WI - Cormorant Research Group Scientific Literature - European shag updated on 02-08-2010

Scientific literature on the
Shag - Phalacrocorax aristotelis

Before 1960 - 1960/1969 - 1970/1979 - 1980 - 1981 - 1982 - 1983 - 1984 - 1985 - 1986 - 1987 - 1988 - 1989 - 1990 - 1991 - 1992 - 1993 - 1994 - 1995 - 1996 - 1997 - 1998 - 1999 - 2000 - 2001 - 2002 - 2003 - 2004 - 2005 - 2006 - 2007 - 2008 - 2009 - 2010


HOLLAND B.R., SPENCER H.G., WORTHY T.H. & KENNEDY M. 2010. Identifying Cliques of Convergent Characters: Concerted Evolution in the Cormorants and Shags. Syst. Biol., 59(4): 433–445. [Abstract. — A phylogenetic tree comprising clades with high bootstrap values or other strong measures of statistical support is usually interpreted as providing a good estimate of the true phylogeny. Convergent evolution acting on groups of characters in concert, however, can lead to highly supported but erroneous phylogenies. Identifying such groups of phylogenetically misleading characters is obviously desirable. Here we present a procedure that uses an independent data source to identify sets of characters that have undergone concerted convergent evolution. We examine the problematic case of the cormorants and shags, for which trees constructed using osteological and molecular characters both have strong statistical support and yet are fundamentally incongruent. We find that the osteological characters can be separated into those that fit the phylogenetic history implied by the molecular data set and those that do not. Moreover, these latter nonfitting osteological characters are internally consistent and form groups of mutually compatible characters or “cliques,” which are significantly larger than cliques of shuffled characters.We suggest, therefore, that these cliques of characters are the result of similar selective pressures and are a signature of concerted convergence.]




Álvarez D. & Velando A. 2007. El cormorán moñudo en España. Población en 2006-2007 y método de censo. SEO/BirdLife. Madrid.


Martínez-Abraín A., Velando A., Genovart M., Gerique C., Bartolomé M. A., Villuendas E., Sarzo B. & Oro, D. 2006. Sex-specific mortality of European shags during an oil spill: demographic implications for the recovery of colonies. Mar. Ecol. Progr. Ser., 318: 271-276.


Velando A., Álvarez D., Mouriño J. et al. 2005 Population trends and reproductive success of the European shag Phalacrocorax aristotelis on the Iberian Peninsula following the Prestige oil spill. Journal of Ornithology, 146(2): 116-120.

Velando A., Munilla I. & Leyenda P. M. 2005. Short-term indirect effects of the Prestige oil spill on a marine top predator: changes in prey availability for European shags. Mar. Ecol. Progr. Ser., 302: 263–274.


Wanless S. & Harris M. P. 2004. European Shag Phalacrocorax aristotelis. In> P. I. Mitchell, S. F. Newton, N. Ratcliffe & T. E. Dunn (eds.). Seabird populations of Britain and Ireland. T & D Poyser. London.


Álvarez D. 2003. Reproducción del cormorán moñudo (Phalacrocorax aristotelis) en Asturies en 1997: censo de colonias, distribución espacial y fenología reproductiva. El Draque, 4: 3-11.


Velando A. & Freire J. 2002. Population modelling of European shag at their southern limit: conservation implications. Biological Conservation, 107: 59-69.

Velando A., Graves J. & Ortega-Ruano. 2002. Sex ratio in relation to timing of breeding, and laying sequence in a dimorphic seabird. Ibis, 144: xx-xx.


Abollo E., Gestal C. & Pascual S. 2001. Anisakid infection in the European shag Phalacrocorax aristotelis aristotelis. JOURNAL OF HELMINTHOLOGY, 75(3): 209-214.

Daunt F., Monahan P., Wanless S. & Harris M. P. 2001. Parental age and offspring ectoparasite load in European Shags Stictocarbo aristotelis. Ardea, 89(3): 449-455.

Velando A. 2001 Postfledging crèche behaviour in European shag. J. Ethology, 19: 116-122.

Velando A. & Freire J. 2001. Can the central-periphery distribution become general in seabird colonies? Nest spatial pattern in the European Shag. Condor, 103: 544-554.


Debout G. 2000. Les conséquences de la nidification du Grand Cormoran Phalacrocorax carbo sur celle du Cormoran huppé Phalacrocorax aristotelis. Alauda, 68 (1): 1-9.

