Seasonal productivity drives aggregations of killer whales and other cetaceans over submarine canyons of the Bremer Sub-Basin, south-western Australia
Chandra Salgado Kent
A B C H , Phil Bouchet D E , Rebecca Wellard A F , Iain Parnum A , Leila Fouda A G and Christine Erbe A
+ Author Affiliations
- Author Affiliations
A Centre for Marine Science and Technology, Curtin University, Kent Street, Bentley, WA 6102, Australia.
B Oceans Blueprint, Coogee, WA 6166, Australia.
C Centre for Marine Ecosystems Research, Edith Cowan University, Joondalup Drive, Joondalup, WA 6027, Australia.
D School of Biological Sciences, University of Western Australia, Crawley, WA 6009, Australia.
E Centre for Research into Ecological & Environmental Modelling, The Observatory, Buchanan Gardens, University of St Andrews, Fife KY16 9LZ, Scotland, UK.
F Project ORCA, Perth, WA 6026, Australia.
G School of Biological and Chemical Sciences, Queen Mary University of London, Mile End Road, London E1 4NS, UK.
H Corresponding author. Email: c.salgado@oceansblueprint.com
Australian Mammalogy 43(2) 168-178 https://doi.org/10.1071/AM19058
Submitted: 2 October 2019 Accepted: 3 May 2020 Published: 25 June 2020
Journal Compilation © Australian Mammal Society 2021 Open Access CC BY-NC-ND
Abstract
Cetaceans are iconic predators that serve as important indicators of marine ecosystem health. The Bremer Sub-Basin, south-western Australia, supports a diverse cetacean community including the largest documented aggregation of killer whales (Orcinus orca) in Australian waters. Knowledge of cetacean distributions is critical for managing the area’s thriving ecotourism industry, yet is largely sporadic. Here we combined aerial with opportunistic ship-borne surveys during 2015–2017 to describe the occurrence of multiple cetacean species on a regional scale. We used generalised estimating equations to model variation in killer whale relative density as a function of both static and dynamic covariates, including seabed depth, slope, and chlorophyll a concentration, while accounting for autocorrelation. Encountered cetacean groups included: killer (n = 177), sperm (n = 69), long-finned pilot (n = 29), false killer (n = 2), and strap-toothed beaked (n = 1) whales, as well as bottlenose (n = 12) and common (n = 5) dolphins. Killer whale numbers peaked in areas of low temperatures and high primary productivity, likely due to seasonal upwelling of nutrient-rich waters supporting high prey biomass. The best predictive model highlighted potential killer whale ‘hotspots’ in the Henry, Hood, Pallinup and Bremer Canyons. This study demonstrates the value of abundance data from platforms of opportunity for marine planning and wildlife management in the open ocean.
Additional keywords: generalised estimating equations, habitat modelling, submarine canyons, temporal autocorrelation, whale watching.
References
Akhir, M. F. M. (2010). Physical processes along the southern continental shelf and slope of Western Australia. PhD thesis, School of Environmental Systems Engineering, University of Western Australia, Perth, WA, Australia.Ashe, E., Noren, D. P., and Williams, R. (2010). Animal behaviour and marine protected areas: incorporating behavioural data into the selection of marine protected areas for an endangered killer whale population. Animal Conservation 13, 196–203.
| Animal behaviour and marine protected areas: incorporating behavioural data into the selection of marine protected areas for an endangered killer whale population.Crossref | GoogleScholarGoogle Scholar |
Baird, R. W., and Dill, L. M. (1995). Occurrence and behaviour of transient killer whales: seasonal and pod-specific variability, foraging behaviour, and prey handling. Canadian Journal of Zoology 73, 1300–1311.
| Occurrence and behaviour of transient killer whales: seasonal and pod-specific variability, foraging behaviour, and prey handling.Crossref | GoogleScholarGoogle Scholar |
Bouchet, P., Meeuwig, J., Erbe, C., Salgado Kent, C., Wellard, R., and Pattiaratchi, C. (2018). Bremer Canyon Emerging Priorities Project EP2: Final report for the National Environmental Science Programme, Marine Biodiversity Hub. University of Western Australia, Perth, WA, Australia.
