<?xml version="1.0" encoding="utf-8" standalone="no"?>
<!DOCTYPE article SYSTEM "http://www.clim-past-discuss.net/inc/cpd/copernicus.dtd">
<article language="en">
	<journal>
		<journal_title>Climate of the Past Discussions</journal_title>
		<journal_url>www.clim-past-discuss.net</journal_url>
		<issn>1814-9340</issn>
		<eissn>1814-9359</eissn>
		<volume_number>6</volume_number>
		<issue_number>4</issue_number>
		<publication_year>2010</publication_year>
	</journal>
	<doi>10.5194/cpd-6-1351-2010</doi>
	<article_url>http://www.clim-past-discuss.net/6/1351/2010/</article_url>
	<abstract_html>http://www.clim-past-discuss.net/6/1351/2010/cpd-6-1351-2010.html</abstract_html>
	<fulltext_pdf>http://www.clim-past-discuss.net/6/1351/2010/cpd-6-1351-2010.pdf</fulltext_pdf>
	<start_page>1351</start_page>
	<end_page>1389</end_page>
	<publication_date>2010-07-16</publication_date>
	<article_title content_type="html">Refugia of marine fish in the Northeast Atlantic during the Last Glacial Maximum: concordant  assessment from archaeozoology and palaeotemperature reconstructions</article_title>
	<authors>
		<author numeration="1" affiliations="1">
			<name>A. J. Kettle</name>
			<email>kettle@oswego.edu</email>
		</author>
		<author numeration="2" affiliations="2">
			<name>A. Morales-Muñiz</name>
		</author>
		<author numeration="3" affiliations="2">
			<name>E. Roselló-Izquierdo</name>
		</author>
		<author numeration="4" affiliations="3">
			<name>D. Heinrich</name>
		</author>
		<author numeration="5" affiliations="4">
			<name>L. A. Vøllestad</name>
		</author>
	</authors>
	<affiliations>
		<affiliation numeration="1" content_type="html">Department of Earth Science, SUNY-Oswego, Oswego, New York, USA</affiliation>
		<affiliation numeration="2" content_type="html">Laboratorio de Arqueozoologia, Universidad Autonoma de Madrid, Madrid, Spain</affiliation>
		<affiliation numeration="3" content_type="html">Boesselstrasse 9, 24937 Flensburg, Germany</affiliation>
		<affiliation numeration="4" content_type="html">Center for Ecological and Evolutionary Synthesis, Department of Biology, University of Oslo, Oslo,  Norway</affiliation>
	</affiliations>
	<abstract content_type="html">Archaeozoological finds of the remains of marine and amphihaline fish from the Last Glacial
      Maximum (LGM) ca. 21 ka ago show evidence of very different species ranges compared to the
      present. We show how an ecological niche model (ENM) based on palaeoclimatic reconstructions
      of sea surface temperature and bathymetry can be used to effectively predict the spatial
      range of marine fish during the LGM. The results indicate that the ranges of marine fish
      species that are now in Northwestern Europe were almost completely displaced southward from
      the modern distribution. Significantly, there is strong evidence that there was an invasion of fish of current economic importance
      into the Western Mediterranean through the Straits of
      Gibraltar, where they were exploited by Palaeolithic human populations. There has been much
      recent interest in the marine glacial refugia to understand how the ranges of the
      economically important fish species will be displaced with the future climate
      warming. Recent ENM studies have suggested that species ranges may not have been displaced
      far southward during the coldest conditions of the LGM. However, archaeozoological evidence
      and LGM ocean temperature reconstructions indicate that there were large range changes, and
      certain marine species were able invade the Western Mediterranean. These findings are
      important for ongoing studies of molecular ecology that aim to assess marine glacial refugia
      from the genetic structure of living populations, and they pose questions about the genetic
      identity of vanished marine populations during the LGM. The research presents a  challenge
      for future archaeozoological work to verify palaeoclimatic reconstructions and delimit the
    glacial refugia.</abstract>
	<references>
