ganias 1990, Simboura et al. 1995, Morri et al. 1999

Fleming, 1828

Ellis & Solander, 1786

(Linnaeus, 1758)

Ganias 1990

(Hastings, 1947)

Harmelin 1969

Lamouroux, 1816

Hayward 1974, Castritsi-Catharios and Kiortis 1984, Castritsi-Catharios and Kiortis 1985, Ganias 1990

Johnston, 1838

Heller, 1867

(Hincks, 1862)

Ganias 1990

Heller, 1867

Harmelin 1969

Linnaeus, 1767

(Hayward, 1975)

Ganias 1990

(Pallas, 1766)

Morri et al. 1999

Gray, 1848

(Waters, 1879)

Gautier 1962, Harmelin 1969, Hayward 1974, Castritsi-Catharios and Kiortis 1984, Castritsi-Catharios et al. 1985a, Castritsi-Catharios et al. 1986a, Ganias 1990, Morri et al. 1999

(Johnston, 1847)

​Castritsi-Catharios and Kiortis 1984 (as ), Castritsi-Catharios et al. 1985a, Castritsi-Catharios et al. 1985b, Castritsi-Catharios et al. 1986a, Ganias 1990, Simboura et al. 1995, Morri et al. 1999

(Hincks, 1880)

Morri et al. 1999

(Hincks, 1880)

Harmelin 1969, Morri et al. 1999

Alder, 1864

Alder, 1864

Harmelin 1969, Ganias 1990

Ryland, 1963

(Hincks, 1860)

Ganias 1990

Hayward, 1978

Hayward 1978, Ganias 1990, Morri et al. 1999

(Wood, 1844)

Hayward 1974, Ganias 1990, Gerovasileiou et al. 2015

Hayward, 1978

Ganias 1990

(Barroso, 1919)

Ganias 1990

Busk, 1884

Lamouroux, 1824

(Bertoloni, 1810)

​Harmelin 1969, Hayward 1974, Castritsi-Catharios and Kiortis 1984, Castritsi-Catharios et al. 1986a, Castritsi-Catharios et al. 1986b, Ganias 1990, Morri et al. 1999

Vigneaux, 1949

Hincks, 1879

(Audouin, 1826)

​Gautier 1962, Harmelin 1969, Hayward 1974, Castritsi-Catharios and Marcopoulou-Diacantoni 1983, Castritsi-Catharios and Kiortis 1984, Castritsi-Catharios et al. 1985a, Castritsi-Catharios et al. 1986a, Castritsi-Catharios and Ganias 1989, Ganias 1990, Simboura et al. 1995, Morri et al. 1999, Antoniadou and Chintiroglou 2005

Winston, 2005

Osburn, 1940

Ayari & Taylor, 2008

Harmelin 1969 (as )

Levinsen, 1909

Levinsen, 1909

(Audouin, 1826)

NIB

Ganias 1990​

Hincks, 1879

Jullien, 1886

(Audouin, 1826)

​Hayward 1974, Castritsi-Catharios and Marcopoulou-Diacantoni 1983, Castritsi-Catharios and Kiortis 1984, Castritsi-Catharios and Kiortis 1985, Castritsi-Catharios et al. 1986a, Castritsi-Catharios and Ganias 1989, Ganias 1990

Jullien, 1886

(Johnston, 1847)

​Harmelin 1969, Hayward 1974, Ganias 1990

Jullien, 1886

(Couch, 1844)

Harmelin 1969, Hayward 1974, Castritsi-Catharios et al. 1986a, Castritsi-Catharios et al. 1986b, Castritsi-Catharios and Ganias 1989, Ganias 1990

(Friedl, 1917)

Gautier 1962, Morri et al. 1999

(Moll, 1803)

​Harmelin 1969, Hayward 1974, Castritsi-Catharios and Marcopoulou-Diacantoni 1983, Castritsi-Catharios and Kiortis 1984, Castritsi-Catharios and Kiortis 1985, Castritsi-Catharios et al. 1986a, Castritsi-Catharios and Ganias 1989, Ganias 1990, Simboura et al. 1995, Morri et al. 1999

(Canu & Bassler, 1925)

Ganias 1990

Harmelin & Aristegui, 1987

​Morri et al. 1999

(Gautier, 1956)

Harmelin 1969, Hayward 1974

(Landsborough, 1852)