Butler, R. W., MacVicar, R. S., and Hollick-Kenyon, S. (2017). Observation of a super pod of Pacific Harbor porpoises (Phocoena phocoena vomerina) in the Salish Sea. Northwestern Naturalist 98, 137–138.
| Observation of a super pod of Pacific Harbor porpoises (Phocoena phocoena vomerina) in the Salish Sea.Crossref | GoogleScholarGoogle Scholar |
Carrillo, M., Pérez-Vallazza, C., and Álvarez-Vázquez, R. (2010). Cetacean diversity and distribution off Tenerife (Canary Islands). Marine Biodiversity Records 3, e97.
| Cetacean diversity and distribution off Tenerife (Canary Islands).Crossref | GoogleScholarGoogle Scholar |
Carroll, G., Hedley, S., Bannister, J., Ensor, P., and Harcourt, R. (2014). No evidence for recovery in the population of sperm whale bulls off Western Australia, 30 years post-whaling. Endangered Species Research 24, 33–43.
| No evidence for recovery in the population of sperm whale bulls off Western Australia, 30 years post-whaling.Crossref | GoogleScholarGoogle Scholar |
Cui, J. (2007). QIC program and model selection in GEE analyses. The Stata Journal 7, 209–220.
| QIC program and model selection in GEE analyses.Crossref | GoogleScholarGoogle Scholar |
Dallas, T. A., and Hastings, A. (2018). Habitat suitability estimated by niche models is largely unrelated to species abundance. Global Ecology and Biogeography 27, 1448–1456.
| Habitat suitability estimated by niche models is largely unrelated to species abundance.Crossref | GoogleScholarGoogle Scholar |
Davidson, A. D., Boyer, A. G., Kim, H., Pompa-Mansilla, S., Hamilton, M. J., Costa, D. P., Ceballos, G., and Brown, J. H. (2012). Drivers and hotspots of extinction risk in marine mammals. Proceedings of the National Academy of Sciences of the United States of America 109, 3395–3400.
| Drivers and hotspots of extinction risk in marine mammals.Crossref | GoogleScholarGoogle Scholar | 22308490PubMed |
de Bruyn, P. N., Tosh, C. A., and Terauds, A. (2013). Killer whale ecotypes: Is there a global model? Biological Reviews of the Cambridge Philosophical Society 88, 62–80.
| Killer whale ecotypes: Is there a global model?Crossref | GoogleScholarGoogle Scholar |
Donnelly, D., McInnes, J., Morrice, M., and Andrews, C. (2016). ‘Killer whales of eastern Australia: photo-identification catalogue.’ (Killer Whales Australia: Box Hill South, Vic., Australia)
Elith, J., Phillips, S. J., Hastie, T., Dudík, M., Chee, Y. E., and Yates, C. J. (2011). A statistical explanation of MaxEnt for ecologists. Diversity & Distributions 17, 43–57.
| A statistical explanation of MaxEnt for ecologists.Crossref | GoogleScholarGoogle Scholar |
Esteban, R., Verborgh, P., Gauffier, P., Giménez, J., Afán, I., Cañadas, A., García, P., Murcia, J. L., Magalhães, S., and Andreu, E. (2014). Identifying key habitat and seasonal patterns of a critically endangered population of killer whales. Journal of the Marine Biological Association of the United Kingdom 94, 1317–1325.
| Identifying key habitat and seasonal patterns of a critically endangered population of killer whales.Crossref | GoogleScholarGoogle Scholar |
Evans, P. G. H., and Hammond, P. S. (2004). Monitoring cetaceans in European waters. Mammal Review 34, 131–156.
| Monitoring cetaceans in European waters.Crossref | GoogleScholarGoogle Scholar |
Ferguson, M. C., Barlow, J., Fiedler, P., Reilly, S. B., and Gerrodette, T. (2006). Spatial models of delphinid (family Delphinidae) encounter rate and group size in the eastern tropical Pacific Ocean. Ecological Modelling 193, 645–662.
| Spatial models of delphinid (family Delphinidae) encounter rate and group size in the eastern tropical Pacific Ocean.Crossref | GoogleScholarGoogle Scholar |
Fernandez-Arcaya, U., Ramirez-Llodra, E., Aguzzi, J., Allcock, A. L., Davies, J. S., Dissanayake, A., Harris, P., Howell, K., Huvenne, V. A. I., Macmillan-Lawler, M., Martín, J., Menot, L., Nizinski, M., Puig, P., Rowden, A. A., Sanchez, F., and Van den Beld, I. M. J. (2017). Ecological role of submarine canyons and need for canyon conservation: a review. Frontiers in Marine Science 4, 5.