Harris M. P., S. Wanless & Rothery P. 2000. Adult survival rates of Shag Phalacrocorax aristotelis, Common Guillemot Uria aalge, Razorbill Alca torda, Puffin Fratercula arctica and kittiwake Rissa tridactyla. Atlantic Seabirds 2: 133-150.

Velando A. 2000. The importance of the hatching date on dominace of young shags. Anim. Behav. 60:181-185.

Velando A., J.Graves & Freire J. 2000. Sex-specific growth in the European Shag Stictocarbo aristoteles, a sexually dimorphic seabird. Ardea, (88)2: 127-136.


Murvoll K.M., J.U. Shake, V.H. Nilssen, C. Bech, J.E. Ostnes & Jenssen B.M. 1999. Yolk PCB and plasma retinol concentration in Shag (Phalacrocorax aristotelis) hatchlings. Archives of Environmental Contamination and Toxicology, 36: 308-315.

Velando A., F. Docampo & Álvarez D. 1999. Status of European shag population on the Atlantic coast of the Iberian Peninsula. Atlantic Seabirds, 1(3): 105-114.

 Velando A. & Freire J. 1999. Intercolony and seasonal differences in the breeding diet of European shags on the galician coast (NW Spain). Marine Ecology Progress Series, 188: pp. 225

Velando A. & Freire J. 1999. Coloniabilidad y conservación de aves marinas: el caso del cormorán moñudo. Etología, 7: 55–62.

Velando A., Ortega-Ruano J. E. & Freire J. 1999. Chick mortality in European shag Stictocarbo aristotelis related to food limitations during adverse weather events. Ardea, 87: 51-59.


Álvarez D. 1998. The diet of Shags Phalacrocorax aristotelis in the Cantabrian Sea, northern Spain, during the breeding season. Seabird, 20: 22-30.

 Beskaravainyi, M.M., Kostin & Yu S. 1998. [Distribution, numbers and some peculiarities of the breeding ecology of the Shag and the Herring Gull in the South-Eastern Crimea]. // Berkut. 7 (1-2). 25-29. Tabl.2. Bibl. 7 titl. In Russian with Engl. summary. [rif. 1998-1]

Grémillet D. & Argentin G. 1998. Cormorants, Shags and fisheries in the Chausey Islands area. Le Cormoran, 10 : 196-202.

Grémillet D., G. Argentin, B. Schulte & Culik B. M. 1998. Flexible foraging techniques in breeding cormorants Phalacrocorax carbo and shags Phalacrocorax aristotelis: benthic or pelagic feeding? Ibis, 140: 113-119.

Grémillet D., L. Tuschy & Kierspel M. 1998. Body temperature and insulation in diving Great Cormorants and European Shags. Funct. Ecol, 12: 386-394.

Harris M.P., S. Wanless & Elston D.A. 1998. Age-related effects of a non -breeding event and a winter wreck on the the survival of shags Phalacrocorax aristotelis. Ibis, 140: 310-314.

Wanless S., D. Grémillet & Harris M.P. 1998. Foraging activity and performance of Shag Phalacrocorax aristotelis in relation to environmental characteristics. Journal of Avian Biology, 29: 49-54. --- Using radiotelemetry and automatic nest balance system in four pairs of shags in Scotland. Distribution of available food resources may influence choice of foraging areas. ---


Calvo B. & Bolton M. 1997. Sexing Shags Phalacrocorax aristotelis from external measurements using discriminant analysis. Ringing & Migration, 18: 50-56.

Østnes J. E. & Bech C. 1997. The early emergence of cold sensation in Shag nestlings Phalocrocorax aristotelis. Journal Avian Biology, 28: 24-30.

Neto J. 1997. [Contribution to the breeding biology of the Shag Phalacrocorax aristotelis at Berlenga Islands.] Airo, 8: 16-24. (In Portuguese with English abstract).

Wanless S., Grémillet D., Hilton G. M. & Harris M. P. 1997. Estimation of digestion and defaecation rate in the Shag Phalacrocorax aristotelis. Seabird, 19: 47-50.

Wanless S. & Harris M. P. 1997. Successful double-brooding in European Shags. Colonial Waterbirds, 20(2): 291-294.

