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This study investigates effects of subtle methodological choices on the estimation and biological interpretation of age, growth and reproductive parameters for harbour porpoises. The core analyses are based on a focal Norwegian data set built on samples from 134 harbour porpoises caught incidentally in gillnet fisheries along the Norwegian coast during autumn 2016 and spring 2017. Two contrasting practices for interpretation of seasonal and ontogenetic characteristics of tooth growth layer formation resulted in significant age differences among spring samples of young porpoises and for older animals across seasons. In turn, these differences affected estimates of age at maturity and asymptotic lengths, respectively. We also found significant differences in male age at maturity between two well-documented maturity criteria and between mathematical estimators of age at maturity for both sexes. Two different criteria for corpus albicans classification furthermore resulted in different patterns of ovarian corpora accumulation, which may affect some estimates of fecundity rates and contaminant loads. Both corpora accumulation patterns were also found in reanalysed data from German and Greenlandic porpoises. Based on tabulated overviews of methodological choices made in previous harbour porpoise studies, we argue that several of the issues mentioned above have wider relevance and may affect the validity of meta-analyses as a tool for estimating harbour porpoise sensitivity to extrinsic pressures. Differences in cause of death (COD) composition between data sets can have a similar effect. We demonstrate this in a meta-analysis of published harbour porpoise pregnancy rates, showing significantly higher values for trauma-killed samples compared to samples comprising mixed COD categories. COD also affected the estimated impacts of three previously analysed extrinsic predictors as well as an added predictor for vessel noise levels. We discuss the potential contributions of methodological, biological and anthropogenic factors in shaping observed regional differences in estimates of harbour porpoise life history parameters.

Many large terrestrial mammalian predators use energy-intensive, high-risk, high-gain strategies to pursue large, high-quality prey. However, similar-sized marine mammal predators with even higher field metabolic rates (FMRs) consistently target prey three to six orders of magnitude smaller than themselves. Here, we address the question of how these active and expensive marine mammal predators can gain sufficient energy from consistently targeting small prey during breath-hold dives. Using harbor porpoises as model organisms, we show that hunting small aquatic prey is energetically cheap (<20% increase in FMR) for these marine predators, but it requires them to spend a large proportion (>60%) of time foraging. We conclude that this grazing foraging strategy on small prey is viable for marine mammal predators despite their high FMR because they can hunt near continuously at low marginal expense. Consequently, cessation of foraging due to human disturbance comes at a high cost, as porpoises must maintain their high thermoregulation costs with a reduced energy intake.

Shipping is one of the largest industries globally, with well-known negative impacts on the marine environment. Despite the known negative short-term (minutes to hours) impact of shipping on individual animal behavioural responses, very little is understood about the long-term (months to years) impact on marine species presence and area use. This study took advantage of a planned rerouting of a major shipping lane leading into the Baltic Sea, to investigate the impact on the presence and foraging behaviour of a marine species known to be sensitive to underwater noise, the harbour porpoise (Phocoena phocoena). Passive acoustic monitoring data were collected from 15 stations over two years. Against predictions, no clear change occurred in monthly presence or foraging behaviour of the porpoises, despite the observed changes in noise and vessel traffic. However, long-term heightened noise levels may still impact communication, echolocation, or stress levels of individuals, and needs further investigation.

Predator-prey interactions in marine ecosystems are dynamically influenced by light, as demonstrated by diel vertical migrations of low-trophic level organisms. At high latitudes, the long winter nights can provide foraging opportunities for marine predators targeting vertically migrating prey closer to the surface at night. However, there is limited documentation of such diel patterns in marine predators under extreme light regimes. To address this, we recorded the diving behavior of 17 harbour porpoises just south of the Arctic circle in West Greenland, from summer to winter. Unlike classical diel vertical migration, the porpoises dove 24-37% deeper at night and the frequency of deep dives (> 100 m) increased tenfold as they entered the darkest months. The daily mean depth was negatively correlated with daylength, suggesting an increased diving activity when approaching the polar night. Our findings suggest a light-mediated strategy in which harbour porpoises would either target (i) benthic prey, (ii) pelagic prey migrating seasonally towards the seafloor, or (iii) vertically migrating prey that may be otherwise inaccessible in deeper waters at night, therefore maximizing feeding activity during extended periods of darkness. Extreme light regimes observed at high latitudes are therefore critical in structuring pelagic communities and food webs.

Many man-made marine structures (MMS) will have to be decommissioned in the coming decades. While studies on the impacts of construction of MMS on marine mammals exist, no research has been done on the effects of their decommissioning. The complete removal of an oil and gas platform in Scotland in 2021 provided an opportunity to investigate the response of harbour porpoises to decommissioning. Arrays of broadband noise recorders and echolocation detectors were used to describe noise characteristics produced by decommissioning activities and assess porpoise behaviour. During decommissioning, sound pressure spectral density levels in the frequency range 100 Hz to 48 kHz were 30-40 dB higher than baseline, with vessel presence being the main source of noise. The study detected small-scale (< 2 km) and short-term porpoise displacement during decommissioning, with porpoise occurrence increasing immediately after this. These findings can inform the consenting process for future decommissioning projects.

