Phylogeography of shy and white-capped albatrosses inferred from mitochondrial DNA sequences: implications for population history and taxonomy

The evolutionary relationship between shy (Thalassarche cauta) and white-capped (T. steadi) albatrosses was examined using mitochondrial control region sequences. Results were interpreted in the context of a recent and contentious taxonomic revision that recommended full species status for shy and white-capped albatrosses. Low sequence divergence between shy and white-capped albatrosses (1.8%) and between their close relatives, Salvin's and Chatham albatrosses (2.9%), was observed. Much higher sequence divergence was found between the shy/white-capped pair and the Salvin's/Chatham pair (7.0%). Phylogenetic analyses confirmed the separation of the shy/white-capped pair from the Salvin's/Chatham pair but did not provide species-level resolution. Phylogeographic analyses, including a nested clade analysis, FST estimates and an analysis of molecular variance, indicated unambiguous genetic structuring between shy and white-capped albatrosses, thus confirming the demographic isolation of the species, but showed little to no structuring within each species. The geographical distribution of mtDNA haplotypes and other evidence suggest that shy albatrosses arose through range expansion by white-capped albatrosses.     

Genetic structure, conservation genetics and evidence of speciation by range expansion in shy and white-capped albatrosses

Six variable microsatellite loci were used to examine genetic structuring in the closely related shy albatross (Thalassarche cauta) and white-capped albatross (T. steadi). First, levels of genetic differentiation between the species, and among three populations within each species, were analysed using amova, FST and RST. We found high levels of genetic structuring and detected many unshared alleles between the species, which provide strong evidence against any contemporary gene flow between them. Within each species, shy albatross populations were found to be genetically distinct whereas white-capped albatross populations were undifferentiated, which implies that dispersal events are much rarer in the former than in the latter. These results formed the basis for the recommendation that the three white-capped albatross populations (as a whole) and each shy albatross population be treated as separate units for conservation. Second, levels of genetic diversity and allelic patterns in shy and white-capped albatrosses were assessed for whether they support earlier mtDNA results suggesting that shy albatrosses arose through range expansion of white-capped albatrosses. All measures indicated lower genetic diversity within shy albatrosses than within white-capped albatrosses and upheld the hypothesis that shy albatrosses were founded by white-capped albatrosses.     

Effects of Climate Change and Fisheries Bycatch on Shy Albatross (Thalassarche cauta) in Southern Australia

The impacts of climate change on marine species are often compounded by other stressors that make direct attribution and prediction difficult. Shy albatrosses (Thalassarche cauta) breeding on Albatross Island, Tasmania, show an unusually restricted foraging range, allowing easier discrimination between the influence of non-climate stressors (fisheries bycatch) and environmental variation. Local environmental conditions (rainfall, air temperature, and sea-surface height, an indicator of upwelling) during the vulnerable chick-rearing stage, have been correlated with breeding success of shy albatrosses. We use an age-, stage- and sex-structured population model to explore potential relationships between local environmental factors and albatross breeding success while accounting for fisheries bycatch by trawl and longline fisheries. The model uses time-series of observed breeding population counts, breeding success, adult and juvenile survival rates and a bycatch mortality observation for trawl fishing to estimate fisheries catchability, environmental influence, natural mortality rate, density dependence, and productivity. Observed at-sea distributions for adult and juvenile birds were coupled with reported fishing effort to estimate vulnerability to incidental bycatch. The inclusion of rainfall, temperature and sea-surface height as explanatory variables for annual chick mortality rate was statistically significant. Global climate models predict little change in future local average rainfall, however, increases are forecast in both temperatures and upwelling, which are predicted to have detrimental and beneficial effects, respectively, on breeding success. The model shows that mitigation of at least 50% of present bycatch is required to offset losses due to future temperature changes, even if upwelling increases substantially. Our results highlight the benefits of using an integrated modeling approach, which uses available demographic as well as environmental data within a single estimation framework, to provide future predictions. Such predictions inform the development of management options in the face of climate change.     

Environmental heterogeneity and the evolution of foraging behaviour in long ranging greater albatrosses