​Gautier 1962, Hayward 1974, Castritsi-Catharios and Marcopoulou-Diacantoni 1983, Castritsi-Catharios and Kiortis 1984, Castritsi-Catharios et al. 1986a, Castritsi-Catharios et al. 1986b, Castritsi-Catharios and Ganias 1989, Ganias 1990, Morri et al. 1999

(Waters, 1899)

Harmelin 1969, Ganias 1990, Morri et al. 1999

Vigneaux, 1949

Canu & Bassler, 1925

(Moll, 1803)

Hayward 1974, Castritsi-Catharios and Marcopoulou-Diacantoni 1983, Castritsi-Catharios et al. 1985a, Castritsi-Catharios et al. 1986a, Castritsi-Catharios et al. 1986b, Ganias 1990, Antoniadou and Chintiroglou 2005

d'Orbigny, 1851

Gray, 1848

(Canu, 1928)

Ganias 1990

Lamouroux, 1816

(Linnaeus, 1767)

Castritsi-Catharios et al. 1986a, Castritsi-Catharios and Ganias 1989, Ganias 1990, Antoniadou and Chintiroglou 2005

Gautier, 1954

Hayward 1974, Castritsi-Catharios and Marcopoulou-Diacantoni 1983, Castritsi-Catharios and Kiortis 1984, Castritsi-Catharios and Kiortis 1985, Castritsi-Catharios et al. 1985a, Castritsi-Catharios et al. 1986a, Castritsi-Catharios and Ganias 1989, Ganias 1990

Gregory, 1893

Pieper, 1881

(Audouin, 1826)

​Harmelin 1969, Hayward 1974, Ganias 1990

Tilbrook, 2006

Milne Edwards, 1836

Zabala, Maluquer & Harmelin, 1993

​Harmelin 1969, Hayward 1974, Morri et al. 1999

(Quelch, 1884)

Ganias 1990

(Moll, 1803)

Harmelin 1969, Hayward 1974, Castritsi-Catharios et al. 1986a (as ), Castritsi-Catharios et al. 1986b (as ), Ganias 1990, ​Morri et al. 1999

Gray, 1848

(Johnston, 1847)

Harmelin 1969, Morri et al. 1999

Canu & Bassler, 1917

(Waters, 1899)

Harmelin 1969, Hayward 1974, Ganias 1990

David & Pouyet, 1978

Berning, Tilbrook & Rosso, 2008

​Hayward 1974 (as ), Ganias 1990 (as ), Berning et al. 2008

Harmer, 1957

Smitt, 1868

(Busk, 1860)

Harmelin 1969, Hayward 1974, Ganias 1990, Morri et al. 1999

Duvergier, 1924

(Osburn, 1927)

NIB

Hayward 1974

Bassler, 1935

Milne Edwards, 1836

(Abildgaard, 1806)

Harmelin 1969, Hayward 1974, Castritsi-Catharios and Marcopoulou-Diacantoni 1983, Castritsi-Catharios and Kiortis 1984 (as ), Castritsi-Catharios et al. 1986a, Castritsi-Catharios and Ganias 1989, Ganias 1990, Morri et al. 1999​, Antoniadou and Chintiroglou 2005

Fleming, 1828

Gray, 1848

(Pallas, 1766)

Hayward 1974

Barroso, 1949

(Heller, 1867)

​Harmelin 1969, Hayward 1974, Castritsi-Catharios and Kiortis 1984

Norman, 1903

(Hincks, 1877)

​Hayward 1974, Ganias 1990, Morri et al. 1999

Gordon in De Blauwe, 2009

Levinsen, 1909

(Audouin, 1826)

​Harmelin 1969, Hayward 1974, Ganias 1990, Morri et al. 1999

Winston, 2005

Canu, 1919

(Busk, 1854)

Busk 1854, Harmelin 1969, Hayward 1974, Castritsi-Catharios and Marcopoulou-Diacantoni 1983, Castritsi-Catharios et al. 1985a, Castritsi-Catharios et al. 1985b, Castritsi-Catharios et al. 1986a, Castritsi-Catharios and Ganias 1989, Ganias 1990, Morri et al. 1999, Gerovasileiou et al. 2013, Gerovasileiou et al. 2015

Levinsen, 1909

Levinsen, 1909

(Hayward, 1974)

​Harmelin 1969 (as sp.), Hayward 1974, Ganias 1990

(Busk, 1884)

NIB

Morri et al. 1999

Vigneaux, 1949

Bishop & Hayward, 1989

(Busk, 1856)

