Ontogenetic niche partitioning in southern elephant seals from Argentine Patagonia

Elephant seals, Mirounga spp., are highly dimorphic, having different energetic requirements according to age and sex, and foraging in various ecological and oceanographic contexts. Resource partitioning has been shown for the sub-Antarctic populations of southern elephant seals, M. leonina, where colonies are surrounded by narrow shelves that deepen abruptly. In contrast, seals from Península Valdés (Argentina), in the northernmost extent of the breeding range, face an extended, shallow, temperate, and productive continental shelf. We integrated tracking data from 98 animals (juveniles and adults, males and females) gathered over more than two decades, and found that although all available habitats were used, individuals segregated by age and sex. Juvenile males favored shelf habitats, whereas subadult and adult males also used the shelf break. Juvenile females preferred the shelf and the more distant Argentine Basin used by postbreeding and postmolt adult females. Males showed the highest proportion of area-restricted search locations, suggesting more spatially concentrated feeding activity, and likely reflecting a preference for foraging habitat and prey. Our results are consistent with those from other populations, implying that elephant seals show remarkable similarities in habitat use by age and sex classes, despite broad differences in the offshore habitats between sub-Antarctic and temperate ecosystems.     

Segregation in space and time explains the coexistence of two sympatric sub‐Antarctic petrels

Biological communities are shaped by competition between and within species. Competition is often reduced by inter‐ and intraspecific specialization on resources, such as differencet foraging areas or time, allowing similar species to coexist and potentially contributing to reproductive isolation. Here, we examine the simultaneous role of temporal and spatial foraging segregation within and between two sympatric sister species of seabirds, Northern Macronectes halli and Southern Macronectes giganteus Giant Petrels. These species show marked sexual size dimorphism and allochrony (with earlier breeding by Northern Giant Petrels) but this is the first study to test for differences in foraging behaviours and areas across the entire breeding season both between the two species and between the sexes. We tracked males and females of both species in all breeding stages at Bird Island, South Georgia, to test how foraging distribution, behaviour and habitat use vary between and within species in biological time (incubation, brood‐guard or post‐brood stages) and in absolute time (calendar date). Within each breeding stage, both species took trips of comparable duration to similar areas, but due to breeding allochrony they segregated temporally. Northern Giant Petrels had a somewhat smaller foraging range than Southern Giant Petrels, reflecting their greater exploitation of local carrion and probably contributing to their recent higher population growth. Within species, segregation was spatial, with females generally taking longer, more pelagic trips than males, although both sexes of both species showed unexpectedly plastic foraging behaviour. There was little evidence of interspecific differences in habitat use. Thus, in giant petrels, temporal segregation reduces interspecific competition and sexual segregation reduces intraspecific competition. These results demonstrate how both specialization and dynamic changes in foraging strategies at different scales underpin resource division within a community.     

The ecology of 3-D space use in a sexually dimorphic mammal

The distribution of animals is the result of habitat selection according to sex, reproductive status and resource availability. Little is known about how marine predators investigate their 3-dimensional space along both the horizontal and vertical axes and how temporal variation affects space use. In this study, we assessed the spatio-temporal movement of a sexually dimorphic marine mammal, the grey seal Halichoerus grypus by 1) determining seasonal home range size, 2) testing whether space use of seals was affected by water depth, and 3) investigating the vertical movement of seals according to the maximum depth of each dive. Between 1993 and 2005, we fitted 49 grey seals in the Gulf of St. Lawrence with satellite transmitters. We estimated seasonal 95% fixed-kernel home ranges for each individual. For each seal, we tested for selectivity and preference for 4 water depth classes at the home range scale and within the home range. We also evaluated the proportional number of dives made in each water depth classes according to the maximum depth of each dive. Home ranges were 10 times larger in winter than in summer. Seals generally selected habitats <50 m deep. They also mainly dove to depths of 40 m or less. At both scales of selection, preference for shallow areas decreased in winter. We also observed that adults used shallow habitats more than juveniles to establish their home range. A spatial segregation based on sex also occurred at the finer scale of selection where females were more concentrated in the shallowest parts of their home range than males. Segregation in space use according to age and sex classes occurred at both the horizontal and vertical scales. Our results emphasise the importance of studying habitat selection of marine predators in 3-dimensional space, in addition to the temporal scale.     

