Assessing stress in Western Hudson Bay polar bears using hair cortisol concentration as a biomarker

The development of novel biomarkers to help assess whether polar bear (Ursus maritimus) health is impacted by long-term physiological stress associated with climate change represents an emerging area of research. Reductions in sea ice cover and food availability are potentially stressful, and chronic stress can have deleterious effects that may impair individual and population level health. Cortisol is the principal effector hormone of the stress response and has previously been linked to aspects of polar bear life history (e.g., reproduction and growth) known to be negatively influenced by environmental change. Understanding stress is important for polar bears at the southern limit of their range, such as those in Western Hudson Bay (WH), where rapidly changing sea ice phenology threatens population viability. We examined the relationships between age, reproductive status, and body condition (fatness) and hair cortisol concentration (HCC) in 729 polar bear hair samples collected in WH from 2004–2013. Overall, there was a negative relationship between fatness and HCC, suggesting that bears in poorer body condition experienced higher levels of stress. However, when reproductive status was included in our analysis, this relationship only held for male and lone female bears. Females with dependent offspring had consistently low fatness and elevated HCC, likely because of the high cost of maternal care. We also found a positive correlation between HCC and age for: (1) bears in poor body condition, possibly due to nutritional stress compounding effects of aging; and (2) male bears, potentially due to stress and injury associated with intrasexual mate competition. These findings support the use of HCC as a biomarker for polar bear health. Furthermore, we have established a HCC benchmark against which future population-level effects of climate change in WH polar bears can be compared.     

Movements of female polar bears (<Emphasis Type="Italic">Ursus maritimus</Emphasis>) in the East Greenland pack ice

The movements of two adult female polar bears ( Ursus maritimus) in East Greenland and the Greenland Sea area were studied by use of satellite telemetry between the fall of 1994 and the summer of 1998. One female was tracked for 621 days, the other for 1,415 days. During this time the females used maternity dens on land. If denning periods on land were excluded, the two females used between 73% and 100% of the tracking time offshore where they were able to navigate in the dynamic pack ice and counteract the fast southward movement of the ice (up to 30 km/h) in the East Greenland Current. Mean monthly movement rates varied between 0.32 and 0.76km/h. Both bears had very large home ranges (242,000 and 468,000 km ²) within the dynamic pack ice of the Greenland Sea. The facts that the bears made extensive use of the offshore sea ice and that there is a marked reduction of the Greenland Sea ice call for a closer monitoring of the effects of this change on the East Greenland polar bear population.     

Field Metabolic Rate and PCB Adipose Tissue Deposition Efficiency in East Greenland Polar Bears Derived from Contaminant Monitoring Data

Climate change will increasingly affect the natural habitat and diet of polar bears (Ursus maritimus). Understanding the energetic needs of polar bears is therefore important. We developed a theoretical method for estimating polar bear food consumption based on using the highly recalcitrant polychlorinated biphenyl (PCB) congener, 2,2′,4,4′,55-hexaCB (CB153) in bear adipose tissue as an indicator of food intake. By comparing the CB153 tissue concentrations in wild polar bears with estimates from a purposely designed individual-based model, we identified the possible combinations of field metabolic rates (FMR) and CB153 deposition efficiencies in East Greenland polar bears. Our simulations indicate that if 30% of the CB153 consumed by polar bear individuals were deposited into their adipose tissue, the corresponding FMR would be only two times the basal metabolic rate. In contrast, if the modelled CB153 deposition efficiency were 10%, adult polar bears would require six times more energy than that needed to cover basal metabolism. This is considerably higher than what has been assumed for polar bears in previous studies though it is similar to FMRs found in other marine mammals. An implication of this result is that even relatively small reductions in future feeding opportunities could impact the survival of East Greenland polar bears.     

