The Ecological Role of Loligo forbesii in the Moray Firth Ecosystem, Northeast Scotland
The squid Loligo forbesii is suspected to be an important species in marine food webs, as it can strongly impact its prey and be impacted upon by predation, competition, fishing and/or climate variability. To quantify these impacts in the food web, the measurement of its trophic position and ecological role within well-studied ecosystems is essential. An Ecopath model was balanced and run for the Moray Firth ecosystem and was used to investigate the significance of this squid’s trophic roles. The network analysis routine included in Ecopath with Ecosim (EwE) was used to estimate trophic interaction, system indicators (health condition and developmental stage) and food web features. Results indicated that within the Moray Firth squid occupy a top trophic position in the food web and also a major prey item for many other species. Results from Omnivory Index (OI) showed that squid is a generalized feeder transferring energy across wide trophic levels and is more important as a predator than that as a prey in the Moray Firth ecosystem. The results highlight the importance of taking squid into account in the management of Europe’s living marine resources.
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 J. S. Link, C. A. Griswold, E. Methratta, and J. Gunnard, Documentation for the energy modeling and analysis exercise (EMAX). Northeast Fisheries Science Center Reference Document, 2006, pp. 06–15.
 F. R. B. Canada, and A. Vovk, Method of Determining Maturing Stages in Gonads of the Squid, Loligo Pealei. Department of the Environment, Fisheries Research Board of Canada, 1973
 A. Tibbetts, Squid fisheries (Loligo pealei and Illex illecebrosus) off the northeastern coast of the United States of America, 1963-1974. Int. Comm. Northwest Atl. Fish., Sel. Pap, vol. 2, 1977, pp. 85–109.
 H. I. Daly, G. J. Pierce, M. B. Santos, J. Royer, S. K. Cho, G. Stowasser, J. Robin, et al. Cephalopod consumption by trawl caught fish in Scottish and English Channel waters. Fisheries Research, vol, 52, 2001, pp. 51–64.
 C. M. Nigmatullin, K. Nesis, and A. Arkhipkin, A review of the biology of the jumbo squid Dosidicus gigas (Cephalopoda: Ommastrephidae). Fisheries Research, vol. 54, 2001, pp. 9–19.
 P. Boyle, and P. Rodhouse, Cephalopods: ecology and fisheries. Wiley. com. 2008.
 M. Coll, J. Navarro, R. J. Olson, and V. Christensen, Assessing the trophic position and ecological role of squids in marine ecosystems by means of food-web models. Deep Sea Research Part II: Topical Studies in Oceanography, 2012.
 T. Amarantunga, The role of cephalopods in the marine ecosystem: In assessment of world cephalopod resources, Pp 379-415 Ed by J. Caddy. FAO Fisheries Technical Paper, 1983, pp. 231–452.
 S. Wangvoralak, L. C. Hastie, and G. J. Pierce, Temporal and ontogenetic variation in the diet of squid (Loligo forbesii Streenstrup) in Scottish waters. Hydrobiologia, vol. 54, 2011, pp. 223–240.
 I. Young, G. Pierce, G. Stowasser, M. Santos, J. Wang, P. Boyle, P. Shaw, et al. The Moray Firth directed squid fishery. Fisheries Research, vol. 78, 2006, pp. 39–43.
 S. Wangvoralak, Life history and ecological importance of veined squid Loligo forbesii in Scottish waters. PhD Theses, University of Aberdeen, Aberdeen, UK, 2012. 227p. Unpublished.
 M. Smale, Cephalopods as prey. IV. Fishes. Philosophical Transactions of the Royal Society of London. Series B: Biological Sciences, vol. 351, 1996, pp 1067–1081.
 M. R. Clarke, The role of cephalopods in the world’s oceans: an introduction. Philosophical Transactions: Biological Sciences, vol. 351, 1996, pp. 979–983.
 P. Battaglia, F. Andaloro, P. Consoli, V. Esposito, D. Malara, S. Musolino, C. Pedà, et al. Feeding habits of the Atlantic bluefin tuna, Thunnus thynnus (L. 1758), in the central Mediterranean Sea (Strait of Messina). Helgoland Marine Research, vol. 67, 2013, pp. 97–107.
 W. Bowen, Reconstruction of pinniped diets: accounting for complete digestion of otoliths and cephalopod beaks. Canadian Journal of Fisheries and Aquatic Sciences, vol.67, 1996., 2000, pp. 898–905.
 W. K. Macy, Feeding Patterns of the Long-Finned Squid, Loligo pealei, in New England Waters. The Biological Bulletin, vol. 162, 1982, pp. 28–38.