Sexual segregation in ungulates: a comparative test of three hypotheses

In most social ungulate species, males are larger than females and the sexes live in separate groups outside the breeding season. It is important for our understanding of the evolution of sociality to find out why sexual segregation is so widespread not only in ungulates but also in other mammals. Sexual body size dimorphism was proposed as a central factor in the evolution of sexual segregation in ungulates. We tested three hypotheses put forward to explain sexual segregation: the predation-risk, the forage-selection, and the activity budget hypothesis. We included in our analyses ungulate species ranging from non-dimorphic to extremely dimorphic in body size. We observed oryx, zebra, bighorn sheep and ibex in the field and relied on literature data for 31 additional species. The predation-risk hypothesis predicts that females will use relatively predator-safe habitats, while males are predicted to use habitats with higher predation risk but better food quality. Out of 24 studies on different species of ungulates, females and their offspring chose poorer quality but safer habitat in only eight cases. The forage-selection hypothesis predicts that females would select habitat based on food quality, while males should prefer high forage biomass. In fact, females selected higher quality food in only six out of 18 studies where males and females segregated, in eight studies there was no difference in forage quality and in four studies males were in better quality habitat. The activity budget hypothesis predicts that with increasing dimorphism in body size males and females will increasingly differ in the time spent in different activities. Differences in activity budgets would make it difficult for males and females to stay in mixed-sex groups due to increased costs of synchrony to maintain group cohesion. The predictions of the activity budget hypothesis were confirmed in most cases (22 out of 23 studies). The heavier males were compared to females, the more time females spent foraging compared to males. The bigger the dimorphism in body mass, the more males spent time walking compared to females. Lactating females spent more time foraging than did non-lactating females or males. Whether species were mainly bulk or intermediate feeders did not affect sexual differences in time spent foraging. We conclude that sexual differences in activity budgets are most likely driving sexual segregation and that sexual differences in predation risk or forage selection are additive factors.     

Sexual Segregation in Juvenile New Zealand Sea Lion Foraging Ranges: Implications for Intraspecific Competition,Population Dynamics and Conservation

Sexual segregation (sex differences in spatial organisation and resource use) is observed in a large range of taxa. Investigating causes for sexual segregation is vital for understanding population dynamics and has important conservation implications, as sex differences in foraging ecology may affect vulnerability to area-specific human activities. Although behavioural ecologists have proposed numerous hypotheses for this phenomenon, the underlying causes of sexual segregation are poorly understood. We examined the size-dimorphism and niche divergence hypotheses as potential explanations for sexual segregation in the New Zealand (NZ) sea lion (Phocarctos hookeri), a nationally critical, declining species impacted by trawl fisheries. We used satellite telemetry and linear mixed effects models to investigate sex differences in the foraging ranges of juvenile NZ sea lions. Male trip distances and durations were almost twice as long as female trips, with males foraging over the Auckland Island shelf and in further locations than females. Sex was the most important variable in trip distance, maximum distance travelled from study site, foraging cycle duration and percent time at sea whereas mass and age had small effects on these characteristics. Our findings support the predictions of the niche divergence hypothesis, which suggests that sexual segregation acts to decrease intraspecific resource competition. As a consequence of sexual segregation in foraging ranges, female foraging grounds had proportionally double the overlap with fisheries operations than males. This distribution exposes female juvenile NZ sea lions to a greater risk of resource competition and bycatch from fisheries than males, which can result in higher female mortality. Such sex-biased mortality could impact population dynamics, because female population decline can lead to decreased population fecundity. Thus, effective conservation and management strategies must take into account sex differences in foraging behaviour, as well as differential threat-risk to external impacts such as fisheries bycatch.     