Cetaceans are well-regarded as sentinels for toxin exposure. Emerging studies suggest that cetaceans can also develop neuropathological changes associated with neurodegenerative disease. The occurrence of neuropathology makes cetaceans an ideal species for examining the impact of marine toxins on the brain across the lifespan. Here, we describe TAR DNA-binding protein 43 (TDP-43) proteinopathy and Alzheimer\'s disease (AD) neuropathological changes in a beached harbor porpoise (Phocoena phocoena) that was exposed to a toxin produced by cyanobacteria called β-N-methylamino-L-alanine (BMAA). We found pathogenic TDP-43 cytoplasmic inclusions in neurons throughout the cerebral cortex, midbrain and brainstem. P62/sequestosome-1, responsible for the autophagy of misfolded proteins, was observed in the amygdala, hippocampus and frontal cortex. Genes implicated in AD and TDP-43 neuropathology such as APP and TARDBP were expressed in the brain. AD neuropathological changes such as amyloid-β plaques, neurofibrillary tangles, granulovacuolar degeneration and Hirano bodies were present in the hippocampus. These findings further support the development of progressive neurodegenerative disease in cetaceans and a potential causative link to cyanobacterial toxins. Climate change, nutrient pollution and industrial waste are increasing the frequency of harmful cyanobacterial blooms. Cyanotoxins like BMAA that are associated with neurodegenerative disease pose an increasing public health risk.

When they are exposed to loud fatiguing sounds in the oceans, marine mammals are susceptible to hearing damage in the form of temporary hearing threshold shifts (TTSs) or permanent hearing threshold shifts. We compared the level-dependent and frequency-dependent susceptibility to TTSs in harbor seals and harbor porpoises, species with different hearing sensitivities in the low- and high-frequency regions. Both species were exposed to 100% duty cycle one-sixth-octave noise bands at frequencies that covered their entire hearing range. In the case of the 6.5 kHz exposure for the harbor seals, a pure tone (continuous wave) was used. TTS was quantified as a function of sound pressure level (SPL) half an octave above the center frequency of the fatiguing sound. The species have different audiograms, but their frequency-specific susceptibility to TTS was more similar. The hearing frequency range in which both species were most susceptible to TTS was 22.5-50 kHz. Furthermore, the frequency ranges were characterized by having similar critical levels (defined as the SPL of the fatiguing sound above which the magnitude of TTS induced as a function of SPL increases more strongly). This standardized between-species comparison indicates that the audiogram is not a good predictor of frequency-dependent susceptibility to TTS.

Human activities in the oceans are increasing and can result in additional mortality on many marine Protected, Endangered or Threatened Species (PETS). It is necessary to implement ambitious measures that aim to restore biodiversity at all nodes of marine food webs and to manage removals resulting from anthropogenic activities. We developed a stochastic surplus production model (SPM) linking abundance and removal processes under the assumption that variations in removals reflect variations in abundance. We then consider several \'harvest\' control rules, included two candidate ones derived from this SPM (which we called \'Anthropogenic Removals Threshold\', or ART), to manage removals of PETS. The two candidate rules hinge on the estimation of a stationary removal rate. We compared these candidate rules to other existing control rules (e.g. potential biological removal or a fixed percentage rule) in three scenarios: (i) a base scenario whereby unbiased but noisy data are available, (ii) scenario whereby abundance estimates are overestimated and (iii) scenario whereby abundance estimates are underestimated. The different rules were tested on a simulated set of data with life-history parameters close to a small-sized cetacean species of conservation interest in the North-East Atlantic, the harbour porpoise (Phocoena phocoena), and in a management strategy evaluation framework. The effectiveness of the rules were assessed by looking at performance metrics, such as time to reach the conservation objectives, the removal limits obtained with the rules or temporal autocorrelation in removal limits. Most control rules were robust against biases in data and allowed to reach conservation objectives with removal limits of similar magnitude when averaged over time. However, one of the candidate rule (ART) displayed greater alignment with policy requirements for PETS such as minimizing removals over time.

An unusual mass mortality event (MME) of harbour seals (Phoca vitulina) and harbour porpoises (Phocoena phocoena) occurred in Denmark and Sweden in June 2007. Prior to this incident, the region had experienced two MMEs in harbour seals caused by Phocine Distemper Virus (PDV) in 1988 and 2002. Although epidemiology and symptoms of the 2007 MME resembled PDV, none of the animals examined for PDV tested positive. Thus, it has been speculated that another - yet unknown - pathogen caused the June 2007 MME. To shed new light on the likely cause of death, we combine previously unpublished veterinary examinations of harbour seals with novel analyses of algal toxins and algal monitoring data. All harbour seals subject to pathological examination showed pneumonia, but were negative for PDV, influenza and coronavirus. Histological analyses revealed septicaemia in multiple animals, and six animals tested positive for Klebsiella pneumonia. Furthermore, we detected the algal Dinophysis toxin DTX-1b (1-115 ng g-1) in five seals subject to toxicology, representing the first time DTX-1b has been detected in marine vertebrates. However, no animals tested positive for both Klebsiella and toxins. Thus, while our relatively small sample size prevent firm conclusions on causative agents, we speculate that the unexplained MME may have been caused by a chance incidence of multiple pathogens acting in parallel in June 2007, including Dinophysis toxin and Klebsiella. Our study illustrates the complexity of wildlife MMEs and highlights the need for thorough sampling during and after MMEs, as well as additional research on and monitoring of DTX-1b and other algal toxins in the region.