​Hayward 1974, Ganias 1990, Morri et al. 1999

Canu & Bassler, 1927

Gautier, 1962

Gautier 1962, Morri et al. 1999

Busk, 1859

Lamouroux, 1821

Manzoni, 1870

Ganias 1990

Jullien, 1888

Levinsen, 1909

(Audouin, 1826)

Hayward 1974, Ganias 1990

Harmer, 1957

Harmer, 1957

Hayward, 1974

Hayward 1974, Ganias 1990

Harmer, 1957

Gray, 1843

(Ellis & Solander, 1786)

Harmelin 1969, Hayward 1974, Castritsi-Catharios et al. 1985a, Ganias 1990

Hincks, 1879

Hincks, 1879

(Peach, 1868)

Harmelin 1969, Ganias 1990

Jullien, 1888

(Audouin, 1826)

Harmelin 1969, Hayward 1974, Castritsi-Catharios et al. 1986b, Castritsi-Catharios and Ganias 1989, Ganias 1990, Morri et al. 1999

Hincks, 1877

(Audouin, 1826)

NIB

Harmelin et al. (2011) suggested that specimens from the Aegean Sea reported as could belong to (followed by Rosso and Di Martino 2016). Recorded by Hayward 1974, Morri et al. 1999.

(Pallas, 1766)

This taxon corresponds to a complex of species (Rosso et al. 2010, Rosso and Di Martino 2016). Recorded by Harmelin 1969, Castritsi-Catharios and Marcopoulou-Diacantoni 1983, Castritsi-Catharios et al. 1986a, Ganias 1990, Antoniadou and Chintiroglou 2005.

(Busk, 1860)

Hayward 1974

Gray, 1848

Jullien, 1888

(Esper, 1796)

Busk 1854, Hayward 1974, Castritsi-Catharios and Marcopoulou-Diacantoni 1983, Castritsi-Catharios and Kiortis 1984, Castritsi-Catharios and Kiortis 1985, Castritsi-Catharios and Ganias 1989, Ganias 1990, Morri et al. 1999

Prenant & Bobin, 1966

(Gautier, 1962)

Harmelin 1969, Hayward 1974

Gray, 1848

(Johnston, 1847)

Hayward 1974, Ganias 1990

Lamouroux, 1821

(Waters, 1879)

Hayward 1974, Morri et al. 1999

(Moll, 1803)

Gautier 1962, Harmelin 1969, Hayward 1974, Ganias 1990, Morri et al. 1999

Hincks, 1882

Hincks, 1881

(Audouin, 1826)

Mediterranean specimens of and actually belong to this species, as suggested by Harmelin (2014a) and followed by Rosso and Di Martino (2016). Recorded by Harmelin 1969, Hayward 1974, Ganias 1990, Morri et al. 1999.

Gray, 1841

de Blainville, 1830

(Pallas, 1766)

​Harmelin 1969, Hayward 1974, Morri et al. 1999, Gerovasileiou et al. 2015

Jullien, 1882

Jullien, 1882

(Reuss, 1847)

Hayward 1974, Ganias 1990

(Jullien, 1881)

​Harmelin 1969, Morri et al. 1999, Gerovasileiou et al. 2015

Neviani, 1895

​Hayward 1974, Castritsi-Catharios and Marcopoulou-Diacantoni 1983, Ganias 1990

Jullien, 1882

(Canu & Bassler, 1930)

is a synonym of

Harmelin 1969, Hayward 1974, Ganias 1990

Gabb & Horn, 1862

Barroso, 1926

(Waters, 1879)

Gautier 1962, Harmelin 1969

Soule, Soule & Chaney, 1991

Chimenz-Gusso & Soule, 2003

Harmelin 1969 (as sp.), Hayward 1974 (as )

Busk, 1884

(Waters, 1894)

Ganias 1990

(Hincks, 1878)

Harmelin 1969, Ganias 1990

(Jullien & Calvet, 1903)

Synonym valid for

​Hayward 1974, Ganias 1990, Morri et al. 1999, Antoniadou and Chintiroglou 2005, Chimenz Gusso et al. 2014

(Hass, 1948)

Hayward 1974, Ganias 1990

Harmer, 1933

Hayward, 1974

Hayward 1974, Harmelin 1976, Ganias 1990

Hincks, 1895

(Johnston, 1847)

Hayward 1974, Ganias 1990

Gautier, 1962

​Harmelin 1969, Hayward 1974, Ganias 1990, Morri et al. 1999, Gerovasileiou et al. 2015

