Variable trends in the distribution of Atlantic cod (Gadus morhua) in the Celtic seas.

Despite decades of active fisheries management, many stocks of Atlantic cod in its southern range are in a depleted state and mortality estimates remain high. Recovery of these stocks, as defined by management areas, could be confounded by cod distributions shifting outside of these areas. Here, we assess data from internationally coordinated trawl surveys to investigate the distribution of three cod stocks in the Celtic Seas ecoregion, Irish Sea, Celtic Sea, and West of Scotland, from 1985 to 2021. We mapped cod densities, analyzed trends in mean weighted depth and bottom temperature, and calculated the center of gravity and equivalent area of the stocks. The distribution of the West of Scotland stock shifted north and east, spilling into the North Sea, while the Irish Sea and Celtic Sea stocks shifted west. Each stock showed decreasing trends in equivalent area, but there were no clear trends in the average depth occupied by the fish. There was no apparent relationship between temperature and the distribution of cod, as bottom temperature varied little from 1993 to 2021. Although Irish Sea cod showed a shift into warmer water, this was due to changes in survey distribution. The shift in distribution of the West of Scotland cod stock towards the North Sea whilst impairing local recovery provides further justification for the recent definition of its incorporation into a larger stock unit that includes the northwest of the North Sea. The Irish Sea and Celtic Sea cod stocks are neither shifting northwards, nor into deeper waters, but remained within current boundaries. This suggests that recent temperature conditions did not affect their distribution, but this may change as temperatures increase towards the limit for reproduction.

Examination of head versus body heading may help clarify the extent to which animal movement pathways are structured by environmental cues?

Understanding the processes that determine how animals allocate time to space is a major challenge, although it is acknowledged that summed animal movement pathways over time must define space-time use. The critical question is then, what processes structure these pathways? Following the idea that turns within pathways might be based on environmentally determined decisions, we equipped Arabian oryx with head- and body-mounted tags to determine how they orientated their heads - which we posit is indicative of them assessing the environment - in relation to their movement paths, to investigate the role of environment scanning in path tortuosity. After simulating predators to verify that oryx look directly at objects of interest, we recorded that, during routine movement, > 60% of all turns in the animals' paths, before being executed, were preceded by a change in head heading that was not immediately mirrored by the body heading: The path turn angle (as indicated by the body heading) correlated with a prior change in head heading (with head heading being mirrored by subsequent turns in the path) twenty-one times more than when path turns occurred due to the animals adopting a body heading that went in the opposite direction to the change in head heading. Although we could not determine what the objects of interest were, and therefore the proposed reasons for turning, we suggest that this reflects the use of cephalic senses to detect advantageous environmental features (e.g. food) or to detect detrimental features (e.g. predators). The results of our pilot study suggest how turns might emerge in animal pathways and we propose that examination of points of inflection in highly resolved animal paths could represent decisions in landscapes and their examination could enhance our understanding of how animal pathways are structured.

Impact of test, vaccinate or remove protocol on home ranges and nightly movements of badgers a medium density population.