		<reference numeration="1" content_type="text"> Acolas,~M L., Veron,~V., Jourdan,~H., Begout,~M L., Sabatie,~M R., and Baglinière,~J L.: Upstream migration and reproductive patterns of a~population of allis shad in a~small river (L&apos;Aulne, Brittany, France), ICES~J. Mar. Sci., 63, 476–484, \doi10.1016/j.icesjms.2004.07.023, 2006. </reference>
		<reference numeration="2" content_type="text"> Addison,~J A. and Hart,~M W.: Colonization, dispersal, and hybridization influence phylogeography of North Atlantic sea urchins (\textitStrongylocentrotus droebachiensis), Evolution, 59, 532–543, \doi10.1554/04-238, 2005. </reference>
		<reference numeration="3" content_type="text"> Aura Tortosa,~J E., Jordá Pardo,~J F., Pérez Ripoll,~M., Rodrigo García,~M J., Badal García,~E., and Guillem Calatayud,~P.: The far south: the Pleistocene-Holocene transition in Nerja Cave (Andalucía, Spain), Quatern. Int., 93–94, 19–30, \doi10.1016/S1040-6182(02)00004-6, 2002. </reference>
		<reference numeration="4" content_type="text"> Beerenhout,~B.: What conclusions can be drawn from mature haddock bones in a~Neolithic site in the Netherlands, Offa, 51, 341–347, 1994. </reference>
		<reference numeration="5" content_type="text"> Bigg,~G R., Cunningham,~C W., Ottersen,~G., Pogson,~G H., Wadley,~M R., and Williamson,~P.: Ice-age survival of Atlantic cod: agreement between palaeoecology and genetics,~P. Roy. Soc. B-Biol. Sci., 275, 163–172, \doi10.1098/rspb.2007.1153, 2008. </reference>
		<reference numeration="6" content_type="text"> Bradley,~R S.: Paleoclimatology. Reconstructing Climates of the Quaternary, Academic Press, San Diego, 1999. </reference>
		<reference numeration="7" content_type="text"> Brander,~K.: Effects of climate change on cod (\textitGadus morhua) stocks, in: Global Warming: Implications for Freshwater and Marine Fish, edited by: Wood,~C M. and McDonald,~D G., Society for Experimental Biology Seminar Series 61, Cambridge University Press, 255–278, 1996. </reference>
		<reference numeration="8" content_type="text"> Brodziak,~J K T.: Haddock, \textitMelanogrammus aeglefinus, Life History and Habitat Characteristics, NOAA Technical Memorandum NMFS-NE-196, US Department of Commerce, December 2005. </reference>
		<reference numeration="9" content_type="text"> Butzer,~K W.: Environment and Archaeology, Aldine Publishing Company, Chicago, 1964 </reference>
		<reference numeration="10" content_type="text"> Campo,~D., Molares,~J., Garcia,~L., Fernandez-Rueda,~P., Garcia-Gonzalez,~C., and Garcia-Vazquez,~E.: Phylogeography of the European stalked barnacle (\textitPollicipes pollicipes): identification of glacial refugia, Mol. Biol., 157, 147–156, \doi10.1007/s00227-009-1305-z, 2010. </reference>
		<reference numeration="11" content_type="text"> Chevolot,~M., Hoarau,~G., Rijnsdorp,~A D., and Stam,~W T.: Phylogeography and population structure of thornback rays (\textitRaja clavata~L., Rajidae), Mol. Ecol., 15, 3693–3705, \doi10.1111/j.1365-294X.2006.03043.x, 2006. </reference>
		<reference numeration="12" content_type="text"> Clark,~J G D.: The development of fishing in prehistoric Europe, Antiq J., 28, 45–85, 1948. </reference>
		<reference numeration="13" content_type="text"> CLIMAP Project Members: The surface of the ice-age Earth, Science, 191, 1131–1137, \doi10.1126/science.191.4232.1131, 1976. </reference>
		<reference numeration="14" content_type="text"> Consuegra,~S., García~de Leániz,~C., Serdio,~A., González Morales,~M., Straus,~L G., Knox,~D., and Verspoor,~E.: Mitochondrial DNA variation in Pleistocene and modern Atlantic salmon from the Iberian glacial refugium, Mol. Ecol., 11, 2037–2048, \doi10.1046/j.1365-294X.2002.01592.x, 2002. </reference>
		<reference numeration="15" content_type="text"> Cortés-Sánchez,~M., Morales-Muñiz,~A., Simón-Vallejo,~M D., Bergadà-Zapata,~M M., Delgado-Huertas,~A., López-García,~P., López-Sáez,~J A., Lozano-Francisco,~M C., Riquelme-Cantal,~J A., Roselló-Izquierdo,~E., Sánchez-Marco,~A., and Vera-Peláez,~J L.: Palaeoenvironmental and cultural dynamics of the coast of Málaga (Andalusia, Spain) during the Upper Pleistocene and early Holocene, Quaternary Sci. Rev., 27, 2176–2193, \doi10.1016/j.quascirev.2008.03.010, 2008. </reference>