Zabala & Maluquer, 1988

Morri et al. 1999

sensu Hayward, 1974

Hayward 1974

Gregory, 1893

(Busk, 1884)

Chimenz Gusso et al. 2014

(Hincks, 1886)

Gautier 1962, Harmelin 1969, Ganias 1990

(Risso, 1826)

Harmelin 1969, Ganias 1990

Hincks, 1877

(Busk, 1856)

Harmelin 1969, Hayward 1974, Ganias 1990

Duvergier, 1921

(Hincks, 1861)

Harmelin 1969, Hayward 1974, Castritsi-Catharios and Marcopoulou-Diacantoni 1983, Castritsi-Catharios et al. 1985a, Castritsi-Catharios et al. 1985b, Ganias 1990

Vigneaux, 1949

Jullien in Jullien & Calvet, 1903

(Heller, 1867)

Harmelin 1969, Hayward 1974, Ganias 1990

Jullien, 1888

Gray, 1848

Geraci, 1974

Ganias 1990

Norman, 1894

(Busk, 1861)

Ganias 1990, Morri et al. 1999

Canu & Bassler, 1925

(Manzoni, 1870)

Harmelin 1969

Levinsen, 1909

Levinsen, 1909

(Audouin, 1826)

Gautier 1962, Hayward 1974, Morri et al. 1999

Jullien, 1883

Canu & Bassler, 1920

(Norman, 1868)

Gautier 1962, Harmelin 1969, Ganias 1990, Morri et al. 1999

Hincks, 1877

(Reuss, 1848)

Harmelin 1969, Hayward 1974, Castritsi-Catharios and Marcopoulou-Diacantoni 1983, Castritsi-Catharios et al. 1986a, Morri et al. 1999, Ganias 1990

(Waters, 1878)

Hayward 1974, Castritsi-Catharios et al. 1985a, Ganias 1990

Hincks, 1886

Harmelin 1969, Hayward 1974, Ganias 1990

(Johnston in Wood, 1844)

Castritsi-Catharios et al. 1986a, Ganias 1990, Morri et al. 1999, Antoniadou and Chintiroglou 2005

Levinsen, 1909

Hincks, 1877

(Busk, 1860)

Hayward 1974, Ganias 1990

Levinsen, 1909

Osburn, 1952

(Audouin, 1826)

NIB

Ganias 1990

(Calvet, 1902)

Harmelin 1969, Hayward 1974, Ganias 1990

Gautier, 1962

(Manzoni, 1869)

Hayward 1974, Ganias 1990

(Calvet, 1906)

Harmelin 1969

Norman, 1903

(Pallas, 1766)

​Gautier 1962 (as ), Ganias 1990 (as ), Morri et al. 1999, Chimenz Gusso et al. 2014

(Jullien & Calvet, 1903)

Harmelin 1969, Ganias 1990

(Johnston, 1847)

Ganias 1990, Antoniadou and Chintiroglou 2005

(Canu & Bassler, 1928)

Hayward 1974

Osburn, 1952

(Hincks, 1877)

Hayward 1974, Ganias 1990

(Waters, 1879)

Harmelin 1969, Hayward 1974

(MacGillivray, 1842)

Gautier 1962, Morri et al. 1999, Antoniadou and Chintiroglou 2005

Jullien, 1903

Norman, 1869

(Busk, 1860)

Ganias 1990

Gordon, Tilbrook & Winston, 2005

Levinsen, 1909

(Norman, 1864)

Ganias 1990

Canu, 1904

Hincks, 1880

Hastings, 1944

Hayward 1974, Castritsi-Catharios and Kiortis 1984, Castritsi-Catharios et al. 1986a, Ganias 1990, Morri et al. 1999

Vigneaux, 1949

Neviani, 1896

(Norman, 1864)

Ganias 1990

(Busk, 1854)

Busk 1854, Hayward 1974, Castritsi-Catharios et al. 1986b, Castritsi-Catharios and Ganias 1989, Ganias 1990. Records of (d'Orbigny, 1852) probably belong to .

