Population connectivity and phylogeography of the Mediterranean endemic skate Raja polystigma and evidence of its hybridization with the parapatric sibling R. montagui

MARINE ECOLOGY PROGRESS SERIES Mar Ecol Prog Ser

Vol. 554: 99–113, 2016

`œˆ\Ê£ä°ÎÎx{ɓi«Ã££Ç™™ Published July 28

INTRODUCTION

iëˆÌiÊÀi«ÀiÃi˜Ìˆ˜}ʜ˜ÞÊä°ÎӯʜvÊ̅iÊÃÕÀv>Viʜv ̅iÊ œVi>˜Ã]Ê Ì…iÊ i`ˆÌiÀÀ>˜i>˜Ê -i>Ê ˆÃÊ iÃ̈“>Ìi`Ê Ìœ Vœ˜Ì>ˆ˜Êœ˜Ê>ÛiÀ>}iÊÈ°{¯ÊœvÊ̅iÊܜÀ`½Ãʓ>Àˆ˜iÊëi‡

VˆiÃ]ʜvÊ܅ˆV…Ê>««ÀœÝˆ“>ÌiÞÊÓä°Ó¯Ê>ÀiÊi˜`i“ˆVÊ­œ iÌÊ>°ÊÓä£ä®°Ê/…iÊi`ˆÌiÀÀ>˜i>˜Ê-i>ʈÃÊ>ÃœÊÀi«œÀÌi`Ê̜ LiÊ £Ê œvÊ ÎÊ Ài}ˆœ˜ÃÊ Ü…iÀiÊ V…œ˜`ÀˆV…Ì…Þ>˜ÃÊ >ÀiÊ “œÃÌ Ì…Ài>Ìi˜i`Ê­ˆi`ÊiÌÊ>°ÊÓä䙮°ÊˆvÌii˜ÊVœ>ÃÌ>ÊëiVˆià œvÊÎ>ÌiÊ­,>ˆ`>i®Ê…>ÛiÊLii˜Êˆ`i˜Ìˆvˆi`ʈ˜Ê̅iÊi`ˆÌiÀ‡

^ʘÌiÀ‡,iÃi>ÀV…ÊÓä£ÈʱÊÜÜÜ°ˆ˜Ì‡ÀiðVœ“ I/…iÃiÊ>Õ̅œÀÃÊVœ˜ÌÀˆLÕÌi`ÊiµÕ>ÞÊ̜Ê̅ˆÃÊܜÀŽ

IIœÀÀi뜘`ˆ˜}Ê>Õ̅œÀ\Êv>ÕÃ̜°Ìˆ˜ÌˆJ՘ˆLœ°ˆÌ

Population connectivity and phylogeography of the

Mediterranean endemic skate Raja polystigma and

evidence of its hybridization with the parapatric

sibling R. montagui

N. Frodella

_{*, R. Cannas}

_{*, A. Velonà}

_{, P. Carbonara}

_{, E. D. Farrell}

_,

F. Fiorentino

, M. C. Follesa

, G. Garofalo

, F. Hemida

, C. Mancusi

,

M. Stagioni

_{, N. Ungaro}

_{, F. Serena}

_{, F. Tinti}

_{**, A. Cariani}

1_{Department of Biological, Geological & Environmental Sciences (BiGeA), Laboratory of Genetics & Genomics of Marine}

Resources and Environment (GenoDREAM), University of Bologna − Campus of Ravenna, via Sant’Alberto 163, 48123 Ravenna, Italy

2_{Department of Life & Environmental Sciences (DISVA), University of Cagliari, via Fiorelli 1, 09126 Cagliari, Italy} 3_{COISPA Tecnologia & Ricerca, Via Lama Di Giotta, 70126 Bari, Italy}

4_{School of Biology & Environmental Science, Science Centre West, University College Dublin, Belfield, Dublin 4, Ireland} 5_{National Research Council (CNR) − Institute for Coastal Marine Environment (IAMC), Via L. Vaccara, 61,}

91026 Mazara del Vallo (TP), Italy

6_{Ecole Nationale Supérieure des Sciences de la Mer et de l’Aménagement du Littoral (ENSSMAL),}

Campus Universitaire de Dely Ibrahim Bois des Cars B.P. 19, 16320 Alger, Algeria

7_{Regional Agency for the Environmental Protection of Tuscany (ARPAT), Via Marradi 114, 57126 Livorno, Italy} 8_{Department of Biological, Geological & Environmental Sciences (BiGeA), Laboratory of Marine Biology & Fishery,}

University of Bologna, viale Adriatico 1/n, 61032 Fano (PU), Italy

9_{Regional Agency for the Environmental Prevention & Protection of Apulia (ARPA), RU Marine & Coastal Biology, 70126 Bari, Italy} 10_{ELASMOMED Consortium, Barcoding of Life Project, c/o Department of Biological,}