Population substructure and space use of Foxe Basin polar bears

Climate change has been identified as a major driver of habitat change, particularly for sea ice-dependent species such as the polar bear (Ursus maritimus). Population structure and space use of polar bears have been challenging to quantify because of their circumpolar distribution and tendency to range over large areas. Knowledge of movement patterns, home range, and habitat is needed for conservation and management. This is the first study to examine the spatial ecology of polar bears in the Foxe Basin management unit of Nunavut, Canada. Foxe Basin is in the mid-Arctic, part of the seasonal sea ice ecoregion and it is being negatively affected by climate change. Our objectives were to examine intrapopulation spatial structure, to determine movement patterns, and to consider how polar bear movements may respond to changing sea ice habitat conditions. Hierarchical and fuzzy cluster analyses were used to assess intrapopulation spatial structure of geographic position system satellite-collared female polar bears. Seasonal and annual movement metrics (home range, movement rates, time on ice) and home-range fidelity (static and dynamic overlap) were compared to examine the influence of regional sea ice on movements. The polar bears were distributed in three spatial clusters, and there were differences in the movement metrics between clusters that may reflect sea ice habitat conditions. Within the clusters, bears moved independently of each other. Annual and seasonal home-range fidelity was observed, and the bears used two movement patterns: on-ice range residency and annual migration. We predict that home-range fidelity may decline as the spatial and temporal predictability of sea ice changes. These new findings also provide baseline information for managing and monitoring this polar bear population.     

Distribution,abundance and biology of polar cod, <Emphasis Type="Italic">Boreogadus saida</Emphasis>, in Iceland–East Greenland waters

Recent warming has caused a northern extension in the distribution of many southern fish species in Icelandic waters. Polar cod (Boreogadus saida) around Iceland are near the southern limit of their distribution, but are poorly studied in the area. Therefore, material sampled during demersal fish surveys in March 1985–2013 and in pelagic 0-group surveys in Iceland–East Greenland waters in August–September 1974–2003 was used to investigate their distribution, abundance and biology. Demersal polar cod were most often caught on the outer shelf to the north-west and north of Iceland, but during years of widest distribution and highest abundance, they were caught farther to the east on the northern shelf. Pelagic 0-group polar cod were only caught sporadically and mainly confined to the waters off the north-west shelf of Iceland and the East Greenland shelf (southern Denmark Strait). In demersal hauls, the number of stations with polar cod decreased with increasing bottom temperature and polar cod were most widely distributed in the years 1989, 1994 and 1995. Highest numbers of demersal polar cod per haul were caught at temperatures of ?0.5 to 2.5 °C and at 200–450 m depth. The length of demersal polar cod ranged from 5 to 32 cm, while the fish caught in the pelagic trawl ranged from 2.2 to 19 cm. The polar cod in the subarctic waters north of Iceland most likely originate from the waters off East Greenland and further warming and decline in sea ice may eventually lead to the disappearance of polar cod from Icelandic waters.     

Effects of Earlier Sea Ice Breakup on Survival and Population Size of Polar Bears in Western Hudson Bay

ABSTRACT Some of the most pronounced ecological responses to climatic warming are expected to occur in polar marine regions, where temperature increases have been the greatest and sea ice provides a sensitive mechanism by which climatic conditions affect sympagic (i.e., with ice) species. Population-level effects of climatic change, however, remain difficult to quantify. We used a flexible extension of Cormack-Jolly-Seber capture-recapture models to estimate population size and survival for polar bears (Ursus maritimus), one of the most ice-dependent of Arctic marine mammals. We analyzed data for polar bears captured from 1984 to 2004 along the western coast of Hudson Bay and in the community of Churchill, Manitoba, Canada. The Western Hudson Bay polar bear population declined from 1,194 (95% CI = 1,020-1,368) in 1987 to 935 (95% CI = 794-1,076) in 2004. Total apparent survival of prime-adult polar bears (5–19 yr) was stable for females (0.93; 95% CI = 0.91-0.94) and males (0.90; 95% CI = 0.88-0.91). Survival of juvenile, subadult, and senescent-adult polar bears was correlated with spring sea ice breakup date, which was variable among years and occurred approximately 3 weeks earlier in 2004 than in 1984. We propose that this correlation provides evidence for a causal association between earlier sea ice breakup (due to climatic warming) and decreased polar bear survival. It may also explain why Churchill, like other communities along the western coast of Hudson Bay, has experienced an increase in human-polar bear interactions in recent years. Earlier sea ice breakup may have resulted in a larger number of nutritionally stressed polar bears, which are encroaching on human habitations in search of supplemental food. Because western Hudson Bay is near the southern limit of the species' range, our findings may foreshadow the demographic responses and management challenges that more northerly polar bear populations will experience if climatic warming in the Arctic continues as projected.     

