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This article should be cited as: Quigley, D.T.G. (2014) Did melting ice sheets create temporary low-salinity corridors that facilitated the natural colonization of Ireland by stenohaline fishes? In: Sleeman, D.P., Carlsson, J. and Carlsson, J.E.L. Mind the Gap II: new insights into the Irish postglacial. Irish Naturalists' Journal, Belfast.

Date of publication: 27 November 2014

The facilitated natural colonisation of Ireland by stenohaline fishes

Did melting ice sheets create temporary low-salinity corridors that facilitated the natural colonisation of Ireland by stenohaline fishes? *Declan Quigley

Dingle Oceanworld (Mara Beo Teo), The Wood, Dingle, Co. Kerry Ireland’s current freshwater fish fauna is represented by a relatively small amalgam of species, including pre-glacial relict populations, post-glacial colonisers, species introduced and spread by man, seasonally occurring euryhaline and diadromous species and occasional vagrants from other biogeographical zones. The traditional consensus promulgated by many Irish ichthyologists over the last century suggests that Ireland’s indigenous freshwater fish could only have been derived from diadromous (or previously diadromous) euryhaline species which entered from the sea after the end of the last Ice Age, c.15 000 years BP. Although there is clear evidence that some of Ireland’s freshwater fish species were introduced by man during historical times, for many species the evidence has often been based on rather sketchy anecdotal references (and in some cases, the absence of any evidence) and/or on the assumption that these species are strictly stenohaline and therefore could not have naturally colonised (or re-colonised) Ireland via the sea after the last Ice Age. Many of these so-called stenohaline species currently live and reproduce in cold saline waters (up to 15 psu) in the Baltic and other boreal/arctic regions in the Northern Hemisphere. Is it possible that some of these species may have naturally migrated into Irish waters via low salinity surface water corridors created by melting ice sheets c.15 000 years BP? The current review considers evidence from several disciplines in the formulation of an alternative hypothesis which proposes that Irish freshwater habitats may have been naturally colonised (or re-colonised) by several stenohaline fish species via low salinity water corridors following the last Ice Age. Keywords: fish, freshwater, Ice Age, Ireland and natural colonisation Introduction Fish species reported from Irish freshwater habitats Ireland’s current freshwater fish fauna is represented by a relatively small amalgam of species, including pre-glacial relict populations, post-glacial colonisers, species introduced and spread by man, seasonally occurring euryhaline and diadromous species and occasional vagrants from other biogeographical zones (Table 1). Although a least 57 species of fish have been reported from Irish freshwater habitats in the literature, anecdotal references to the occurrence of twelve of these species has never been formally verified and most of these reports are likely erroneous: Beluga (Huso huso (Linnaeus, 1758)) (Humphreys 1845); Barbel (Barbus barbus (Linnaeus, 1758)) (Browne 1774, Thompson 1856, Went 1949); Goldfish (Carassius auratus (Linnaeus, 1758)) (Thompson 1856, Ogilby 1885); Crucian Carp (Carassius carassius *Corresponding author – [email protected] 124 Irish Naturalists’ Journal Mind the Gap II

(Linnaeus, 1758)) (J. Caffrey pers. comm.); White Catfish (Ameiurus catus (Linnaeus, 1758)) (Holmes 1994); Brown Bullhead (Ameiurus nebulosus (Lesueur, 1819)) (Welcomme 1988, Holcik 1991); Wels Catfish (Silurus glanis Linnaeus, 1758) (Thompson 1841, 1856); Chinook Salmon (Oncorhynchus tshawytscha (Walbaum, 1792)) (Welcomme 1988, Holcik 1991); Grayling (Thymallus thymallus (Linnaeus, 1758) (Rutty 1772, Browne 1774, Thompson 1856, Went 1949); Flathead Grey Mullet (Mugil cephalus Linnaeus, 1758) (Browne 1774, Sampson 1802, M’Skimin 1909, Colby 1837, Templeton 1837); Common Bullhead (Cottus gobio Linnaeus, 1758) (Browne 1774, Sampson 1802, M’Skimin 1909, Thompson 1856, Ogilby 1885, Went 1949) and Zander (Sander lucioperca (Linnaeus, 1758)) (Browne 1774). Over the last century it has consistently been argued that Ireland’s native freshwater fish fauna is exclusively composed of a relatively small group (12) of pre- and post-glacial amphihaline colonisers: sea lamprey Petromyzon marinus L., river lamprey Lampetra fluviatilis (L.), eel Anguilla anguilla (L.), allis shad Alosa alosa (L.), twaite

shad A. fallax (Lacepede, 1803), Killarney shad A. killarnensis Regan, 1916, smelt Osmerus eperlanus (L.), pollan Coregonus pollan Thompson, 1835, sea trout/brown trout Salmo trutta L., Atlantic salmon S. salar L., Arctic charr Salvelinus alpinus (L.) and three-spined stickleback Gasteroseus aculeatus L. (Wheeler, 1977; Went 1980; Wilson 1986; Maitland and Campbell 1992; King et al. 2011). Some of these species e.g. Atlantic salmon, sea trout and Arctic charr, are also known to have been supplemented by the introduction of nonnative stocks from other European countries (Fahy 1985; Cross 1996; Quigley and Flannery 1997a). At least nine other native euryhaline species also occur on a seasonal basis in Irish freshwater habitats, albeit some e.g. thick-lipped grey mullet Chelon labrosus (Risso, 1826) (Kennedy and Fitzmaurice 1969; Quigley 2012a), golden grey mullet Liza aurata (Risso, 1810) (Walsh 1993; Quigley 2012b), thin-lipped grey mullet L. ramada (Risso, 1826) (McCoy 1841; Harvey 1845; Thompson 1856; Ogilby 1885; Kennedy and Fitzmaurice 1972a; Walsh 1993; Quigley 2012b), big-scale sand smelt Atherina boyeri Risso, 1810 (Went and Kennedy 1976), European bass Dicentrarchus labrax (L.) (Kennedy and Fitzmaurice 1972c), transparent goby Aphia minuta (Risso, 1810) (Quigley 1996) and flounder Pleuronectes flesus L. (Rutty 1772; O’Neill et al. 2011; Quigley 2011) occur more frequently and consistently than others e.g. Atlantic herring Clupea harengus L. (Warren 1898; Barrett-Hamilton 1899) and ballan wrasse Labrus bergylta Ascanius, 1767 (Briggs and McCurdy 1977). In addition, four naturally occurring amphihaline vagrant species have been recorded, either infrequently e.g. European sturgeon Acipenser sturio L. (Rutty 1772; Browne 1774; Sampson 1802; Templeton 1837; Harvey 1845; Ogilby 1885; Went 1948; Quigley 2014) and anchovy Engraulis encrasicolus (L.) (Quigley and Flannery 1997b), or rarely e.g. pink salmon Oncorhynchus gorbuscha (Walbaum, 1792) (Went 1974; Moriarty and Fitzmaurice 2002a, 2002b; Minchin 2007) and tarpon Megalops atlanticus Valenciennes, 1847 (Twomey and Byrne 1985). There is clear evidence that the following seven species were introduced by man during historical times: Common Carp (Cyprinus carpio Linnaeus, 1758) (Rutty 1772, Browne 1774, Templeton 1837, Thompson 1856, Holcik 1991, Brazier et al. 2012); Chub (Squalius cephalus (Linnaeus, 1758) (Browne 1774, Thompson 1856, Went 1949, Caffrey et al. 2008); Dace Leuciscus leuciscus (Linnaeus, 1758) (Holt 1911, Went 1946, 1950, 1978, Healy 1956, Holcik 1991); Roach (Rutilus rutilus (Linnaeus, 1758) (Browne 1774, Went

1946, 1949, 1950, 1978, Holcik 1991); Tench (Tinca tinca (Linnaeus, 1758) (Rutty 1772, Templeton 1837, Harvey 1845, Thompson 1856, Holcik 1991, Brazier et al. 2012); Huchen (Hucho hucho (Linnaeus, 1758) (Moriarty and Fitzmaurice 2002a, 2002b, Minchin 2007) and Rainbow Trout (Oncorhynchus mykiss (Walbaum, 1792)) (Ravenel 1889, Went 1946, Fahy 1983, Minchin 2007). In addition, two tropical aquarium species, most likely discarded by their owners, were recently found dead in Irish waters: Pacu (Colossoma sp.) (P. McCluskey pers. comm.) and Piranha (Serrasalmus sp.) (Minchin 2007). However, the evidence for human-mediated introductions of the following nine species has often been based on rather sketchy anecdotal references and in some cases, the absence of any evidence: Brook Lamprey (Lampetra planeri (Bloch, 1784)) (Igoe et al. 2004); Common Bream (Abramis brama (Linnaeus, 1758) (O’Flaherty 1684, Browne 1774, Templeton 1837, Thompson 1856, Went 1946, 1978, Hayden et al. 2011); Gudgeon (Gobio gobio (Linnaeus, 1758)) (Browne 1774, Templeton 1837, Thompson 1856, Ogilby 1885, Went 1949, 1978, Kennedy and Fitzmaurice 1972b); Minnow (Phoxinus phoxinus (Linnaeus, 1758)) (Thompson 1856, Ogilby 1885, Scharff 1905, Went 1949, 1978); Rudd (Scardinius erythrophthalmus (Linnaeus, 1758)) (O’Flaherty 1684, Rutty 1772, Templeton 1837, Harvey 1845, Thompson 1856, Ogilby 1885, Went 1946, 1978, Kennedy and Fitzmaurice 1974); Stone Loach (Barbatula barbatula (Linnaeus, 1758)) (Rutty 1772, Browne 1774, Templeton 1837, Thompson 1856, Ogilby 1885, Went 1946, 1949, 1978); Pike (Esox lucius Linnaeus, 1758) (O’Flaherty 1684, Rutty 1772, Browne 1774, Sampson 1802, Colby 1837, Templeton 1837, Harvey 1845, Thompson 1856, Went 1949, 1957, Holcik 1991, Pedreschi et al. 2013); Nine-spined Stickleback (Pungitus pungitus (Linnaeus, 1758)) (Rutty 1772, Browne 1774, Templeton 1837, Thompson 1841, Harvey 1845, Went 1946, 1978, Hayden 2002) and Perch (Perca fluviatilis Linnaeus, 1758) (Rutty 1772, Browne 1774, Sampson 1802, Templeton 1837, Thompson 1856, Harvey 1845, Went 1946, 1949, 1978). Indeed, it has frequently and consistently been argued that these freshwater species are strictly stenohaline and therefore could not have naturally colonised (or re-colonised) Ireland based on the assumption that the island of Ireland was fully isolated by high salinity marine sea conditions at end of the last Ice Age (Went 1946, 1950, 1980, Wheeler, 1977, Fitzmaurice 1981, Wilson 1986, Maitland and Campbell 1992, Griffiths 1997, Moriarty and Fitzmaurice 2000a, King et Irish Naturalists’ Journal Mind the Gap II 125


Did melting ice sheets create temporary low-salinity corridors that facilitated the natural colonisation of Ireland by stenohaline fishes? *Declan Quigley

Dingle Oceanworld (Mara Beo Teo), The Wood, Dingle, Co. Kerry Ireland’s current freshwater fish fauna is represented by a relatively small amalgam of species, including pre-glacial relict populations, post-glacial colonisers, species introduced and spread by man, seasonally occurring euryhaline and diadromous species and occasional vagrants from other biogeographical zones. The traditional consensus promulgated by many Irish ichthyologists over the last century suggests that Ireland’s indigenous freshwater fish could only have been derived from diadromous (or previously diadromous) euryhaline species which entered from the sea after the end of the last Ice Age, c.15 000 years BP. Although there is clear evidence that some of Ireland’s freshwater fish species were introduced by man during historical times, for many species the evidence has often been based on rather sketchy anecdotal references (and in some cases, the absence of any evidence) and/or on the assumption that these species are strictly stenohaline and therefore could not have naturally colonised (or re-colonised) Ireland via the sea after the last Ice Age. Many of these so-called stenohaline species currently live and reproduce in cold saline waters (up to 15 psu) in the Baltic and other boreal/arctic regions in the Northern Hemisphere. Is it possible that some of these species may have naturally migrated into Irish waters via low salinity surface water corridors created by melting ice sheets c.15 000 years BP? The current review considers evidence from several disciplines in the formulation of an alternative hypothesis which proposes that Irish freshwater habitats may have been naturally colonised (or re-colonised) by several stenohaline fish species via low salinity water corridors following the last Ice Age. Keywords: fish, freshwater, Ice Age, Ireland and natural colonisation Introduction Fish species reported from Irish freshwater habitats Ireland’s current freshwater fish fauna is represented by a relatively small amalgam of species, including pre-glacial relict populations, post-glacial colonisers, species introduced and spread by man, seasonally occurring euryhaline and diadromous species and occasional vagrants from other biogeographical zones (Table 1). Although a least 57 species of fish have been reported from Irish freshwater habitats in the literature, anecdotal references to the occurrence of twelve of these species has never been formally verified and most of these reports are likely erroneous: Beluga (Huso huso (Linnaeus, 1758)) (Humphreys 1845); Barbel (Barbus barbus (Linnaeus, 1758)) (Browne 1774, Thompson 1856, Went 1949); Goldfish (Carassius auratus (Linnaeus, 1758)) (Thompson 1856, Ogilby 1885); Crucian Carp (Carassius carassius *Corresponding author – [email protected] 124 Irish Naturalists’ Journal Mind the Gap II