In the British Isles, the European badger (Meles meles) is thought to be the primary wildlife reservoir of bovine tuberculosis (bTB), an endemic disease in cattle. Test, vaccinate or remove ('TVR') of bTB test-positive badgers, has been suggested to be a potentially useful protocol to reduce bTB incidence in cattle. However, the practice of removing or culling badgers is controversial both for ethical reasons and because there is no consistent observed effect on bTB levels in cattle. While removing badgers reduces population density, it may also result in disruption of their social behaviour, increase their ranging, and lead to greater intra- and inter-species bTB transmission. This effect has been recorded in high badger density areas, such as in southwest England. However, little is known about how TVR affects the behaviour and movement of badgers within a medium density population, such as those that occur in Northern Ireland (NI), which the current study aimed to examine. During 2014-2017, badger ranging behaviours were examined prior to and during a TVR protocol in NI. Nightly distances travelled by 38 individuals were determined using Global Positioning System (GPS) measurements of animal tracks and GPS-enhanced dead-reckoned tracks. The latter was calculated using GPS, tri-axial accelerometer and tri-axial magnetometer data loggers attached to animals. Home range and core home range size were measured using 95% and 50% autocorrelated kernel density estimates, respectively, based on location fixes. TVR was not associated with measured increases in either distances travelled per night (mean = 3.31 ± 2.64 km) or home range size (95% mean = 1.56 ± 0.62 km2, 50% mean = 0.39 ± 0.62 km2) over the four years of study. However, following trapping, mean distances travelled per night increased by up to 44% eight days post capture. Findings differ from those observed in higher density badger populations in England, in which badger ranging increased following culling. Whilst we did not assess behaviours of individual badgers, possible reasons why no differences in home range size were observed include higher inherent 'social fluidity' in Irish populations whereby movements are less restricted by habitat saturation and/or that the numbers removed did not reach a threshold that might induce increases in ranging behaviour. Nevertheless, short-term behavioural disruption from trapping was observed, which led to significant increases in the movements of individual animals within their home range. Whether or not TVR may alter badger behaviours remains to be seen, but it would be better to utilise solutions such as oral vaccination of badgers and/or cattle as well as increased biosecurity to limit bTB transmission, which may be less likely to cause interference and thereby reduce the likelihood of bTB transmission.

Orexinergic neurons in the hypothalami of an Asiatic lion, an African lion, and a Southeast African cheetah.

Employing orexin-A immunohistochemistry, we describe the distribution, morphology, and nuclear parcellation of orexinergic neurons within the hypothalami of an Asiatic lion (Panthera leo subsp. persica), an African lion (Panthera leo subsp. melanochaita), and a Southeast African cheetah (Acinonyx jubatus subsp. jubatus). In all three felids, the clustering of large, bipolar, and multipolar hypothalamic orexinergic neurons primarily follows the pattern observed in other mammals. The orexinergic neurons were found, primarily, to form three distinct clusters-the main, zona incerta, and optic tract clusters. In addition, large orexinergic neurons were observed in the ventromedial supraoptic region of the hypothalamus, where they are not typically observed in other species. As has been observed in cetartiodactyls and the African elephant, a cluster of small, multipolar orexinergic neurons, the parvocellular cluster, was observed in the medial zone of the hypothalamus in all three felids, although this parvocellular cluster has not been reported in other carnivores. In both subspecies of lions, but not the cheetah, potential orexin-immunopositive neurons were observed in the paraventricular hypothalamic nucleus, supraoptic nucleus, the lateral part of the retrochiasmatic area, and the inner layer of the median eminence. The distribution and parcellation of orexinergic neurons in the hypothalami of the three felids studied appear to be more complex than observed in many other mammals and for the two subspecies of lion may be even more complex. These findings are discussed in terms of potential technical concerns, phylogenetic variations of this system, and potentially associated functional aspects of the orexinergic system.

Decision rules for determining terrestrial movement and the consequences for filtering high-resolution global positioning system tracks: a case study using the African lion (Panthera leo).

The combined use of global positioning system (GPS) technology and motion sensors within the discipline of movement ecology has increased over recent years. This is particularly the case for instrumented wildlife, with many studies now opting to record parameters at high (infra-second) sampling frequencies. However, the detail with which GPS loggers can elucidate fine-scale movement depends on the precision and accuracy of fixes, with accuracy being affected by signal reception. We hypothesized that animal behaviour was the main factor affecting fix inaccuracy, with inherent GPS positional noise (jitter) being most apparent during GPS fixes for non-moving locations, thereby producing disproportionate error during rest periods. A movement-verified filtering (MVF) protocol was constructed to compare GPS-derived speed data with dynamic body acceleration, to provide a computationally quick method for identifying genuine travelling movement. This method was tested on 11 free-ranging lions (Panthera leo) fitted with collar-mounted GPS units and tri-axial motion sensors recording at 1 and 40 Hz, respectively. The findings support the hypothesis and show that distance moved estimates were, on average, overestimated by greater than 80% prior to GPS screening. We present the conceptual and mathematical protocols for screening fix inaccuracy within high-resolution GPS datasets and demonstrate the importance that MVF has for avoiding inaccurate and biased estimates of movement.