Dietary habits of polar bears in Foxe Basin,Canada: possible evidence of a trophic regime shift mediated by a new top predator

Polar bear (Ursus maritimus) subpopulations in several areas with seasonal sea ice regimes have shown declines in body condition, reproductive rates, or abundance as a result of declining sea ice habitat. In the Foxe Basin region of Nunavut, Canada, the size of the polar bear subpopulation has remained largely stable over the past 20 years, despite concurrent declines in sea ice habitat. We used fatty acid analysis to examine polar bear feeding habits in Foxe Basin and thus potentially identify ecological factors contributing to population stability. Adipose tissue samples were collected from 103 polar bears harvested during 2010–2012. Polar bear diet composition varied spatially within the region with ringed seal (Pusa hispida) comprising the primary prey in northern and southern Foxe Basin, whereas polar bears in Hudson Strait consumed equal proportions of ringed seal and harp seal (Pagophilus groenlandicus). Walrus (Odobenus rosmarus) consumption was highest in northern Foxe Basin, a trend driven by the ability of adult male bears to capture large‐bodied prey. Importantly, bowhead whale (Balaena mysticetus) contributed to polar bear diets in all areas and all age and sex classes. Bowhead carcasses resulting from killer whale (Orcinus orca) predation and subsistence harvest potentially provide an important supplementary food source for polar bears during the ice‐free period. Our results suggest that the increasing abundance of killer whales and bowhead whales in the region could be indirectly contributing to improved polar bear foraging success despite declining sea ice habitat. However, this indirect interaction between top predators may be temporary if continued sea ice declines eventually severely limit on‐ice feeding opportunities for polar bears.     

Projected Polar Bear Sea Ice Habitat in the Canadian Arctic Archipelago

Background

Sea ice across the Arctic is declining and altering physical characteristics of marine ecosystems. Polar bears (Ursus maritimus) have been identified as vulnerable to changes in sea ice conditions. We use sea ice projections for the Canadian Arctic Archipelago from 2006 – 2100 to gain insight into the conservation challenges for polar bears with respect to habitat loss using metrics developed from polar bear energetics modeling.

Principal Findings

Shifts away from multiyear ice to annual ice cover throughout the region, as well as lengthening ice-free periods, may become critical for polar bears before the end of the 21^st century with projected warming. Each polar bear population in the Archipelago may undergo 2–5 months of ice-free conditions, where no such conditions exist presently. We identify spatially and temporally explicit ice-free periods that extend beyond what polar bears require for nutritional and reproductive demands.

Conclusions/Significance

Under business-as-usual climate projections, polar bears may face starvation and reproductive failure across the entire Archipelago by the year 2100.     

Differences in growth,size and sexual dimorphism in skulls of East Greenland and Svalbard polar bears (<Emphasis Type="Italic">Ursus maritimus</Emphasis>)

Size, growth and sexual dimorphism of nine skull traits was studied in 300 East Greenland and 391 Svalbard polar bears (Ursus maritimus). Two traits were significantly larger in bears from East Greenland compared to Svalbard bears, and trait size was smaller after 1960 in five traits. For both localities and both age groups (subadult, adult), mean trait size values were higher in males than females (all: P < 0.05). Gompertz growth models showed trait size increasing with age in seven traits. Depending on the trait, males reached 95% asymptotic trait size at age 3–10, females at age 2–6. The females of both localities matured at approximately the same age, whereas the Svalbard males generally matured years later than their East Greenland peers. The differences found in the present study between the two polar bear subpopulations support the notion that East Greenland and Svalbard polar bears probably should be managed as separate units.     

Longer ice-free seasons increase the risk of nest depredation by polar bears for colonial breeding birds in the Canadian Arctic

Northern polar regions have warmed more than other parts of the globe potentially amplifying the effects of climate change on biological communities. Ice-free seasons are becoming longer in many areas, which has reduced the time available to polar bears (Ursus maritimus) to hunt for seals and hampered bears’ ability to meet their energetic demands. In this study, we examined polar bears’ use of an ancillary prey resource, eggs of colonial nesting birds, in relation to diminishing sea ice coverage in a low latitude region of the Canadian Arctic. Long-term monitoring reveals that bear incursions onto common eider (Somateria mollissima) and thick-billed murre (Uria lomvia) nesting colonies have increased greater than sevenfold since the 1980s and that there is an inverse correlation between ice season length and bear presence. In surveys encompassing more than 1000 km of coastline during years of record low ice coverage (2010–2012), we encountered bears or bear sign on 34% of eider colonies and estimated greater egg loss as a consequence of depredation by bears than by more customary nest predators, such as foxes and gulls. Our findings demonstrate how changes in abiotic conditions caused by climate change have altered predator–prey dynamics and are leading to cascading ecological impacts in Arctic ecosystems.     