| Ecological role of submarine canyons and need for canyon conservation: a review.Crossref | GoogleScholarGoogle Scholar |
Fiedler, P. C., Redfern, J. V., Forney, K. A., Palacios, D. M., Sheredy, C., Rasmussen, K., García-Godos, I., Santillán, L., Tetley, M. J., Félix, F., and Ballance, L. T. (2018). Prediction of large whale distributions: a comparison of presence–absence and presence-only modeling techniques. Frontiers in Marine Science 5, 419.
| Prediction of large whale distributions: a comparison of presence–absence and presence-only modeling techniques.Crossref | GoogleScholarGoogle Scholar |
Findlay, K. P., Seakamela, S. M., Meÿer, M. A., Kirkman, S. P., Barendse, J., Cade, D. E., Hurwitz, D., Kennedy, A. S., Kotze, P. G., and McCue, S. A. (2017). Humpback whale ‘super-groups’ – a novel low-latitude feeding behaviour of Southern Hemisphere humpback whales (Megaptera novaeangliae) in the Benguela Upwelling System. PLoS One 12, e0172002.
| Humpback whale ‘super-groups’ – a novel low-latitude feeding behaviour of Southern Hemisphere humpback whales (Megaptera novaeangliae) in the Benguela Upwelling System.Crossref | GoogleScholarGoogle Scholar | 28249036PubMed |
Fletcher, W. J., Shaw, J., Metcalf, S. J., and Gaughan, D. J. (2010). An ecosystem based fisheries management framework: the efficient, regional-level planning tool for management agencies. Marine Policy 34, 1226–1238.
| An ecosystem based fisheries management framework: the efficient, regional-level planning tool for management agencies.Crossref | GoogleScholarGoogle Scholar |
Gill, P. C., Pirzl, R., Morrice, M. G., and Lawton, K. (2015). Cetacean diversity of the continental shelf and slope off southern Australia. The Journal of Wildlife Management 79, 672–681.
| Cetacean diversity of the continental shelf and slope off southern Australia.Crossref | GoogleScholarGoogle Scholar |
Halpern, B. S., Frazier, M., Afflerbach, J., Lowndes, J. S., Micheli, F., O’Hara, C., Scarborough, C., and Selkoe, K. A. (2019). Recent pace of change in human impact on the world’s ocean. Scientific Reports 9, 11609.
| Recent pace of change in human impact on the world’s ocean.Crossref | GoogleScholarGoogle Scholar | 31406130PubMed |
Higby, L. K., Stafford, R., and Bertulli, C. G. (2012). An evaluation of ad hoc presence-only data in explaining patterns of distribution: cetacean sightings from whale-watching vessels. International Journal of Zoology 2012, art428752.
| An evaluation of ad hoc presence-only data in explaining patterns of distribution: cetacean sightings from whale-watching vessels.Crossref | GoogleScholarGoogle Scholar |
Howard, C., Stephens, P. A., Pearce-Higgins, J. W., Gregory, R. D., and Willis, S. G. (2014). Improving species distribution models: the value of data on abundance. Methods in Ecology and Evolution 5, 506–513.
| Improving species distribution models: the value of data on abundance.Crossref | GoogleScholarGoogle Scholar |
Hu, C., Lee, Z., and Franz, B. (2012). Chlorophyll a algorithms for oligotrophic oceans: a novel approach based on three-band reflectance difference. Journal of Geophysical Research 117, C01011.
| Chlorophyll a algorithms for oligotrophic oceans: a novel approach based on three-band reflectance difference.Crossref | GoogleScholarGoogle Scholar |
Huang, Z., Nichol, S., Harris, P., and Caley, J. (2014). Classification of submarine canyons of the Australian continental margin. Marine Geology 357, 362–383.
| Classification of submarine canyons of the Australian continental margin.Crossref | GoogleScholarGoogle Scholar |
Huang, Z., Schlacher, T. A., Nichol, S., Williams, A., Althaus, F., and Kloser, R. (2018). A conceptual surrogacy framework to evaluate the habitat potential of submarine canyons. Progress in Oceanography 169, 199–213.
| A conceptual surrogacy framework to evaluate the habitat potential of submarine canyons.Crossref | GoogleScholarGoogle Scholar |
Isojunno, S., Matthiopoulos, J., and Evans, P. G. (2012). Harbour porpoise habitat preferences: robust spatio-temporal inferences from opportunistic data. Marine Ecology Progress Series 448, 155–170.