Geological & Environmental Sciences (BiGeA), University of Bologna, via Selmi 3, 40126 Bologna, Italy

-/,
/\Ê /…iÊ }i˜ïVÊ ÃÌÀÕVÌÕÀiÊ >˜`Ê «œ«Õ>̈œ˜Ê Vœ˜˜iV̈ۈÌÞÊ œvÊ Ì…iÊ i`ˆÌiÀÀ>˜i>˜Ê i˜`i“ˆV ëiVŽi`ÊÎ>ÌiÊRaja polystigma ÜiÀiʈ˜ÛiÃ̈}>Ìi`ʈ˜Ê£äÊ«œ«Õ>̈œ˜ÊÃ>“«iÃÊ­ ÊrÊÓÎÓ®Ê>ÌÊÇÊiݜ˜‡ «Àˆ“i`Ê ˜ÕVi>ÀÊ “ˆVÀœÃ>ÌiˆÌiÃÊ >˜`Ê >ÌÊ ÎÊ “ˆÌœV…œ˜`Àˆ>Ê 
Ê ÃiµÕi˜ViÊ “>ÀŽiÀÃ°Ê /…iÊ «…ޏœ}iœ‡ }À>«…ˆVÊ>˜`Ê«œ«Õ>̈œ˜Ê}i˜ïVÊ>˜>ÞÃiÃÊÀiÛi>i`Ê̅>ÌÊR. polystigma ˆ˜Ê̅iÊÜiÃÌiÀ˜Ê>˜`ÊVi˜ÌÀ> i`ˆÌiÀÀ>˜i>˜ÊÀi«ÀiÃi˜ÌÃÊ>ʘi>À‡«>˜“ˆV̈VÊ«œ«Õ>̈œ˜]Ê܈̅Ê>ÊÃÕL̏iÊLÕÌÊÈ}˜ˆvˆV>˜ÌʓˆÌœV…œ˜Ê‡ `Àˆ>Ê`ˆÛiÀ}i˜ViʜvÊ̅iÊ
`Àˆ>̈VÊ`i“i°Ê ÕVi>ÀÊ}i˜œÌÞ«iÃÊÀiÛi>i`Ê̅>ÌÊÓ°x¯ÊœvÊ̅iÊ̜Ì>Êˆ˜`ˆÛˆ`‡ Õ>ÃÊi݅ˆLˆÌi`Ê>˜Ê>`“ˆÝi`Ê>˜ViÃÌÀÞÊ܈̅Ê̅iÊÈLˆ˜}ÊëiVˆiÃÊR. montagui ­Ã«œÌÌi`ÊÀ>Þ®°Ê˜`ˆÛˆ`Õ>Ã ܈̅Ê>`“ˆÝi`Ê>˜ViÃÌÀÞÊÜiÀiÊ`iÌiVÌi`Ê>œ˜}Ê܈̅ʫÕÀiLÀi`ʈ˜`ˆÛˆ`Õ>Ãʈ˜Ê̅iÊ
}iÀˆ>˜]ÊÜÕ̅iÀ˜ /ÞÀÀ…i˜ˆ>˜]Ê-ˆVˆˆ>˜Ê>˜`Ê
`Àˆ>̈VÊR. polystigma «œ«Õ>̈œ˜ÊÃ>“«iÃ]ÊLÕÌÊ̅iÞÊÜiÀiÊ>LÃi˜ÌʜÀÊÀ>Ài ˆ˜Ê ->À`ˆ˜ˆ>˜Ê >˜`Ê ˜œÀ̅iÀ˜Ê /ÞÀÀ…i˜ˆ>˜Ê œ˜iÃ°Ê -ˆ˜ViÊ Ì…iÊ ÓÊ Ã«iVˆiÃÊ Vœ‡œVVÕÀÊ ˆ˜Ê ̅iÊ ÃœÕ̅ÜiÃÌiÀ˜ i`ˆÌiÀÀ>˜i>˜]ÊÜiÊÃÕ}}iÃÌi`Ê̅>ÌÊ̅ˆÃÊ>Ài>ʓ>ÞÊ>VÌÊ>ÃÊ>ÊÃiVœ˜`>ÀÞʅÞLÀˆ`Ê✘i°

 9Ê7",-\ÊÊ-«iVŽi`ÊÎ>Ìiʱ -«œÌÌi`ÊÀ>Þʱ -«iVˆiÃʓˆÃˆ`i˜ÌˆvˆV>̈œ˜Ê± *œ«Õ>̈œ˜ÊVœ˜˜iV̈ۈÌÞʱ
`“ˆÝi`Ê>˜ViÃÌÀÞʱ i`ˆÌiÀÀ>˜i>˜Ê± ˆÌœV…œ˜`Àˆ>Ê
ʱ ˆVÀœÃ>ÌiˆÌiÃʱ -iVœ˜`>ÀÞʅÞLÀˆ`Ê✘i