A tale of two polar bear populations: ice habitat, harvest, and body condition

One of the primary mechanisms by which sea ice loss is expected to affect polar bears is via reduced body condition and growth resulting from reduced access to prey. To date, negative effects of sea ice loss have been documented for two of 19 recognized populations. Effects of sea ice loss on other polar bear populations that differ in harvest rate, population density, and/or feeding ecology have been assumed, but empirical support, especially quantitative data on population size, demography, and/or body condition spanning two or more decades, have been lacking. We examined trends in body condition metrics of captured bears and relationships with summertime ice concentration between 1977 and 2010 for the Baffin Bay (BB) and Davis Strait (DS) polar bear populations. Polar bears in these regions occupy areas with annual sea ice that has decreased markedly starting in the 1990s. Despite differences in harvest rate, population density, sea ice concentration, and prey base, polar bears in both populations exhibited positive relationships between body condition and summertime sea ice cover during the recent period of sea ice decline. Furthermore, females and cubs exhibited relationships with sea ice that were not apparent during the earlier period (1977–1990s) when sea ice loss did not occur. We suggest that declining body condition in BB may be a result of recent declines in sea ice habitat. In DS, high population density and/or sea ice loss, may be responsible for the declines in body condition.     

Polar bear predatory behaviour reveals seascape distribution of ringed seal lairs

Ringed seal (Pusa hispida) breeding distribution has been extensively studied across near-shore habitats, but has received limited attention at a seascape scale due to the difficulty in accessing offshore sea ice environments. Employing highly visible predation attempts by polar bears (Ursus maritimus) on ringed seals in subnivean lairs observed by helicopter, the spatial relationship between predatory behaviour and ringed seal breeding habitat was examined. Resource selection functions were used to determine the relative probability of predation attempts on ringed seals in lairs as a function of habitat during a period of low ringed seal natality (2004–2006). Ringed seal pup kill locations were compared between years of low (2003–2006) and high (2007–2011) natality to assess the effect of reproductive output on habitat use. During years of low natality, polar bear hunting attempts were more likely in near-shore fast ice, and pup kills were observed predominately in fast ice (fast = 65 %, pack = 29 %, P = 0.002) at a median distance of 36 km from shore. In years of high natality, pup kills were observed farther from shore (median = 46 km, P = 0.03), and there was no difference in the proportion of observations in fast ice and pack ice (fast = 43 %, pack = 52 %, P = 0.29). These results suggest that the facultative use of adjacent offshore pack ice by breeding ringed seals may be influenced by natality. This study illustrates how documenting the behaviour of a predator can facilitate insight into the distribution of a cryptic prey.     

Are polar bear habitat resource selection functions developed from 1985–1995 data still useful?

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Consequences of long-distance swimming and travel over deep-water pack ice for a female polar bear during a year of extreme sea ice retreat

Polar bears (Ursus maritimus) prefer to live on Arctic sea ice but may swim between ice floes or between sea ice and land. Although anecdotal observations suggest that polar bears are capable of swimming long distances, no data have been available to describe in detail long distance swimming events or the physiological and reproductive consequences of such behavior. Between an initial capture in late August and a recapture in late October 2008, a radio-collared adult female polar bear in the Beaufort Sea made a continuous swim of 687 km over 9 days and then intermittently swam and walked on the sea ice surface an additional 1,800 km. Measures of movement rate, hourly activity, and subcutaneous and external temperature revealed distinct profiles of swimming and walking. Between captures, this polar bear lost 22% of her body mass and her yearling cub. The extraordinary long distance swimming ability of polar bears, which we confirm here, may help them cope with reduced Arctic sea ice. Our observation, however, indicates that long distance swimming in Arctic waters, and travel over deep water pack ice, may result in high energetic costs and compromise reproductive fitness.     