Busk, 1852

Johnston, 1838

Lamouroux, 1813

(Calvet, 1900)

Castritsi-Catharios and Kiortis 1984, Castritsi-Catharios and Kiortis 1985

Prouho, 1892

Ganias 1990

(Hassall, 1841)

Ganias 1990

Hincks, 1880

Alder, 1857

Alder, 1857

​Castritsi-Catharios and Kiortis 1984, Castritsi-Catharios and Kiortis 1985, Ganias 1990

d'Hondt, 1983

Ehrenberg, 1838

(Farre, 1837)

Castritsi-Catharios et al. 1986a, Castritsi-Catharios et al. 1986b

Hincks, 1877

Jebram, 1973

(Heller, 1867)

​Castritsi-Catharios and Marcopoulou-Diacantoni 1983, Castritsi-Catharios and Kiortis 1984, Castritsi-Catharios and Kiortis 1985, Castritsi-Catharios et al. 1985a, Castritsi-Catharios et al. 1986a, Castritsi-Catharios et al. 1986b, Castritsi-Catharios and Ganias 1989, Ganias 1990, Morri et al. 1999

Hincks, 1851

Hincks, 1851

Harmelin 1969, Castritsi-Catharios and Kiortis 1984, Castritsi-Catharios and Kiortis 1985, Castritsi-Catharios et al. 1985a, Castritsi-Catharios et al. 1986a, Castritsi-Catharios and Ganias 1989, Ganias 1990, Morri et al. 1999

Harmer, 1915

Gosse, 1851

(Hincks, 1860)

Harmelin 1969, Ganias 1990

Gosse, 1855

Castritsi-Catharios and Kiortis 1984, Castritsi-Catharios et al. 1986a, Castritsi-Catharios and Ganias 1989, Ganias 1990, Morri et al. 1999

Osburn & Soule, 1953

Soule, 1951

(Ellis & Solander, 1786)

Castritsi-Catharios and Kiortis 1984, Castritsi-Catharios and Kiortis 1985, Castritsi-Catharios et al. 1986a, Ganias 1990, Morri et al. 1999

Hincks, 1880

Lamouroux, 1812

(Leidy, 1855)

Ganias 1990, Morri et al. 1999

(Hincks, 1877)

NIB

Castritsi-Catharios and Kiortis 1984, Castritsi-Catharios and Kiortis 1985, Ganias 1990

(Adams, 1798)

Castritsi-Catharios and Kiortis 1984, Castritsi-Catharios and Kiortis 1985, Castritsi-Catharios et al. 1986a, Castritsi-Catharios et al. 1986b, Castritsi-Catharios and Ganias 1989, Ganias 1990

(Linnaeus, 1758)

Castritsi-Catharios and Ganias 1989, Ganias 1990, Morri et al. 1999

Calvet, 1911

Castritsi-Catharios and Marcopoulou-Diacantoni 1983, Castritsi-Catharios and Kiortis 1984, Castritsi-Catharios and Kiortis 1985, Castritsi-Catharios et al. 1986a, Castritsi-Catharios and Ganias 1989, Ganias 1990, Morri et al. 1999

(Ellis & Solander, 1786)

Castritsi-Catharios et al. 1986a, Castritsi-Catharios et al. 1986b

Lamouroux, 1824

Castritsi-Catharios and Ganias 1989, Ganias 1990

(Delle Chiaje, 1822)

NIB

Castritsi-Catharios and Kiortis 1984, Castritsi-Catharios and Kiortis 1985, Castritsi-Catharios et al. 1986a, Castritsi-Catharios et al. 1986b, Castritsi-Catharios and Ganias 1989, Ganias 1990, Conides et al. 1999

Hincks, 1880

Saville-Kent, 1870

Saville-Kent, 1870

Castritsi-Catharios et al. 1986a, Castritsi-Catharios et al. 1986b

Hincks, 1880

Fleming, 1823

Heller, 1867

Castritsi-Catharios and Kiortis 1984, Castritsi-Catharios et al. 1985a, Castritsi-Catharios et al. 1986a, Castritsi-Catharios and Ganias 1989, Ganias 1990, Morri et al. 1999

(Linnaeus, 1758)

Castritsi-Catharios and Kiortis 1984, Castritsi-Catharios et al. 1986a, Castritsi-Catharios and Ganias 1989, Ganias 1990

Borg, 1926

Busk, 1852

Hayward & Ryland, 1985

Hayward & Ryland, 1985

(Harmelin, 1976)

Ganias 1990

(Johnston, 1847)

​Castritsi-Catharios et al. 1985a (as ), Castritsi-Catharios et al. 1986a, Castritsi-Catharios et al. 1986b, Castritsi-Catharios and Ganias 1989 (as ), Ganias 1990, Simboura et al. 1995, Morri et al. 1999