		<reference numeration="16" content_type="text"> Costamagno,~S. and Laroulandie,~V.: L&apos;exploitation des petits vertébrés dans les Pyrénées françaises du Paléolithique au Mésolithique: un inventaire taphonomique et archéozoologique, in: Petits Animaux et Sociétés Humaines, du Complement Alimentaire aux Ressources Utilitaires, edited by: Brugal,~J.-P. and Desse,~J., XXIVe rencontres internationales d&apos;archéologie et d&apos;histoire d&apos;Antibes, Éditions APDCA, Antibes, 403–416, 2004. </reference>
		<reference numeration="17" content_type="text"> Coyer,~J A., Peters,~A F., Stam,~W T., and Olsen,~J L.: Post-ice age recolonization and differentiation of \textitFucus serratus~L. (\textitPhaeophyceae; Fucuceae) populations in Northern Europe, Mol. Ecol., 12, 1817–1829, \doi10.1046/j.1365-294X.2003.01850.x, 2003. </reference>
		<reference numeration="18" content_type="text"> Dahlgren,~T G., Weinberg,~J R., and Halanych,~K M.: Phylogeography of the ocean quahog (\textitArctica islandica): influences of paleoclimate on genetic diversity and species range, Mar. Biol., 137, 487–495, \doi10.1007/s002270000342, 2000. </reference>
		<reference numeration="19" content_type="text"> Desse,~G. and Granier,~J.: Les poissons, in: La Préhistoire française, I, 1, CNRS, Paris, 437–441, 1976. </reference>
		<reference numeration="20" content_type="text"> Desse,~J. and Desse-Berset,~N.: Pêche et surpêche en Méditerranée: le témoignage des os, in: Exploitation des animaux sauvages à travers le temps, actes des XIII$^e$ Rencontres internationals d&apos;archéologie et d&apos;histoire d&apos;Antibes, octobre 1992, Antibes, Éditions APDCA, 327–339, 1993. </reference>
		<reference numeration="21" content_type="text"> Desse,~J. and Desse-Berset,~N.: Le cortège de Neptune: les poissons de la Mediterranée durant l&apos;Holocène, in: Mouvements ou déplacements de populations animales en Mediterranée au cours de l&apos;Holocène, edited by: Gardeisen,~A., Séminaire de recherche du thème 15. Archéologie de l&apos;animal (UMR-154-CNRS), Lattes, Montpellier (France) 29 Septembre, 2000, BAR International Series 1017, 83–96, 2002. </reference>
		<reference numeration="22" content_type="text"> Desse-Berset,~N.: Les poissons, in: Les grottes Jean-Pierre I et II à Saint-Thibaud-de-Couz (Savoie), edited by: Bintz,~P., Gallia Préhistoire, 36, 218–224, 1994. </reference>
		<reference numeration="23" content_type="text"> Domingues,~V S., Bucciarelli,~G., Almada,~V C., and Bernardi,~G.: Historical colonization and demography of the Mediterranean damselfish, \textitChromis chromis, Mol. Ecol., 14, 4051–4063, \doi10.1111/j.1365-294X.2005.02723.x, 2005. </reference>
		<reference numeration="24" content_type="text"> Elder,~K L., Jones,~G A., and Bolz,~G.: Distribution of otoliths in surficial sediments of the US Atlantic continental shelf and slope and potential for reconstructing Holocene climate, Paleoceanography, 11, 359–367, \doi10.1029/96PA00042, 1996. </reference>
		<reference numeration="25" content_type="text"> Enghoff,~I B., MacKenzie,~B R., and Nielsen,~E E.: The Danish fish fauna during the warm Atlantic period (ca. 7000–3900 BC): forerunner of future changes? Fish. Res., 87, 167–180, \doi10.1016/j.fishres.2007.03.004, 2007. </reference>
		<reference numeration="26" content_type="text"> Francisco,~S M., Castilho,~R., Soares,~M., Congiu,~L., Brito,~A., Vieira,~M N., and Almada,~V C.: Phylogeography and demographic history of \textitAtherina presbyter (Pisces: Atherinidae) in the North-Eastern Atlantic based on mitochondrial DNA, Mar. Biol., 156, 1421–1432, \doi10.1007/s00227-009-1182-5, 2009. </reference>
		<reference numeration="27" content_type="text"> Friedland,~K D., Reddin,~D G., and Castonguay,~M.: Oceanic thermal conditions in the post-smolt nursery of North Atlantic salmon, ICES~J. Mar. Sci., 60, 343–355, \doi10.1016/S1054-3139(03)00022-5, 2003. </reference>
		<reference numeration="28" content_type="text"> Froget,~C., Thommeret,~J., and Thommeret,~Y.: Mollusques septentrionaux en Méditerranée occidentale: datation par le $^14$C, Palaeogeogr. Palaeocl., 12, 285–293, 1972. </reference>