Soil δ13C and δ15N baselines clarify biogeographic heterogeneity in isotopic discrimination of European badgers (Meles meles).

Isotopic techniques have been used to study phenomena in the geological, environmental, and ecological sciences. For example, isotopic values of multiple elements elucidate the pathways energy and nutrients take in the environment. Isoscapes interpolate isotopic values across a geographical surface and are used to study environmental processes in space and time. Thus, isoscapes can reveal ecological shifts at local scales, and show distribution thresholds in the wider environment at the macro-scale. This study demonstrates a further application of isoscapes, using soil isoscapes of 13C/12C and 15N/14N as an environmental baseline, to understand variation in trophic ecology across a population of Eurasian badgers (Meles meles) at a regional scale. The use of soil isoscapes reduced error, and elevated the statistical signal, where aggregated badger hairs were used, and where individuals were identified using genetic microarray analysis. Stable isotope values were affected by land-use type, elevation, and meteorology. Badgers in lowland habitats had diets richer in protein and were adversely affected by poor weather conditions in all land classes. It is concluded that soil isoscapes are an effective way of reducing confounding biases in macroscale, isotopic studies. The method elucidated variation in the trophic and spatial ecology of economically important taxa at a landscape level. These results have implications for the management of badgers and other carnivores with omnivorous tendencies in heterogeneous landscapes.

Animal lifestyle affects acceptable mass limits for attached tags.

Animal-attached devices have transformed our understanding of vertebrate ecology. To minimize any associated harm, researchers have long advocated that tag masses should not exceed 3% of carrier body mass. However, this ignores tag forces resulting from animal movement. Using data from collar-attached accelerometers on 10 diverse free-ranging terrestrial species from koalas to cheetahs, we detail a tag-based acceleration method to clarify acceptable tag mass limits. We quantify animal athleticism in terms of fractions of animal movement time devoted to different collar-recorded accelerations and convert those accelerations to forces (acceleration × tag mass) to allow derivation of any defined force limits for specified fractions of any animal's active time. Specifying that tags should exert forces that are less than 3% of the gravitational force exerted on the animal's body for 95% of the time led to corrected tag masses that should constitute between 1.6% and 2.98% of carrier mass, depending on athleticism. Strikingly, in four carnivore species encompassing two orders of magnitude in mass (ca 2-200 kg), forces exerted by '3%' tags were equivalent to 4-19% of carrier body mass during moving, with a maximum of 54% in a hunting cheetah. This fundamentally changes how acceptable tag mass limits should be determined by ethics bodies, irrespective of the force and time limits specified.

Quantifying intraherd cattle movement metrics: Implications for disease transmission risk.

There are a paucity of data quantifying on-farm management practices such as the frequency of intraherd cattle movements, use of consolidated or spatially fragmented grazing pastures, and duration of time cattle spend at grass with respect to biosecurity and disease transmission. Such movement dynamics are important when attempting to understand the maintenance of chronic infectious disease, such as bovine tuberculosis (bTB). We captured empirical data on daily cattle movements for a sample of eighteen farms throughout one complete grazing season (n = 18,988 grazing days) and assessed these attributes in relation to herd bTB risk. Dairy herds were stocked at significantly higher densities compared to beef production systems (6.6 animals/ha, 95 % confidence intervals (CI) 6.5-6.7 and 4.1 animals/ha, 95 %CI 4.1 - 4.1 respectively, p < 0.001). Most notably milking cows, were grazed at higher densities than other life stages (e.g. calves, heifers and bullocks) (p < 0.001) and experienced four times the number of movements between pastures. Beef cattle were more likely to be grazed across multiple (rather than single) fields (p < 0.001), with greater time spent on fragmented land away from the main/home farm (p < 0.001). None of the farm or herd attributes analysed (e.g. stocking density, frequency of movement, movement distances or land fragmentation) were associated with herd bovine tuberculosis (bTB) breakdowns during this study. However, there was a weak positive association between bTB breakdowns during the 3 years prior to the study and cattle movement distances (p = 0.05) and time spent on fragmented land (p = 0.08). After a bTB breakdown occurs, restrictions on animals moving out of these herds are implemented to control disease spread, yet we argue that more attention is needed on the role of intraherd grazing patterns in modelling disease transmission risk between herds.