(Linnaeus, 1758)) (J. Caffrey pers. comm.); White Catfish (Ameiurus catus (Linnaeus, 1758)) (Holmes 1994); Brown Bullhead (Ameiurus nebulosus (Lesueur, 1819)) (Welcomme 1988, Holcik 1991); Wels Catfish (Silurus glanis Linnaeus, 1758) (Thompson 1841, 1856); Chinook Salmon (Oncorhynchus tshawytscha (Walbaum, 1792)) (Welcomme 1988, Holcik 1991); Grayling (Thymallus thymallus (Linnaeus, 1758) (Rutty 1772, Browne 1774, Thompson 1856, Went 1949); Flathead Grey Mullet (Mugil cephalus Linnaeus, 1758) (Browne 1774, Sampson 1802, M’Skimin 1909, Colby 1837, Templeton 1837); Common Bullhead (Cottus gobio Linnaeus, 1758) (Browne 1774, Sampson 1802, M’Skimin 1909, Thompson 1856, Ogilby 1885, Went 1949) and Zander (Sander lucioperca (Linnaeus, 1758)) (Browne 1774). Over the last century it has consistently been argued that Ireland’s native freshwater fish fauna is exclusively composed of a relatively small group (12) of pre- and post-glacial amphihaline colonisers: sea lamprey Petromyzon marinus L., river lamprey Lampetra fluviatilis (L.), eel Anguilla anguilla (L.), allis shad Alosa alosa (L.), twaite

shad A. fallax (Lacepede, 1803), Killarney shad A. killarnensis Regan, 1916, smelt Osmerus eperlanus (L.), pollan Coregonus pollan Thompson, 1835, sea trout/brown trout Salmo trutta L., Atlantic salmon S. salar L., Arctic charr Salvelinus alpinus (L.) and three-spined stickleback Gasteroseus aculeatus L. (Wheeler, 1977; Went 1980; Wilson 1986; Maitland and Campbell 1992; King et al. 2011). Some of these species e.g. Atlantic salmon, sea trout and Arctic charr, are also known to have been supplemented by the introduction of nonnative stocks from other European countries (Fahy 1985; Cross 1996; Quigley and Flannery 1997a). At least nine other native euryhaline species also occur on a seasonal basis in Irish freshwater habitats, albeit some e.g. thick-lipped grey mullet Chelon labrosus (Risso, 1826) (Kennedy and Fitzmaurice 1969; Quigley 2012a), golden grey mullet Liza aurata (Risso, 1810) (Walsh 1993; Quigley 2012b), thin-lipped grey mullet L. ramada (Risso, 1826) (McCoy 1841; Harvey 1845; Thompson 1856; Ogilby 1885; Kennedy and Fitzmaurice 1972a; Walsh 1993; Quigley 2012b), big-scale sand smelt Atherina boyeri Risso, 1810 (Went and Kennedy 1976), European bass Dicentrarchus labrax (L.) (Kennedy and Fitzmaurice 1972c), transparent goby Aphia minuta (Risso, 1810) (Quigley 1996) and flounder Pleuronectes flesus L. (Rutty 1772; O’Neill et al. 2011; Quigley 2011) occur more frequently and consistently than others e.g. Atlantic herring Clupea harengus L. (Warren 1898; Barrett-Hamilton 1899) and ballan wrasse Labrus bergylta Ascanius, 1767 (Briggs and McCurdy 1977). In addition, four naturally occurring amphihaline vagrant species have been recorded, either infrequently e.g. European sturgeon Acipenser sturio L. (Rutty 1772; Browne 1774; Sampson 1802; Templeton 1837; Harvey 1845; Ogilby 1885; Went 1948; Quigley 2014) and anchovy Engraulis encrasicolus (L.) (Quigley and Flannery 1997b), or rarely e.g. pink salmon Oncorhynchus gorbuscha (Walbaum, 1792) (Went 1974; Moriarty and Fitzmaurice 2002a, 2002b; Minchin 2007) and tarpon Megalops atlanticus Valenciennes, 1847 (Twomey and Byrne 1985). There is clear evidence that the following seven species were introduced by man during historical times: Common Carp (Cyprinus carpio Linnaeus, 1758) (Rutty 1772, Browne 1774, Templeton 1837, Thompson 1856, Holcik 1991, Brazier et al. 2012); Chub (Squalius cephalus (Linnaeus, 1758) (Browne 1774, Thompson 1856, Went 1949, Caffrey et al. 2008); Dace Leuciscus leuciscus (Linnaeus, 1758) (Holt 1911, Went 1946, 1950, 1978, Healy 1956, Holcik 1991); Roach (Rutilus rutilus (Linnaeus, 1758) (Browne 1774, Went

1946, 1949, 1950, 1978, Holcik 1991); Tench (Tinca tinca (Linnaeus, 1758) (Rutty 1772, Templeton 1837, Harvey 1845, Thompson 1856, Holcik 1991, Brazier et al. 2012); Huchen (Hucho hucho (Linnaeus, 1758) (Moriarty and Fitzmaurice 2002a, 2002b, Minchin 2007) and Rainbow Trout (Oncorhynchus mykiss (Walbaum, 1792)) (Ravenel 1889, Went 1946, Fahy 1983, Minchin 2007). In addition, two tropical aquarium species, most likely discarded by their owners, were recently found dead in Irish waters: Pacu (Colossoma sp.) (P. McCluskey pers. comm.) and Piranha (Serrasalmus sp.) (Minchin 2007). However, the evidence for human-mediated introductions of the following nine species has often been based on rather sketchy anecdotal references and in some cases, the absence of any evidence: Brook Lamprey (Lampetra planeri (Bloch, 1784)) (Igoe et al. 2004); Common Bream (Abramis brama (Linnaeus, 1758) (O’Flaherty 1684, Browne 1774, Templeton 1837, Thompson 1856, Went 1946, 1978, Hayden et al. 2011); Gudgeon (Gobio gobio (Linnaeus, 1758)) (Browne 1774, Templeton 1837, Thompson 1856, Ogilby 1885, Went 1949, 1978, Kennedy and Fitzmaurice 1972b); Minnow (Phoxinus phoxinus (Linnaeus, 1758)) (Thompson 1856, Ogilby 1885, Scharff 1905, Went 1949, 1978); Rudd (Scardinius erythrophthalmus (Linnaeus, 1758)) (O’Flaherty 1684, Rutty 1772, Templeton 1837, Harvey 1845, Thompson 1856, Ogilby 1885, Went 1946, 1978, Kennedy and Fitzmaurice 1974); Stone Loach (Barbatula barbatula (Linnaeus, 1758)) (Rutty 1772, Browne 1774, Templeton 1837, Thompson 1856, Ogilby 1885, Went 1946, 1949, 1978); Pike (Esox lucius Linnaeus, 1758) (O’Flaherty 1684, Rutty 1772, Browne 1774, Sampson 1802, Colby 1837, Templeton 1837, Harvey 1845, Thompson 1856, Went 1949, 1957, Holcik 1991, Pedreschi et al. 2013); Nine-spined Stickleback (Pungitus pungitus (Linnaeus, 1758)) (Rutty 1772, Browne 1774, Templeton 1837, Thompson 1841, Harvey 1845, Went 1946, 1978, Hayden 2002) and Perch (Perca fluviatilis Linnaeus, 1758) (Rutty 1772, Browne 1774, Sampson 1802, Templeton 1837, Thompson 1856, Harvey 1845, Went 1946, 1949, 1978). Indeed, it has frequently and consistently been argued that these freshwater species are strictly stenohaline and therefore could not have naturally colonised (or re-colonised) Ireland based on the assumption that the island of Ireland was fully isolated by high salinity marine sea conditions at end of the last Ice Age (Went 1946, 1950, 1980, Wheeler, 1977, Fitzmaurice 1981, Wilson 1986, Maitland and Campbell 1992, Griffiths 1997, Moriarty and Fitzmaurice 2000a, King et Irish Naturalists’ Journal Mind the Gap II 125

Quigley, D.

al. 2011). However, it is interesting to note that several of these and other so-called stenohaline species currently live and reproduce in cold saline waters (up to 15 psu) in the Baltic and other boreal/arctic regions in the Northern Hemisphere e.g. Common Bream, Roach, Rudd, Nine-spined Stickleback, Pike, Perch and Zander (Kottelat and Freyhof 2007). Is it possible that some of these species may have naturally migrated into Irish waters via low salinity corridors created by melting ice sheets c.15 000 years BP? The current review considers evidence from several disciplines in formulating an alternative hypothesis which proposes that Ireland may have been naturally colonised (or re-colonised) by several stenohaline fish species via low salinity corridors following the end of the last Ice Age c.15 000 years BP. Although it is acknowledged that the hypothesis is highly speculative and raises more questions than answers, it identifies significant knowledge gaps where a multidisciplinary approach to further research is required, particularly in relation to those species for which there is little or no documented evidence for human-mediated introductions. Absence of evidence may not always be evidence of absence (Sagan 1983). Potential post-glacial low-salinity colonisation routes for stenohaline fishes Pedreschi et al. (2013) recently published the results of a molecular genetic study which strongly suggested the possibility that pike, contrary to long-held and promulgated popular opinion (Went 1957), may have naturally colonised Ireland, most likely from refuge populations in mainland Europe c.8 000 years BP, i.e. c.7 000 years after the final retreat of the glaciers from Ireland c.15 000 years BP (Edwards and Brooks 2006) and c.1 000 years after the first archaeological evidence of human habitation c.9 000 years BP (Mitchell 1994). Pedreschi et al. (2013) indicated that that around 3 500-4 000 years BP, Irish and British Pike populations became isolated. However, Ensing (2014) recently challenged the veracity of the results and conclusions presented by Pedreschi et al. (2013). If salinity levels around Ireland were low enough for Pike to colonise Irish freshwater habitats from European refugia c.8 000 years BP, is it possible that these same conditions may also have facilitated the natural colonisation of Ireland by other species of stenohaline fish during the same period? Unlike terrestrial animals, stenohaline fish would have required relatively low salinity water corridors rather than land bridges in order to gain access to and colonise Ireland’s evolving post-glacial freshwater habitats. 126 Irish Naturalists’ Journal Mind the Gap II