Inter‐ and intrasex habitat partitioning in the highly dimorphic southern elephant seal

Partitioning resources is a key mechanism for avoiding intraspecific competition and maximizing individual energy gain. However, in sexually dimorphic species it is difficult to discern if partitioning is due to competition or the different resource needs of morphologically distinct individuals. In the highly dimorphic southern elephant seal, there are intersexual differences in habitat use; at Iles Kerguelen, males predominantly use shelf waters, while females use deeper oceanic waters. There are equally marked intrasexual differences, with some males using the nearby Kerguelen Plateau, and others using the much more distant Antarctic continental shelf (~2,000 km away). We used this combination of inter and intrasexual behavior to test two hypotheses regarding habitat partitioning in highly dimorphic species. (a) that intersexual differences in habitat use will not appear until the seals diverge in body size and (b) that some habitats have higher rates of energy return than others. In particular, that the Antarctic shelf would provide higher energy returns than the Kerguelen Shelf, to offset the greater cost of travel. We quantified the habitat use of 187 southern elephant seals (102 adult females and 85 subadult males). The seals in the two groups were the same size (~2.4 m) removing the confounding effect of body size. We found that the intersexual differences in habitat use existed before the divergence in body size. Also, we found that the amount of energy gained was the same in all of the major habitats. This suggests that the use of shelf habitats by males is innate, and a trade‐off between the need to access the large benthic prey available on shelf waters, against the higher risk of predation there. Intrasexual differences in habitat use are another trade‐off; although there are fewer predators on the Antarctic shelf, it is subject to considerable interannual fluctuations in sea‐ice extent. In contrast, the Kerguelen Plateau presents more consistent foraging opportunities, but contains higher levels of predation. Habitat partitioning in this highly dimorphic species is therefore the result of complex interplay of life history strategies, environmental conditions and predation pressure.     

Sex promotes spatial and dietary segregation in a migratory shorebird during the non-breeding season

Several expressions of sexual segregation have been described in animals, especially in those exhibiting conspicuous dimorphism. Outside the breeding season, segregation has been mostly attributed to size or age-mediated dominance or to trophic niche divergence. Regardless of the recognized implications for population dynamics, the ecological causes and consequences of sexual segregation are still poorly understood. We investigate the foraging habits of a shorebird showing reversed sexual dimorphism, the black-tailed godwit Limosa limosa, during the winter season, and found extensive segregation between sexes in spatial distribution, microhabitat use and dietary composition. Males and females exhibited high site-fidelity but differed in their distributions at estuary-scale. Male godwits (shorter-billed) foraged more frequently in exposed mudflats than in patches with higher water levels, and consumed more bivalves and gastropods and fewer polychaetes than females. Females tended to be more frequently involved and to win more aggressive interactions than males. However, the number of aggressions recorded was low, suggesting that sexual dominance plays a lesser role in segregation, although its importance cannot be ruled out. Dimorphism in the feeding apparatus has been used to explain sex differences in foraging ecology and behaviour of many avian species, but few studies confirmed that morphologic characteristics drive individual differences within each sex. We found a relationship between resource use and bill size when pooling data from males and females. However, this relationship did not hold for either sex separately, suggesting that differences in foraging habits of godwits are primarily a function of sex, rather than bill size. Hence, the exact mechanisms through which this segregation operates are still unknown. The recorded differences in spatial distribution and resource use might expose male and female to distinct threats, thus affecting population dynamics through differential mortality. Therefore, population models and effective conservation strategies should increasingly take sex-specific requirements into consideration.     

Looking at the unseen: combining animal bio‐logging and stable isotopes to reveal a shift in the ecological niche of a deep diving predator

Understanding how marine top predators exploit their environment is a central topic in marine ecology. Among all methodologies used to investigate this part of ecology, electronic devices are very useful to track animals' movements and foraging habitats, but they do not provide any dietary information. Stable isotopes provide information on trophic levels but remain imprecise to identify small spatial‐scale habitats. In this study, we combined the two approaches to obtain a synoptic view of the foraging behaviour variability of southern elephant seals Mirounga leonina. Our results suggested marked differences in distribution, diving behaviour, foraging habitats, trophic levels, and dietary habits of elephant seals according to their sex and age. Thus, we characterized main foraging habitats over the Kerguelen‐Heard Plateau and the Antarctic shelf for juvenile males, while females foraged mainly in oceanic waters of the Polar Frontal Zone and the Antarctic Zone. In addition, we highlighted the ontogeny of niche partitioning in this sexually dimorphic species. While females did not exhibit a major dietary shift in relation to their age and their breeding status, a different picture emerged for males. Young males had a trophic level identical to that of all females. However, at 3–4 yr of age, males showed a progressive increase in trophic level. The inter‐annual combination of bio‐logging and stable isotopes could provide a powerful tool to investigate possible shifts in ecological niche between years according to environmental changes.     