		<reference numeration="29" content_type="text"> Gómez,~A., Hughes,~R N., Wright,~P J., Carvalho,~G R., and Lunt,~D H.: Mitochondrial DNA phylogeography and mating compatibility reveal marked genetic structuring and speciation in the NE Atlantic bryozoans \textitCelleporella hyaline, Mol. Ecol., 16, 2173–2188, \doi10.1111/j.1365-294X.2007.03308.x, 2007. </reference>
		<reference numeration="30" content_type="text"> Gysels,~E S., Hellemans,~B., Patarnello,~T., and Volckaert,~F A M.: Current and historic gene flow of the sand goby \textitPomastoschistus minutus on the European Continental Shelf and in the Mediterranean Sea, Biol J. Linn. Soc., 83, 561–576, \doi10.1111/j.1095-8312.2004.00411.x,2004a. </reference>
		<reference numeration="31" content_type="text"> Gysels,~E S., Hellemans,~B., Pampoulie,~C., and Volckaert,~F A M.: Phylogeography of the common goby, \textitPomatoschistus microps, with particular emphasis on the colonization of the Mediterranean and the North Sea, Mol. Ecol., 13, 403–417, \doi10.1046/j.1365-294X.2003.02087.x, 2004b. </reference>
		<reference numeration="32" content_type="text"> Heinrich,~D.: Bemerkungen zu den Tierknochenfunden von den submarinen Siedlungsplätzen der Ertebølle-Kultur bei Timmendorf/Poel und bei Neustadt (Marienbad) – ein Vorbericht, Beitr. Archäozool. Prähist. Anthropol., 3, 39–43, 2001. </reference>
		<reference numeration="33" content_type="text"> Hewitt,~G.: The genetic legacy of the Quaternary ice ages, Nature, 405, 907–913, \doi10.1038/35016000, 2000. </reference>
		<reference numeration="34" content_type="text"> Hoarau,~G., Coyer,~J A., Veldsink,~J H., Stam,~W T., and Olsen,~J L.: Glacial refugia and recolonization pathways in the brown seaweed \textitFucus serratus, Mol. Ecol., 16, 3606–3616, \doi10.1111/j.1365-294X.2007.03408.x, 2007. </reference>
		<reference numeration="35" content_type="text"> Hvidsten,~N A., Heggberget,~T G., and Jensen,~A J.: Sea water temperatures at Atlantic salmon smolt enterance, Nordic J. Freshwater Res., 74, 79–86, 1998. </reference>
		<reference numeration="36" content_type="text"> ICES-FishMap: Haddock (WWW document): available at: http://www.ices.dk/marineworld/fishmap/ices/pdf/haddock.pdf, 2009. </reference>
		<reference numeration="37" content_type="text"> Jolly,~M T., Jollivet, D, Gentil,~F., Thiébaut,~E., and Viard,~F.: Sharp genetic break between Atlantic and English Channel populations of the polychaete \textitPectinaria koreni, along the North coast of France, Heredity, 94, 23–32, \doi10.1038/sj.hdy.6800543, 2005. </reference>
		<reference numeration="38" content_type="text"> Jolly,~M T., Viard,~F., Gentil,~F., Thiébaut,~E., and Jollivet,~D.: Comparative phylogeography of two coastal polychaete tubeworms in the North East Atlantic supports shared history and vicariant events, Mol. Ecol., 15, 1841–1855, \doi10.1111/j.1365-294X.2006.02910.x, 2006. </reference>
		<reference numeration="39" content_type="text"> Juan-Muns i Plans,~N.: La ictiofauna dels jaciments arqueològics Catalans, Cypsela (Girona), V, 21–33, 1985. </reference>
		<reference numeration="40" content_type="text"> Juan-Muns,~N.: La ictiofauna de la cova de l&apos;Arbreda (Serinyà, Girona), Cypsela (Girona), VI, 97–100, 1987. </reference>
		<reference numeration="41" content_type="text"> Juan-Muns i Plans,~N., Rodrigo García,~M J., and Rodríguez Santana,~C G.: La ictiofauna de los yacimientos arqueológicos. Sus posilidades en la reconstrucción paleoecológica y de interpretación paleoeconómica, Arqueología, Madrid, 83–99, 1991. </reference>
		<reference numeration="42" content_type="text"> Kaschner,~K., Watson,~R., Trites,~A W., and Pauly,~D.: Mapping world-wide distributions of marine mammal species using a~relative environmental suitability (RES) model, Mar. Ecol.-Prog. Ser., 316, 285–310, \doi10.3354/meps316285, 2006. </reference>
		<reference numeration="43" content_type="text"> Kettle,~A J., Heinrich,~D., Barrett,~J H., Benecke,~N., and Locker,~A.: Past distributions of the European freshwater eel from archaeological and palaeontological evidence, Quaternary Sci. Rev., 27, 1309–1334, \doi10.1016/j.quascirev.2008.03.005, 2008. </reference>
