Journal Articles

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    A contribution to the anatomy of two rare cetacean species: The hourglass dolphin (Cephalorhynchus cruciger) and the spectacled porpoise (Phocoena dioptrica)
    (Wiley Periodicals LLC on behalf of American Association for Anatomy, 2025-10-22) Graïc J-M; Gerussi T; Cozzi B; Boys RM; Kot BCW; Perrott MR; Fleury K; Chung TYT; Tsui HCL; Burns E; Webster T; Hunter S; Betty EL; Howarth O; Loch C; White S; Dawson S; Rayment W; Cole R; Cox D; Waterhouse T; Hendriks H; van Helden A; Johnstone M; Stewart RO; Fordyce RE; Stockin KA
    The anatomical description of the hourglass dolphin (Cephalorhynchus cruciger) and the spectacled porpoise (Phocoena dioptrica) remains largely unexplored, due to limited specimen availability and preservation challenges. This study employed digital imaging techniques, conventional histology, and computed tomography to provide visualization of anatomical structures for a detailed analysis. We present a comprehensive analysis of the gross macroscopical and microscopical morphology of two hourglass dolphins and four spectacled porpoises. Morphometric measurements and skeletal characteristics aligned with the literature, while internal anatomy (organs and systems) was similar to other odontocetes. Precise and consistent lung measurements were challenging; qualitative assessments indicated relatively large lungs with respect to body size. The spectacled porpoise dorsal fin was uniquely large with a well-developed blood supply; this is hypothesized to act as a thermoregulatory window, though it may also play a role in sexual display in the case of males. Overall, this study provides new data on the anatomy of the hourglass dolphin and spectacled porpoise, contributing insights that may influence future research on these rare species. The findings highlight the importance of anatomical studies as a basis for explaining evolutionary relationships within cetaceans and their ecological roles in the Southern Ocean ecosystems.
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    A contribution to the anatomy of two rare cetacean species: the hourglass dolphin (Lagenorhynchus cruciger) and the spectacled porpoise (Phocoena dioptrica)
    (2025-03-26) Graïc J-M; Gerussi T; Cozzi B; Boys RM; Kot BCW; Perrott MR; Fleury K; Chung TYT; Tsui HCL; Burns E; Webster T; Hunter S; Betty EL; Howarth O; Loch C; White S; Dawson S; Rayment W; Cole R; Cox D; Waterhouse T; Hendriks H; van Helden A; Johnstone M; Stewart RO; Fordyce RE; Stockin KA
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    Using post-mortem computed tomography to identify traumatic cranial lesions in small stranded odontocetes euthanised via ballistics
    (Taylor and Francis Group on behalf of the New Zealand Veterinary Association, 2025-07-29) Boys RM; Stewart RO; Kot BCW; Lye G; Beausoleil NJ; Stockin KA
    Aims: To describe the pathophysiology caused by ballistics applied to the head of stranded cetaceans that required euthanasia and use this information to infer the likely welfare implications of these real-life euthanasia events. Methods: Post-mortem computed tomography (PMCT) imaging was conducted on six cetacean carcasses of five different species that were submitted for dissection following ballistics euthanasia. No animals were euthanised for the purpose of this study. All data were collected opportunistically at stranding events where euthanasia was a priori deemed necessary by the Department of Conservation (DOC) in partnership with indigenous Māori (iwi). The cranial damage assessed included osseous lesions at entry and exit wounds, intracranial bone fractures and intracranial projectile fragments. Potential welfare implications were inferred based on the likelihood of instantaneous insensibility occurring due to direct physical disruption to the brainstem. Additionally, basic stranding data were collected from DOC incident reports including whether animals were refloated prior to the decision for euthanasia. We also gathered information from the marksperson involved on how the ballistics method was employed, equipment used, and animal insensibility assessed at each stranding event. Results: PMCT results suggest that two animals were likely instantaneously insensible, three had a moderate chance of being instantly insensible and one had a low chance of instantaneous insensibility. In one case, the marksperson reported a faster time to insensibility/death than was judged likely to have occurred from PMCT. In only two cases were the criteria for verifying insensibility reported as being checked. The likelihood of instantaneous insensibility occurring was affected by shot placement/aim and characteristics of the projectile used. Conclusions: Euthanasia is a critical option for severely compromised cetaceans. While most animals examined were probably rapidly insensible following shooting, cranial trauma and thus confidence in the efficacy of ballistics seemed to vary with shot placement, aim, equipment and cranial anatomy. Clinical relevance: Verification of insensibility immediately following shooting, by assessing reflex responses as recommended, is crucial to ensure humane emergency management. PMCT provides an effective non-destructive tool to validate field methods for assessing the efficacy of euthanasia.
