Oceanic Drivers of Sei Whale Distribution in the North Atlantic
This study investigated the oceanic drivers of sei whale (Balaenoptera borealis) distribution in the central and eastern North Atlantic, and explored how distribution may have changed over almost three decades. Cetacean sightings data were available from Icelandic, Faroese and Norwegian surveys conducted throughout the central and eastern North Atlantic during summer between 1987 and 2015. Effective strip half width was estimated from the data to take account of variation in detection probability. Spatially-referenced environmental variables used as predictors in generalised additive models of sei whale relative density included: relief-related variables seabed depth, slope and aspect; monthly-varying physical oceanographic variables sea surface temperature (SST), mixed layer depth, bottom temperature, salinity, and sea surface height anomaly (SSH); and monthly-varying biological oceanographic variables chlorophyll-a concentration and primary productivity. Preliminary analysis considered which month (March-August) in the dynamic oceanographic variables explained most variability in sei whale density. Models including all variables (“full models”) could only be run for 1998-2015 because data for several variables were missing in earlier years. “Simple models" including only relief-related variables and SST were therefore run for 1987-89, and also for 1998-2015 for comparison. The best-fitting full model for 1998-2015 retained the covariates depth, May SST, May bottom temperature, July salinity, July SSH and July primary productivity. Of these, depth, May SST and July SSH were the strongest predictors of sei whale density. In the simple models for both 1987-89 and 1998-2015, depth (especially), May SST and seabed slope were the strongest predictors of sei whale density. The highest densities of sei whales were predicted in the Irminger Sea and over the Charles-Gibbs Fracture Zone; a pattern driven by large negative SSH, deep water (>1500m) and polar-temperate SST (5-12oC). There was some inter-annual variability in predicted distribution and there appears to be a northward expansion in distribution consistent with prey species responding to ocean warming. The models could be used to predict future distribution of sei whales based on future environmental conditions predicted by climate models.
Akaike, H. (1973). Information Theory and an Extension of the Maximum Likelihood Principle. In B. Petran and F. Csaaki (Eds.), Proceeding of the Second International Symposium on Information Theory (pp. 267–281). Budapest: Akadeemiai Kiadi.
Biggs, D.C., Zimmerman, R.A., Gasca, R., Suarez-Morales, E., Castellanos, I. and Leben, R.R. (1997). Note on plankton and cold-core rings in the Gulf of Mexico. Fishery Bulletin, 95(2), 369-375.
Borchers, D. and Burt, M. (1997). Sei and fin whale abundance in the North Atlantic, estimated from NASS-95 shipboard survey data. NAMMCO Scientific Committee Working Paper, SC/5/AE/1.
Brodie, P. and Víkingsson, G. (2009). On the feeding mechanism of the sei whale (Balaenoptera borealis). Journal of Northwest Atlantic Fishery Science, 42, 49–54. https://doi.org/10.2960/J.v42.m646
Buckland, S.T., Rexstad, E.A., Marques, T.A. and Oedekoven, C.S. (2015). Distance sampling: methods and applications. New York, USA: Springer. https://doi.org/10.1007/978-3-319-19219-2
Cattanach, K.L., Sigurjónsson, J., Buckland, S.T. and Gunnlaugsson, T. (1993). Sei whale abundance in the North Atlantic, estimated from NASS-87 and NASS-89 data. Reports of the International Whaling Commission, 43, 315-321.
Christensen, I., Haug, T. and Øien, N. (1992). A review of feeding and reproduction in large baleen whales (Mysticeti) and sperm whales Physeter macrocephalus in Norwegian and adjacent waters. Fauna Norvegica Series A, 13, 39-48.
Collet, R. (1886). On the External Characteristics of the Rudolphi’s Rorqual (Balaenoptera borealis). Proceedings of the Zoological Society of London, 243–265.
Cooke, J.G. (2018). Balaenoptera borealis. The IUCN Red List of Threatened Species 2018: e.T2475A130482064.