When, where and how did these low salinity events occur and for how long did they persist? During the global Last Glacial Maximum (gLGM), which occurred c.21 000 years BP, large ice sheets existed in the mid to high latitudes of the Northern Hemisphere, sufficient in volume to lower eustatic sea levels by up to 130 m (Clark et al. 2012). The British-Irish Ice Sheet (BIIS), which constituted a relatively small part of the overall volume of ice in the Northern Hemisphere during the gLGM, attained its maximum extent c.27 000 years BP when it is estimated to have covered an area of c.840 000 km2, including extensive areas of the current North Sea and continental shelves of Britain and Ireland (Clarke et al. 2012). When the BIIS finally melted c.15 000 years BP, it’s total freshwater runoff is estimated to have raised global sea levels by c.2.5 m (Clark et al. 2012). The release of this huge volume of fresh water into the ocean is likely to have had a significant impact, albeit temporary, on the evolving post glacial ecology of coastal and continental shelf areas by, inter alia, lowering salinity levels and thus providing potential dispersal and colonisation routes for refuge populations of stenohaline fishes. Southern North Sea Lake and the Fleuve Manche (English Channel Palaeo-River) Between 20 000 and 18 000 years BP, it is thought that a large freshwater pro-glacial lake occupied an area (c.1 300 km2) of the southern North Sea (Clark et al. 2012). The lake was dammed to the north by an ice bridge extending between Britain and Norway and to the south by a low ridge, the Weald-Artois, stretching from SE England to NE France. A number of large rivers discharged into this lake, including the Thames, Rhine-Meuse and Scheldt. The lake over-spilled at the Straits of Dover into a large palaeo-valley (currently the English Channel) where it joined with several other rivers (e.g. Somme and Seine) to form the English Channel Palaeo-River System or Fleuve Manche which discharged into the northern Bay of Biscay between SW Britain and NW France (Gibbard 2007). Between 18 000 and 17 500 years BP, the Fleuve Manche palaeo-river is estimated to have carried half of the freshwater drainage of Western Europe to the Atlantic Ocean (Gibbard 2007); the estimated discharge rate ranged from 400 000 to 800 000 m3/second (Toucanne et al. 2010). Sedimentary deposits of freshwater origin derived from the discharge of the Fleuve Manche have been found in the Bay of Biscay up to 200 km off the original mouth of this palaeo-river (Auffret et al. 2002, Toucanne et al. 2008, 2010). Toucanne et al. (2010) suggested that the

massive runoff of melt-water through the Fleuve Manche, combined with the massive iceberg releases from the circum-North Atlantic ice sheets may have had a profound destabilising effect on the strength of the Atlantic Ocean’s thermohaline circulation. Indeed, the analysis of dinocyst and foramiferal assemblages in sediment cores from both sides of the North Atlantic during this period indicated that melting glaciers maintained an extensive zone of relatively cold and slightly dilute seawater on the continental shelves (Vilks 1981, Auffret et al. 2002, Van Nieuwenhove et al. 2008, Eynaud et al. 2012). Is it possible that the massive freshwater runoff and offshore plume created by the Fleuve Manche may have facilitated the migration of stenohaline fishes from European refugia and the colonisation of suitable freshwater habitats in Ireland during this period? ‘Fleuve Hibernia’ (Waterford Harbour) It is interesting to note that Gallagher (2002) discovered what appears to be an exceptionally large submerged palaeo-river channel extending over a distance of 22 km from the mouth of Waterford Harbour (SE Ireland) onto the nearshore continental shelf of the Celtic Sea to a depth of 56 m below present sea level. Gallagher estimated that this palaeo-river could have discharged up to 46 832 m3/sec of freshwater, several orders of magnitude greater than those of the present river channel. Is it possible that the low salinity plume derived from this ‘Fleuve Hibernia’ may have coalesced with the freshwater plume of the Fleuve Manche, thereby facilitating the migration and colonisation of Ireland by stenohaline fishes? ‘Lake Hibernia’ (Irish Sea) Between c.20 000 and 14 000 years BP, the present Irish Sea is thought to have been occupied by a pro-glacial freshwater lake (‘Lake Hibernia’) which was closed to marine influence in the north by an ice sheet extending between Ireland, Scotland and the Isle of Man and in the south by a low-level (15 m) emerged Celtic Sea seafloor extending from Land’s End (SW Britain) to Cork (Lambeck 1995, Clark et al. 2012). Lambeck (1995) remarked that the volumes of melt-water draining into the Irish Sea at this time would have been very substantial such that the emerged sea floor would be a swampy and inhospitable region at best. Nevertheless, it may have been possible for some stenohaline fish populations to survive and perhaps colonise suitable freshwater habitats on either side of this pro-glacial lake. However, between 15 000 and 14 000 years

BP, a large pulse of melt-water, equivalent to the wasting of almost three times the amount of ice currently stored in the Greenland Ice Sheet, resulted in a rapid increase in global sea levels giving rise to the inundation of the Irish Sea and ultimately the separation of Ireland from Britain (Edwards and Brooks 2008). However, considering that this rapid increase in sea level was caused by glacial melt-water, perhaps the Irish Sea was inundated, at least temporarily, by low salinity rather than fully marine water? Perhaps the latter short-lived event may also have facilitated the migration of stenohaline fishes from European refugia and the colonisation of suitable freshwater habitats in Ireland? The catastrophic final Doggerland (North Sea)

flooding

of

During the late glacial period (c.18 000-14 000 years BP) it is estimated that large areas of the present North Sea were exposed above sea level and constituted a low-lying land connection between Britain and mainland Europe known as Doggerland (Coles 2000). During this period the area was characterized by low-lying tundra with a complex drainage system of meandering rivers, marshes and lakes (Coles 2000). However, by 10 000 years BP, the southward advance of the North Sea had started consistent with evidence of brackish marine incursions into the southern North Sea during the early Holocene (Lambeck 1995). Soon afterwards, the Dogger Bank, an upland area of Doggerland situated in the central North Sea, became isolated as the only remaining emerged platform (Lambeck 1995, Coles 2000). Then, c.8 200-8 000 years BP, it is thought that the Storegga Tsunami, one of the largest known during the Holocene, and generated by a massive submarine landslide off the Norwegian coast, led to the final catastrophic flooding of Doggerland and the North Sea (Weninger et al. 2008). Is it possible that this catastrophic event may have literally washed out any remaining stenohaline fishes from the Dogger Bank into the English Channel and onwards via a low-salinity plume to the Irish coast? It is interesting to recall that the timing of this catastrophic event corresponds very closely with the earliest proposed date for the presence of Pike in Irish waters (Pedreschi et al. 2013). Conclusions If Pike naturally colonised Ireland c.8 000 years BP (Pedreschi et al. 2013), sufficiently low salinity conditions must have existed around and off the Irish coast prior to this time in order for successful immigration to have occurred. It Irish Naturalists’ Journal Mind the Gap II 127

Quigley, D.

al. 2011). However, it is interesting to note that several of these and other so-called stenohaline species currently live and reproduce in cold saline waters (up to 15 psu) in the Baltic and other boreal/arctic regions in the Northern Hemisphere e.g. Common Bream, Roach, Rudd, Nine-spined Stickleback, Pike, Perch and Zander (Kottelat and Freyhof 2007). Is it possible that some of these species may have naturally migrated into Irish waters via low salinity corridors created by melting ice sheets c.15 000 years BP? The current review considers evidence from several disciplines in formulating an alternative hypothesis which proposes that Ireland may have been naturally colonised (or re-colonised) by several stenohaline fish species via low salinity corridors following the end of the last Ice Age c.15 000 years BP. Although it is acknowledged that the hypothesis is highly speculative and raises more questions than answers, it identifies significant knowledge gaps where a multidisciplinary approach to further research is required, particularly in relation to those species for which there is little or no documented evidence for human-mediated introductions. Absence of evidence may not always be evidence of absence (Sagan 1983). Potential post-glacial low-salinity colonisation routes for stenohaline fishes Pedreschi et al. (2013) recently published the results of a molecular genetic study which strongly suggested the possibility that pike, contrary to long-held and promulgated popular opinion (Went 1957), may have naturally colonised Ireland, most likely from refuge populations in mainland Europe c.8 000 years BP, i.e. c.7 000 years after the final retreat of the glaciers from Ireland c.15 000 years BP (Edwards and Brooks 2006) and c.1 000 years after the first archaeological evidence of human habitation c.9 000 years BP (Mitchell 1994). Pedreschi et al. (2013) indicated that that around 3 500-4 000 years BP, Irish and British Pike populations became isolated. However, Ensing (2014) recently challenged the veracity of the results and conclusions presented by Pedreschi et al. (2013). If salinity levels around Ireland were low enough for Pike to colonise Irish freshwater habitats from European refugia c.8 000 years BP, is it possible that these same conditions may also have facilitated the natural colonisation of Ireland by other species of stenohaline fish during the same period? Unlike terrestrial animals, stenohaline fish would have required relatively low salinity water corridors rather than land bridges in order to gain access to and colonise Ireland’s evolving post-glacial freshwater habitats. 126 Irish Naturalists’ Journal Mind the Gap II


When, where and how did these low salinity events occur and for how long did they persist? During the global Last Glacial Maximum (gLGM), which occurred c.21 000 years BP, large ice sheets existed in the mid to high latitudes of the Northern Hemisphere, sufficient in volume to lower eustatic sea levels by up to 130 m (Clark et al. 2012). The British-Irish Ice Sheet (BIIS), which constituted a relatively small part of the overall volume of ice in the Northern Hemisphere during the gLGM, attained its maximum extent c.27 000 years BP when it is estimated to have covered an area of c.840 000 km2, including extensive areas of the current North Sea and continental shelves of Britain and Ireland (Clarke et al. 2012). When the BIIS finally melted c.15 000 years BP, it’s total freshwater runoff is estimated to have raised global sea levels by c.2.5 m (Clark et al. 2012). The release of this huge volume of fresh water into the ocean is likely to have had a significant impact, albeit temporary, on the evolving post glacial ecology of coastal and continental shelf areas by, inter alia, lowering salinity levels and thus providing potential dispersal and colonisation routes for refuge populations of stenohaline fishes. Southern North Sea Lake and the Fleuve Manche (English Channel Palaeo-River) Between 20 000 and 18 000 years BP, it is thought that a large freshwater pro-glacial lake occupied an area (c.1 300 km2) of the southern North Sea (Clark et al. 2012). The lake was dammed to the north by an ice bridge extending between Britain and Norway and to the south by a low ridge, the Weald-Artois, stretching from SE England to NE France. A number of large rivers discharged into this lake, including the Thames, Rhine-Meuse and Scheldt. The lake over-spilled at the Straits of Dover into a large palaeo-valley (currently the English Channel) where it joined with several other rivers (e.g. Somme and Seine) to form the English Channel Palaeo-River System or Fleuve Manche which discharged into the northern Bay of Biscay between SW Britain and NW France (Gibbard 2007). Between 18 000 and 17 500 years BP, the Fleuve Manche palaeo-river is estimated to have carried half of the freshwater drainage of Western Europe to the Atlantic Ocean (Gibbard 2007); the estimated discharge rate ranged from 400 000 to 800 000 m3/second (Toucanne et al. 2010). Sedimentary deposits of freshwater origin derived from the discharge of the Fleuve Manche have been found in the Bay of Biscay up to 200 km off the original mouth of this palaeo-river (Auffret et al. 2002, Toucanne et al. 2008, 2010). Toucanne et al. (2010) suggested that the

massive runoff of melt-water through the Fleuve Manche, combined with the massive iceberg releases from the circum-North Atlantic ice sheets may have had a profound destabilising effect on the strength of the Atlantic Ocean’s thermohaline circulation. Indeed, the analysis of dinocyst and foramiferal assemblages in sediment cores from both sides of the North Atlantic during this period indicated that melting glaciers maintained an extensive zone of relatively cold and slightly dilute seawater on the continental shelves (Vilks 1981, Auffret et al. 2002, Van Nieuwenhove et al. 2008, Eynaud et al. 2012). Is it possible that the massive freshwater runoff and offshore plume created by the Fleuve Manche may have facilitated the migration of stenohaline fishes from European refugia and the colonisation of suitable freshwater habitats in Ireland during this period? ‘Fleuve Hibernia’ (Waterford Harbour) It is interesting to note that Gallagher (2002) discovered what appears to be an exceptionally large submerged palaeo-river channel extending over a distance of 22 km from the mouth of Waterford Harbour (SE Ireland) onto the nearshore continental shelf of the Celtic Sea to a depth of 56 m below present sea level. Gallagher estimated that this palaeo-river could have discharged up to 46 832 m3/sec of freshwater, several orders of magnitude greater than those of the present river channel. Is it possible that the low salinity plume derived from this ‘Fleuve Hibernia’ may have coalesced with the freshwater plume of the Fleuve Manche, thereby facilitating the migration and colonisation of Ireland by stenohaline fishes? ‘Lake Hibernia’ (Irish Sea) Between c.20 000 and 14 000 years BP, the present Irish Sea is thought to have been occupied by a pro-glacial freshwater lake (‘Lake Hibernia’) which was closed to marine influence in the north by an ice sheet extending between Ireland, Scotland and the Isle of Man and in the south by a low-level (15 m) emerged Celtic Sea seafloor extending from Land’s End (SW Britain) to Cork (Lambeck 1995, Clark et al. 2012). Lambeck (1995) remarked that the volumes of melt-water draining into the Irish Sea at this time would have been very substantial such that the emerged sea floor would be a swampy and inhospitable region at best. Nevertheless, it may have been possible for some stenohaline fish populations to survive and perhaps colonise suitable freshwater habitats on either side of this pro-glacial lake. However, between 15 000 and 14 000 years