The Long and the Short of It: No Dietary Specialisation between Male and Female Western Sandpipers Despite Strong Bill Size Dimorphism

Many bird species show spatial or habitat segregation of the sexes during the non-breeding season. One potential ecological explanation is that differences in bill morphology favour foraging niche specialisation and segregation. Western sandpipers Calidris mauri have pronounced bill size dimorphism, with female bills averaging 15% longer than those of males. The sexes differ in foraging behaviour and exhibit partial latitudinal segregation during the non-breeding season, with males predominant in the north and females in the south. Niche specialisation at a local scale might account for this broad geographic pattern, and we investigated whether longer-billed females and shorter-billed males occupy different foraging niches at 16 sites across the non-breeding range. We used stable-nitrogen (δ¹⁵N) isotope analysis of whole blood to test for dietary specialisation according to bill length and sex. Stable-nitrogen isotope ratios increase with trophic level. We predicted that δ¹⁵N values would increase with bill length and would be higher for females, which use a greater proportion of foraging behaviour that targets higher-trophic level prey. We used stable-carbon (δ¹³C) isotope analysis to test for habitat segregation according to bill length and sex. Stable-carbon isotope ratios vary between marine- and freshwater-influenced habitats. We predicted that δ¹³C values would differ between males and females if the sexes segregate between habitat types. Using a model selection approach, we found little support for a relationship between δ¹⁵N and either bill length or sex. There was some indication, however, that more marine δ¹³C values occur with shorter bill lengths. Our findings provide little evidence that male and female western sandpipers exhibit dietary specialisation as a function of their bill size, but indicate that the sexes may segregate in different habitats according to bill length at some non-breeding sites. Potential ecological factors underlying habitat segregation between sexes include differences in preferred habitat type and predation risk.     

Habitat segregation in ungulates: are males forced into suboptimal foraging habitats through indirect competition by females?

Sex differences in habitat use (termed `habitat segregation') are widespread in sexually dimorphic ungulate species. They are a puzzling phenomenon, particularly when females use better foraging habitats than males. It has been suggested that males, owing to their larger body size and higher forage requirements, are inferior in indirect competition to females and are forced by female grazing pressure into marginal habitats (`indirect-competition hypothesis'). This hypothesis has been widely cited and has important theoretical and practical implications. However, evidence for it is inconclusive. The present paper presents the results of the first experimental test of the indirect-competition hypothesis. We manipulated female and male numbers of red deer (Cervus elaphus L.) on a large scale on the Isle of Rum, Scotland, and tested the influence of this manipulation on deer habitat use. We predicted that where female numbers were reduced, male use of preferred habitat should increase and sex differences in habitat use should decrease, while a reduction in male numbers should not have such effects. In contrast, we found that the manipulation of female and male numbers did not affect habitat use, and conclude that the indirect-competition hypothesis does not explain habitat segregation on Rum. Received: 19 August 1998 / Accepted: 13 February 1999     

Intersexual segregation in foraging microhabitat use by Magellanic Woodpeckers (Campephilus magellanicus): Seasonal and habitat effects at the world's southernmost forests

Animals facing seasonal food shortage and habitat degradation may adjust their foraging behaviour to reduce intraspecific competition. In the harsh environment of the world's southernmost forests in the Magellanic sub‐Antarctic ecoregion in Chile, we studied intersexual foraging differences in the largest South American woodpecker species, the Magellanic Woodpecker (Campephilus magellanicus). We assessed whether niche overlap between males and females decrease when food resources are less abundant or accessible, that is, during winter and in secondary forests, compared to summer and in old‐growth forests, respectively. We analysed 421 foraging microhabitat observations from six males and six females during 2011 and 2012. As predicted, the amount of niche overlap between males and females decreased during winter, when provisioning is more difficult. During winter, males and females (i) used trees with different diameter at breast height (DBH); (ii) fed in trunk sections with different diameters; and (iii) fed at different heights on tree trunks or branches. Vertical niche partitioning between sexes was found in both old‐growth and secondary forests. Such a niche partitioning during winter may be a seasonal strategy to avoid competition between sexes when prey resources are less abundant or accessible. Our results suggest that the conservation of this forest specialist, dimorphic and charismatic woodpecker species requires considering differences in habitat use between males and females.     