		<reference numeration="44" content_type="text"> King,~T L., Verspoor,~E., Spidle,~A P., Gross,~R., Phillips,~R B., Koljonen,~M.-L., Sanchez,~J A., and Morrison,~C L.: 5. Biodiversity and population structure, in: The Atlantic Salmon. Genetics, Conservation and Management, edited by: Verspoor,~E., Stradmeyer,~L., and Nielsen,~J L., Blackwell Publishing, Oxford, 117–166, 2007. </reference>
		<reference numeration="45" content_type="text"> Langefors,~A H.: Adaptive and neutral genetic variation and colonization history of Atlantic salmon, \textitSalmo salar, Environ. Biol. Fish., 74, 297–308, \doi10.1007/s10641-005-0501-z, 2005. </reference>
		<reference numeration="46" content_type="text"> Larmuseau,~M H D., Van Houdt,~J K J., Guelinckx,~J., Hellemans,~B., and Volckaert,~F A M.: Distributional and demographic consequences of Pleistocene climate fluctuations for a~marine demersal fish in the North-Eastern Atlantic,~J. Biogeogr., 36, 1138–1151, \doi10.1111/j.1365-2699.2008.02072.x, 2009. </reference>
		<reference numeration="47" content_type="text"> Lassalle,~G. and Rochard,~E.: Impact of twenty-first century climate change on diadromous fish spread over Europe, North Africa and the Middle East, Glob. Change Biol., 15, 1072–1089, \doi10.1111/j.1365-2486.2008.01794.x, 2009. </reference>
		<reference numeration="48" content_type="text"> Le Gall,~O.: Presence de vestiges de saumon Atlantique dans un site Epipaleolithique et Mesolithique du bassin versant Mediterranean, Bulletin du Société d&apos;Anthropologie de Sud-Ouest, XVIII, 53–55, 1983. </reference>
		<reference numeration="49" content_type="text"> Le Gall,~O.: L&apos;Exploitation de l&apos;ichthyofaune par les paléolithiques quelques exemples., in: 2nd Fish Osteoarchaeology Meeting, edited by: Desse-Berset,~N., Table Rond, Sophia Antipolis – Valbonne, 14–16 Octobre, 1983, CNRS, Centres de Recherches Archéologiques, Notes et Monographies Techniques No. 16, 89–112, 1984. </reference>
		<reference numeration="50" content_type="text"> Le Gall,~O.: Poissons et pêches au Paléolithique (Quelques données de l&apos;Europe occidentale), L&apos;Anthopologie (Paris), 96, 121–134, 1992. </reference>
		<reference numeration="51" content_type="text"> Le Gall,~O.: Évolution des pêches de l&apos;Épipaléolithique au Néolithique Ancien, in: Préhistoire Anthropologie Mediterranéenes, LAPMO, Université de Provence, CNRS, 2, 135–142, 1993. </reference>
		<reference numeration="52" content_type="text"> Le Gall,~O.: Eléments de réflexion sur la pêche dans le bassin méditerranéen nord-occidental pendant le développement des faciès leptolithiques, XXIVe Congrès Préhistorique de France – Carcassonne 26–30 septembre 1994 – Les faciès leptolithiques du nord-ouest méditerranéen: milieux naturels et culturels, 251–265, 1994a. </reference>
		<reference numeration="53" content_type="text"> Le Gall,~O.: Quelques remarques sur l&apos;adaptation à court et à long termes chez les poissons d&apos;eau douce du sud de la France, in: Fish Exploitation in the Past, edited by: Van Neer,~W., Proceedings of the 7th meetings of the ICAZ Fish Remains Working Group, Annales du Musée Royal de l&apos;Afrique Centrale, Sciences Zoologiques, no. 274, Turvuren, 91–98, 1994b. </reference>
		<reference numeration="54" content_type="text"> Le Gall,~O.: Études des Poissons, in: La Grotte de Pégourié, Caniac-du-Causse (Lot), edited by: Séronie-Vivien,~M.-R., Préhistoire Quercinoise, Supplément no. 2, 149–154, 1995. </reference>
		<reference numeration="55" content_type="text"> Le Gall,~O.: 6. Origine et histoire des aloses, in: Les Aloses (\textitAlosa alosa et \textitAlosa fallax spp.), edited by: Baglinière,~J L. and Elie,~P., Écobiologie et variabilité des populations, INRA, Paris, 127–136, 2000. </reference>
		<reference numeration="56" content_type="text"> Le Gall,~O.: Les représentations de poissons dans l&apos;art mobilier magdalenien. Une expression de l&apos;importance culturelle de la pêche, Bulletin Préhistoire du Sud-Ouest, no. 8, 55–69, 2001. </reference>
		<reference numeration="57" content_type="text"> Le Gall,~O.: Les poissons des eaux douces pléistocènes sont-ils des indicateurs paleoclimatiques? Une approche élargie à l&apos;Europe, in: Archéologie du Poisson. 30 Ans d&apos;Archéo-Ichtyologie au CNRS, edited by: Béarez,,~P., Grouard,~S., and Clavel,~B., Hommage aux travaux de Jean Desse et Nathalie Desse-Berset, XIVth ICAZ Fish remains working group meeting, Éditions APDCA, Antibes, 311–326, 2008. </reference>