BP, a large pulse of melt-water, equivalent to the wasting of almost three times the amount of ice currently stored in the Greenland Ice Sheet, resulted in a rapid increase in global sea levels giving rise to the inundation of the Irish Sea and ultimately the separation of Ireland from Britain (Edwards and Brooks 2008). However, considering that this rapid increase in sea level was caused by glacial melt-water, perhaps the Irish Sea was inundated, at least temporarily, by low salinity rather than fully marine water? Perhaps the latter short-lived event may also have facilitated the migration of stenohaline fishes from European refugia and the colonisation of suitable freshwater habitats in Ireland? The catastrophic final Doggerland (North Sea)

flooding

of

During the late glacial period (c.18 000-14 000 years BP) it is estimated that large areas of the present North Sea were exposed above sea level and constituted a low-lying land connection between Britain and mainland Europe known as Doggerland (Coles 2000). During this period the area was characterized by low-lying tundra with a complex drainage system of meandering rivers, marshes and lakes (Coles 2000). However, by 10 000 years BP, the southward advance of the North Sea had started consistent with evidence of brackish marine incursions into the southern North Sea during the early Holocene (Lambeck 1995). Soon afterwards, the Dogger Bank, an upland area of Doggerland situated in the central North Sea, became isolated as the only remaining emerged platform (Lambeck 1995, Coles 2000). Then, c.8 200-8 000 years BP, it is thought that the Storegga Tsunami, one of the largest known during the Holocene, and generated by a massive submarine landslide off the Norwegian coast, led to the final catastrophic flooding of Doggerland and the North Sea (Weninger et al. 2008). Is it possible that this catastrophic event may have literally washed out any remaining stenohaline fishes from the Dogger Bank into the English Channel and onwards via a low-salinity plume to the Irish coast? It is interesting to recall that the timing of this catastrophic event corresponds very closely with the earliest proposed date for the presence of Pike in Irish waters (Pedreschi et al. 2013). Conclusions If Pike naturally colonised Ireland c.8 000 years BP (Pedreschi et al. 2013), sufficiently low salinity conditions must have existed around and off the Irish coast prior to this time in order for successful immigration to have occurred. It Irish Naturalists’ Journal Mind the Gap II 127

Quigley, D.

also suggests that suitable freshwater habitats and other palaeo-environmental and ecological conditions must have existed for Pike to have become successfully established. Indeed, it seems reasonable to speculate that the same conditions may also have facilitated the natural colonisation of Ireland by other stenohaline fish, and particularly those species for which there is little or no documented evidence of humanmediated introductions e.g. Brook Lamprey, Common Bream, Gudgeon, Minnow, Rudd, Stone Loach, Nine-spined Stickleback and Perch. Although genetic studies may indicate when, where and whence stenohaline fish species may have colonised Irish waters, the elucidation of colonisation routes and mechanisms requires a multidisciplinary approach, involving, inter alia, various branches of palaeo and quaternary science. Acknowledgments I am grateful to the following for their assistance: Professor Michael A. Bell (Department of Ecology and Evolution, Stony Brook University, New York, USA); Dr Joe Caffrey (Inland Fisheries Ireland, Dublin); Dr Kim M. Cohen (Faculty of Geosciences, Utrecht University, Netherlands); Professor Peter Coxon (Department of Geography, Trinity College, Dublin); Professor Phillip L. Gibbard (Department of Geography, University of Cambridge, UK); Dr Sheila Hamilton-Dyer (SHD Archaeozoology, UK); Professor Stefano Mariani (School of Environment and Life Sciences, University of Salford, UK); Mr Peter McCluskey (West End House, Drogheda, Co Louth); Dr Gillian Scott (National University of Ireland, Department of Geography, Maynooth, Co. Kildare) and two anonymous referees. References Anon (1901) Special investigations and inspection. Report of the U.S. Commission of Fish and Fisheries. Part 26: 31. Auffret, G., Zaragosi, S., Dennielou, B., Cortijo, E., Van Rooij, D., Grousset, F., Pujol, C., Eynaud, F. and Siegert, M. (2002) Terrigenous fluxes at the Celtic margin during the last glacial cycle. Marine Geology 188: 79-108. Barrett-Hamilton, G.E. (1899) Extraordinary abundance of herrings in the Rivers Suir and Barrow. Irish Naturalist 8: 165. Brazier, B., Caffrey, J.M., Cross, T.F. and Chapman, D.V. (2012) A history of common carp Cyprinus carpio (L.) in Ireland: a review. Irish Fisheries Investigations No. 25. Briggs, R.P. and McCurdy, W.J. (1977) Marine fishes taken in waters off the north coast of 128 Irish Naturalists’ Journal Mind the Gap II


Ireland during 1977. Irish Naturalists’ Journal 19: 267-268. Browne, P. (1774) Monthly Chronologer for Ireland. A catalogue of fishes observed on our coasts, and in our lakes and rivers, classed and disposed according to Linnaeus. Exshaw’s Magazine (August 1774): 515-516. Caffrey, J.M., Acevedo, S., Gallagher, K. and Britton, R. (2008) Chub (Leuciscus cephalus): a new potentially invasive fish species in Ireland. Aquatic Invasions 3: 197-2005. Colby, E.E. (1837) Ordnance Survey of the County of Londonderry. 1. Memoir of the City and North Western Liberties of Londonderry, Parish of Templemore: Notices pp1-16. Published for Her Majesty’s Government, Hodges and Smith, Dublin 336pp. Coles, B.J. (2000) Doggerland: the cultural dynamics of a shifting coastline. In Pye, K. and Allen, J.R.L. (eds.) Coastal and Estuarine Environments: sedimentology, geomorphology and geoarchaeology: 393-401. Geological Society, London. Special Publications 175. Coscia, I., Rountree, V., King, J.J., Roche, W.K. and Mariani, S. (2010) A highly permeable species boundary between two anadromous fishes. Journal of Fish Biology 77: 1137-1149. Coscia, I., McDevitt, A.D., King, J.J., Roche, W.K., McLoughlin, C. and Mariani, S. (2013) A species to be? The genetic status and colonization history of the critically endangered Killarney Shad. Molecular Phylogenetics and Evolution 69: 1190-1195. Clark, C.D., Hughes, A.L.C., Greenwood, S.L., Jordan, C. and Sejrup, H.P. (2012) Pattern and timing of retreat of the last British-Irish Ice Sheet. Quaternary Science Reviews 44: 112146. Cross, T.F. (1996) Irish research on molecular genetics of Atlantic Salmon. Occasional Papers in Irish Science and Technology No. 13. Royal Dublin Society. Edwards, R. and Brooks, A. (2008) The island of Ireland: Drowning the myth of an Irish landbridge? In Davenport, J.J., Sleeman, D.P and Woodman, P.C. (eds) Mind the gap: postglacial colonisation of Ireland: 19-34. Irish Naturalists’ Journal, Belfast. Ensing, D. (2014) Pike (Esox lucius) could have been an exclusive human introduction to Ireland after all: a comment on Pedreschi et al. (2014). Journal of Biogeography DOI: 10.1111/jbi.12410 Eynaud, F., Malaize, B., Zaragosi, S., de Vernal, A., Scourse, J., Pujol, C., Cortijo, E., Grousset, F.E., Penaud, A., Toucanne, S., Turon, J-L. and Auffret, G. (2012) New constraints on European glacial freshwater releases to the North Atlantic Ocean. Geophysical Research

Letters 39: 1-6. Fahy, E. (1983) Feeding ecology of feral rainbow trout Salmo gairdneri Richardson in Mulroy Bay, an Irish Atlantic Sea Lough. Irish Naturalists’ Journal 21: 103-107. Fahy, E. (1985) Child of the Tides: a sea trout handbook. Glendale Press, Dun Laoghaire. Ferguson, A. (1981) Systematics of Irish Charr as indicated by electrophoretic analysis of tissue proteins. Biochemical Systematics and Ecology 9: 225-232. Fitzmaurice, P. (1984) The effects of freshwater fish introductions into Ireland. EIFAC Technical Paper 42 (Supplement 2): 449-457. Gallagher, C. (2002) The morphology and palaeohydrology of a submerged glaciofluvial channel emerging from Waterford Harbour onto the nearshore continental shelf of the Celtic Sea. Irish Geography 35: 111-132. Gibbard, P. (2007) Europe cut adrift. Nature 448: 259-260. Griffiths, D. (1997) The status of the Irish freshwater fish fauna: a review. Journal of Applied Ichthyology 13: 9-13. Harrison, A.J., Connor, L., Morrissey, E. and Kelly, F. (2012) Current status of pollan Coregonus autumnalis pollan in Lough Ree, Ireland. Biology and Environment: Proceedings of the Royal Irish Academy 112B: 1-9. Harvey, J.R. (1845) The Vertebrata – Pisces. In Cuvierian Society of Cork. Contributions towards a Fauna and Flora of the County of Cork. John Van Voorst, London and George Purcell, Cork. Hayden, B., Coscia, I. and Mariani, S. (2011) Low cytochrome b variation in bream Abramis brama. Journal of Fish Biology 78: 1579-1587. Hayden, T.J. (2002) The assembly of Ireland’s freshwater fish, amphibian, reptiles and mammals: tempo and mode. In Moriarty, C. and Murray, D. (eds) Biological Invaders: the impact of exotic species: 13-29. Royal Irish Academy, Dublin. Healy, A. (1956) Roach and dace in the Cork Blackwater. Department of Lands Report on the Sea and Inland Fisheries for the year 1956: 3-14. Department of Lands and Fisheries, Dublin. Holcik, J. (1991) Fish introductions in Europe with particular reference to its central and eastern part. Canadian Journal of Fisheries and Aquatic Science 48 (Supplement 1): 13-23. Holmes, J.M.C. (1994) Catalogue of the reserve collection of fish. Unpublished Manuscript, National Museum of Ireland, Dublin. Holt, E.W.L. (1911) Dace in Ireland. Irish Naturalist 20: 116. Humphreys, J. (1845) Capture of Acipenser huso. Annals and Magazine of Natural History 16: 213.

Igoe, F., O’Grady, M.F., Tierney, D. and Fitzmaurice, P. (2003) Arctic Char Salvelinus alpinus (L.) in Ireland – a millennium review of its distribution and status with conservation recommendations. Biology and Environment: Proceedings of the Royal Irish Academy 103B (1): 9-22. Igoe, F., Quigley, D.T.G., Marnell, F., Meskell, E., O’Connor, W. and Byrne, C. (2004) The sea lamprey Petromyzon marinus (L.), river lamprey Lampetra fluviatilis (L.) and brook lamprey Lampetra planeri (Bloch) in Ireland: general biology, ecology, distribution and status with recommendations for conservation. Biology and Environment: Proceedings of the Royal Irish Academy 104B: 43-56. Kennedy, M. and Fitzmaurice, P. (1969) Age and growth of thick-lipped grey mullet Crenimugil labrosus in Irish waters. Journal of the Marine Biological Association of the UK 49: 683-699. Kennedy, M. and Fitzmaurice, P. (1972a) Occurrence of Thin-Lipped Grey Mullet, Liza ramada (Risso) in County Mayo. Irish Naturalists’ Journal 17: 242-243. Kennedy, M. and Fitzmaurice, P. (1972b) Some aspects of the biology of gudgeon Gobio gobio (L.) in Irish waters. Journal of Fish Biology 4: 425-440. Kennedy, M. and Fitzmaurice, P. (1972c) The biology of the bass, Dicentrarchus labrax, in Irish waters. Journal of the Marine Biological Association of the UK 52: 557-597. Kennedy, M. and Fitzmaurice, P. (1974) Biology of the rudd Scardinius erythrophthalmus (L.) in Irish waters. Proceedings of the Royal Irish Academy 74B: 245-303. King, J.L., Marnell, F., Kingston, N., Rosell, R., Boylan, P., Caffrey, J.M., Fitzpatrick, U., Gargan, P.G., Kelly, F.L., O’Grady, M.F., Poole, R., Roche, W.K. and Cassidy, D. (2011) Ireland Red List No. 5: Amphibians, reptiles and freshwater fish. National Parks and Wildlife Service, Department of Arts, Heritage and the Gaeltacht, Dublin. Kottelat. M. and Freyhof, J. (2007) Handbook of European Freshwater Fishes. Kottelat, Cornol, Switzerland and Freyhof, Berlin, Germany. Lambeck, K. (1995) Late Devensian and Holocene shorelines of the British Isles and North Sea from models of glacio-hydroisostatic rebound. Journal of the Geological Society, London 152: 437-448. Maitland, P.S. and Campbell, R.N. (1992) Freshwater Fishes of the British Isles. HarperCollins, London. Maitland, P.S., Winfield, I.J., McCarthy, I.D. and Igoe, F. (2007) The status of Arctic charr Salvelinus alpinus in Britain and Ireland. Ecology of Freshwater Fish 16: 6-19. Irish Naturalists’ Journal Mind the Gap II 129