| Harbour porpoise habitat preferences: robust spatio-temporal inferences from opportunistic data.Crossref | GoogleScholarGoogle Scholar |
Johnson, C. M., Beckley, L. E., Kobryn, H., Johnson, G. E., Kerr, I., and Payne, R. (2016). Crowdsourcing modern and historical data identifies sperm whale (Physeter macrocephalus) habitat offshore of South-Western Australia. Frontiers in Marine Science 3, 167.
| Crowdsourcing modern and historical data identifies sperm whale (Physeter macrocephalus) habitat offshore of South-Western Australia.Crossref | GoogleScholarGoogle Scholar |
Johnston, A., Fink, D., Reynolds, M. D., Hochachka, W. M., Sullivan, B. L., Bruns, N. E., Hallstein, E., Merrifield, M. S., Matsumoto, S., and Kelling, S. (2015). Abundance models improve spatial and temporal prioritization of conservation resources. Ecological Applications 25, 1749–1756.
| Abundance models improve spatial and temporal prioritization of conservation resources.Crossref | GoogleScholarGoogle Scholar | 26591443PubMed |
Jones, A., Bruce, E., Davies, K. P., Blewitt, M., and Sheehan, S. (2019). Assessing potential environmental influences on killer whale (Orcinus orca) distribution patterns in the Bremer Canyon, south-west Australia. The Australian Geographer 50, 381–405.
| Assessing potential environmental influences on killer whale (Orcinus orca) distribution patterns in the Bremer Canyon, south-west Australia.Crossref | GoogleScholarGoogle Scholar |
Kiszka, J., Ersts, P., and Ridoux, V. (2007a). Cetacean diversity around the Mozambique Channel Island of Mayotte (Comoros archipelago). The Journal of Cetacean Research and Management 9, 105.
Kiszka, J., Macleod, K., Van Canneyt, O., Walker, D., and Ridoux, V. (2007b). Distribution, encounter rates, and habitat characteristics of toothed cetaceans in the Bay of Biscay and adjacent waters from platform-of-opportunity data. ICES Journal of Marine Science 64, 1033–1043.
| Distribution, encounter rates, and habitat characteristics of toothed cetaceans in the Bay of Biscay and adjacent waters from platform-of-opportunity data.Crossref | GoogleScholarGoogle Scholar |
Laran, S., Authier, M., Van Canneyt, O., Dorémus, G., Watremez, P., and Ridoux, V. (2017). A comprehensive survey of pelagic megafauna: their distribution, densities, and taxonomic richness in the tropical Southwest Indian Ocean. Frontiers in Marine Science 4, 139.
| A comprehensive survey of pelagic megafauna: their distribution, densities, and taxonomic richness in the tropical Southwest Indian Ocean.Crossref | GoogleScholarGoogle Scholar |
Lück, M. (2015). Education on marine mammal tours – but what do tourists want to learn? Ocean and Coastal Management 103, 25–33.
| Education on marine mammal tours – but what do tourists want to learn?Crossref | GoogleScholarGoogle Scholar |
Lusseau, D., Bain, D. E., Williams, R., and Smith, J. C. (2009). Vessel traffic disrupts the foraging behavior of southern resident killer whales Orcinus orca. Endangered Species Research 6, 211–221.
| Vessel traffic disrupts the foraging behavior of southern resident killer whales Orcinus orca.Crossref | GoogleScholarGoogle Scholar |
Mannocci, L., Catalogna, M., Dorémus, G., Laran, S., Lehodey, P., Massart, W., Monestiez, P., Van Canneyt, O., Watremez, P., and Ridoux, V. (2014). Predicting cetacean and seabird habitats across a productivity gradient in the South Pacific gyre. Progress in Oceanography 120, 383–398.
| Predicting cetacean and seabird habitats across a productivity gradient in the South Pacific gyre.Crossref | GoogleScholarGoogle Scholar |
McClellan, C. M., Brereton, T., Dell’Amico, F., Johns, D. G., Cucknell, A.-C., Patrick, S. C., Penrose, R., Ridoux, V., Solandt, J.-L., Stephan, E., Votier, S. C., Williams, R., and Godley, B. J. (2014). Understanding the distribution of marine megafauna in the English Channel region: identifying key habitats for conservation within the busiest seaway on Earth. PLoS One 9, e89720.