 S. Mackinson, and G. Daskalov, The ecosystem model of the North Sea to support an ecosyetm approach to fisheries management:discription and parameterisation. Technical report, Cepas Lowestoft, 2007, 142p.
 G. J. Pierce, P. R., Boyle, L. C. Hastie, and L. Key, The life history of Loligo forbesi (Cephalopoda: Loliginidae) in Scottish waters. Fisheries Research, vol. 21, 1994, pp. 17–41.
 J. C. Xavier, Y. Cherel, J. Roberts, and U. Piatkowski, How do cephalopods become available to seabirds: Can fish gut contents from tuna fishing vessels be a major food source of deep-dwelling cephalopods? ICES Journal of Marine Science, vol. 70, 2013, pp. 46–49.
 J. F. Bromaghin, M. M. Lance, E. W. Elliott, S. J. Jeffries, A. Acevedo-Gutiérrez, and J. M. Kennish, New insights into the diets of harbor seals (Phoca vitulina) in the Salish Sea revealed by analysis of fatty acid signatures. Fishery Bulletin, vol. 111, 2013, pp. 13–26.
 M. Santos, G. Pierce, A. López, R. Reid, V. Ridoux, and E. Mente, Pygmy sperm whales Kogia breviceps in the Northeast Atlantic: New information on stomach contents and strandings. Marine Mammal Science, vol. 22, 2006, pp. 600–616.
 R. W. Furness, and M. L. Tasker, Diet of seabirds and consequences of changes in food supply. ICES Co-operative Research Reports, vol. 232, 1999, pp. 1- 66.
 G. Pierce, A. Miller, P. Thompson, and J. Hislop, Prey remains in grey seal (Halichoerus grypus) faeces from the Moray Firth, north-east Scotland. Journal of Zoology, vol. 224, 1991, pp. 337–341.
 G. Pierce, N. Bailey, Y. Stratoudakis, and A. Newton, Distribution and abundance of the fished population of Loligo forbesi in Scottish waters: analysis of research cruise data. ICES Journal of Marine Science, vol. 55, 1998, pp. 14–33.
 M. Santos, G. Pierce, P. Boyle, R. Reid, H. Ross, I. Patterson, C. Kinze, et al. Stomach contents of sperm whales Physeter macrocephalus stranded in the North Sea 1990-1996. Marine Ecology Progress Series, vol. 183, 1999, pp. 281–294.
 P. R. Boyle, and G. J. Pierce, Fishery biology of northeast Atlantic squid: an overview. Fisheries Research, vol. 21, 1994, pp. 1–15.
 G. J. Pierce, P. R. Boyle, L. C. Hastie, and A. M. Shanks, Distribution and abundance of the fished population of Loligo forbesii in UK waters: analysis of fishery data. Fisheries Research, vol. 21, 1994, pp. 193–216.
 Nixon, M. “Cephalopod Diet”. Cephalopod Life Cycles, vo. 2, 1987, pp. 201 – 219.
 M. Collins, S. De Grave, C. Lordan, G. Burnell, and P. Rodhouse, Diet of the squid Loligo forbesi Steenstrup (Cephalopoda: Loliginidae) in Irish waters. ICES Journal of Marine Science: Journal du Conseil, vol. 51, 1994, pp. 337–344.
 Scottish government: http://www.scotland.gov.uk/Topics/marine/Sea-Fisheries/InshoreFisheries/ScallopReview. 23rd Dec. 2014.
 L. C. Hastie, G. J. Pierece, C. Pita, M. Viana, J. M. Smith, and S. Wangvoralak, Squid fishing in UK waters. A Report to SEAFISH Industry Authority, 2009.
 I. D. Tuck, S. J. Hall, M. R. Robertson, E. Armstrong, and D. J. Basford, Effects of physical trawling disturbance in a previously unfished sheltered Scottish sea loch. Marine Ecology Progress Series, vol. 162, 1998, pp. 227–242.
 J. H. Churchill, The effect of commercial trawling on sediment resuspension and transport over the Middle Atlantic Bight continental shelf. Continental Shelf Research, vol. 9, 1989, pp. 841–865.
 S. Jennings, T. A. Dinmore, D. E. Duplisea, K. J. Warr, and J. E. Lancaster, Trawling disturbance can modify benthic production processes. Journal of Animal Ecology, vol. 70, 2001, pp. 459–475.
 G. Pierce, and P. Boyle, Empirical modelling of inter annual trends in abundance of squid (Loligo forbesii) in Scottish waters. Fisheries Research, vol. 59, 2003, pp. 305–326.