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also suggests that suitable freshwater habitats and other palaeo-environmental and ecological conditions must have existed for Pike to have become successfully established. Indeed, it seems reasonable to speculate that the same conditions may also have facilitated the natural colonisation of Ireland by other stenohaline fish, and particularly those species for which there is little or no documented evidence of humanmediated introductions e.g. Brook Lamprey, Common Bream, Gudgeon, Minnow, Rudd, Stone Loach, Nine-spined Stickleback and Perch. Although genetic studies may indicate when, where and whence stenohaline fish species may have colonised Irish waters, the elucidation of colonisation routes and mechanisms requires a multidisciplinary approach, involving, inter alia, various branches of palaeo and quaternary science. Acknowledgments I am grateful to the following for their assistance: Professor Michael A. Bell (Department of Ecology and Evolution, Stony Brook University, New York, USA); Dr Joe Caffrey (Inland Fisheries Ireland, Dublin); Dr Kim M. Cohen (Faculty of Geosciences, Utrecht University, Netherlands); Professor Peter Coxon (Department of Geography, Trinity College, Dublin); Professor Phillip L. Gibbard (Department of Geography, University of Cambridge, UK); Dr Sheila Hamilton-Dyer (SHD Archaeozoology, UK); Professor Stefano Mariani (School of Environment and Life Sciences, University of Salford, UK); Mr Peter McCluskey (West End House, Drogheda, Co Louth); Dr Gillian Scott (National University of Ireland, Department of Geography, Maynooth, Co. Kildare) and two anonymous referees. References Anon (1901) Special investigations and inspection. Report of the U.S. Commission of Fish and Fisheries. Part 26: 31. Auffret, G., Zaragosi, S., Dennielou, B., Cortijo, E., Van Rooij, D., Grousset, F., Pujol, C., Eynaud, F. and Siegert, M. (2002) Terrigenous fluxes at the Celtic margin during the last glacial cycle. Marine Geology 188: 79-108. Barrett-Hamilton, G.E. (1899) Extraordinary abundance of herrings in the Rivers Suir and Barrow. Irish Naturalist 8: 165. Brazier, B., Caffrey, J.M., Cross, T.F. and Chapman, D.V. (2012) A history of common carp Cyprinus carpio (L.) in Ireland: a review. Irish Fisheries Investigations No. 25. Briggs, R.P. and McCurdy, W.J. (1977) Marine fishes taken in waters off the north coast of 128 Irish Naturalists’ Journal Mind the Gap II


Ireland during 1977. Irish Naturalists’ Journal 19: 267-268. Browne, P. (1774) Monthly Chronologer for Ireland. A catalogue of fishes observed on our coasts, and in our lakes and rivers, classed and disposed according to Linnaeus. Exshaw’s Magazine (August 1774): 515-516. Caffrey, J.M., Acevedo, S., Gallagher, K. and Britton, R. (2008) Chub (Leuciscus cephalus): a new potentially invasive fish species in Ireland. Aquatic Invasions 3: 197-2005. Colby, E.E. (1837) Ordnance Survey of the County of Londonderry. 1. Memoir of the City and North Western Liberties of Londonderry, Parish of Templemore: Notices pp1-16. Published for Her Majesty’s Government, Hodges and Smith, Dublin 336pp. Coles, B.J. (2000) Doggerland: the cultural dynamics of a shifting coastline. In Pye, K. and Allen, J.R.L. (eds.) Coastal and Estuarine Environments: sedimentology, geomorphology and geoarchaeology: 393-401. Geological Society, London. Special Publications 175. Coscia, I., Rountree, V., King, J.J., Roche, W.K. and Mariani, S. (2010) A highly permeable species boundary between two anadromous fishes. Journal of Fish Biology 77: 1137-1149. Coscia, I., McDevitt, A.D., King, J.J., Roche, W.K., McLoughlin, C. and Mariani, S. (2013) A species to be? The genetic status and colonization history of the critically endangered Killarney Shad. Molecular Phylogenetics and Evolution 69: 1190-1195. Clark, C.D., Hughes, A.L.C., Greenwood, S.L., Jordan, C. and Sejrup, H.P. (2012) Pattern and timing of retreat of the last British-Irish Ice Sheet. Quaternary Science Reviews 44: 112146. Cross, T.F. (1996) Irish research on molecular genetics of Atlantic Salmon. Occasional Papers in Irish Science and Technology No. 13. Royal Dublin Society. Edwards, R. and Brooks, A. (2008) The island of Ireland: Drowning the myth of an Irish landbridge? In Davenport, J.J., Sleeman, D.P and Woodman, P.C. (eds) Mind the gap: postglacial colonisation of Ireland: 19-34. Irish Naturalists’ Journal, Belfast. Ensing, D. (2014) Pike (Esox lucius) could have been an exclusive human introduction to Ireland after all: a comment on Pedreschi et al. (2014). Journal of Biogeography DOI: 10.1111/jbi.12410 Eynaud, F., Malaize, B., Zaragosi, S., de Vernal, A., Scourse, J., Pujol, C., Cortijo, E., Grousset, F.E., Penaud, A., Toucanne, S., Turon, J-L. and Auffret, G. (2012) New constraints on European glacial freshwater releases to the North Atlantic Ocean. Geophysical Research

Letters 39: 1-6. Fahy, E. (1983) Feeding ecology of feral rainbow trout Salmo gairdneri Richardson in Mulroy Bay, an Irish Atlantic Sea Lough. Irish Naturalists’ Journal 21: 103-107. Fahy, E. (1985) Child of the Tides: a sea trout handbook. Glendale Press, Dun Laoghaire. Ferguson, A. (1981) Systematics of Irish Charr as indicated by electrophoretic analysis of tissue proteins. Biochemical Systematics and Ecology 9: 225-232. Fitzmaurice, P. (1984) The effects of freshwater fish introductions into Ireland. EIFAC Technical Paper 42 (Supplement 2): 449-457. Gallagher, C. (2002) The morphology and palaeohydrology of a submerged glaciofluvial channel emerging from Waterford Harbour onto the nearshore continental shelf of the Celtic Sea. Irish Geography 35: 111-132. Gibbard, P. (2007) Europe cut adrift. Nature 448: 259-260. Griffiths, D. (1997) The status of the Irish freshwater fish fauna: a review. Journal of Applied Ichthyology 13: 9-13. Harrison, A.J., Connor, L., Morrissey, E. and Kelly, F. (2012) Current status of pollan Coregonus autumnalis pollan in Lough Ree, Ireland. Biology and Environment: Proceedings of the Royal Irish Academy 112B: 1-9. Harvey, J.R. (1845) The Vertebrata – Pisces. In Cuvierian Society of Cork. Contributions towards a Fauna and Flora of the County of Cork. John Van Voorst, London and George Purcell, Cork. Hayden, B., Coscia, I. and Mariani, S. (2011) Low cytochrome b variation in bream Abramis brama. Journal of Fish Biology 78: 1579-1587. Hayden, T.J. (2002) The assembly of Ireland’s freshwater fish, amphibian, reptiles and mammals: tempo and mode. In Moriarty, C. and Murray, D. (eds) Biological Invaders: the impact of exotic species: 13-29. Royal Irish Academy, Dublin. Healy, A. (1956) Roach and dace in the Cork Blackwater. Department of Lands Report on the Sea and Inland Fisheries for the year 1956: 3-14. Department of Lands and Fisheries, Dublin. Holcik, J. (1991) Fish introductions in Europe with particular reference to its central and eastern part. Canadian Journal of Fisheries and Aquatic Science 48 (Supplement 1): 13-23. Holmes, J.M.C. (1994) Catalogue of the reserve collection of fish. Unpublished Manuscript, National Museum of Ireland, Dublin. Holt, E.W.L. (1911) Dace in Ireland. Irish Naturalist 20: 116. Humphreys, J. (1845) Capture of Acipenser huso. Annals and Magazine of Natural History 16: 213.

Igoe, F., O’Grady, M.F., Tierney, D. and Fitzmaurice, P. (2003) Arctic Char Salvelinus alpinus (L.) in Ireland – a millennium review of its distribution and status with conservation recommendations. Biology and Environment: Proceedings of the Royal Irish Academy 103B (1): 9-22. Igoe, F., Quigley, D.T.G., Marnell, F., Meskell, E., O’Connor, W. and Byrne, C. (2004) The sea lamprey Petromyzon marinus (L.), river lamprey Lampetra fluviatilis (L.) and brook lamprey Lampetra planeri (Bloch) in Ireland: general biology, ecology, distribution and status with recommendations for conservation. Biology and Environment: Proceedings of the Royal Irish Academy 104B: 43-56. Kennedy, M. and Fitzmaurice, P. (1969) Age and growth of thick-lipped grey mullet Crenimugil labrosus in Irish waters. Journal of the Marine Biological Association of the UK 49: 683-699. Kennedy, M. and Fitzmaurice, P. (1972a) Occurrence of Thin-Lipped Grey Mullet, Liza ramada (Risso) in County Mayo. Irish Naturalists’ Journal 17: 242-243. Kennedy, M. and Fitzmaurice, P. (1972b) Some aspects of the biology of gudgeon Gobio gobio (L.) in Irish waters. Journal of Fish Biology 4: 425-440. Kennedy, M. and Fitzmaurice, P. (1972c) The biology of the bass, Dicentrarchus labrax, in Irish waters. Journal of the Marine Biological Association of the UK 52: 557-597. Kennedy, M. and Fitzmaurice, P. (1974) Biology of the rudd Scardinius erythrophthalmus (L.) in Irish waters. Proceedings of the Royal Irish Academy 74B: 245-303. King, J.L., Marnell, F., Kingston, N., Rosell, R., Boylan, P., Caffrey, J.M., Fitzpatrick, U., Gargan, P.G., Kelly, F.L., O’Grady, M.F., Poole, R., Roche, W.K. and Cassidy, D. (2011) Ireland Red List No. 5: Amphibians, reptiles and freshwater fish. National Parks and Wildlife Service, Department of Arts, Heritage and the Gaeltacht, Dublin. Kottelat. M. and Freyhof, J. (2007) Handbook of European Freshwater Fishes. Kottelat, Cornol, Switzerland and Freyhof, Berlin, Germany. Lambeck, K. (1995) Late Devensian and Holocene shorelines of the British Isles and North Sea from models of glacio-hydroisostatic rebound. Journal of the Geological Society, London 152: 437-448. Maitland, P.S. and Campbell, R.N. (1992) Freshwater Fishes of the British Isles. HarperCollins, London. Maitland, P.S., Winfield, I.J., McCarthy, I.D. and Igoe, F. (2007) The status of Arctic charr Salvelinus alpinus in Britain and Ireland. Ecology of Freshwater Fish 16: 6-19. Irish Naturalists’ Journal Mind the Gap II 129

Quigley, D.

McCoy, F. (1841) On some new or rare fish occurring on the coast of Ireland. Annals and Magazine of Natural History 6: 402-408. McKeown, N.J., Hynes, R.A., Duguid, R.A., Ferguson, A., and Prodohl, P.A. (2010) Phylogeographic structure of brown trout Salmo trutta in Britain and Ireland: glacial refugia, postglacial colonization and origins of sympatric populations. Journal of Fish Biology 76: 319-347. M’Skimin, S. (1909) The History and Antiquities of the County of the Town of Carrickfergus from the earliest records till 1839. Also, a statistical survey of said county. New Edition. E.J. M’Crum (ed.), Belfast. Minchin, D. (2007) A checklist of alien and cryptogenic aquatic species in Ireland. Aquatic Invasions 2: 341-366. Mitchell, F. (1994) Where has Ireland come from? Country House, Dublin. Moriarty, C. (1988) The Eel in Ireland. Went Memorial Lecture 1987. Occasional Papers in Irish Science and Technology No. 4. Royal Dublin Society. Moriarty, C. and Fitzmaurice, P. (2000a) Origin and diversity of freshwater fishes in Ireland. Verhandlungen internatonale Vereinigung fur Theoretische und Angewandte Limnologie 27: 128-130. Moriarty, C. and Fitzmaurice, P. (2002b) The impact of the introduction of freshwater fishes to Ireland. In Moriarty, C. and Murray, D. (eds) Biological Invaders: the impact of exotic species:41-46. Royal Irish Academy, Dublin. O’Flaherty, R. (1684) A chorographical description of West or H-Iar Connaught. Hardiman, J. and Irish Archaeological Society (1978). Kenny’s Bookshops and Art Galleries, Galway. O’Neill, B., De Raedemaecker, F., McGrath, D. and Brophy, D. (2011) An experimental investigation of salinity effects on growth, development and condition in the European flounder (Platichthys flesus L.). Journal of Experimental Marine Biology and Ecology 410: 39-44. Ogilby, J.D. (1885) Notes on some Irish fishes. Scientific Proceedings of the Royal Dublin Society 4 (N.S.) (9): 510-535. Pedreschi, D., Kelly-Quinn, M., Caffrey, J., O’Grady, M. and Mariani, S. (2013) Genetic structure of pike (Esox lucius) reveals a complex and previously unrecognized colonization history of Ireland. Journal of Biogeography 41: 548-560. Quigley, D. (2011) Flounder Facts, Feats, Follies and Fallacies. Irish Angler’s Digest 13: 34-37. Quigley, D. (2012a) Ireland’s Mugil Empire – The Grey Ghosts. Irish Angler’s Digest 14: 3135. 130 Irish Naturalists’ Journal Mind the Gap II