| Understanding the distribution of marine megafauna in the English Channel region: identifying key habitats for conservation within the busiest seaway on Earth.Crossref | GoogleScholarGoogle Scholar | 24586985PubMed |
Merow, C., Smith, M. J., and Silander, J. A. (2013). A practical guide to MaxEnt for modeling species’ distributions: what it does, and why inputs and settings matter. Ecography 36, 1058–1069.
| A practical guide to MaxEnt for modeling species’ distributions: what it does, and why inputs and settings matter.Crossref | GoogleScholarGoogle Scholar |
Moors-Murphy, H. B. (2014). Submarine canyons as important habitat for cetaceans, with special reference to the Gully: a review. Deep-sea Research. Part II, Topical Studies in Oceanography 104, 6–19.
| Submarine canyons as important habitat for cetaceans, with special reference to the Gully: a review.Crossref | GoogleScholarGoogle Scholar |
Morrice, M. G. (2004). Killer whales (Orcinus orca) in Australian territorial waters. Report for the Whale and Dolphin Conservation Society. Deakin University, Warrnambool, Vic., Australia.
Nielsen, S. E., Johnson, C. J., Heard, D. C., and Boyce, M. S. (2005). Can models of presence-absence be used to scale abundance? Two case studies considering extremes in life history. Ecography 28, 197–208.
| Can models of presence-absence be used to scale abundance? Two case studies considering extremes in life history.Crossref | GoogleScholarGoogle Scholar |
Pan, W. (2001). Akaike’s information criterion in generalized estimating equations. Biometrics 57, 120–125.
| Akaike’s information criterion in generalized estimating equations.Crossref | GoogleScholarGoogle Scholar | 11252586PubMed |
Parrish, J. K., and Edelstein-Keshet, L. (1999). Complexity, pattern, and evolutionary trade-offs in animal aggregation. Science 284, 99–101.
| Complexity, pattern, and evolutionary trade-offs in animal aggregation.Crossref | GoogleScholarGoogle Scholar | 10102827PubMed |
Passadore, C., Domingo, A., Szephegyi, M., and Secchi, E. R. (2014). Influence of environmental and longline fishing operational variables on the presence of killer whales (Orcinus orca) in south-western Atlantic. Journal of the Marine Biological Association of the United Kingdom 94, 1267–1276.
| Influence of environmental and longline fishing operational variables on the presence of killer whales (Orcinus orca) in south-western Atlantic.Crossref | GoogleScholarGoogle Scholar |
Pearce, J., and Ferrier, S. (2001). The practical value of modelling relative abundance of species for regional conservation planning: a case study. Biological Conservation 98, 33–43.
| The practical value of modelling relative abundance of species for regional conservation planning: a case study.Crossref | GoogleScholarGoogle Scholar |
Pirotta, E., Booth, C. G., Costa, D. P., Fleishman, E., Kraus, S. D., Lusseau, D., Moretti, D., New, L. F., Schick, R. S., and Schwarz, L. K. (2018). Understanding the population consequences of disturbance. Ecology and Evolution 8, 9934–9946.
| Understanding the population consequences of disturbance.Crossref | GoogleScholarGoogle Scholar | 30386587PubMed |
Pitman, R. L., Totterdell, J. A., Fearnbach, H., Ballance, L. T., Durban, J. W., and Kemps, H. (2015). Whale killers: prevalence and ecological implications of killer whale predation on humpback whale calves off Western Australia. Marine Mammal Science 31, 629–657.
| Whale killers: prevalence and ecological implications of killer whale predation on humpback whale calves off Western Australia.Crossref | GoogleScholarGoogle Scholar |
Pitman, R. L., Totterdell, J., Wellard, R., Cullen, P., and de Boer, M. (2019). Long in the tooth: biological observations from at-sea sightings of strap-toothed beaked whales (Mesoplodon layardii). Marine Mammal Science 35, 1141–1161.
| Long in the tooth: biological observations from at-sea sightings of strap-toothed beaked whales (Mesoplodon layardii).Crossref | GoogleScholarGoogle Scholar |
Prieto, R., Tobeña, M., and Silva, M. A. (2017). Habitat preferences of baleen whales in a mid-latitude habitat. Deep-sea Research. Part II, Topical Studies in Oceanography 141, 155–167.