 G. J. Pierce, and M. B. Santos, Trophic interactions of squid Loligo forbesi in Scottish waters. , 1996, pp. 58-64. http://www.scopus.com/inward/record.url?eid=2-s2.0-0030453972&partnerID=40&md5=8d9a9a8b0d6e07cc7ec70e956e2c6791.
 V. Christensen, and C. J. Walters, Ecopath with Ecosim: methods, capabilities and limitations. Ecological modelling, vol. 172, 2004, pp. 109–139.
 A. Bundy, Structure and functioning of the eastern Scotian Shelf ecosystem before and after the collapse of ground fish stocks in the early 1990s. Canadian Journal of Fisheries and Aquatic Sciences, vol. 62, 2005, pp. 1453–1473.
 A., M. Hernando, Jr. and E. Flores, The Phillipines Squid Fishery: A Review. Marine Fesheries Review: pp. 13–20, 1981.
 J. Cornejo-Donoso, and T. Antezana, Priliminary trophic model of the Antartic Peninsula Ecosystem (Sub-area CCAMLR 48.1), 2008.
 S. P. Cox, T. E. Essington, J. F. Kitchell, S. J. Martell, C. J. Walters, C. Boggs, and I. Kaplan, Reconstructing ecosystem dynamics in the central Pacific Ocean, 1952 1998. II. A preliminary assessment of the trophic impacts of fishing and effects on tuna dynamics. Canadian Journal of Fisheries and Aquatic Sciences, vol. 59, 2002, pp. 1736–1747.
 T. Kooten, and C. Kok, The Mackinson-Daskalov North Sea EcoSpace model as a simulation tool for spatial planning scenarios, 2011.
 I. Young, G. Pierce, H. Daly, M. Santos, L. Key, N. Bailey, J.-P. Robin, et al. Application of depletion methods to estimate stock size in the squid Loligo forbesii in Scottish waters (UK). Fisheries Research, vol. 69, 2004, pp. 211–227.
 Scottish government, Scottish sea fisheries statistics 2009. Edinburgh, Scottish government. Scottish Natural Heritage, 2006. Moray Firth Special Area of Conservation. Scottish Natural Heritage. 1-16 pp. Available from: http://www.ukmpas.org/mpareferences/viewreference.php?refid=317, 2009, on 29 Jun 2013.
 Scottish Natural Heritage, Moray Firth Special Area of Conservation. Scottish Natural Heritage. 1-16 pp. Available from: http://www.ukmpas.org/mpareferences/viewreference.php?refid=317, 2006, on 19 Aug 2013.
 D. Eisma, The North Sea: an overview. Phil. Trans. R. Sco. Lond. B vol. 316 no. 1181, 1987, pp. 461-485.
 J. Araújo, S. Mackinson, J. Ellis, and P. Hart, An Ecopath model of the western English Channel ecosystem with an exploration of its dynamic properties. Centre for Environment, Fisheries and Aquaculture Science, 2005.
 K. Lees, and S. Mackinson, An Ecopath model of the Irish Sea: ecosystems properties and sensitivity analysis. CEFAS Lowestoft. Science Series Technical Report, 138p, 2007.
 V. Christensen, C. J. Walters, D. Pauly, et al., Ecopath with Ecosim: a user’s guide. Fisheries centre University of British Columbia, Vancouver. November 2005 edition, 154p.
 V. Christensen, C. J. Walters, D. Pauly, and R. Forrest, Ecopath with Ecosim version 6: User Guide. November-2008. Lenfest Ocean Futures Project 2008, 235p.
 D. Pauly, V. Christensen, and C. Walters, Ecopath, Ecosim, and Ecospace as tools for evaluating ecosystem impact of fisheries. ICES Journal of Marine Science: Journal du Conseil, vol. 57, 2000, pp. 697–706.
 J. J. Polovina, Model of a coral reef ecosystem. Coral reefs, vol.3, 1984, pp. 1–11.
 V. Christensen, C. Walters, and D. Pauly, Ecopath with Ecosim Version 4. Help system, 2000.
 W. Odum, E. J. Heald, and L. Cronin, Estuarine research. Estuarine research, 1975.
 V. Christensen, and D. Pauly, ECOPATH II—a software for balancing steady-state ecosystem models and calculating network characteristics. Ecological modelling, vol. 61, 1992, pp. 169–185.
 J. Chesson, The Estimation and Analysis of Preference and Its Relationship to Foraging Models. Ecology: 1983, pp. 1297–1304.