Quigley, D. (2012b) Ireland’s Mugil Empire – A New Liza Life. Irish Angler’s Digest 14: 32-34. Quigley, D.T. (2014) Sturgeons (Family: Acipenseridae) in Irish & NW European Waters. Sherkin Comment 57: 10-11. Quigley, D. T. G. (1996) Status and conservation of euryhaline fish in Irish waters. In Costello, M.J., Mitchell, R. and Emblow, C.S. (eds) Strategies and methods in coastal and estuarine management. Aquatic Conservation: Marine and Freshwater Ecosystems 6: 313-318. Quigley, D.T.G. and Flannery, K. (1997a) Arctic Charr Salvelinus alpinus L.: First record from Lough Anscaul, Co Kerry; further records from Bunaveela, Kindrum and Coomasaharn; and notice of an introduction to Lough Owel, Co Westmeath. Irish Naturalists’ Journal 25: 435-439. Quigley, D.T.G. and Flannery, K. (1997b) Anchovy Engraulis encrasicolus (L.) (Pisces: Engraulidae) in Irish and UK Waters. Irish Naturalists’ Journal 25: 439-442. Quigley, D.T.G., Igoe, F. and O’Connor, W. (2004) The European smelt Osmerus eperlanus L. in Ireland: general biology, ecology, distribution and status with conservation recommendations. Biology and Environment: Proceedings of the Royal Irish Academy 104B: 57-66. Ravenel, W.de C. (1899) Report on the propagation and distribution of food-fishes, 1897-99 – steelhead and rainbow trout. Report of the U.S. Commission of Fish and Fisheries Part 25. Rutty, J. (1772) An Essay towards a Natural History of the County of Dublin. 1: 345-388. W. Sleater, Dublin. Sagan, C. (1983) Cosmos: The Story of Cosmic Evolution, Science and Civilisation. Futura, London. Sampson, G.V. (1802) Statistical survey of the county of Londonderry. Graisberry and Campbell, Dublin. Scharff, R.F. (1900) Attempted introduction of the American Shad into Irish waters. Irish Naturalist 9: 185. Scharff, R.F. (1905) Is the minnow a native of Ireland? Irish Naturalist 14: 225-228. Templeton, R. (1837) Irish Vertebrate Animals: selected from the Papers of John Templeton. Magazine of Natural History 1: 403-413. Thompson, W. (1841) Additions to the fauna of Ireland. Annals and Magazine of Natural History 7: 477-482. Thompson, W. (1856) The natural history of Ireland 4. Henry G. Bohn, London. Toucanne, S., Zaragosi, S., Bourillet, J.F., Naughton, F., Cremer, M., Eynaud, F. and Dennielou, B. (2008) Activity of the turbidite

levees of the Celtic-Armoricanm margin (Bay of Biscay) during the last 30,000 years: Imprints of the last European deglaciation and Heinrich events. Marine Geology 247: 84-103. Toucanne, S., Zaragosi, S., Bourillet, J.F., Marieu, V., Cremer, M., Kageyama, M., Van Vliet-Lanoe, B., Eynaud, F., Turon, J-L. and Gibbard, P.L. (2010) The first estimation of Fleuve Manche palaeoriver discharge during the last deglaciation: Evidence for Fennoscandian ice sheet melt water flow in the English Channel ca 20-18 ka ago. Earth and Planetary Science Letters 290: 459-473. Twomey, E. and Byrne, P. (1985) A new record for the Tarpon, Tarpon atlanticus Valenciennes (Osteichthyes-Elopiformes-Elopidae), in the eastern North Atlantic. Journal of Fish Biology 26: 359-362. Van Nieuwenhove, N., Bauch, H.A. and Matthiessen, J. (2008) Last interglacial surface water conditions in the eastern Nordic Seas inferred from dinocyst and foraminiferal assemblages. Marine Micropaleontology 66: 247-263. Vilks, G. (1981) Late glacial-postglacial foraminiferal boundary in sediments of eastern Canada, Denmark and Norway. Geoscience Canada 8: 48-55. Walsh, A.R. (1993) The occurrence of Liza species (Teleostei: Mugilidae) in Irish coastal waters. Irish Naturalists’ Journal 25: 255-256. Warren, R. (1898) Extraordinary run of herrings in the Moy estuary. Irish Naturalist 7: 19. Welcomme R.L. (1988) International introductions of inland aquatic species. FAO Fisheries Technical Paper 294: 1-318. Weninger, B., Schulting, R., Bradtmoller, M., Clare, L., Collard, M., Edinborough, K., Hilpert, J., Joris, O., Niekus, M., Rohling, E.J.

and Wagner, B. (2008) The catastrophic final flooding of Doggerland by the Storegga Slide tsunami. Documenta Praehistorica 35: 1-24. Went, A.E.J. (1946) Irish Freshwater Fish – some notes on their introduction. Salmon and Trout Magazine 118: 248-256. Went. A.E.J. (1948) The status of the sturgeon Acipenser sturio L. in Irish waters now and in former days. Irish Naturalists’ Journal 9: 172174. Went, A.E.J. (1949) Giraldus Cambrensis’ notes on Irish fish. Irish Naturalists’ Journal 9: 221224. Went, A.E.J. (1950) Notes on the introduction of some freshwater fish into Ireland. Journal of the Department of Agriculture 47: 3-7. Went, A.E.J. (1957) The pike in Ireland. Irish Naturalists’ Journal 12: 177-182. Went. A.E.J. (1974) Some interesting fishes taken from Irish waters in 1973. Irish Naturalists’ Journal 18: 57-65. Went, A.E.J. (1978) The zoogeography of some fishes in Irish waters. Fishery Leaflet No. 93. Went, A.E.J. (1980) Freshwater Fishes. In Kernan, R.P., Mooney O.V. and Went, A.E.J. (eds) The Introduction of Exotic Species – Advantages and Problems. Royal Irish Academy, Dublin. Went, A.E.J. and Kennedy, M. (1976) List of Irish Fishes (3rd Edition). Stationery Office, Dublin Wheeler, A. (1977) The origin and distribution of the freshwater fishes of the British Isles. Journal of Biogeography 4: 1-24. Wilson, J.P.F. (1986) The record from fish, amphibians and reptiles. In Sleeman, D.P., Devoy, R.J. and Woodman, P.C. (eds) Proceedings of the Post Glacial Colonization Conference (UCC, 15-16 October 1983). Occasional Publication of the Irish Biogeographical Society 1: 53-58.

Irish Naturalists’ Journal Mind the Gap II 131

Quigley, D.

McCoy, F. (1841) On some new or rare fish occurring on the coast of Ireland. Annals and Magazine of Natural History 6: 402-408. McKeown, N.J., Hynes, R.A., Duguid, R.A., Ferguson, A., and Prodohl, P.A. (2010) Phylogeographic structure of brown trout Salmo trutta in Britain and Ireland: glacial refugia, postglacial colonization and origins of sympatric populations. Journal of Fish Biology 76: 319-347. M’Skimin, S. (1909) The History and Antiquities of the County of the Town of Carrickfergus from the earliest records till 1839. Also, a statistical survey of said county. New Edition. E.J. M’Crum (ed.), Belfast. Minchin, D. (2007) A checklist of alien and cryptogenic aquatic species in Ireland. Aquatic Invasions 2: 341-366. Mitchell, F. (1994) Where has Ireland come from? Country House, Dublin. Moriarty, C. (1988) The Eel in Ireland. Went Memorial Lecture 1987. Occasional Papers in Irish Science and Technology No. 4. Royal Dublin Society. Moriarty, C. and Fitzmaurice, P. (2000a) Origin and diversity of freshwater fishes in Ireland. Verhandlungen internatonale Vereinigung fur Theoretische und Angewandte Limnologie 27: 128-130. Moriarty, C. and Fitzmaurice, P. (2002b) The impact of the introduction of freshwater fishes to Ireland. In Moriarty, C. and Murray, D. (eds) Biological Invaders: the impact of exotic species:41-46. Royal Irish Academy, Dublin. O’Flaherty, R. (1684) A chorographical description of West or H-Iar Connaught. Hardiman, J. and Irish Archaeological Society (1978). Kenny’s Bookshops and Art Galleries, Galway. O’Neill, B., De Raedemaecker, F., McGrath, D. and Brophy, D. (2011) An experimental investigation of salinity effects on growth, development and condition in the European flounder (Platichthys flesus L.). Journal of Experimental Marine Biology and Ecology 410: 39-44. Ogilby, J.D. (1885) Notes on some Irish fishes. Scientific Proceedings of the Royal Dublin Society 4 (N.S.) (9): 510-535. Pedreschi, D., Kelly-Quinn, M., Caffrey, J., O’Grady, M. and Mariani, S. (2013) Genetic structure of pike (Esox lucius) reveals a complex and previously unrecognized colonization history of Ireland. Journal of Biogeography 41: 548-560. Quigley, D. (2011) Flounder Facts, Feats, Follies and Fallacies. Irish Angler’s Digest 13: 34-37. Quigley, D. (2012a) Ireland’s Mugil Empire – The Grey Ghosts. Irish Angler’s Digest 14: 3135. 130 Irish Naturalists’ Journal Mind the Gap II


Quigley, D. (2012b) Ireland’s Mugil Empire – A New Liza Life. Irish Angler’s Digest 14: 32-34. Quigley, D.T. (2014) Sturgeons (Family: Acipenseridae) in Irish & NW European Waters. Sherkin Comment 57: 10-11. Quigley, D. T. G. (1996) Status and conservation of euryhaline fish in Irish waters. In Costello, M.J., Mitchell, R. and Emblow, C.S. (eds) Strategies and methods in coastal and estuarine management. Aquatic Conservation: Marine and Freshwater Ecosystems 6: 313-318. Quigley, D.T.G. and Flannery, K. (1997a) Arctic Charr Salvelinus alpinus L.: First record from Lough Anscaul, Co Kerry; further records from Bunaveela, Kindrum and Coomasaharn; and notice of an introduction to Lough Owel, Co Westmeath. Irish Naturalists’ Journal 25: 435-439. Quigley, D.T.G. and Flannery, K. (1997b) Anchovy Engraulis encrasicolus (L.) (Pisces: Engraulidae) in Irish and UK Waters. Irish Naturalists’ Journal 25: 439-442. Quigley, D.T.G., Igoe, F. and O’Connor, W. (2004) The European smelt Osmerus eperlanus L. in Ireland: general biology, ecology, distribution and status with conservation recommendations. Biology and Environment: Proceedings of the Royal Irish Academy 104B: 57-66. Ravenel, W.de C. (1899) Report on the propagation and distribution of food-fishes, 1897-99 – steelhead and rainbow trout. Report of the U.S. Commission of Fish and Fisheries Part 25. Rutty, J. (1772) An Essay towards a Natural History of the County of Dublin. 1: 345-388. W. Sleater, Dublin. Sagan, C. (1983) Cosmos: The Story of Cosmic Evolution, Science and Civilisation. Futura, London. Sampson, G.V. (1802) Statistical survey of the county of Londonderry. Graisberry and Campbell, Dublin. Scharff, R.F. (1900) Attempted introduction of the American Shad into Irish waters. Irish Naturalist 9: 185. Scharff, R.F. (1905) Is the minnow a native of Ireland? Irish Naturalist 14: 225-228. Templeton, R. (1837) Irish Vertebrate Animals: selected from the Papers of John Templeton. Magazine of Natural History 1: 403-413. Thompson, W. (1841) Additions to the fauna of Ireland. Annals and Magazine of Natural History 7: 477-482. Thompson, W. (1856) The natural history of Ireland 4. Henry G. Bohn, London. Toucanne, S., Zaragosi, S., Bourillet, J.F., Naughton, F., Cremer, M., Eynaud, F. and Dennielou, B. (2008) Activity of the turbidite

levees of the Celtic-Armoricanm margin (Bay of Biscay) during the last 30,000 years: Imprints of the last European deglaciation and Heinrich events. Marine Geology 247: 84-103. Toucanne, S., Zaragosi, S., Bourillet, J.F., Marieu, V., Cremer, M., Kageyama, M., Van Vliet-Lanoe, B., Eynaud, F., Turon, J-L. and Gibbard, P.L. (2010) The first estimation of Fleuve Manche palaeoriver discharge during the last deglaciation: Evidence for Fennoscandian ice sheet melt water flow in the English Channel ca 20-18 ka ago. Earth and Planetary Science Letters 290: 459-473. Twomey, E. and Byrne, P. (1985) A new record for the Tarpon, Tarpon atlanticus Valenciennes (Osteichthyes-Elopiformes-Elopidae), in the eastern North Atlantic. Journal of Fish Biology 26: 359-362. Van Nieuwenhove, N., Bauch, H.A. and Matthiessen, J. (2008) Last interglacial surface water conditions in the eastern Nordic Seas inferred from dinocyst and foraminiferal assemblages. Marine Micropaleontology 66: 247-263. Vilks, G. (1981) Late glacial-postglacial foraminiferal boundary in sediments of eastern Canada, Denmark and Norway. Geoscience Canada 8: 48-55. Walsh, A.R. (1993) The occurrence of Liza species (Teleostei: Mugilidae) in Irish coastal waters. Irish Naturalists’ Journal 25: 255-256. Warren, R. (1898) Extraordinary run of herrings in the Moy estuary. Irish Naturalist 7: 19. Welcomme R.L. (1988) International introductions of inland aquatic species. FAO Fisheries Technical Paper 294: 1-318. Weninger, B., Schulting, R., Bradtmoller, M., Clare, L., Collard, M., Edinborough, K., Hilpert, J., Joris, O., Niekus, M., Rohling, E.J.