Seasonal home ranges and fidelity to breeding sites among ringed seals

Population structure and patterns of habitat use among ringed seals (Phoca hispida) are poorly known, in part because seasonal movements have not been adequately documented. We monitored the movements of 98 ringed seals in the Beaufort and Chukchi seas between 1990 and 2006 using three forms of telemetry. In the winter—spring period (when the seals were occupying shorefast ice), we used radio and ultra-sonic tags to track movements above and below the ice, respectively. We used satellite-linked transmitters in summer and fall (when the seals ranged away from their winter sites) to track at-sea movements. In the shorefast ice habitat, the home ranges of 27 adult males ranged from <1 to 13.9 km² (median = 0.628) while the home ranges of 28 adult females ranged from <1 to 27.9 km² (median = 0.652). The 3-dimensional volumes used by 9 seals tracked acoustically under the ice averaged 0.07 (SD = 0.04) km³ for subadults and adult males and 0.13 (SD = 0.04) km³ for adult females. Three of the radio-tracked seals and 9 tracked by satellite ranged up to 1,800 km from their winter/spring home ranges in summer but returned to the same small (1–2 km²) sites during the ice-bound months in the following year. The restricted movements of ringed seals during the ice-bound season—including the breeding season—limits their foraging activities for most of the year and may minimize gene flow within the species.     

Resource partitioning through oceanic segregation of foraging juvenile southern elephant seals (<Emphasis Type="Italic">Mirounga leonina</Emphasis>)

In highly dynamic and unpredictable environments such as the Southern Ocean, species that have evolved behaviors that reduce the effects of intra-specific competition may have a selective advantage. This is particularly true when juveniles face disadvantages when foraging due to morphological or physiological limitation, which is the case for many marine mammals. We tracked the at-sea movements of 48 juvenile southern elephant seals (Mirounga leonina) between the ages of 1 and 4 years from the population at Macquarie Island using locations derived from recorded light levels. There were significant differences in the total amount of the Southern Ocean covered by the different age-groups. The younger seals used a smaller area than the older seals. On average, the younger individuals also made more trips to sea than the older seals and did not travel as far on each trip. Females spent more time at sea than males and there were no significant differences between the total areas used by male and females. In summary, younger seals remained closer to the island at all times, and they spent more time in more northerly regions that older seals. These differences in behavior created temporal and spatial segregation between juveniles of different ages. Therefore, we suggest that these temporal and spatial separations help to avoid intra-specific competition for resources on land, space on beaches, and at-sea foraging areas. Such modifications of haul-out timing and behavior enable them to exploit a patchy and unpredictable environment.     

Sexual Niche Segregation and Gender-Specific Individual Specialisation in a Highly Dimorphic Marine Mammal

While sexual segregation is expected in highly dimorphic species, the local environment is a major factor driving the degree of resource partitioning within a population. Sexual and individual niche segregation was investigated in the Australian fur seal (Arctocephalus pusillus doriferus), which is a benthic foraging species restricted to the shallow continental shelf region of south-eastern Australia. Tracking data and the isotopic values of plasma, red blood cells and whiskers were combined to document spatial and dietary niche segregation throughout the year. Tracking data indicated that, in winter, males and females overlapped in their foraging habitat. All individuals stayed within central Bass Strait, relatively close (< 220 km) to the breeding colony. Accordingly, both genders exhibited similar plasma and red cell δ¹³C values. However, males exhibited greater δ¹³C intra-individual variation along the length of their whisker than females. This suggests that males exploited a greater diversity of foraging habitats throughout the year than their female counterparts, which are restricted in their foraging grounds by the need to regularly return to the breeding colony to suckle their pup. The degree of dietary sexual segregation was also surprisingly low, both sexes exhibiting a great overlap in their δ¹⁵N values. Yet, males displayed higher δ¹⁵N values than females, suggesting they fed upon a higher proportion of higher trophic level prey. Given that males and females exploit different resources (mainly foraging habitats), the degree of individual specialisation might differ between the sexes. Higher degrees of individual specialisation would be expected in males which exploit a greater range of resources. However, comparable levels of inter-individual variation in δ¹⁵N whisker values were found in the sampled males and females, and, surprisingly, all males exhibited similar seasonal and inter-annual variation in their δ¹³C whisker values, suggesting they all followed the same general dispersion pattern throughout the year.     