Habitat selection in heterogeneous environments is assumed to allow diversification. Wide‐ranging species like pelagic seabirds present a paradox, in that their diversity appears difficult to reconcile with a frequent lack of geographical isolation between populations. We studied the foraging strategies of three closely related species of greater albatrosses, wandering albatross, Diomedea exulans, Amsterdam albatrosses D. amsterdamensis and royal albatross, D. epomophora, in relation to environmental heterogeneity at coarse‐grained and fine‐grained scales. During the incubation period the three species foraged at long distances from their colonies. We observed significant differences between the species in the duration of foraging trips and the distance travelled per day. There were significant differences in preference for habitat types in relation to bathymetric features, and in chlorophyll a concentrations in the waters traversed. Royal albatross preferred shallower waters (<1500 m depth), which were rich in chlorophyll (>0.5 mg/m³), while the other species spent on average 80% of their time in waters deeper than this, where chlorophyll levels were lower. Wandering albatrosses foraged in colder waters than Amsterdam albatrosses. Patterns of activity divided the species into two groups: those exploiting oceanic habitats (wandering and Amsterdam albatrosses) spent high proportions of time on the water (49%), and had on average 1.35 takeoffs and landings per hour, while royal albatross, which foraged mainly over neritic waters spent only 35% of their time sitting on the water, and made on average 2.6 takeoff per hour. Further, royal albatross showed a similar pattern of activity during all periods of the day, while wandering and Amsterdam albatrosses were mostly inactive during the night. We link these differences in activity to prey patch availability in two contrasting habitats – continental shelf areas compared to open ocean habitats. The divergent styles of foraging observed in this study suggest that these closely‐related and wide‐ranging species could effectively co‐exist by dividing the resources available to them by different modes of exploitation.     

Predator–prey interactions: why do larger albatrosses eat bigger squid?

The relationship between predator sizes and prey sizes is well documented for terrestrial but rarely for marine ecosystems. We show that wandering albatrosses, the biggest albatross species, feed on larger cephalopod prey than those consumed by smaller albatrosses (grey-headed and black-browed albatrosses). This reflects differences in timing of breeding, foraging ecology and their feeding methods. Wandering albatrosses breed later in the year, during the austral winter, than smaller albatrosses (therefore catching older squid) and forage most of the year in Antarctic open waters, sub-Antarctic, subtropical and tropical waters, overlapping minimally with the smaller albatrosses' foraging range while breeding. Also, wandering albatrosses mostly scavenge whereas smaller albatrosses feed more on live prey. Prey ecology may also play a key role because many squid species might experience post-spawning mortality during the austral winter, becoming easily available to wandering albatrosses. Spawning in winter can be linked to predator avoidance (i.e. reduction in mortality in winter by avoiding pelagic predators) and would allow squid larvae to develop and take advantage of the high productivity (i.e. Antarctic phytoplankton bloom) in spring and at the beginning of summer. Thus, aspects of prey and predator ecology may combine to generate observed differences in prey size.     

Use and exploitation of channel waters by the black-browed albatross

Black-browed albatrosses are the most abundant albatross species of the southern hemisphere, breeding on sub-Antarctic and Antarctic oceanic islands around the globe. Their foraging habitat during the breeding season is reasonably well known along its distributional range, indicating a preferred use of waters <500 m deep. The discovery of a colony inserted within the Admiralty Sound, Tierra del Fuego, poses an interesting challenge to the known precepts on foraging behavior for the species. In this study, we present the first record on the foraging distribution of the only known inner-channel colony of albatrosses in the world, using high-resolution GPS loggers. Black-browed albatrosses breeding at the Albatross Islet used exclusively inner-channel waters, at least during the chick-guard stage. Our results indicate a significant smaller foraging range during chick-guard compared with conspecifics from Diego Ramirez and Falklands/Malvinas Islands. Implications for the conservation of this colony are discussed.     

Satellite tracking of light-mantled sooty albatrosses

Five light-mantled sooty albatrosses (Phoebetria palpebrata) breeding at Macquarie Island were tracked with miniaturised satellite transmitters during foraging trips of the incubation period. Birds moved rapidly to specific sectors of the Southern Ocean, where they spent several days foraging before returning to their nests. These specific sectors were at an average distance of 1516 km from Macquarie Island and located in pelagic Antarctic waters, mostly along the Antarctic continent. The maximum foraging range was in average 1721 km and the total distance covered by two birds for which there were complete tracks was 6463 and 6975 km. This study confirms previous suggestions that light-mantled sooty albatrosses are able to forage in the waters of the high Antarctic while breeding in the sub-Antarctic. The implications of the extreme separation of feeding zones from nesting grounds, in terms of conservation and life-history strategies, are discussed.     

Copulation behaviour and paternity in shy albatrosses (Thalassarche cauta)

We investigated the mating system of shy albatrosses Thalassarche cauta by combining behavioural observations during the pre-laying period with genetic paternity analysis. Genetic data on the mating systems of several procellariiform seabirds have recently become available, but data on the reproductive behaviour of these species are rarely obtained. Our main aims were to describe the copulatory behaviour of this species and identify how males achieve within-pair and extra-pair paternity (EPP). Most copulations occurred on the nest, were unforced and were within-pair. Females controlled the success of copulations and were observed soliciting extra-pair matings. Within-pair and extra-pair copulations were behaviourally similar. A low frequency (7–10%, n =29 chicks) of EPP was detected despite male use of frequent copulation as a paternity guard. The pre-laying foraging exodus of female shy albatrosses differed from that in other albatrosses: it was relatively short in length, lasting c . 2 days, and within-pair copulations occurred after the female's return 2 days before laying. This may reflect the close proximity of feeding grounds to the breeding colony.     