and Wagner, B. (2008) The catastrophic final flooding of Doggerland by the Storegga Slide tsunami. Documenta Praehistorica 35: 1-24. Went, A.E.J. (1946) Irish Freshwater Fish – some notes on their introduction. Salmon and Trout Magazine 118: 248-256. Went. A.E.J. (1948) The status of the sturgeon Acipenser sturio L. in Irish waters now and in former days. Irish Naturalists’ Journal 9: 172174. Went, A.E.J. (1949) Giraldus Cambrensis’ notes on Irish fish. Irish Naturalists’ Journal 9: 221224. Went, A.E.J. (1950) Notes on the introduction of some freshwater fish into Ireland. Journal of the Department of Agriculture 47: 3-7. Went, A.E.J. (1957) The pike in Ireland. Irish Naturalists’ Journal 12: 177-182. Went. A.E.J. (1974) Some interesting fishes taken from Irish waters in 1973. Irish Naturalists’ Journal 18: 57-65. Went, A.E.J. (1978) The zoogeography of some fishes in Irish waters. Fishery Leaflet No. 93. Went, A.E.J. (1980) Freshwater Fishes. In Kernan, R.P., Mooney O.V. and Went, A.E.J. (eds) The Introduction of Exotic Species – Advantages and Problems. Royal Irish Academy, Dublin. Went, A.E.J. and Kennedy, M. (1976) List of Irish Fishes (3rd Edition). Stationery Office, Dublin Wheeler, A. (1977) The origin and distribution of the freshwater fishes of the British Isles. Journal of Biogeography 4: 1-24. Wilson, J.P.F. (1986) The record from fish, amphibians and reptiles. In Sleeman, D.P., Devoy, R.J. and Woodman, P.C. (eds) Proceedings of the Post Glacial Colonization Conference (UCC, 15-16 October 1983). Occasional Publication of the Irish Biogeographical Society 1: 53-58.


132 Irish Naturalists’ Journal Mind the Gap II stenohaline? anadromous/ euryhaline euryhaline

Alosa killarnensis Regan, 1916 Alosa sapidissima (Wilson, 1811) Clupea harengus L.

Killarney Shad American Shad Atlantic Herring

Engraulis encrasicolus (L.)

euryhaline

vagrant

native

introduced

native

native

native

native

vagrant

native/vagrant?

native/vagrant?

native/introduced?

native

native

Biogeographical Origin

vagrant

established

not established

established

established

established

established

not established

extinct

extinct?

established

established

established

Current Status

post-glacial

post glacial

1899

post glacial

post glacial

post glacial

post glacial

1981

c.1845

post glacial

unknown

post glacial

post glacial

Date of Introduction/ Colonisation

Quigley and Flannery (1997b)

Warren (1898); Barrett-Hamilton (1899)

Scharff (1900); Anon (1901)

Coscia et al. (2013)



Moriarty (1988)

Twomey and Byrne (1985)

Humphreys (1845)

Rutty (1772); Browne (1774); Sampson (1802); Templeton (1837); Harvey (1845); Ogilby (1885); Went (1948); Quigley (2014)

Igoe et al. (2004)

Igoe et al. (2004)

Igoe et al. (2004)

References

stenohaline

stenohaline

Phoxinus phoxinus (L.) Minnow

stenohaline

Gobio gobio (L.) Gudgeon

Leuciscus leuciscus (L.)

stenohaline

Cyprinus carpio L. Common Carp

Dace

stenohaline

Carassius carassius (L.) Crucian Carp

stenohaline

stenohaline

Carassius auratus (L.) Goldfish

Leuciscus cephalus (L.)

stenohaline

Barbus barbus (L.) Barbel

Chub

stenohaline

Abramis brama (L.) Common Bream

Salinity Tolerance

Cyprinidae

Scientific Name

Common Name

Family Name

native/introduced?

introduced

introduced

native/introduced?

introduced

introduced?

introduced?

introduced/ erroreous record?

native/introduced ?


established

established

established?

established

established

established?

established?

not established

established

Current Status

pre 1856

c.1873

2004

pre 1774

pre 1634

unknown

pre 1856

pre 1774

pre 1774


Thompson (1856); Ogilby (1885); Scharff (1905); Went (1949, 1978)

Holt (1911); Went (1946, 1950, 1978); Healy (1956); Holcik (1991)

Browne (1774); Thompson (1856); Went (1949); Caffrey et al. (2008)

Browne (1774); Templeton (1837); Thompson (1856); Ogilby (1885); Went (1949, 1978); Kennedy and Fitzmaurice (1972b)

Rutty (1772); Browne (1774); Templeton (1837); Thompson (1856); Holcik (1991); Brazier et al. (2012)

J. Caffrey (pers. comm.)

Thompson (1856); Ogilby (1885)

Browne (1774); Thompson (1856); Went (1949)

O'Flaherty (1684); Browne (1774); Templeton (1837); Thompson (1856); Went (1946, 1978); Hayden et al. (2011)

References

Appendix 1. Table 1. (cont.) List of fish species reported from Irish freshwater habitats along with details on salinity tolerance, biogeographical origin, current status, and date of introduction and/or natural colonisation.

Anchovy

anadromous/ euryhaline

Alosa fallax (Lacepede, 1803)

Twaite Shad

Engraulidae


Alosa alosa (L.)

Allis Shad

catadromous/ euryhaline

Clupidae

Anguilla anguilla (L.)

Eel

euryhaline

Anguillidae

Megalops atlanticus Valenciennes, 1847

Tarpon


Huso huso (L.)

Beluga Megalopidae


Acipenser sturio L.

stenohaline

Lampetra planeri (Bloch, 1784)

Brook Lamprey European Atlantic Sturgeon


Lampetra fluviatilis (L.)

River Lamprey (Lampern)

Acipenseridae


Petromyzon marinus L.

Sea Lamprey

Salinity Tolerance

Petromyzonidae

Scientific Name

Common Name

Family Name

Appendix 1. Table 1. List of fish species reported from Irish freshwater habitats along with details on salinity tolerance, biogeographical origin, current status, and date of introduction and/or natural colonisation.

Quigley, D. The facilitated natural colonisation of Ireland by stenohaline fishes


132 Irish Naturalists’ Journal Mind the Gap II stenohaline? anadromous/ euryhaline euryhaline

Alosa killarnensis Regan, 1916 Alosa sapidissima (Wilson, 1811) Clupea harengus L.

Killarney Shad American Shad Atlantic Herring

Engraulis encrasicolus (L.)

euryhaline

vagrant

native

introduced

native

native

native

native

vagrant

native/vagrant?

native/vagrant?

native/introduced?

native

native


vagrant

established

not established

established

established

established

established

not established

extinct

extinct?

established

established

established

Current Status

post-glacial

post glacial

1899

post glacial

post glacial

post glacial

post glacial

1981

c.1845

post glacial

unknown

post glacial

post glacial


Quigley and Flannery (1997b)

Warren (1898); Barrett-Hamilton (1899)

Scharff (1900); Anon (1901)




Moriarty (1988)

Twomey and Byrne (1985)

Humphreys (1845)

Rutty (1772); Browne (1774); Sampson (1802); Templeton (1837); Harvey (1845); Ogilby (1885); Went (1948); Quigley (2014)

Igoe et al. (2004)

Igoe et al. (2004)

Igoe et al. (2004)

References

stenohaline

stenohaline

Phoxinus phoxinus (L.) Minnow

stenohaline

Gobio gobio (L.) Gudgeon

Leuciscus leuciscus (L.)

stenohaline

Cyprinus carpio L. Common Carp

Dace

stenohaline

Carassius carassius (L.) Crucian Carp

stenohaline

stenohaline

Carassius auratus (L.) Goldfish

Leuciscus cephalus (L.)

stenohaline

Barbus barbus (L.) Barbel

Chub

stenohaline

Abramis brama (L.) Common Bream

Salinity Tolerance

Cyprinidae

Scientific Name

Common Name

Family Name

native/introduced?

introduced

introduced

native/introduced?

introduced

introduced?

introduced?

introduced/ erroreous record?

native/introduced ?


established

established

established?

established

established

established?

established?

not established

established

Current Status

pre 1856

c.1873

2004

pre 1774

pre 1634

unknown

pre 1856

pre 1774

pre 1774


Thompson (1856); Ogilby (1885); Scharff (1905); Went (1949, 1978)

Holt (1911); Went (1946, 1950, 1978); Healy (1956); Holcik (1991)

Browne (1774); Thompson (1856); Went (1949); Caffrey et al. (2008)

Browne (1774); Templeton (1837); Thompson (1856); Ogilby (1885); Went (1949, 1978); Kennedy and Fitzmaurice (1972b)

Rutty (1772); Browne (1774); Templeton (1837); Thompson (1856); Holcik (1991); Brazier et al. (2012)

J. Caffrey (pers. comm.)

Thompson (1856); Ogilby (1885)

Browne (1774); Thompson (1856); Went (1949)

O'Flaherty (1684); Browne (1774); Templeton (1837); Thompson (1856); Went (1946, 1978); Hayden et al. (2011)

References


Anchovy


Alosa fallax (Lacepede, 1803)

Twaite Shad

Engraulidae


Alosa alosa (L.)

Allis Shad

catadromous/ euryhaline

Clupidae

Anguilla anguilla (L.)

Eel

euryhaline

Anguillidae

Megalops atlanticus Valenciennes, 1847

Tarpon


Huso huso (L.)

Beluga Megalopidae


Acipenser sturio L.

stenohaline

Lampetra planeri (Bloch, 1784)

Brook Lamprey European Atlantic Sturgeon


Lampetra fluviatilis (L.)

River Lamprey (Lampern)

Acipenseridae


Petromyzon marinus L.

Sea Lamprey

Salinity Tolerance

Petromyzonidae

Scientific Name

Common Name

Family Name

Appendix 1. Table 1. List of fish species reported from Irish freshwater habitats along with details on salinity tolerance, biogeographical origin, current status, and date of introduction and/or natural colonisation.



134 Irish Naturalists’ Journal Mind the Gap II stenohaline

stenohaline

Ameiurus melas (Rafinesque, 1820)

Ameiurus nebulosus (LeSueur, 1819)

Black Bullhead

Brown Bullhead

introduced/ erroneous record?

introduced

introduced

native/introduced?

discarded aquarium fish


introduced

native/introduced?

introduced


established?

established?

established?

established

not established

not established

established

established

established

Current Status

unknown

pre 1984

c. 1889

pre 1772

1988

2008

pre 1634

pre 1772

1889


Welcomme (1988); Holcik (1991)

Fitzmaurice (1984); Welcomme (1988); Holcik (1991); Minchin (2007)

Holmes (1994)

Rutty (1772); Browne (1774); Templeton (1837); Thompson (1856); Ogilby (1885); Went (1946, 1949, 1978)

Minchin (2007)

P. McCluskey (pers. comm.)

Rutty (1772); Templeton (1837); Harvey (1845); Thompson (1856); Holcik (1991); Brazier et al. (2012)

O'Flaherty (1684); Rutty (1772); Templeton (1837); Harvey (1845); Thompson (1856); Ogilby (1885); Went (1946, 1978); Kennedy and Fitzmaurice (1974)

Browne (1774); Went (1946, 1949, 1950, 1978); Holcik (1991)

References

stenohaline


anadromous/ euryhaline euryhaline

anadromous/ euryhaline anadromous/ euryhaline

Hucho hucho (L.)