Grazing cattle exposure to neighbouring herds and badgers in relation to bovine tuberculosis risk.

Bovine tuberculosis (bTB) can be spread between and among cattle and wildlife hosts e.g. European badger (Meles meles). The majority of cattle in the UK and Ireland are grazed during the summer, potentially exposing them to Mycobacterium bovis. 18 farms were surveyed (39% dairy, 61% beef; fields n = 697) for one grazing season (May-November 2016, n = 148,461 field days) to quantify the co-occurrence of cattle with badger setts and latrines and adjacency to neighbouring cattle herds. 3% (n = 24) of the fields had a badger sett or latrine recorded, dairy cattle were significantly more likely to co-occur with badger setts and latrines than beef cattle. Most farms (89%) grazed cattle adjacent to a neighbouring herd, which accounted for 18% of the grazing season. Potential exposure to neighbouring herds did not differ between production systems but did vary between life stages. A significant positive association between the proportion of time cattle spent grazing fields with setts present and the historic 1-, 3- and 5- year bTB status (p = 0.007, p = 0.013 and p = 0.013 respectively) was found. However, when cattle were grazed in fields with latrines, a significant negative association was found between the proportion of time cattle spent grazing fields with latrines present and the historic 3- and 5- year bTB status (p = 0.033 and p = 0.012 respectively). Historic bTB status and percentage of days spent beside a neighbouring herd was unrelated. Idiosyncrasies at farm-level and between risk factors indicated that individual farm assessments would be beneficial to understand potential exposure risk.

Surviving in steep terrain: a lab-to-field assessment of locomotor costs for wild mountain lions (Puma concolor).

BACKGROUND: Under current scenarios of climate change and habitat loss, many wild animals, especially large predators, are moving into novel energetically challenging environments. Consequently, changes in terrain associated with such moves may heighten energetic costs and effect the decline of populations in new localities. METHODS: To examine locomotor costs of a large carnivorous mammal moving in mountainous habitats, the oxygen consumption of captive pumas (Puma concolor) was measured during treadmill locomotion on level and incline (6.8°) surfaces. These data were used to predict energetic costs of locomotor behaviours of free-ranging pumas equipped with GPS/accelerometer collars in California's Santa Cruz Mountains. RESULTS: Incline walking resulted in a 42.0% ± 7.2 SEM increase in the costs of transport compared to level performance. Pumas negotiated steep terrain by traversing across hillsides (mean hill incline 17.2° ± 0.3 SEM; mean path incline 7.3° ± 0.1 SEM). Pumas also walked more slowly up steeper paths, thereby minimizing the energetic impact of vertical terrains. Estimated daily energy expenditure (DEE) based on GPS-derived speeds of free-ranging pumas was 18.3 MJ day- 1 ± 0.2 SEM. Calculations show that a 20 degree increase in mean steepness of the terrain would increase puma DEE by less than 1% as they only spend a small proportion (10%) of their day travelling. They also avoided elevated costs by utilizing slower speeds and shallower path angles. CONCLUSIONS: While many factors influence survival in novel habitats, we illustrate the importance of behaviours which reduce locomotor costs when traversing new, energetically challenging environments, and demonstrate that these behaviours are utilised by pumas in the wild.

Estimates for energy expenditure in free-living animals using acceleration proxies: A reappraisal.