Friedland, K.D., Record, N.R., Asch, R.G., Kristiansen, T., Saba, V.S., Drinkwater, K.F., Henson, S., Leaf, R.T., Morse, R.E., Johns, D.G. and Large, S.I. (2016). Seasonal phytoplankton blooms in the North Atlantic linked to the overwintering strategies of copepods. Elementa: Science of the Anthropocene, 4. https://doi.org/10.12952/journal.elementa.000099
Gavrilchuk, K., Lesage, V., Ramp, C., Sears, R., Bérubé, M., Bearhop, S. and Beauplet, G. (2014). Trophic niche partitioning among sympatric baleen whale species following the collapse of groundfish stocks in the Northwest Atlantic. Marine Ecology Progress Series, 497, 285-301. https://doi.org/10.3354/meps10578
Häkkinen, S., Rhines, P.B. and Worthen, D.L. (2013). Northern North Atlantic sea surface height and ocean heat content variability. Journal of Geophysical Research Oceans, 118, 3670–3678. https://doi.org/10.1002/jgrc.20268
Hátún, H., Payne, M.R., Beaugrand, G., Reid, P.C., Sandø, A.B., Drange, H., Hansen, B., Jacobsen, J.A. and Bloch, D. (2009). Large bio-geographical shifts in the north-eastern Atlantic Ocean: From the subpolar gyre, via plankton, to blue whiting and pilot whales. Progress in Oceanography, 80, 149–162. https://doi.org/10.1016/j.pocean.2009.03.001
Horwood, J. (1987). The Sei Whale: Population Biology, Ecology and Management. London, UK: Croon Helm.
Horwood, J. (2009). Sei whale: Balaenoptera borealis. In W.F. Perrin, B. Würsig, and J.G.M. Thewissen (Eds.), Encyclopaedia of Marine Mammals Second Edition (pp. 1001-1003). San Diego: Elsevier. https://doi.org/10.1016/B978-0-12-373553-9.00231-5
Huijser, L.A., Bérubé, M., Cabrera, A.A., Prieto, R., Silva, M.A., Robbins, J., Kanda, N., Pastene, L.A., Goto, M., Yoshida, H. and Víkingsson, G.A. (2018). Population structure of North Atlantic and North Pacific sei whales (Balaenoptera borealis) inferred from mitochondrial control region DNA sequences and microsatellite genotypes. Conservation Genetics, 19(4), 1007-1024. https://doi.org/10.1007/s10592-018-1076-5
IJsseldijk, L.L., Brownlow, A., Davison, N.J., Deaville, R., Haelters, J., Keijl, G., Siebert, U. and ten Doeschate, M.T.I. (2018). Spatiotemporal Trends in White-Beaked Dolphin Strandings along the North Sea Coast from 1991-2017. Lutra, 61, 153–164. https://zoogdierwinkel.nl/content/lutra-61-1-2018-0
Jonsgård, Å. and Darling, K. (1977). On the biology of the eastern North Atlantic sei whale, Balaenoptera borealis. Reports of the International Whaling Commission Special Issue, 1, 124–129.
Kim, Y.J. and Gu, C. (2004). Smoothing spline Gaussian regression: more scalable computation via efficient approximation. Journal of the Royal Statistical Society: Series B (Statistical Methodology), 66(2), 337-356. https://doi.org/10.1046/j.1369-7412.2003.05316.x
Laake, J.L. (1999). Distance sampling with independent observers: reducing bias from heterogeneity by weakening the conditional independence assumption. In G.W. Garner, S.C. Armstrup, J.L. Laake, B.F.J. Manly, L.L. McDonald, and D.G. Robertson (Eds.), Marine Mammal Survey and Assessment Methods (pp. 137–148). Rotterdam: Balkema.
Murase, H., Hakamada, T., Matsuoka, K., Nishiwaki, S., Inagake, D., Okazaki, M., Tojo, N. and Kitakado, T. (2014). Distribution of sei whales (Balaenoptera borealis) in the subarctic–subtropical transition area of the western North Pacific in relation to oceanic fronts. Deep Sea Research Part II: Topical Studies in Oceanography, 107, 22-28. https://doi.org/10.1016/j.dsr2.2014.05.002
Nøttestad, L., Krafft, B.A., Anthonypillai, V., Bernasconi, M., Langård, L., Mørk, H.L. and Fernö, A. (2015). Recent changes in distribution and relative abundance of cetaceans in the Norwegian Sea and their relationship with potential prey. Frontiers in Ecology and Evolution, 2. https://doi.org/10.3389/fevo.2014.00083
Nøttestad, L., Sivle, L.D., Krafft, B.A., Langård, L., Anthonypillai, V., Bernasconi, M., Langøy, H. and Axelsen, B.E. (2014). Ecological aspects of fin whale and humpback whale distribution during summer in the Norwegian Sea. Marine Ecology, 35, pp.221–232. https://doi.org/10.1111/maec.12075