| Habitat preferences of baleen whales in a mid-latitude habitat.Crossref | GoogleScholarGoogle Scholar |
R Core Team (2013). R: a language and environment for statistical computing. (R Foundation for Statistical Computing: Vienna, Austria) Available at http://www.R-project.org/ [Verified 15 May 2020]
Reisinger, R. R., Keith, M., Andrews, R. D., and de Bruyn, P. J. N. (2015). Movement and diving of killer whales (Orcinus orca) at a Southern Ocean archipelago. Journal of Experimental Marine Biology and Ecology 473, 90–102.
| Movement and diving of killer whales (Orcinus orca) at a Southern Ocean archipelago.Crossref | GoogleScholarGoogle Scholar |
Rennie, S., Hanson, C. E., McCauley, R. D., Pattiaratchi, C., Burton, C., Bannister, J., Jenner, C., and Jenner, M. N. (2009). Physical properties and processes in the Perth Canyon, Western Australia: links to water column production and seasonal pygmy blue whale abundance. Journal of Marine Systems 77, 21–44.
| Physical properties and processes in the Perth Canyon, Western Australia: links to water column production and seasonal pygmy blue whale abundance.Crossref | GoogleScholarGoogle Scholar |
Ryan, J. P., Chavez, F. P., and Bellingham, J. G. (2005). Physical–biological coupling in Monterey Bay, California: topographic influences on phytoplankton ecology. Marine Ecology Progress Series 287, 23–32.
| Physical–biological coupling in Monterey Bay, California: topographic influences on phytoplankton ecology.Crossref | GoogleScholarGoogle Scholar |
Salgado Kent, C., Jenner, K. C. S., Jenner, M.-N., Bouchet, P., and Rexstad, E. (2012). Southern Hemisphere breeding stock ‘D’ humpback whale population estimates from north west cape, Western Australia. The Journal of Cetacean Research and Management 12, 29–38.
Santora, J. A., Zeno, R., Dorman, J. G., and Sydeman, W. J. (2018). Submarine canyons represent an essential habitat network for krill hotspots in a large marine ecosystem. Scientific Reports 8, 7579.
| Submarine canyons represent an essential habitat network for krill hotspots in a large marine ecosystem.Crossref | GoogleScholarGoogle Scholar | 29765085PubMed |
Scott-Hayward, L., Oedekoven, C. S., Mackenzie, M. L., Walker, C. G., and Rexstad, E. (2013). User guide for the MRSea package: statistical modelling of bird and cetacean distributions in offshore renewables development areas. University of St Andrews, Report for Marine Scotland No. CR/2012/05, St Andrews, Fife, UK.
Scott-Hayward, L. A. S., MacKenzie, M. L., Donovan, C. R., Walker, C., and Ashe, E. (2014). Complex region spatial smoother (CReSS). Journal of Computational and Graphical Statistics 23, 340–360.
| Complex region spatial smoother (CReSS).Crossref | GoogleScholarGoogle Scholar |
Scott-Hayward, L. A., Mackenzie, M. L., Ashe, E., and Williams, R. (2015). Modelling killer whale feeding behaviour using a spatially adaptive complex region spatial smoother (CReSS) and generalised estimating equations (GEEs). Journal of Agricultural Biological & Environmental Statistics 20, 305–322.
| Modelling killer whale feeding behaviour using a spatially adaptive complex region spatial smoother (CReSS) and generalised estimating equations (GEEs).Crossref | GoogleScholarGoogle Scholar |
Smith, J. N., Grantham, H. S., Gales, N., Double, M. C., Noad, M. J., and Paton, D. (2012). Identification of humpback whale breeding and calving habitat in the Great Barrier Reef. Marine Ecology Progress Series 447, 259–272.
| Identification of humpback whale breeding and calving habitat in the Great Barrier Reef.Crossref | GoogleScholarGoogle Scholar |
Thomas, L., Buckland, S. T., Rexstad, E. A., Laake, J. L., Strindberg, S., Hedley, S. L., Bishop, J. R. B., Marques, T. A., and Burnham, K. P. (2010). Distance software: design and analysis of distance sampling surveys for estimating population size. Journal of Applied Ecology 47, 5–14.
| Distance software: design and analysis of distance sampling surveys for estimating population size.Crossref | GoogleScholarGoogle Scholar | 20383262PubMed |
Torres, L. G., Read, A. J., and Halpin, P. (2008). Fine-scale habitat modeling of a top marine predator: do prey data improve predictive capacity. Ecological Applications 18, 1702–1717.
| Fine-scale habitat modeling of a top marine predator: do prey data improve predictive capacity.Crossref | GoogleScholarGoogle Scholar | 18839765PubMed |
Totterdell, J. A. (2015). WA Orca Research Report - Ningaloo ID Catalogue. Marine Information and Research Group (MIRG).