It is fundamentally important for many animal ecologists to quantify the costs of animal activities, although it is not straightforward to do so. The recording of triaxial acceleration by animal-attached devices has been proposed as a way forward for this, with the specific suggestion that dynamic body acceleration (DBA) be used as a proxy for movement-based power. Dynamic body acceleration has now been validated frequently, both in the laboratory and in the field, although the literature still shows that some aspects of DBA theory and practice are misunderstood. Here, we examine the theory behind DBA and employ modelling approaches to assess factors that affect the link between DBA and energy expenditure, from the deployment of the tag, through to the calibration of DBA with energy use in laboratory and field settings. Using data from a range of species and movement modes, we illustrate that vectorial and additive DBA metrics are proportional to each other. Either can be used as a proxy for energy and summed to estimate total energy expended over a given period, or divided by time to give a proxy for movement-related metabolic power. Nonetheless, we highlight how the ability of DBA to predict metabolic rate declines as the contribution of non-movement-related factors, such as heat production, increases. Overall, DBA seems to be a substantive proxy for movement-based power but consideration of other movement-related metrics, such as the static body acceleration and the rate of change of body pitch and roll, may enable researchers to refine movement-based metabolic costs, particularly in animals where movement is not characterized by marked changes in body acceleration.

Towards informed metrics for examining the role of human-induced animal responses in tag studies on wild animals.

Two prime issues can detrimentally affect animals that have been equipped with tags: (i) the effect of the capture and restraint process; and (ii) the effect of the tag itself. This work examines some of the issues surrounding quantification of tag effects on wild animals for both restrained and free-living animals. A new method to quantify stress effects based on monitoring ventilation rates in relation to activity is suggested for restrained animals which may help improve the practice of handling animals. It is also suggested that various metrics, many derived from accelerometers, can be examined in tagged wild animals to examine the change in behaviors over time with a view to having a better understanding of welfare issues, assuring the quality of recorded data and informing best practice.

Luck in Food Finding Affects Individual Performance and Population Trajectories.

Energy harvesting by animals is important because it provides the power needed for all metabolic processes. Beyond this, efficient food finding enhances individual fitness [1] and population viability [2], although rates of energy accumulation are affected by the environment and food distribution. Typically, differences between individuals in the rate of food acquisition are attributed to varying competencies [3], even though food-encounter rates are known to be probabilistic [4]. We used animal-attached technology to quantify food intake in four disparate free-living vertebrates (condors, cheetahs, penguins, and sheep) and found that inter-individual variability depended critically on the probability of food encounter. We modeled this to reveal that animals taking rarer food, such as apex predators and scavengers, are particularly susceptible to breeding failure because this variability results in larger proportions of the population failing to accrue the necessary resources for their young before they starve and because even small changes in food abundance can affect this variability disproportionately. A test of our model on wild animals indicated why Magellanic penguins have a stable population while the congeneric African penguin population has declined for decades. We suggest that such models predicting probabilistic ruin can help predict the fortunes of species operating under globally changing conditions.

Prying into the intimate secrets of animal lives; software beyond hardware for comprehensive annotation in 'Daily Diary' tags.

BACKGROUND: Smart tags attached to freely-roaming animals recording multiple parameters at infra-second rates are becoming commonplace, and are transforming our understanding of the way wild animals behave. Interpretation of such data is complex and currently limits the ability of biologists to realise the value of their recorded information. DESCRIPTION: This work presents Framework4, an all-encompassing software suite which operates on smart sensor data to determine the 4 key elements considered pivotal for movement analysis from such tags (Endangered Species Res 4: 123-37, 2008). These are; animal trajectory, behaviour, energy expenditure and quantification of the environment in which the animal moves. The program transforms smart sensor data into dead-reckoned movements, template-matched behaviours, dynamic body acceleration-derived energetics and position-linked environmental data before outputting it all into a single file. Biologists are thus left with a single data set where animal actions and environmental conditions can be linked across time and space. CONCLUSIONS: Framework4 is a user-friendly software that assists biologists in elucidating 4 key aspects of wild animal ecology using data derived from tags with multiple sensors recording at high rates. Its use should enhance the ability of biologists to derive meaningful data rapidly from complex data.