Perry, A.L., Low, P.J., Ellis, J.R. and Reynolds, J.D. (2005). Climate change and distribution shifts in marine fishes. Science, 308 (5730), 1912-1915. https://doi.org/10.1126/science.1111322
Olsen, E., Budgell, P., Head, E., Kleivane, L., Nøttestad, L., Prieto, R., Silva., M. A., Skov, H., Víkingsson, G.A., Waring, G. and Øien., N. (2009). First satellite-tracked long-distance movement of a sei whale (Balaenoptera borealis) in the North Atlantic. Aquatic Mammals, 35(3), 313-318. https://doi.org/10.1578/AM.35.3.2009.313
Pike, D.G., Gunnlaugsson, T., Mikkelsen, B., Halldórsson, S.D. and Víkingsson, G. A. (2019a). Estimates of the abundance of cetaceans in the central North Atlantic based on the NASS Icelandic and Faroese shipboard surveys conducted in 2015. NAMMCO Scientific Publications, 11. https://doi.org/10.7557/3.4941
Pike, D.G., Gunnlaugsson, T., Mikkelsen, B. and Vikingsson, G. A. (2019b). Estimates of the abundance of cetaceans from the T-NASS Icelandic and Faroese ship surveys conducted in 2007. NAMMCO Scientific Publications, 11 (in press). https://doi.org/10.7557/3.11
Prieto, R., Janiger, D., Silva, M.A., Waring, G.T. and Goncalves, J. (2012). The forgotten whale: a bibliometric analysis and literature review of the North Atlantic sei whale Balaenoptera borealis. Mammal Review, 42, 235–272. https://doi.org/10.1111/j.1365-2907.2011.00195.x
Prieto, R., Silva, M.A., Waring, G.T. and Gonçalves, J.M. (2014). Sei whale movements and behaviour in the North Atlantic inferred from satellite telemetry. Endangered Species Research, 26(2), 103-113. https://doi.org/10.3354/esr00630
QGIS Development Team. (2018). QGIS geographic information system. Open Source Geospatial Foundation Project.
R Core Team. (2018). R: A language and environment for statistical computing. R Foundation for Statistical Computing, Vienna, Austria. URL: https://www.R-project.org/.
Ramirez-Martinez, N. (2020). "Factors influencing decadal-scale changes in cetacean distributions and habitat use in the North Atlantic". Unpublished Ph.D. Thesis, University of St Andrews, Scotland.
Ressler, P.H., Dalpadado, P., Macaulay, G.J., Handegard, N. and Skern-Mauritzen, M. (2015). Acoustic surveys of euphausiids and models of baleen whale distribution in the Barents Sea. Marine Ecology Progress Series, 527, 13-29. https://doi.org/10.3354/meps11257
Roberts, J., Best, B., Mannocci, L., Fujioka, E., Halpin, P., L Palka, D., … G Lockhart, G. (2016). Habitat-based cetacean density models for the U.S. Atlantic and Gulf of Mexico. Scientific Reports, 6(22615). https://doi.org/10.1038/srep22615
Sasaki, H., Murase, H., Kiwada, H., Matsuoka, K., Mitani, Y. and Saitoh, S. (2013). Habitat differentiation between sei (Balaenoptera borealis) and Bryde’s whales (B. brydei) in the western North Pacific. Fisheries Oceanography, 22(6), 496-508. https://doi.org/10.1111/fog.12037
Schleimer, A., Ramp, C., Plourde, S., Lehoux, C., Sears, R. and Hammond, P.S. (2019). Spatio-temporal patterns in fin whale Balaenoptera physalus habitat use in the northern Gulf of St. Lawrence. Marine Ecology Progress Series, 623, 221-234. https://doi.org/10.3354/meps13020
Shuntov, V.P. and Ivanov, O.A. (2015). Marine Mammals in Macro-Ecosystems of the Far Eastern Seas and Adjacent Waters of the North Pacific. Russian Journal of Marine Biology, 41(7), 548-564. https://doi.org/10.1134/S1063074015070056
Sigurjónsson, J. (1995). On the life history and autecology of North Atlantic rorquals. Developments in Marine Biology, 4, 425-441. https://doi.org/10.1016/S0163-6995(06)80044-2
Sigurjónsson, J., Gunnlaugsson, T., Ensor, P., Newcomer, M.W. and Víkingsson, G.A. (1991) North Atlantic sightings survey 1989 (NASS-89): shipboard surveys in Icelandic and adjacent waters July−August 1989 (Balaenoptera borealis, Balaenoptera physalus, Balaenoptera acutorostrata). Reports of the International Whaling Commission, 41, 559−572.