(Busk, 1875)

Ganias 1990

Harmelin, 1976

(Couch, 1844)

Castritsi-Catharios et al. 1986a, Castritsi-Catharios et al. 1986b, Castritsi-Catharios and Ganias 1989, Ganias 1990

Harmelin, 1976

Ganias 1990

Harmelin, 1976

Ganias 1990

Johnston, 1838

Lamouroux, 1812

(Lamarck, 1816)

Ganias 1990

(Linnaeus, 1758)

Castritsi-Catharios and Kiortis 1984, Castritsi-Catharios et al. 1986a, Castritsi-Catharios et al. 1986b, Castritsi-Catharios and Ganias 1989, Ganias 1990

Milne Edwards, 1838

Castritsi-Catharios et al. 1986a, Castritsi-Catharios et al. 1986b

(Heller, 1867)

Castritsi-Catharios and Marcopoulou-Diacantoni 1983, Castritsi-Catharios et al. 1986a, Ganias 1990

Harmelin, 1990

Harmelin 1969 (as sp. II), Ganias 1990​

Harmer, 1891

Castritsi-Catharios and Marcopoulou-Diacantoni 1983, Castritsi-Catharios et al. 1986a, Castritsi-Catharios and Ganias 1989, Ganias 1990, Simboura et al. 1995

Waters, 1916

Harmelin 1969, Castritsi-Catharios et al. 1986a, Castritsi-Catharios and Ganias 1989, Ganias 1990, Simboura et al. 1995

Reuss, 1869

Canu, 1918

(Busk, 1875)

Ganias 1990

Busk, 1875

Link, 1807

(Lamouroux, 1821)

Harmelin 1969 (as ), Ganias 1990, Gerovasileiou et al. 2015

Smitt, 1867

Lamouroux, 1821

Lamouroux, 1821

Harmelin 1969, Ganias 1990, Simboura et al. 1995

Smitt, 1867

Gray, 1848

(Fleming, 1828)

Rosso and Di Martino (2016) maintained this taxon but it is possible that these records actually belong to a different species, Alvarez, 1992; all records need to be validated. Recorded by Harmelin 1969, Castritsi-Catharios and Marcopoulou-Diacantoni 1983, Castritsi-Catharios et al. 1986a, Castritsi-Catharios and Ganias 1989 (as ), Ganias 1990, Simboura et al. 1995, Morri et al. 1999.

Gray, 1848

(Audouin, 1826)

​Castritsi-Catharios and Marcopoulou-Diacantoni 1983, Castritsi-Catharios and Kiortis 1984, Castritsi-Catharios et al. 1985a, Castritsi-Catharios et al. 1986a, Castritsi-Catharios and Ganias 1989, Ganias 1990, Simboura et al. 1995, Morri et al. 1999, Antoniadou and Chintiroglou 2005

(Linnaeus, 1758)

Castritsi-Catharios et al. 1986a, Castritsi-Catharios and Ganias 1989, Ganias 1990

Canu, 1918

Canu, 1918

(Harmelin, 1976)

See

Ganias 1990

Canu, 1918

Canu & Bassler, 1922

(O'Donoghue & de Watteville, 1939)

Harmelin 1969 (as ), Harmelin 1976, Castritsi-Catharios et al. 1986b, Castritsi-Catharios and Ganias 1989, Ganias 1990

Canu & Bassler, 1920

Harmelin, 1976

Ganias 1990

Harmelin, 1976

Ganias 1990

Canu, 1918

(Johnston, 1838)

The use of the name

Harmelin 1969, Ganias 1990, Morri et al. 1999

Canu, 1918

(Waters, 1879)

Ganias 1990

(Jullien, 1882)

Ganias 1990

(Lamarck, 1816)

Harmelin 1969, Ganias 1990, Antoniadou and Chintiroglou 2005

Harmelin, 1976

Ganias 1990

(Norman, 1864)

Harmelin 1976, Ganias 1990

Canu & Bassler, 1920

Jullien, 1883

(Meneghini, 1844)

Ganias 1990

Johnston, 1838

Brood, 1976

(Canu and Bassler, 1929)

See

Castritsi-Catharios and Marcopoulou-Diacantoni 1983, Ganias 1990

Canu & Bassler, 1920

Harmelin, 1976

See

Ganias 1990

(Heller, 1867)

See

Harmelin 1969 (as ), Ganias 1990 (as )