		<reference numeration="58" content_type="text"> Lepiksaar,~J.: Die spät- und postglaziale Faunengeschichte der Süßwasserfische Schwedens, Oetker-Voges Verlag, Kiel, 2001. </reference>
		<reference numeration="59" content_type="text"> Levitus, S.: World Ocean Atlas 2005, NOAA Atlas NESDIS, US Government Printing Office, Washington, DC, http://www.nodc.noaa.gov/OC5/WOA05/pr_woa05.html, 2001. </reference>
		<reference numeration="60" content_type="text"> Luttikhuizen,~P C., Drent,~J., and Baker,~A J.: Disjunct distribution of highly diverged mitochondrial lineage clade and population subdivision in a~marine bivalve with pelagic larval dispersal, Mol. Ecol., 12, 2215–2229, \doi10.1046/j.1365-294X.2003.01872.x, 2003. </reference>
		<reference numeration="61" content_type="text"> Maggio,~T., Brutto,~S L., Garoia,~F., Tinti,~F., and Arculeo,~M.: Microsatellite analysis of red mullet \textitMullus barbatus (\textitPerciformes, Mullidae) reveals the isolation of the Adriatic Basin in the Mediterranean Sea, ICES~J. Mar. Sci., 66, 1883–1891, \doi10.1093/icesjms/fsp160, 2009. </reference>
		<reference numeration="62" content_type="text"> Maggs,~C A., Castilho,~R., Foltz,~D., Henzler,~C., Jolly,~M T., Kelly,~J., Olsen,~J., Perez,~K E., Stam,~W., Väinölä,~R., Viard,~F., and Wares,~J.: Evaluating signatures of glacial refugia for North Atlantic benthic marine taxa, Ecology, 89 Suppl., S108–S122, \doi10.1890/08-0257.1, 2008. </reference>
		<reference numeration="63" content_type="text"> Makhrov,~A A., Verspoor,~E., Artamonova,~V S., and O&apos;Sullivan,~M.: Atlantic salmon colonization of the Russian Arctic coast: pioneers from North America,~J. Fish Biol., 67(Suppl. A), 68–79, \doi10.1111/j.0022-1112.2005.00840.x, 2005. </reference>
		<reference numeration="64" content_type="text"> Malatesta,~A. and Zarlenga,~F.: Northern guests in the Pleistocene Mediterranean Sea, Geologica Romana, 25, 91–154, 1986. </reference>
		<reference numeration="65" content_type="text"> MARGO Project Members: Constraints on the magnitude and patterns of ocean cooling at the Last Glacial Maximum, Nature Geosci., 2, 127–132, \doi10.1038/ngeo411, 2009. </reference>
		<reference numeration="66" content_type="text"> Martin,~H. and Le Gall,~O.: La faune: un indicateur des comportements humains. La chasse de cerf au Post-Paléolithique: deux exemples, Anthropologie et Préhistoire, 111, 364–369, 2000. </reference>
		<reference numeration="67" content_type="text"> Morales,~A.: Los peces fosiles del yacimiento achelense de Aridos-1 (Arganda, Madrid), in: Ocupaciones achelenses en el Valle del Jarama, edited by: Santonja,~M. and López,~N., Madrid, 93–104, 1980. </reference>
		<reference numeration="68" content_type="text"> Morales,~A. and Roselló,~E.: Fishing down the food web in Iberian prehistory? A~new look at the fishes from Cueva de Nerja (Málaga, Spain), in: Petits Animaux et Sociétés Humaines. Du Complément Alimentaire aux Ressources Utilitaires, edited by: Brugal,~J.-P. and Desse,~J., XXIVe rencontres internationals d&apos;archéologie et d&apos;histoire d&apos;Antibes, Éditions APDCA, Antibes, 111–123, 2004. </reference>
		<reference numeration="69" content_type="text"> Morales-Muñiz,~A. and Roselló-Izquierdo,~E.: Twenty thousand years of fishing in the Strait, in: Human Impacts on Ancient Marine Ecosystems. A~Global Perspective, edited by: Rick,~T C. and Erlandson,~J M., University of California Press, Berkeley, 243–277, 2008. </reference>
		<reference numeration="70" content_type="text"> Nesbø,~C L., Rueness,~E K., Iversen,~S A., Skagen,~D W., and Jakobsen,~K S.: Phylogeography and population history of Atlantic mackerel (\textitScomber scombrus~L.): a~genealogical approach reveals genetic structuring among the Eastern Atlantic stocks,~P. Roy. Soc. B-Biol. Sci., 267, 281–292, \doi10.1098/rspb.2000.0998, 2000. </reference>
		<reference numeration="71" content_type="text"> Nielsen,~E E. and Hansen,~M M.: Waking the dead: the value of population genetic analyses of historical samples, Fish Fish., 9, 450–461, \doi10.1111/j.1467-2979.2008.00304.x, 2008. </reference>