Oncorhynchus gorbuscha (Walbaum, 1792) Oncorhynchus tshawytscha (Walbaum, 1792) Oncorhynchus mykiss (Walbaum, 1792) Salmo trutta L. Salmo salar L.

Pike

Smelt (Sparling) Pollan (Arctic Cisco) Huchen

Pink Salmon

Chinook Salmon

Rainbow Trout

Sea Trout/Brown Trout Atlantic Salmon

Osmeridae Coregonidae Salmonidae

Coregonus pollan Thompson, 1835

Osmerus eperlanus (L.)

Esox lucius L.

stenohaline


stenohaline

stenohaline

Esocidae

Silurus glanis L. Wels Catfish

Salinity Tolerance

Siluridae

Scientific Name

Common Name

Family Name

native and introduced


introduced

vagrant/erroneous record?

vagrant

introduced

native

native

native/introduced



established

widespread

established

not established

not established

not established

established

established

established

not established

Current Status

post glacial and introduced


1899

unknown

1973

1993

post glacial

post glacial

post glacial

c.1837?


Cross (1996)

Fahy (1985); McKeown et al. (2010)

Ravenel (1889); Went (1946); Fahy (1983); Minchin (2007)


Went (1974); Moriarty and Fitzmaurice (2002a, 2002b); Minchin (2007)

Moriarty and Fitzmaurice (2002a, 2002b); Minchin (2007)

Ogilby (1885); Harrison et al. (2011)

Quigley et al. (2004)

O'Flaherty (1684); Rutty (1772); Browne (1774); Sampson (1802); Colby (1837); Templeton (1837); Harvey (1845); Thompson (1856); Went (1949, 1957); Holcik (1991); Pedreschi et al. (2013)

Thompson (1841, 1856)

References


stenohaline

Ameiurus catus (L.)

White Catfish

stenohaline

Ictaluridae

Barbatula barbatula (L.)

Stone Loach

stenohaline

Serrasalmus sp.

Piranha sp. Nemacheilidae

stenohaline

Colossoma sp.

stenohaline

Tinca tinca (L.)

Tench

Pacu sp.

stenohaline

Scardinius erythrophthalmus (L.)

Rudd

Characidae

stenohaline

Rutilus rutilus (L.)

Roach

Salinity Tolerance

Cyprinidae

Scientific Name

Common Name

Family Name




134 Irish Naturalists’ Journal Mind the Gap II stenohaline

stenohaline

Ameiurus melas (Rafinesque, 1820)

Ameiurus nebulosus (LeSueur, 1819)

Black Bullhead

Brown Bullhead


introduced

introduced

native/introduced?



introduced

native/introduced?

introduced


established?

established?

established?

established

not established

not established

established

established

established

Current Status

unknown

pre 1984

c. 1889

pre 1772

1988

2008

pre 1634

pre 1772

1889



Fitzmaurice (1984); Welcomme (1988); Holcik (1991); Minchin (2007)

Holmes (1994)

Rutty (1772); Browne (1774); Templeton (1837); Thompson (1856); Ogilby (1885); Went (1946, 1949, 1978)

Minchin (2007)

P. McCluskey (pers. comm.)

Rutty (1772); Templeton (1837); Harvey (1845); Thompson (1856); Holcik (1991); Brazier et al. (2012)

O'Flaherty (1684); Rutty (1772); Templeton (1837); Harvey (1845); Thompson (1856); Ogilby (1885); Went (1946, 1978); Kennedy and Fitzmaurice (1974)

Browne (1774); Went (1946, 1949, 1950, 1978); Holcik (1991)

References

stenohaline


anadromous/ euryhaline euryhaline

anadromous/ euryhaline anadromous/ euryhaline

Hucho hucho (L.)

Oncorhynchus gorbuscha (Walbaum, 1792) Oncorhynchus tshawytscha (Walbaum, 1792) Oncorhynchus mykiss (Walbaum, 1792) Salmo trutta L. Salmo salar L.

Pike

Smelt (Sparling) Pollan (Arctic Cisco) Huchen

Pink Salmon

Chinook Salmon

Rainbow Trout

Sea Trout/Brown Trout Atlantic Salmon

Osmeridae Coregonidae Salmonidae

Coregonus pollan Thompson, 1835

Osmerus eperlanus (L.)

Esox lucius L.

stenohaline


stenohaline

stenohaline

Esocidae

Silurus glanis L. Wels Catfish

Salinity Tolerance

Siluridae

Scientific Name

Common Name

Family Name



introduced


vagrant

introduced

native

native

native/introduced



established

widespread

established

not established

not established

not established

established

established

established

not established

Current Status



1899

unknown

1973

1993

post glacial

post glacial

post glacial

c.1837?


Cross (1996)

Fahy (1985); McKeown et al. (2010)

Ravenel (1889); Went (1946); Fahy (1983); Minchin (2007)


Went (1974); Moriarty and Fitzmaurice (2002a, 2002b); Minchin (2007)

Moriarty and Fitzmaurice (2002a, 2002b); Minchin (2007)

Ogilby (1885); Harrison et al. (2011)

Quigley et al. (2004)

O'Flaherty (1684); Rutty (1772); Browne (1774); Sampson (1802); Colby (1837); Templeton (1837); Harvey (1845); Thompson (1856); Went (1949, 1957); Holcik (1991); Pedreschi et al. (2013)

Thompson (1841, 1856)

References


stenohaline

Ameiurus catus (L.)

White Catfish

stenohaline

Ictaluridae

Barbatula barbatula (L.)

Stone Loach

stenohaline

Serrasalmus sp.

Piranha sp. Nemacheilidae

stenohaline

Colossoma sp.

stenohaline

Tinca tinca (L.)

Tench

Pacu sp.

stenohaline

Scardinius erythrophthalmus (L.)

Rudd

Characidae

stenohaline

Rutilus rutilus (L.)

Roach

Salinity Tolerance

Cyprinidae

Scientific Name

Common Name

Family Name




136 Irish Naturalists’ Journal Mind the Gap II Gasteroseus aculeatus L.

Atherina boyeri Risso, 1810

Labrus bergylta Ascanius, 1767

Thymallus thymallus (L.)

euryhaline

euryhaline?

euryhaline

stenohaline

native

native

native


native/recent immigrant?


native




established

established

established

not established

established

established

established

not established

established

Current Status

post glacial

post glacial

post glacial

pre 1774?

pre 1971

pre 1992

post glacial

pre 1772?



Rutty (1772); Browne (1774); Sampson (1802); Templeton (1837); Harvey (1845); Ogilby (1885); Went (1946, 1978)

Went and Kennedy (1976)

Briggs and McCurdy (1977)

Browne (1774); Sampson (1802); M'Skimin (1909); Colby (1837); Templeton (1837)

McCoy (1841); Harvey (1845); Thompson (1856); Ogilby (1885); Kennedy and Fitzmaurice (1972a); Walsh (1993); Quigley (2012b)

Walsh (1993); Quigley (2012b)

Kennedy and Fitzmaurice (1969); Quigley (2012a)

Rutty (1772); Browne (1774); Thompson (1856); Went (1949)

Ferguson (1981); Quigley and Flannery (1997a); Igoe et al. (2003); Maitland et al. (2007)

References

Flounder

Zander (Pike-Perch)

Pleuronectidae

European Perch

Percidae

Transparent Goby

euryhaline

Sander lucioperca (L.)

European Bass

Moronidae

Gobiidae

stenohaline

Perca fluviatilis L.

Common Bullhead

Pleuronectes flesus L.

Aphia minuta (Risso, 1810)

Dicentrarchus labrax (L.)

Cottus gobio L.

euryhaline

euryhaline

euryhaline

stenohaline

stenohaline

Cottidae

Pungitus pungitus (L.) Nine-spined Stickleback

Salinity Tolerance

Gasterosteidae

Scientific Name

Common Name

Family Name

native

native


native/introduced?

native


native/introduced?


established

established

not established

established

established

not established

established

Current Status

post glacial

post glacial

pre 1774?

pre 1772

post glacial

pre 1774?

pre 1772


Rutty (1772); O'Neill et al. (2011); Quigley (2011)

Quigley (1996)

Browne (1774); Brown et al. (2001)

Rutty (1772); Browne (1774); Sampson (1802); Templeton (1837); Thompson (1856); Harvey (1845); Went (1946, 1949, 1978)

Kennedy and Fitzmaurice (1972c)

Browne (1774); Sampson (1802); M'Skimin (1909); Thompson (1856); Ogilby (1885); Went (1949)

Rutty (1772); Browne (1774); Templeton (1837); Thompson (1841); Harvey (1845); Went (1946, 1978); Hayden (2002)

References


Three-spined Stickleback

Gasterosteidae

Flathead Grey Mullet

Big-scale Sand Smelt

euryhaline

Mugil cephalus L.

Thin-lipped Grey Mullet

Atherinidae

euryhaline

Liza ramada (Risso, 1826)

Golden Grey Mullet

Ballan Wrasse

euryhaline

Liza aurata (Risso, 1810)

Thick-lipped Grey Mullet

Mugilidae

Labridae

euryhaline

Chelon labrosus (Risso, 1826)

Grayling

stenohaline

Thymallidae

Salvelinus alpinus (L.)

Arctic Charr

Salinity Tolerance

Salmonidae

Scientific Name

Common Name

Family Name




136 Irish Naturalists’ Journal Mind the Gap II Gasteroseus aculeatus L.

Atherina boyeri Risso, 1810

Labrus bergylta Ascanius, 1767

Thymallus thymallus (L.)

euryhaline

euryhaline?

euryhaline

stenohaline

native

native

native




native




established

established

established

not established

established

established

established

not established

established

Current Status

post glacial

post glacial

post glacial

pre 1774?

pre 1971

pre 1992

post glacial

pre 1772?



Rutty (1772); Browne (1774); Sampson (1802); Templeton (1837); Harvey (1845); Ogilby (1885); Went (1946, 1978)

Went and Kennedy (1976)

Briggs and McCurdy (1977)

Browne (1774); Sampson (1802); M'Skimin (1909); Colby (1837); Templeton (1837)

McCoy (1841); Harvey (1845); Thompson (1856); Ogilby (1885); Kennedy and Fitzmaurice (1972a); Walsh (1993); Quigley (2012b)

Walsh (1993); Quigley (2012b)

Kennedy and Fitzmaurice (1969); Quigley (2012a)

Rutty (1772); Browne (1774); Thompson (1856); Went (1949)

Ferguson (1981); Quigley and Flannery (1997a); Igoe et al. (2003); Maitland et al. (2007)

References

Flounder

Zander (Pike-Perch)

Pleuronectidae

European Perch

Percidae

Transparent Goby

euryhaline

Sander lucioperca (L.)

European Bass

Moronidae

Gobiidae

stenohaline

Perca fluviatilis L.

Common Bullhead

Pleuronectes flesus L.

Aphia minuta (Risso, 1810)

Dicentrarchus labrax (L.)

Cottus gobio L.

euryhaline

euryhaline

euryhaline

stenohaline

stenohaline

Cottidae

Pungitus pungitus (L.) Nine-spined Stickleback

Salinity Tolerance

Gasterosteidae

Scientific Name

Common Name

Family Name

native

native


native/introduced?

native


native/introduced?


established

established

not established

established

established

not established

established

Current Status

post glacial

post glacial

pre 1774?

pre 1772

post glacial

pre 1774?

pre 1772


Rutty (1772); O'Neill et al. (2011); Quigley (2011)

Quigley (1996)

Browne (1774); Brown et al. (2001)

Rutty (1772); Browne (1774); Sampson (1802); Templeton (1837); Thompson (1856); Harvey (1845); Went (1946, 1949, 1978)

Kennedy and Fitzmaurice (1972c)

Browne (1774); Sampson (1802); M'Skimin (1909); Thompson (1856); Ogilby (1885); Went (1949)

Rutty (1772); Browne (1774); Templeton (1837); Thompson (1841); Harvey (1845); Went (1946, 1978); Hayden (2002)

References


Three-spined Stickleback

Gasterosteidae

Flathead Grey Mullet

Big-scale Sand Smelt

euryhaline

Mugil cephalus L.

Thin-lipped Grey Mullet

Atherinidae

euryhaline

Liza ramada (Risso, 1826)

Golden Grey Mullet

Ballan Wrasse

euryhaline

Liza aurata (Risso, 1810)

Thick-lipped Grey Mullet

Mugilidae

Labridae

euryhaline

Chelon labrosus (Risso, 1826)

Grayling

stenohaline

Thymallidae

Salvelinus alpinus (L.)

Arctic Charr

Salinity Tolerance

Salmonidae

Scientific Name

Common Name

Family Name