The diet of polar bears (Ursus maritimus) from Svalbard, Norway, inferred from scat analysis

Polar bears (Ursus maritimus) are heavily dependent on marine prey, in particular ice-associated seals, which they hunt on landfast sea ice or free-floating pack ice. Dramatic current (and predicted) losses of sea ice habitat make it increasingly important to gain more knowledge of the relative use by bears of all types of prey from the marine food web as well as from terrestrial sources. This study uses frequency of occurrence of food items in 119 polar bear scats sampled on the sea ice as well as on shore in coastal areas in the Svalbard Archipelago, mainly in spring, between 2003 and 2010 to explore the diet of bears in the region. Ringed seals (Pusa hispida) occurred in 62.2 % (CI 52.8–70.9 %) of the scat samples examined. Various terrestrial plants (32.8 %, CI 24.4–42.0 %) and marine algae (21.8 %, CI 14.8–30.4 %) also occurred frequently in the scats; the significance of this high occurrence of plants and algae is not clear. Bearded seals (Erignathus barbatus) and various bird species constituted only minor components of the diet, while Svalbard reindeer (Rangifer tarandus platyrhynchus) occurred in 9.2 % (CI 4.7–15.9 %) of the scats, indicating that this species may play a more important role than previously reported. The novel combination of genetic analyses of material in the fecal samples along with detailed exploration of the physical–structural properties of prey hairs and plant parts provided a much fuller picture of the diet of polar bears than would have been possible from observational studies of polar bear predation behavior alone. This approach may provide an important tool for monitoring the responses of polar bears to ongoing ecosystem changes that will result from continued warming in the Arctic.     

Sea ice resource selection models for polar bears in the Barents Sea subpopulation

The extent, thickness and age of Arctic sea ice has dramatically declined since the late 1990s, and these trends are predicted to continue. Exploring the habitat use of sea‐ice‐dependent species can help us understand which resources they use and how their distribution responds to a changing environment. The goal of this study was to develop predictive models of the habitat use of an Arctic apex predator. Polar bear Ursus maritimus habitat use in the Barents Sea subpopulation was modelled with seasonal resource selection functions (RSFs) using satellite‐linked telemetry data from 294 collars deployed on female polar bears between 1991 and 2015. Polar bears selected habitat in the Marginal Ice Zone, with a preference for intermediate sea ice concentrations (40–80%). They spent most time in areas with relatively short travel distances to 15 or 75% ice concentration, and during spring and autumn they exhibited a preference for sea ice areas over the continental shelf or over shallower bathymetry). Predictions of the distribution of polar bears in the Barents Sea area can be made for specific sea ice scenarios using these models. Two such predictive distribution maps based on the autumn seasonal model were made and validated against two independent polar bear survey datasets collected in August 2004 and August 2015. The distribution of optimal polar bear habitat has shifted strongly northwards in all seasons of the year during the 25 yr study period.     

Increased Land Use by Chukchi Sea Polar Bears in Relation to Changing Sea Ice Conditions

Recent observations suggest that polar bears (Ursus maritimus) are increasingly using land habitats in some parts of their range, where they have minimal access to their preferred prey, likely in response to loss of their sea ice habitat associated with climatic warming. We used location data from female polar bears fit with satellite radio collars to compare land use patterns in the Chukchi Sea between two periods (1986–1995 and 2008–2013) when substantial summer sea-ice loss occurred. In both time periods, polar bears predominantly occupied sea-ice, although land was used during the summer sea-ice retreat and during the winter for maternal denning. However, the proportion of bears on land for > 7 days between August and October increased between the two periods from 20.0% to 38.9%, and the average duration on land increased by 30 days. The majority of bears that used land in the summer and for denning came to Wrangel and Herald Islands (Russia), highlighting the importance of these northernmost land habitats to Chukchi Sea polar bears. Where bears summered and denned, and how long they spent there, was related to the timing and duration of sea ice retreat. Our results are consistent with other studies supporting increased land use as a common response of polar bears to sea-ice loss. Implications of increased land use for Chukchi Sea polar bears are unclear, because a recent study observed no change in body condition or reproductive indices between the two periods considered here. This result suggests that the ecology of this region may provide a degree of resilience to sea ice loss. However, projections of continued sea ice loss suggest that polar bears in the Chukchi Sea and other parts of the Arctic may increasingly use land habitats in the future, which has the potential to increase nutritional stress and human-polar bear interactions.     