Wanless S., et al. 1997. Modeling environmental and energetic effects on feeding performance and distribution of shags (Phalocrocorax aristotelis): integrating telemetry, geographical information systems and modeling techniques. ICES J. Marine Sci. 54: 524--544.


Grémillet D., R. Dey, S. Wanless, M. P. Harris & Regel J. 1996. Determining food intake by Great Cormorants and European Shags with electronic balances. J. Field Ornithol., 67: 637-648.

Lea S. E. G. et al. 1996. Diving patterns in shags and cormorants (Phalacrocorax): tests of an optimal breathing model. Ibis, 138: 391--398. (Dept. Psychol., Univ. Exeter, Washington Singer Lab., Exeter EX4 4QG, UK.)---Comparative study between Phalacrocorax aristotelis, Phalacrocorax melanoleucos, Phalacrocorax carbo and Phalacrocorax varius.---


Álvarez Fernández D., A. Fernández Gutiérrez & García-Rovés J.F. 1995. El Cormorán Moñudo en el Concejo de Cudillero. Fundación Selgas-Fagalde & Ayuntamiento de Cudillero. 77 pp. (monograph).

Russell, A.F., S. Wanless & M.P. Harris. 1995. Factors affecting the production of pellets by Shags, Phalacrocorax aristotelis. Seabird 17: 44-49.


Harris M.P., Buckland S.T., Russell S.M. & Wanless S. 1994. Year- and age-related variation in the survival of adult european shag over a 24-year period. Condor, 96: 600-605

Harris M.P., Buckland S.T., Russell S.M. & Wanless S. 1994. Post fledging survival to breeding age of shags Phalacrocorax aristotelis in relation to year, date of fledging and brood size. Journal Avian Biology, 25: 268-274.


Carss D.N. 1993. Shags Phalacrocorax aristotelis at cage fish farms in Argyll, western Scotland. Bird Study, 40: 203-211.

Greenstreet S.P.R., Morgan R.I.G., Barnett S. & Redhead P. 1993. Variation in the numbers of shags, Phalacrocorax aristotelis, and common seals, Phoca vitulina, near the mouth of an Atlantic salmon, Salmo salar, river at the time of the smolt run. J. Anim. Ecol., 62: 565-576.

Harris M.P. & Wanless S. 1993. The diet of Shags Phalacrocorax aristotelis during the chick rearing period assessed by three methods. Bird Study 40: 135-139.

Wanless S., Corfield M.P., Harris M.P., Buckland S.T. & Morris J.A. 1993. Diving behaviour of the shag Phalacrocorax aristotelis (Aves: Pelecaniformes) in relation to water depth and prey size. J. Zool. Lond., 231: 11-25.


Aebischer, N J. & S. Wanless 1992. Relationships between colony size, adult non-breeding and environmental conditions for Shags Phalacrocorax aristotelis on the Isle of May, Scotland. Bird Study, 39: 43-52.

Carss D. N. & Marquiss M. 1992. Cormorants and the Loch Leven Trout Fishery. 37pp. Scottish Natural Heritage.

Graves J., R. T. Hay, M. Scallan & Rowe S. 1992. extra-pair paternity in the Shag, Phalacrocorax aristotelis, as determined by DNA fingerprinting. J. Zool., 226: 399-408.


Harris M. P. &Wanless S. 1991. The importance of the Lesser Sandeel Ammodytes marinus in the diet of the Shag Phalacrocorax aristotelis. Ornis Scandinavica, 22: 375-382.

Wanless S., A. E. Burger & Harris M. P. 1991. Diving depths of Shags Phalacrocorax aristotelis breeding on the Isle of May. Ibis, 133: 37-42.

Wanless S., M. P. Harris & Morris J. A. 1991. Foraging range and feeding locations of Shags Phalacrocorax aristotelis during chick rearing. Ibis, 133: 30-36.


Amundsen T. andStokland J. N. 1990. Egg size and parental quality influence nestling growth in the shag. Auk 107:410-13.

Guyot I. 1990. Le cormoran huppé en Corse: biologie et interactions avec la peche professionnelle. Trav. Sci. Parc. Nat. Rég. Nat. Corse, 28: 1-40.

Johnstone I. P. Harris, S. Wanless and Graves J. A. 1990. The uselfulness of pellets for assessing the diet of adult Shags Phalacrocorax aristotelis. Bird Study, 37: 5-11.

Pasquet E. & Monnat J.Y. 1990. Dispersion géographique des Cormorans huppés juvéniles de la mer Celtique. L´Oiseau et R.F.O., 60: 94-109.