Veloz, S., Salas, L., Altman, B., Alexander, J., Jongsomjit, D., Elliott, N., and Ballard, G. (2015). Improving effectiveness of systematic conservation planning with density data. Conservation Biology 29, 1217–1227.
| Improving effectiveness of systematic conservation planning with density data.Crossref | GoogleScholarGoogle Scholar | 25873240PubMed |
Waples, K., and Raudino, H. (2018). Setting a course for marine mammal research in Western Australia. Pacific Conservation Biology 24, 289–303.
| Setting a course for marine mammal research in Western Australia.Crossref | GoogleScholarGoogle Scholar |
Weber, M. M., Stevens, R. D., Diniz-Filho, J. A. F., and Grelle, C. E. V. (2017). Is there a correlation between abundance and environmental suitability derived from ecological niche modelling? A meta-analysis. Ecography 40, 817–828.
| Is there a correlation between abundance and environmental suitability derived from ecological niche modelling? A meta-analysis.Crossref | GoogleScholarGoogle Scholar |
Wellard, R., and Erbe, C. (2017). ‘Killer whales of the Bremer Sub-Basin: a photo-ID catalogue’, 2nd edn. (Project ORCA and Centre for Marine Science and Technology, Curtin University: Perth, WA, Australia)
Wellard, R., Erbe, C., Fouda, L., and Blewitt, M. (2015). Vocalisations of killer whales (Orcinus orca) in the Bremer Canyon, Western Australia. PLoS One 10, e0136535.
| Vocalisations of killer whales (Orcinus orca) in the Bremer Canyon, Western Australia.Crossref | GoogleScholarGoogle Scholar | 26462069PubMed |
Wellard, R., Lightbody, K., Fouda, L., Blewitt, M., Riggs, D., and Erbe, C. (2016). Killer whale (Orcinus orca) predation on beaked whales (Mesoplodon spp.) in the Bremer Sub-Basin, Western Australia. PLoS One 11, e0166670.
| Killer whale (Orcinus orca) predation on beaked whales (Mesoplodon spp.) in the Bremer Sub-Basin, Western Australia.Crossref | GoogleScholarGoogle Scholar | 27923044PubMed |
Williams, R., Hedley, S. L., and Hammond, P. S. (2006a). Modeling distribution and abundance of Antarctic baleen whales using ships of opportunity. Ecology and Society 11, 1.
| Modeling distribution and abundance of Antarctic baleen whales using ships of opportunity.Crossref | GoogleScholarGoogle Scholar |
Williams, R., Lusseau, D., and Hammond, P. S. (2006b). Estimating relative energetic costs of human disturbance to killer whales (Orcinus orca). Biological Conservation 133, 301–311.
| Estimating relative energetic costs of human disturbance to killer whales (Orcinus orca).Crossref | GoogleScholarGoogle Scholar |
Yackulic, C. B., Chandler, R., Zipkin, E. F., Royle, J. A., Nichols, J. D., Campbell Grant, E. H., and Veran, S. (2013). Presence-only modelling using MAXENT: when can we trust the inferences? Methods in Ecology and Evolution 4, 236–243.
| Presence-only modelling using MAXENT: when can we trust the inferences?Crossref | GoogleScholarGoogle Scholar |
Yan, J., and Fine, J. (2004). Estimating equations for association structures. Statistics in Medicine 23, 859–874.
| Estimating equations for association structures.Crossref | GoogleScholarGoogle Scholar | 15027075PubMed |
Yin, D., and He, F. (2014). A simple method for estimating species abundance from occurrence maps. Methods in Ecology and Evolution 5, 336–343.
| A simple method for estimating species abundance from occurrence maps.Crossref | GoogleScholarGoogle Scholar |
Zorn, C. J. (2001). Generalized estimating equation models for correlated data: a review with applications. American Journal of Political Science 45, 470–490.
| Generalized estimating equation models for correlated data: a review with applications.Crossref | GoogleScholarGoogle Scholar |
Zuur, A. F., Ieno, E. N., Walker, N. J., Saveliev, A. A., and Smith, G. M. (2009). ‘Mixed effects models and extensions in ecology using R.’ (Springer: New York, NY, USA)