Future sea ice conditions in Western Hudson Bay and consequences for polar bears in the 21st century

The primary habitat of polar bears is sea ice, but in Western Hudson Bay (WH), the seasonal ice cycle forces polar bears ashore each summer. Survival of bears on land in WH is correlated with breakup and the ice‐free season length, and studies suggest that exceeding thresholds in these variables will lead to large declines in the WH population. To estimate when anthropogenic warming may have progressed sufficiently to threaten the persistence of polar bears in WH, we predict changes in the ice cycle and the sea ice concentration (SIC) in spring (the primary feeding period of polar bears) with a high‐resolution sea ice‐ocean model and warming forced with 21st century IPCC greenhouse gas (GHG) emission scenarios: B1 (low), A1B (medium), and A2 (high). We define critical years for polar bears based on proposed thresholds in breakup and ice‐free season and we assess when ice‐cycle conditions cross these thresholds. In the three scenarios, critical years occur more commonly after 2050. From 2001 to 2050, 2 critical years occur under B1 and A2, and 4 under A1B; from 2051 to 2100, 8 critical years occur under B1, 35 under A1B and 41 under A2. Spring SIC in WH is high (>90%) in all three scenarios between 2001 and 2050, but declines rapidly after 2050 in A1B and A2. From 2090 to 2100, the mean spring SIC is 84 (±7)% in B1, 56 (±26)% in A1B and 20 (±13)% in A2. Our predictions suggest that the habitat of polar bears in WH will deteriorate in the 21st century. Ice predictions in A1B and A2 suggest that the polar bear population may struggle to persist after ca. 2050. Predictions under B1 suggest that reducing GHG emissions could allow polar bears to persist in WH throughout the 21st century.     

East Greenland and Barents Sea polar bears (Ursus maritimus): adaptive variation between two populations using skull morphometrics as an indicator of environmental and genetic differences

A morphometric study was conducted on four skull traits of 37 male and 18 female adult East Greenland polar bears (Ursus maritimus) collected 1892-1968, and on 54 male and 44 female adult Barents Sea polar bears collected 1950-1969. The aim was to compare differences in size and shape of the bear skulls using a multivariate approach, characterizing the variation between the two populations using morphometric traits as an indicator of environmental and genetic differences. Mixture analysis testing for geographic differentiation within each population revealed three clusters for Barents Sea males and three clusters for Barents Sea females. East Greenland consisted of one female and one male cluster. A principal component analysis (PCA) conducted on the clusters defined by the mixture analysis, showed that East Greenland and Barents Sea polar bear populations overlapped to a large degree, especially with regards to females. Multivariate analyses of variance (MANOVA) showed no significant differences in morphometric means between the two populations, but differences were detected between clusters from each respective geographic locality. To estimate the importance of genetics and environment in the morphometric differences between the bears, a PCA was performed on the covariance matrix derived from the skull measurements. Skull trait size (PC1) explained approx. 80% of the morphometric variation, whereas shape (PC2) defined approx. 15%, indicating some genetic differentiation. Hence, both environmental and genetic factors seem to have contributed to the observed skull differences between the two populations. Overall, results indicate that many Barents Sea polar bears are morphometrically similar to the East Greenland ones, suggesting an exchange of individuals between the two populations. Furthermore, a subpopulation structure in the Barents Sea population was also indicated from the present analyses, which should be considered with regards to future management decisions.     

Bear-hybrids: behaviour and phenotype

The behaviour of a female and a male bear hybrid (brown bear Ursus arctos x polar bear Ursus maritimus) was studied. Behavioural and morphological comparisons between the hybrids and the two parent-species were made.The observation period covered July to November 2007. Different objects were offered to the bears to evoke new patterns of behaviour. While manipulating the offered objects, both bears showed elements of behaviour also found in polar bears, though the male performed those to a larger extend. Not only in the way of manipulating objects but also in the male's type of stereotype parallels to polar bears can be drawn. Phenotypically both bears possess features of both species, brown bear and polar bear. Concerning it lighter colour of fur and the structure of hair the female hybrid resembles more a polar bear than the male. Summarising, the female appears phenotypically more like a polar bear, the male with respect to the behaviour.