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    Out of habitat marine mammals – Identification, causes, and management recommendations
    (Elsevier Ltd, 2025-07) Nunny L; Bossley M; Boys RM; Brakes P; Genov T; Parsons ECM; Peters KJ; Rose NA; Simeone CA; Stockin KA; Vail CS; van der Linde K; Visser IN; Simmonds MP
    An out of habitat (OOH) marine mammal is an individual found outside of their natural range, e.g., an Atlantic walrus (Odobenus rosmarus rosmarus) along the coastline of mainland Europe, or an individual within their natural range in habitat that is not optimal for their health or survival due to a lack of suitable conditions and/or because of potential conflict with humans. As the number of OOH marine mammals appears to be increasing, and following on from two international workshops, here we define what constitutes an OOH marine mammal and provide a detailed assessment of potential drivers. Climate change, habitat loss and/or degradation, disturbance, changes in prey distribution, and morbidity may all be reasons for a marine mammal being OOH. Appropriate management of OOH marine mammals is considered. This may simply involve monitoring to better understand the situation, to protect the welfare of the animal(s) and to ensure that members of the public are kept safe. However, on occasion, further intervention may be required, such as providing safe areas for the OOH marine mammal(s), encouraging the animal(s) to move, capture and translocation, or euthanasia. Regardless of the action taken, educating and communicating with the public are key elements of OOH marine mammal management. Lessons can be learned from human-wildlife conflict and rewilding scenarios. The potential for OOH marine mammals to be pioneers seeking new habitat, contributing to species resilience in the face of rapidly changing environments, and how to include them in conservation legislation and management planning are considered.
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    Admixture Increases Genetic Diversity and Adaptive Potential in Australasian Killer Whales
    (John Wiley and Sons Ltd, 2025-02-28) Reeves IM; Totterdell JA; Sandoval-Castillo J; Betty EL; Stockin KA; Oliphant Stewart R; Johnstone M; Foote AD; Kardos M
    Admixture is the exchange of genetic variation between differentiated demes, resulting in ancestry within a population coalescing in multiple ancestral source populations. Low-latitude killer whales (Orcinus orca) populations typically have higher genetic diversity than those in more densely populated, high productivity and high-latitude regions. This has been hypothesized to be due to episodic admixture between populations with distinct genetic backgrounds. We test this hypothesis by estimating variation in local ancestry of whole genome sequences from three genetically differentiated, low-latitude killer whale populations and comparing them to global genetic variation. We find 'Antarctic-like' ancestry tracts in the genomes of southwestern Australia (SWA) population including recent (within the last 2-4 generations) admixture. Admixed individuals had, on average, shorter and fewer runs of homozygosity than unadmixed individuals and increased effective population size (Ne). Thus, connectivity between demes results in the maintenance of Ne of relatively small demes at a level comparable to the sum of Ne across demes. A subset of the admixed regions was inferred to be evolving under selection in the SWA population, suggesting that this admixed variation may be contributing to the population's adaptive potential. This study provides important and rare empirical evidence that small populations can maintain genetic diversity due to sporadic admixture between different genetic backgrounds and that admixed ancestry can promote the long-term stability of Ne.