Røv N. 1990. Population studies of Shags Phalacrocorax aristotelis in Norway. Nina Forskningsrapport., 7: 1-28.



Amundsen, T. and J N. Stokland 1988. Adaptative significance of asynchronous hayching in the Shag: a test of brood reduction hypothesis. J. Anim. Ecology, 57: 329-344.


Harris, M. P. y Forbes, R. 1987. The effect of date on counts of nests of Shags Phalacrocorax aristotelis. Bird Study, 34: 187-190.


Aebischer, N J. 1986. Retrospective investigation of an ecological disaster in the Shag, Phalacrocorax aristotelis: a general method based on long-term marking. J. Anim. Ecol, 55: 613-629.

Galbraith H., S. Baillie, R. W. Furness & Russell S. 1986. Regional variations in the dispersal patterns of Shags Phalacrocorax aristotelis in northern Europe. Ornis. Scand., 17: 68-74.

Wilson J. G. & Early J. J. 1986. Pesticide and PCB levels in the eggs of shag Phalacrocorax aristotelis and cormorant P. carbo from Ireland. Environ. Pollut. Ser. B, 12: 15-26.


Alström P. 1985. Artbestamning av storskarv Phalacrocorax carbo och toppskarv Ph. aristotelis. Vår Fåglevärld, 44: 325-350.



Godó G. 1983. [The populations of shags Phalacrocorax aristotelis in Sogn and Fjordane, western Norway]. Vár Fuglefauna 6:88-92. (Norwegian, English summary.)



Galbraith H., S. Russell & Furness R. W. 1981. Movements and mortality of isle of May Shags as shown by ringing recoveries. Ringing & Migration, 3: 181-189.


Potts G. R., J. C. Coulson and Deans I. R. 1980. Population dynamics and breeding success of the Shag, Phalacrocorax aristotelis, on the Farne Islands, Northumberland. J. Anim. Ecol., 49: 465-484

1970 - 1979

Cooke A. S. 1979. Egg shell characteristics of Gannets Sula bassana, Shags Phalacrocorax aristotelis and Great Black-backed Gulls Larus marinus exposed to DDE and other environmental pollutants. Environ. Pollut., 19: 47-65.

Gardarsson A. 1979. [A census of breeding great cormorants and European shags in Iceland in 1975.] Ndttúrufraedingurinn 49:126-54. (Icelandic, English summary.)

Johansen H. 1975. The relationship between breeding grounds and wintering grounds in the shag, Phalacrocorax aristotelis, as shown by the ringing recoveries. Sterna 14:1-21.

King B. 1972. Feeding and aerial diving by shags. Brit. Birds 65:480-81.

Potts G R. 1971. Moult in the Shag Phalacrocorax aristotelis, and the ontogeny of the "Staffelmauser". Ibis, 113: 298-305.

1960 - 1969

Coulson J.C. 1961. Movements and seasonal variation in mortality of shags and cormorants ringed on the Farne Islands, Northumberland. Brit. Birds, 54: 225-235.

Coulson, J C., G R. Potts And J. Horobin 1969. Variation in eggs of the Shag (Phalacrocorax aristotelis). Auk, 86: 232-245.

Potts G. R. 1969. The influence of eruptive movement, age, population size and other factors in the survival of the shag, Phalacrocorax aristotelis. J. Anim. Ecol., 38:53-102.

Rae B. B. 1969. The food of cormorants and shags in Scottish estuaries and coastal waters. Mar. Res., 1: 1-16.

Snow B. 1960. The breeding biology of the Shag Phalacrocorax aristotelis on the island of Lundy, Bristol Channel. Ibis, 102: 554-575.

Snow B. 1963. Behaviour of the shag. Brit. Birds 56:77-103, 164-186.

Before 1960

Lack D. 1945. The ecology of closely related species with special reference to Cormorant and Shag. J. Anim. Ecol., 14: 12-16.

Lumsden W. H. R. and Haddow A. J. 1946. The food of the shag (Phalacrocorax aristotelis) in the Clyde Sea area. J. Anim. Ecol., 15:35-42.

Steven C. A. 1933. The foods consumed by shags and cormorants around the shores of Cornwall (England). J. Mar. Biol. Asso. U.K., 19:277-92.

Page create on 10/11/2000
S. Volponi