Canu & Bassler, 1920

Harmelin, 1976

Harmelin 1976, Ganias 1990, Morri et al. 1999

Lamarck, 1816

Harmelin, 1976

Castritsi-Catharios and Kiortis 1984, Castritsi-Catharios et al. 1985a, Castritsi-Catharios et al. 1985b, Castritsi-Catharios et al. 1986a, Castritsi-Catharios et al. 1986b, Ganias 1990

(Pallas, 1766)

Castritsi-Catharios et al. 1985a, Ganias 1990

(Busk, 1875)

Ganias 1990

Thompson in Harmer, 1898

Castritsi-Catharios and Ganias 1989, Ganias 1990, Morri et al. 1999

Harmelin, 1976

Ganias 1990​

Analysis

The bryozoan fauna of the Greek seas consists of 237 species, classified into 127 genera, 66 families, 3 orders, and 2 classes (Suppl. material 1). The vast majority belongs to the class , comprising 198 species (177 and 21 ) and the remaining 39 species are (all ). The families with the highest number of species belong to , and are (16 species), (14 spp.), (12 spp.), (12 spp.), (11 spp.), (10 spp.), and (10 spp.). These families are the most speciose in the Mediterranean Sea as well (Rosso and Di Martino 2016). All other families include less than 10 species each.

All species included in the checklist are recognized in the World List of and the World Register of Marine Species, except for three of them. These are: (Heller, 1967) and Harmelin, 1976, allocated within the genus Canu and Bassler, 1920 and not David, Mongereau and Pouyet, 1972; '' (Harmelin, 1976), provisionally left in its former allocation because the characters of the species differ from those of the genus Canu, 1918 that has been suggested for its allocation (although Canu, 1918 is recognized as a synonym of Canu, 1918). The taxon sp. 1 sensu Hayward, 1974 described from Chios Island, was retained in the checklist following extensive lists by Zabala and Maluquer 1988, Rosso et al. 2010, Rosso and Di Martino 2016 and taxonomic papers with description and illustrations of material from Tyrrhenian and Adriatic localities (Chimenz Gusso et al. 2014).

A number of taxa were omitted from the present checklist (Table 1) for different reasons: (1) taxa now recognized as species complexes and considered absent from the Mediterranean; further examination of specimens is needed to ascertain their correct identification; (2) species whose currently accepted geographic distribution does not include the Mediterranean Sea; the examination of the material with the Scanning Electron Microscope could possibly raise the number of non-indigenous species in the Mediterranean basin; (3) species known exclusively as fossils from Tertiary sediments of Europe, whose persistence in the Mediterranean Sea throughout the Messinian Salinity Crisis needs to be ascertained.

Table 1.

Questionable taxa reported from Greece not included in the present checklist.

Species and authority as given in the source	First report from Greece	Remarks
Adeonella polystomella (Reuss, 1847)	Harmelin (1969)	Omitted following Rosso and Novosel (2010).
Cribrilina annulata (Fabricius, 1780)	Castritsi-Catharios and Kiortis (1984)	Questionable (see Rosso and Di Martino 2016).
Cribrilina punctata (Hassall, 1841)	Castritsi-Catharios and Marcopoulou-Diacantoni (1983)	Possibly confused with Collarina balzaci (see Rosso and Di Martino 2016).
Escharina porosa (Smitt)	Hayward (1974)	Removed after the splitting of a previous complex of species (see Berning et al. 2008). Examination of material is needed.
Hippodiplosia delicatula Manzoni	Castritsi-Catharios et al. (1986)	Possibly an exclusively fossil species.
Microecia suborbicularis (Hincks)	Castritsi-Catharios et al. (1986)	The material for this species needs further examination (see Rosso et al. 2010).
Monoporella fimbriata carinifera Canu & Bassler, 1929 and M. nodulifera (Hincks)	Harmelin (1969), Hayward (1974)	Replaced by M. bouchardii following Harmelin (2014a).
Parasmittina nitida Verrill	Castritsi-Catharios et al. (1986)	Species restricted to the SW Atlantic (Winston and Maturo 2009).
Pentapora foliacea	Hayward (1974)	Species restricted to the North Atlantic (see Lombardi et al. 2010 and Rosso et al. 2010).
? Rhyncozoon lobulatum (Waters, 1879)	Harmelin (1969)	This species name is likely to refer to an aged portion of R. digitatum (see Zabala and Maluquer 1988).
Schizomavella gardensis	Castritsi-Catharios et al. (1986)	A fossil species described from the Burdigalian of France.
Scrupocellaria reptans	Hayward (1974)	Species restricted to the British Isles (see Rosso and Di Martino 2016 and references therein). All records from the Mediterranean (except for specimens from Alexandria attributed to C. aegyptiaca) need to be examined.
Sertella scbuermanni Jullien	Castritsi-Catharios et al. (1986)	This fossil species is reported only from its type locality (Miocene of west Germany).
Tricellaria peachii Osburn	Castritsi-Catharios et al. (1986)	Species restricted to the North Atlantic. Could be misidentified with Tricellaria inopinata.
Tubulipora phalangea (Couch, 1844)	Antoniadou and Chintiroglou (2005)	Species restricted to the Boreo-Arctic Atlantic Ocean (see Rosso and Di Martino 2016).
Turbicellepora armata	Castritsi-Catharios et al. (1985a), Castritsi-Catharios et al. (1985b)	This species is restricted to the North Atlantic.
Turbicellepora redoutei (Audouin, 1826)	Harmelin (1969)	Species seemingly restricted to the Indo-Pacific. It was recorded only once in the Mediterranean.