Sexual dimorphism and foraging niche partitioning in the Middle Spotted Woodpecker Dendrocopos medius

I investigated sexual differences in morphology and foraging behaviour in the nearly monomorphic Middle Spotted Woodpecker Dendrocopos medius. In the northeastern Swiss lowlands, radiotracked birds were observed from 1994 to 1996 between January and June. The sexes overlapped in all morphological characters, with males ( n = 25) being on average larger than females ( n = 17). Bill-length and length of the crimson-red crown patch were most useful to discriminate the sexes. Sex-specific differences in foraging behaviour were found both within and between the prebreeding and breeding periods concerning the use of tree species, foraging techniques, relative height zones and living and dead substrates within a tree. Overall, both sexes had broader niches in the prebreeding than in the breeding period and niche overlap was larger during the latter period. Hence, sex-specific differences were more pronounced in the prebreeding than in the breeding period, indicating different niche use of males and females in the two periods. These varying patterns of sex-specific differences imply that niche segregation results from behavioural plasticity, probably based on dominance relationships between the sexes, rather than from sexual dimorphism.     

The role of foraging behaviour in the sexual segregation of the African elephant

Elephants (Loxodonta africana) exhibit pronounced sexual dimorphism, and in this study we test the prediction that the differences in body size and sociality are significant enough to drive divergent foraging strategies and ultimately sexual segregation. Body size influences the foraging behaviour of herbivores through the differential scaling coefficients of metabolism and gut size, with larger bodied individuals being able to tolerate greater quantities of low-quality, fibrous vegetation, whilst having lower mass-specific energy requirements. We test two distinct theories: the scramble competition hypothesis (SCH) and the forage selection hypothesis (FSH). Comprehensive behavioural data were collected from the Pongola Game Reserve and the Phinda Private Game Reserve in South Africa over a 2.5-year period. The data were analysed using sex as the independent variable. Adult females targeted a wider range of species, adopted a more selective foraging approach and exhibited greater bite rates as predicted by the body size hypothesis and the increased demands of reproductive investment (lactation and pregnancy). Males had longer feeding bouts, displayed significantly more destructive behaviour (31% of observations, 11% for females) and ingested greater quantities of forage during each feeding bout. The independent ranging behaviour of adult males enables them to have longer foraging bouts as they experience fewer social constraints than females. The SCH was rejected as a cause of sexual segregation due to the relative abundance of low quality forage, and the fact that feeding heights were similar for both males and females. However, we conclude that the differences in the foraging strategies of the sexes are sufficient to cause spatial segregation as postulated by the FSH. Sexual dimorphism and the associated behavioural differences have important implications for the management and conservation of elephant and other dimorphic species, with the sexes effectively acting as distinct “ecological species”.     

Residency,Habitat Use and Sexual Segregation of White Sharks,Carcharodon carcharias in False Bay,South Africa

White sharks (Carcharodon carcharias) are threatened apex predators and identification of their critical habitats and how these are used are essential to ensuring improved local and ultimately global white shark protection. In this study we investigated habitat use by white sharks in False Bay, South Africa, using acoustic telemetry. 56 sharks (39 female, 17 male), ranging in size from 1.7–5 m TL, were tagged with acoustic transmitters and monitored on an array of 30 receivers for 975 days. To investigate the effects of season, sex and size on habitat use we used a generalized linear mixed effects model. Tagged sharks were detected in the Bay in all months and across all years, but their use of the Bay varied significantly with the season and the sex of the shark. In autumn and winter males and females aggregated around the Cape fur seal colony at Seal Island, where they fed predominantly on young of the year seals. In spring and summer there was marked sexual segregation, with females frequenting the Inshore areas and males seldom being detected. The shift from the Island in autumn and winter to the Inshore region in spring and summer by females mirrors the seasonal peak in abundance of juvenile seals and of migratory teleost and elasmobranch species respectively. This study provides the first evidence of sexual segregation at a fine spatial scale and demonstrates that sexual segregation in white sharks is not restricted to adults, but is apparent for juveniles and sub-adults too. Overall, the results confirm False Bay as a critical area for white shark conservation as both sexes, across a range of sizes, frequent the Bay on an annual basis. The finding that female sharks aggregate in the Inshore regions when recreational use peaks highlights the need for ongoing shark-human conflict mitigation strategies.     