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    Ethological and procedural assessment of ballistics euthanasia for stranded cetaceans
    (Elsevier B.V., 2025-02-12) Boys RM; Beausoleil NJ; Stockin KA
    Euthanasia may be necessary at cetacean stranding events to end suffering. To ensure a humane death, verification of unconsciousness must be undertaken following application of the killing method. However, only limited information is reported on individual euthanasia events, including how unconsciousness and/or death is assessed. Here, we describe current practices for euthanising stranded cetaceans in New Zealand by analysing video footage of fourteen ballistics events involving various species. Specifically, we detail the ballistics procedures applied and the behavioural responses of the animals. Additionally, we sought to understand how animal unconsciousness was evaluated following each event by surveying the personnel involved. Video recordings ranged in length from 25 to 480 s after the initial shot and involved 14 cetaceans: one Cuvier's beaked whale, one pygmy sperm whale, one bottlenose dolphin, four pilot whales and seven false killer whales. Unconsciousness criteria were typically not observed being verified in the videos analysed, despite markspersons stating that death occurred in < 1 minute for most animals. Number of shots applied to an animal ranged from 1–3. Key behaviours indicative of unconsciousness included a continuously slack lower jaw and epaxial muscle relaxation. Based on behavioural observations, 66.7 % of animals for which an estimate could be made (n = 12) were likely unconscious within 30 s of the initial shot; half of these were likely unconscious within 10 s. The remaining animals appeared conscious for 30 s after the initial shot, and half of these were likely still conscious after one minute. Ten animals displayed post-shot behaviours suggestive of voluntary control, including tail lifting, beyond the point of presumed death reported by markspersons. The lack of verification of unconsciousness in most animals (12/14) hinders our ability to understand how these behaviours may correlate with awareness and thus welfare impacts of shooting. While the application of ballistics appeared to follow current national recommendations, mandated procedures for verifying death were rarely observed. We recommend exploring the reasons for this finding, to ensure that such procedures are appropriately undertaken, and welfare compromise minimised.
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    Estimating abundance of a small population of Bryde's whales: a comparison between aerial surveys and boat-based platforms of opportunity
    (John Wiley & Sons Ltd., on behalf of Zoological Society of London, 2024-08-01) Hamilton ONP; Fewster RM; Low P; Johnson F; Lea C; Stockin KA; van der Linde K; Constantine R; Boersch-Supan P; Williams R
    Accurate abundance estimates are essential for the development of effective conservation management strategies, yet they are difficult to produce for small populations that are elusive and sparsely distributed throughout their range. For such populations it is challenging to collect a representative dataset sufficient for robust estimation of detectability and abundance. Over a one-year study, we used two methods to estimate abundance of a Nationally Critical, widely dispersed Bryde's whale population in the Hauraki Gulf, Aotearoa/New Zealand; (i) distance sampling from systematic line-transect aerial surveys (n = 22 surveys, 9,944 km, total sightings 21–24 whales), and (ii) mark-recapture (MR) using photo-identification images collected from a platform-of-opportunity and small-boat surveys (218 sampling occasions, 27 whales). From the aerial surveys, we estimated an average of 15 whales (95% CI = 6, 30; CV = 37%) at the sea-surface at any time. For the boat-based surveys, we developed a custom MR model to address seasonal and individual heterogeneity in capture probabilities and obtained an estimate of 72 distinct whales (95% CI = 38, 106; CV = 24%) in the population. These two approaches provide different perspectives on the abundance and dynamics of Bryde's whales. The aerial surveys estimate the average number of individuals present at any one time, whereas the MR model estimates the total number of animals that used the Gulf during the study. Although neither sampling method is optimal for estimating the abundance of this small, dispersed population, the use of two complementary approaches informs conservation managers about patterns of abundance and distribution over different temporal and spatial scales. It is common to have limited resources for marine research where model assumptions cannot be met. Here, we highlight pragmatic strategies showing how models can be customized to the population of interest to assist with monitoring species of conservation concern.
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    Multidimensional trace metals and nutritional niche differ between sexually immature and mature common dolphins (Delphinus delphis)
    (Elsevier Ltd., 2023-06-26) Stockin KA; Machovsky-Capuska GE; Palmer EI; Amiot C
    There is a need to understand the links between metals and nutrition for apex marine predators, which may be subject to different ecotoxicological effects at different life stages. We combined stomach content analyses (SCA), prey composition analysis (PCA), the Multidimensional Niche Framework (MNNF) with Bayesian multivariate ellipses, trace metal analysis and nicheROVER to investigate nutrition and trace metals across sex, age, and sexual maturity status in common dolphins (Delphinus delphis) from New Zealand. A broader prey composition niche breadth (SEAc) was estimated for immature compared to mature conspecifics, showing a higher degree of prey and nutrient generalism driven by protein (P) intake. Cd and Zn niche similarities suggests these metals were incorporated through similar prey in both immature and mature dolphins, whereas Hg and Se niche divergence indicates uptake occurred via different prey. Our multidisciplinary assessment demonstrated how nutrients and metal interactions differ in common dolphins depending upon sexual maturity. This approach has relevance when considering how marine pollution, environmental fluctuations and climate change may affect nutritional and trace metal interactions during different reproductive stages within marine predators.