		<reference numeration="72" content_type="text"> Nikula,~R., Strelkov,~P., and Väinölä,~R.: Diversity and trans-Arctic invasion history of mitochondrial lineages in the North Atlantic \textitMacoma balthica complex (bivalvia: tellinidae), Evolution, 61, 928–941, \doi10.1111/j.1558-5646.2007.00066.x, 2007. </reference>
		<reference numeration="73" content_type="text"> Nogués-Bravo,~D.: Predicting the past distribution of species climatic niches, Global Ecol. Biogeogr., 18, 521–531, \doi10.1111/j.1466-8238.2009.00476.x, 2009. </reference>
		<reference numeration="74" content_type="text"> Olsen,~J L., Stam,~W T., Coyer,~J A., Reusch,~T B H., Billingham,~M., Boström,~C., Calvert,~E., Christie,~H., Granger,~S., La Lumière,~R., Milchakova,~N., Oudot-Le Secq,~M.-P., Procaccini,~G., Sanjabi,~B., Serrão,~E., Veldsink,~J., Widdicombe,~S., and Wyllie-Echverria,~S.: North Atlantic phylogeography and large-scale population differentiation of the seagrass \textitZostera marina~L., Mol. Ecol., 13, 1923–1941, \doi10.1111/j.1365-294X.2004.02205.x, 2004. </reference>
		<reference numeration="75" content_type="text"> Olsen,~J L., Zechman,~F W., Hoarau,~G., Coyer,~J A., Stam,~W T., Valero,~M., and Åberg,~P.: The phylogeographic architecture of the fucoid seaweed \textitAscophyllum nodosum: an intertidal \qutmarine tree and survivor of more than one glacial-interglacial cycle,~J. Biogeogr., 37, 842–856, \doi10.1111/j.1365-2699.2009.02262.x, 2010. </reference>
		<reference numeration="76" content_type="text"> Pampoulie,~C., Stefánsson,~M Ö., Jörundsdóttir,~T D., Danilowicz,~B S., and Daníelsdóttir,~A K.: Recolonization history and large-scale dispersal in the open sea: the case study of the North Atlantic cod, \textitGadus morhua~L., Biol J. Linn. Soc., 94, 315–329, \doi10.1111/j.1095-8312.2008.00995.x, 2008. </reference>
		<reference numeration="77" content_type="text"> Paul,~A. and Schäfer-Neth,~C.: Modeling the water masses of the Atlantic Ocean at the Last Glacial Maximum, Paleoceanography, 18, 1058, \doi10.1029/2002PA000783, 2003. </reference>
		<reference numeration="78" content_type="text"> Peck,~M A., Buckley,~L J., Caldarone,~E M., and Bengtson,~D A.: Effects of food consumption and temperature on growth rate and biochemical-based indicators of growth in early juvenile Atlantic cod \textitGadus morhua and haddock \textitMelanogrammus aeglefinus, Mar. Ecol.-Prog. Ser., 251, 233–243, \doi10.3354/meps251233, 2003. </reference>
		<reference numeration="79" content_type="text"> Peck,~M A., Buckley,~L J., and Bengtson,~D A.: Effects of temperature, body size and feeding on rates of metabolism in young-of-the-year haddock,~J. Fish Biol., 66, 911–923, \doi10.1111/j.1095-8649.2005.00633.x, 2005. </reference>
		<reference numeration="80" content_type="text"> Peltier,~W R.: Ice-age paleotopography, Science, 265, 195–201, \doi10.1126/science.265.5169.195, 1994. </reference>
		<reference numeration="81" content_type="text"> Perry,~A L., Low,~P J., Ellis,~J R., and Reynolds,~J D.: Climate change and distribution shifts in marine fishes, Science, 308, 1912–1915, \doi10.1126/science.1111322, 2005. </reference>
		<reference numeration="82" content_type="text"> Person-Le Ruyet,~J., Buchet,~V., Vincent,~B., Le Delliou,~H., and Quemener,~L.: Effects of temperature on the growth of pollack (\textitPollachius pollachius) juveniles, Aquaculture, 251, 340–345, \doi10.1016/j.aquaculture.2005.06.029, 2006. </reference>
		<reference numeration="83" content_type="text"> Pickard,~C. and Bonsall,~C.: Deep-sea fishing in the European Mesolithic: fact for fantasy? Eur. J. Archaeol., 7, 273–290, \doi10.1177/1461957104056504, 2004. </reference>
		<reference numeration="84" content_type="text"> Pörtner,~H O.: Climate change and temperature-dependent biogeography: oxygen limitation of thermal tolerance in animals, Naturwissenschaften, 88, 137–146, \doi10.1007/s001140100216, 2001. </reference>
		<reference numeration="85" content_type="text"> Pörtner,~H O. and Knust,~R.: Climate change affects marine fishes through the oxygen limitation of thermal tolerance, Science, 315, 95–97, \doi10.1126/science.1135471, 2007. </reference>