Sex differences in weather sensitivity can cause habitat segregation: red deer as an example

Sex differences in habitat use (habitat segregation) are widespread in sexually dimorphic ungulates. A possible cause is that males are more sensitive to weather than females, leading to sex differences in sheltering behaviour (the 'weather sensitivity hypothesis'). However, this hypothesis has never been tested. We considered the allometric rates of net energy gain during times of cold weather and food shortage in a model. We argue that the higher absolute heat losses relative to intake rates of larger ungulates should indeed lead to higher weather sensitivity in males than in females. Furthermore, we tested the weather sensitivity hypothesis empirically in red deer, Cervus elaphus, on the Isle of Rum, U.K. We predicted that (1) use of relatively exposed, high-quality forage habitat should be negatively influenced by bad weather; and (2) this influence should be stronger in males. We found that bad weather (strong wind, low temperature, heavy rain) in winter and spring influenced use of high-quality forage habitat negatively in all deer; that adult males responded more strongly to low temperature and strong wind than did females; and that adult males foraged on windy days at better sheltered sites than did females. Thus, the weather sensitivity hypothesis is supported both theoretically and empirically. We suggest that the weather sensitivity hypothesis can potentially explain winter habitat segregation in a large number of ungulate species. Copyright 2000 The Association for the Study of Animal Behaviour.     

Daily selection of habitat in wild boar (Sus scrofa)

Spatial segregation of the sexes is observed in numerous species of ungulates, but are there other forms of segregation in species where spatial segregation is not present, such as in the wild boar? This study investigates the differences in habitat selection between two categories of individuals whose ecological requirements are supposedly divergent: subadult males, and females with dependent piglets. The proportion of the day spent in each category of habitat was used as a parameter representative of daily decisions. The overall results fit with the hypothesis that females with piglets avoid insecure habitats, and that both categories select habitats on the basis of food richness. In addition, it was found that the pattern of habitat selection depends on the total time spent moving slowly. A short duration of slow movement was observed only in males, but the time dedicated to efficient foraging (in rich habitats) was sufficient in all observations of all animals. Subadult males, therefore, do not differ from each other in foraging efficiency, but in the time spent on slow movement in secure habitats. Those spending more time active in secure habitats have the opportunity of performing more potentially beneficial activities such as social interactions and sexual contacts.     

Intersexual variation in the foraging ecology of sexually monochromatic Western Wood‐Pewees

ABSTRACT Investigators generally pool observations of males and females in studies of the foraging behavior of sexually monochromatic songbirds. However, such pooling can obscure possible intersexual differences. We compared the foraging behavior of male and female Western Wood‐Pewees (Contopus sordidulus), a sexually monochromatic species, in the Sierra Nevada Mountains of California during the breeding seasons of 2007 and 2008. We recorded 143 foraging observations (male N= 74, female N= 69). Overall, mean foraging rates of females (2.8 attacks/min) were higher (P < 0.001) than those of males (1.1 attacks/min). In addition, female foraging rates were significantly higher during incubation than during the nest building, nestling, and fledgling periods. When foraging, males perched higher above ground than females (means = 17.1 and 6.7 m, respectively). Differences between male and female Western Wood‐Pewees in foraging rates and perch heights suggest that males may spend more time on vigilance while females focus on foraging quickly during incubation and when feeding nestlings. Because metrics such as foraging attack rates are sometimes used as indicators of habitat quality and we found that rates can differ between the sexes and among nesting stages, investigators should consider the possibility of such differences when assessing habitat quality, especially for sexually monochromatic species of birds.