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    Age- and sex-specific survivorship of the Southern Hemisphere long-finned pilot whale (Globicephala melas edwardii)
    (Oxford University Press on behalf of the American Society of Mammalogists, 2022-10-27) Betty EL; Stockin KA; Hinton B; Bollard BA; Orams MB; Murphy S; Swanson B
    Biodiversity loss is a major global challenge of the 21st century. Ultimately, extinctions of species are determined by birth and death rates; thus, conservation management of at-risk species is dependent on robust demographic data. In this study, data gathered from 381 (227 females, 154 males) long-finned pilot whales (Globicephala melas edwardii) that died in 14 stranding events on the New Zealand coast between 2006 and 2017 were used to construct the first age- and sex-specific life tables for the subspecies. Survivorship curves were fitted to these data using (1) a traditional maximum likelihood approach, and (2) Siler's competing-risk model. Life table construction and subsequent survival curves revealed distinct differences in the age- and sex-specific survival rates, with females outliving males. Both sexes revealed slightly elevated rates of mortality among the youngest age-classes (<2 years) with postweaning mortality rates decreasing and remaining relatively low until the average life expectancy is reached; 11.3 years for males and 14.7 years for females. Overall (total) mortality is estimated to be 8.8% and 6.8% per annum for males and females, respectively. The mortality curve resembles that of other large mammals, with high calf mortality, lower postweaning mortality, and an exponentially increasing risk of senescent mortality. An accelerated mortality rate was observed in mature females, in contrast to the closely related short-finned pilot whale (G. macrorhynchus), which selects for an extension to the postreproductive life span. The reason for the observed differences in the mortality rate acceleration and postreproductive life span between the two pilot whale species have not been established and warrant further investigation. Obtaining robust information on the life history of long-lived species is challenging, but essential to improve our understanding of population dynamics and help predict how future pressures may impact populations. This study illustrates how demographic data from cetacean stranding events can improve knowledge of species survival rates, thus providing essential information for conservation management.
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    Understanding common dolphin and Australasian gannet feeding associations from nutritional and ethological perspectives
    (Oxford University Press on behalf of International Council for the Exploration of the Sea, 2022-09-01) Stockin KA; Amiot C; Meynier L; Purvin C; Machovsky-Capuska GE
    Prey detection and subsequent capture is considered a major hypothesis to explain feeding associations between common dolphins and Australasian gannets. However, a current lack of insight on nutritional strategies with respect to foraging behaviours of both species has until now, prevented any detailed understanding of this conspecific relationship. Here we combine stomach content analysis (SCA), nutritional composition of prey, a multidimensional nutritional niche framework (MNNF) and videography to provide a holistic dietary, nutritional, and behavioural assessment of the feeding association between dolphins and gannets in the Hauraki Gulf, New Zealand. Dolphins consumed ten prey species, including grey mullet (Mugil cephalus) as the most representative by wet mass (33.4%). Gannets preyed upon six species, with pilchards (Sardinops pilchardus) contributing most of the diet by wet mass (32.4%) to their diet. Both predators jointly preyed upon pilchard, jack mackerel (Trachurus spp.), arrow squid (genus Nototodarus), and anchovy (Engraulis australis). Accordingly, the MNNF revealed a moderate overlap in the prey composition niche (0.42) and realized nutritional niche (0.52) between dolphins and gannets. This suggests that both predators coexist in a similar nutritional space, while simultaneously reducing interspecific competition and maximizing the success of both encountering and exploiting patchily distributed prey. Behavioural analysis further indicated that dolphin and gannets feeding associations are likely to be mutually beneficial, with a carouselling foraging strategy and larger pod sizes of dolphins, influencing the diving altitude of gannets. Our approach provides a new, more holistic understanding of this iconic foraging relationship, which until now has been poorly understood.