 R. Ulanowicz, Quantitative Methods for Ecological Network Analysis and Its Application to Coastal Ecosystems, 2011.
 R. Ulanowicz, and C. Puccia, Mixed trophic impacts in ecosystems. Coenoses, vol. 5, 1990, pp. 7–16.
 P. Rodhouse, and C. M. Nigmatullin, Role as consumers. Philosophical Transactions of the Royal Society of London. Series B: Biological Sciences, vol. 351, 1996, pp. 1003–1022.
 M. A. Gasalla, A. R. Rodrigues, and F. A. Postuma, The trophic role of the squid Loligo plei as a keystone species in the South Brazil Bight ecosystem. ICES Journal of Marine Science: Journal du Conseil, vol. 67, 2010, pp. 1413–1424.
 D. Pauly, V. Christensen, S. Guénette, T. J. Pitcher, U. R. Sumaila, C. J. Walters, R., Watson, et al. Towards sustainability in world fisheries. Nature, vol. 418, 2002, pp. 689–695.
 S. Cadrin, and E. Hatfield, Stock assessment of longfin inshore squid, Loligo pealeii. Northeast Fisheries Science Center Reference Document: 1999, pp. 99–12.
 E. M. Hatfield, R. T. Hanlon, J. W. Forsythe, and E. P. Grist, Laboratory testing of a growth hypothesis for juvenile squid Loligo pealeii (Cephalopoda: Loliginidae). Canadian Journal of Fisheries and Aquatic Sciences, vol. 58, 2001, pp. 845–857.
 J. S. Link, and L. P. Garrison, Trophic ecology of Atlantic cod Gadus morhua on the northeast US continental shelf. Marine Ecology Progress Series, vol. 227, 2002, pp. 109–123.
 J. Navarro, M. Coll, M. Louzao, I. Palomera, A. Delgado, and M. G. Forero, Comparison of ecosystem modelling and isotopic approach as ecological tools to investigate food webs in the NW Mediterranean Sea. Journal of Experimental Marine Biology and Ecology, vol. 401, 2011, 97–104.
 S. Neira, and H. Arancibia, Food web and fish stock changes in central Chile: Comparing the roles of jumbo squid (Dosidicus gigas) as predator, the environment, and fishing. Deep-Sea Research Part II: Topical Studies in Oceanography. www.scopus.com, 2013.
 R. Rosas-Luis, C. Salinas-Zavala, V. Koch, P. Luna, and M. Morales-Zárate, Importance of jumbo squid Dosidicus gigas (Orbigny, 1835) in the pelagic ecosystem of the central Gulf of California. Ecological modelling, vol. 218, 2008, pp. 149–161.
 J. Stevens, R. Bonfil, N. Dulvy, and P. Walker, The effects of fishing on sharks, rays, and chimaeras (chondrichthyans), and the implications for marine ecosystems. ICES Journal of Marine Science: Journal du Conseil, vol. 57, 2000, pp. 476–494.
 F. Sergio, I. Newton, and L. Marchesi, Conservation: Top predators and biodiversity. Nature, vol. 436, 2005, pp. 192–192.
 C. F. Roper, M. J. Sweeney, and C. E. Nauen, FAO species catalogue. v. 3: Cephalopods of the world. An annotated and illustrated catalogue of species of interest to fisheries, 1984.
 G. J. Pierce, P. R. Boyle, L. C. Hastie, and A. M. Shanks, Distribution and abundance of the fished population of Loligo forbesii in UK waters: analysis of fishery data. Fisheries Research, vol. 21. 1994, pp. 193–216.
 E.P. Odum, The strategy of ecosystem development. Science vol. 104, 1969, pp. 262– 270,
 S. Neira, and H. Arancibia, Trophic interactions and community structure in the upwelling system off Central Chile (33–39 S). Journal of Experimental Marine Biology and Ecology, 2004, 312: 349–366.
 B. Wilson, P. Thompson, and P. S. Hammond, Habitat use by bottlenose dolphins: Seasonal distribution and stratified movement patterns in the Moray Firth, Scotland. Journal of the Applied Ecology, vol. 34, 1997, pp. 1365–1374.
 W. F. Perrin, B. Wursig, and J. Thewissen, Encyclopedia of marine mammals. Access Online via Elsevier. 22 Jul 2013, 2009.
 J. Caddy, and P. Rodhouse, Cephalopod and groundfish landings: evidence for ecological change in global fisheries? Reviews in Fish Biology and Fisheries, vol. 8, 1998, pp. 431–444.