Discussion

The bryozoan fauna of the Greek seas makes up 42.7% of the species, 59.9% of the genera and 71% of the families of the Mediterranean bryozoan fauna, specifically 41.7% of the , 36.8% of the and 52% of the species, based on the recent update by Rosso and Di Martino (2016). Despite the small number of sporadically distributed studies on this phylum in Greece, a considerable number of bryozoan species has been reported compared to adjacent eastern Mediterranean countries. A total of 185 species has been recorded from Turkey, 139 of which along the Aegean coasts (Koçak and Aydın Önen 2014) and 93 species from Lebanon (Harmelin et al. 2016), while 60 species were estimated to occur along the Mediterranean coasts of Israel (Sokolover et al. 2016). These numbers confirm the observed impoverishment of the bryozoan fauna from the western to the eastern basin (Harmelin et al. 2016).

Among the species reported from Greece, 12 are considered endemic to the eastern Mediterranean basin; these are the cheilostomes , , , , , , , , , , and . Of these, 5 species were first described from the island of Chios based on material collected by Hayward (i.e. , , , , and ), was described from the Gulf of Thessaloniki, while was described from the Turkish coasts of the Aegean Sea. The cheilostome , which was described from the Aegean Sea, is considered more typical of the eastern Mediterranean basin, including the Sicily Strait, the western Ionian and south Adriatic seas (Harmelin et al. 2016 and references therein, AR personal observations). Nevertheless, this species was recorded in a few localities of western Corsica (Calvet 1902, Gautier 1962) and Sardinia (AR and E. Di Martino, personal observations) in the western Mediterranean.

Interestingly, three of the bryozoans recorded from Greece are typical of the North Atlantic (Hayward and Ryland 1998, Hayward and Ryland 1999) and have been rarely reported from the Mediterranean Sea. These are (Hincks, 1860), also reported from northern Catalonia (Madurell et al. 2013), (Quelch, 1884), also reported from the western Ionian Sea (Rosso 1996a, Rosso 1996b, Rosso et al. 2014), and (Busk, 1860).

The recent introduction of non-indigenous species, mostly lessepsian migrants, over the last decades, has considerably increased the number of bryozoans occurring in the Mediterranean Sea (Rosso and Di Martino 2016). Studies in the southeastern Mediterranean basin have brought to light a considerable number of NIBs (Harmelin 2014b, Gerovasileiou et al. 2016, Harmelin et al. 2016, Sokolover et al. 2016), mainly originated from the Red Sea/Indo-Pacific Ocean. Until today, 12 NIBs have been reported from the Greek seas, representing 5% of the bryozoan fauna of Greece and 20% of the Mediterranean NIB fauna; these are the ctenostomes and and the cheilostomes , , , , , , , , , and .

Bryozoans from the Greek seas were reported from a variety of habitats, including soft sediments, seagrass leaves and rhizomes, macroalgae, coralligenous concretions and, to a smaller extent, from marine caves. Further research on bryozoan diversity in understudied habitats, typically species-rich in bryozoans (e.g. marine caves and coralligenous beds), is expected to increase our knowledge, possibly revealing additional new and non-indigenous species.

Checklist of Marine  of Greece

Data type: Taxonomic checklist

Brief description: Taxonomic checklist of known to occur in the Greek seas.

File: oo_104057.xls