Sigurjónsson, J. and Víkingsson, G.A. (1997). Seasonal abundance of and estimated food consumption by cetaceans in Icelandic and adjacent waters. Journal of Northwest Atlantic Fishery Science, 22, 271–287. https://doi.org/10.2960/J.v22.a20
Skern-Mauritzen, M., Skaug, H.J. and Øien, N. (2009). Line transects, environmental data and GIS: Cetacean distribution, habitat and prey selection along the Barents Sea shelf edge. NAMMCO Scientific Publications, 7, 179-200. https://doi.org/10.7557/3.2713
Skov, H., Gunnlaugsson, T., Budgell, W.P., Horne, J., Nøttestad, L., Olsen, E., Søiland, H., Víkingsson, G. and Waring, G. (2008). Small-scale spatial variability of sperm and sei whales in relation to oceanographic and topographic features along the Mid-Atlantic Ridge. Deep Sea Research Part II: Topical Studies in Oceanography, 55(1-2), 254-268. https://doi.org/10.1016/j.dsr2.2007.09.020
Valdimarsson, H., Astthorsson, O.S. and Palsson, J. (2012). Hydrographic variability in Icelandic waters during recent decades and related changes in distribution of some fish species. ICES Journal of Marine Science, 69(5), 816-825. https://doi.org/10.1093/icesjms/fss027
Visser, F., Hartman, K.L., Pierce, G.J., Valavanis, V.D. and Huisman, J. (2011). Timing of migratory baleen whales at the Azores in relation to the North Atlantic spring bloom. Marine Ecology Progress Series, 440, 267-279.
Víkingsson, G.A. (1997). Feeding of fin whales (Balaenoptera physalus) off Iceland - diurnal and seasonal variation and possible rates. Journal of Northwest Atlantic Fishery Science, 22, 77–89. https://doi.org/10.2960/J.v22.a7
Víkingsson, G.A., Pike, D.G., Valdimarsson, H., Schleimer, A., Gunnlaugsson, T., Silva, T., Elvarsson, B.Þ., Mikkelsen, B., Øien, N., Desportes, G., Bogason, V. and Hammond, P.S. (2015). Distribution, abundance, and feeding ecology of baleen whales in Icelandic waters: have recent environmental changes had an effect? Frontiers in Ecology and Evolution, 3. https://doi.org/10.3389/fevo.2015.00006
Waniek, J.J. and Holliday, N.P. (2006). Large-scale physical controls on phytoplankton growth in the Irminger Sea, Part II: Model study of the physical and meteorological preconditioning. Journal of Marine Systems, 59(3), 219–237. https://doi.org/10.1016/j.jmarsys.2005.10.005
Waring, G.T., Nøttestad, L., Olsen, E., Skov, H. and Víkingsson, G. (2008). Distribution and density estimates of cetaceans along the mid-Atlantic Ridge during summer 2004. Journal of Cetacean Research and Management, 10(2), pp.137-146. https://archive.iwc.int/pages/search.php?search=%21collection15&k=
Wickham, H. (2016). ggplot2: Elegant graphics for data analysis. Springer-Verlag, New York. https://doi.org/10.1007/978-3-319-24277-4_9
Wilson, R.J., Heath, M.R. and Speirs, D.C. (2016). Spatial modelling of Calanus finmarchicus and Calanus helgolandicus: parameter differences explain differences in biogeography. Frontiers in Marine Science, 3. https://doi.org/10.3389/fmars.2016.00157
Wood, S.N. (2017). Generalized additive models: an introduction with R. Portland: Taylor and Francis Inc.
Wormuth, J.H., Ressler, P.H., Cady, R.B. and Harris, E.J. (2000). Zooplankton and micronekton in cyclones and anticyclones in the northeast Gulf of Mexico. Gulf of Mexico Science, 18(1), 23-34.
Øien, N.I. (1988). The distribution of killer whales (Orcinus orca) in the North Atlantic based on Norwegian catches, 1938-1981, and incidental sightings, 1967–1987. Rit Fiskideilar, 11, 678.
Copyright (c) 2020 Philip Hammond
This work is licensed under a Creative Commons Attribution 4.0 International License.
Authors who publish with this journal agree to the following terms:
- Authors retain copyright and grant the journal right of first publication with the work simultaneously licensed under a Creative Commons Attribution License that allows others to share the work with an acknowledgement of the work's authorship and initial publication in this journal.
- Authors are able to enter into separate, additional contractual arrangements for the non-exclusive distribution of the journal's published version of the work (e.g., post it to an institutional repository or publish it in a book), with an acknowledgement of its initial publication in this journal.
- Authors are permitted and encouraged to post their work online (e.g., in institutional repositories or on their website) prior to and during the submission process, as it can lead to productive exchanges, as well as earlier and greater citation of published work (See The Effect of Open Access).