		<reference numeration="86" content_type="text"> Provan,~J. and Bennett,~K D.: Phylogeographic insights into cryptic glacial refugia, Trends Ecol. Evol., 23, 564–571, \doi10.1016/j.tree.2008.06.010, 2008. </reference>
		<reference numeration="87" content_type="text"> Provan,~J., Wattier,~R A., and Maggs,~C A.: Phylogeographic analysis of the red seaweed \textitPalmaria palmata reveals a~Pleistocene marine glacial refugium in the English Channel, Mol. Ecol., 14, 793–803, \doi10.1111/j.1365-294X.2005.02447.x, 2005. </reference>
		<reference numeration="88" content_type="text"> Provan,~J., Beatty,~G E., Keating,~S L., Maggs,~C A., and Savidge,~G.: High dispersal potential has maintained long-term population stability in the North Atlantic copepod \textitCalanus finmarchicus,~P. Roy. Soc. B-Biol. Soc., 276, 301–307, \doi10.1098/rspb.2008.1062, 2009. </reference>
		<reference numeration="89" content_type="text"> Renouf,~M A P.: Prehistoric Hunter-Fishers of Varangerfjord, Northeastern Norway, Brit. Archaeol. Rep. In., 478~pp., Oxford, 1989. </reference>
		<reference numeration="90" content_type="text"> Rivière,~E.: Faune des oiseaux, des reptiles et des poissons trouvés dans les cavernes des Baoussé-Roussé (Italie) dites Grottes de Menton, Materiaux pour l&apos;Histoire Primitive et Naturelle de l&apos;Homme, 20, 525–535, 1886. </reference>
		<reference numeration="91" content_type="text"> Rodrigo,~M J.: Remains of \textitMelanogrammus aeglefinus (Linnaeus, 1758) in the Pleistocene – Holocene passage in the cave of Nerja, Málaga/Spain, Offa, 51, 348–351, 1994. </reference>
		<reference numeration="92" content_type="text"> Roman,~J. and Palumbi,~S R.: A~global invader at home: population structure of the green crab, \textitCarcinus maenus, in Europe, Mol. Ecol., 13, 2891–2898, \doi10.1111/j.1365-294X.2004.02255.x, 2004. </reference>
		<reference numeration="93" content_type="text"> Roselló,~E. and Morales,~A.: Lapa dos Coelhos: informe sobre los restos de peces, in: Homenaje a~Francisco Giles Pacheco, edited by: Mata,~E., Publicaciones Junta de Andalucia, Sevilla, 261–276, 2010. </reference>
		<reference numeration="94" content_type="text"> Schmölcke,~U., Endtmann,~E., Kloos,~S., Meyer,~M., Michaelis,~D., Rickert,~B.-H., and Rössler,~D.: Changes of sea level, landscape and culture: a~review of the South-Western Baltic area between 8800 and 4000 BC, Palaeogeogr. Palaeoclim., 240, 423–438, \doi10.1016/j.palaeo.2006.02.009, 2006. </reference>
		<reference numeration="95" content_type="text"> Sejrup,~H P., Hjelstuen,~B O., Dahlgren,~K I T., Haflidason,~H., Kuijpers,~A., Nygård,~A., Praeg,~D., Stoker,~M S., and Vorren,~T O.: Pleistocene glacial history of the NW European continental margin, Mar. Petrol. Geol., 22, 1111–1129, \doi10.1016/j.marpetgeo.2004.09.007, 2005. </reference>
		<reference numeration="96" content_type="text"> Stewart,~J R. and Lister,~A M.: Cryptic northern refugia and the origins of the modern biota, Trends Ecol. Evol., 16, 608–613, \doi10.1016/S0169-5347(01)02338-2, 2001. </reference>
		<reference numeration="97" content_type="text"> Waltari,~E., Hijmans,~R J., Peterson,~A T., Nyári,~A S., Perkins,~S L., and Guralnick,~R P.: Locating Pleistocene refugia: competing phylogeographic and ecological niche model predictions, PLoS ONE, 2, e563. \doi10.1371/journal.pone.0000563, 2007. </reference>
		<reference numeration="98" content_type="text"> Wares,~J P.: Intraspecific variation and geographic isolation in \textitIdotea balthica (Isopoda: valvifera),~J. Crustacean Biol., 21, 1007–1013, \doi10.1651/0278-0372(2001)021[1007:IVAGII]2.0.CO;2, 2001. </reference>
		<reference numeration="99" content_type="text"> Wares,~J P. and Cunningham,~C W.: Phylogeography and historical ecology of the North Atlantic intertidal, Evolution, 55, 2455–2469, \doi10.1554/0014-3820(2001)055[2455:PAHEOT]2.0.CO;2, 2001. </reference>
		<reference numeration="100" content_type="text"> Whitehead,~P J P., Bauchot,~M L., Hureau,~J C., Nielsen,~J., and Tortonese,~E.: Fishes of the North-Eastern Atlantic and the Mediterranean, UNESCO, Paris, 1986. </reference>
	</references>
</article>