Dynamic habitat models: using telemetry data to project fisheries bycatch

Fisheries bycatch is a recognized threat to marine megafauna. Addressing bycatch of pelagic species however is challenging owing to the dynamic nature of marine environments and vagility of these organisms. In order to assess the potential for species to overlap with fisheries, we propose applying dynamic habitat models to determine relative probabilities of species occurrence for specific oceanographic conditions. We demonstrate this approach by modelling habitats for Laysan (Phoebastria immutabilis) and black-footed albatrosses (Phoebastria nigripes) using telemetry data and relating their occurrence probabilities to observations of Hawaii-based longline fisheries in 1997-2000. We found that modelled habitat preference probabilities of black-footed albatrosses were high within some areas of the fishing range of the Hawaiian fleet and such preferences were important in explaining bycatch occurrence. Conversely, modelled habitats of Laysan albatrosses overlapped little with Hawaii-based longline fisheries and did little to explain the bycatch of this species. Estimated patterns of albatross habitat overlap with the Hawaiian fleet corresponded to bycatch observations: black-footed albatrosses were more frequently caught in this fishery despite being 10 times less abundant than Laysan albatrosses. This case study demonstrates that dynamic habitat models based on telemetry data may help to project interactions with pelagic animals relative to environmental features and that such an approach can serve as a tool to guide conservation and management decisions.     

Mercury and cadmium concentrations in the tissues of three species of southern albatrosses

Cadmium and mercury concentrations were measured in the tissues of 64 individual albatrosses [23 wandering albatrosses (Diomedea exulans), 9 royal albatrosses (Diomedea epomophora) and 32 shy albatrosses (Thalassarche cauta)] which were killed as by-catch in longline fishing activities between 1991 and 1994. Mercury concentrations were also determined for 33 shy albatross eggs (excluding shells). The birds were all sexed and assigned to one of two age classes (immature and adult). The three species exhibited differences both in overall concentrations of cadmium and mercury, and also in the pattern of accumulation of metals with age and sex. Wandering albatrosses exhibited the highest mercury concentrations with a mean concentration in adult liver samples of 920.0 ± 794.1 μg g⁻¹ dry weight. Shy albatrosses had the lowest mercury concentrations with mean concentrations in adult livers of 36.3 ± 21.4 mg g⁻¹ dry weight. The highest mercury concentration was 1800 μg g⁻¹ for an adult female wandering albatross. Cadmium concentrations were less variable, with adult royal albatrosses having the highest average concentrations (180.0 ± 165.0 in adult kidneys) and adult shy albatrosses the lowest (40.1 ± 20.0 in adult kidney). The highest individual cadmium concentration was 287 μg g⁻¹ for a juvenile wandering albatross. There was no evidence of increased accumulation of cadmium with age in any of the species, but wandering albatrosses showed higher mercury concentrations in adults than juveniles. Female wandering albatrosses also had significantly higher mercury concentrations than males. The mercury contents of the shy albatross eggs were very low, with a maximum concentration of 5.4 μg g⁻¹. The results of this study are consistent with the findings of previous work on albatrosses and support the notion that the life-history strategy of these species (i.e. long-lived with low reproductive output) may be an important determinant in the concentrations of some metals found in their tissues. Accepted: 15 February 1999     

Unexpectedly high genetic diversity of mtDNA control region through severe bottleneck in vulnerable albatross <Emphasis Type="Italic">Phoebastria albatrus</Emphasis>

In the late part of the nineteenth century and the early part of the last century, the short-tailed albatross Phoebastria albatrus was in danger of extinction owing to feather hunting. In the middle of the last century, the total number of this species was inferred to be approximately 50–60 with breeding occurring only on Torishima Island of the Izu Islands. Recently, the number of individuals has increased to more than 2,000 and that of their breeding islands to three, namely, Torishima Island, and Minami- and Kita-kojima Islands of the Senkaku Islands. Here, we show that the 44 short-tailed albatrosses we examined represent 29 haplotypes in the control region of mitochondrial DNA, and have a considerably higher genetic diversity than most avian species, but not very high in albatross species; the h and π were 0.96 and 0.013, respectively. However, the parsimony network clearly showed that many intermediate haplotypes were lost. It was concluded that the majority of the haplotypes in the founder population have been maintained. Judging from these findings and the exponential increase in the number of individuals, the present population of the short-tailed albatross seems not to be affected by inbreeding depression through a severe bottleneck. The conservation and expansion of their breeding grounds, and effective protection from bycatch mortality in foraging areas are important for the future survival of this species.     

Differential adult survival at close seabird colonies: The importance of spatial foraging segregation and bycatch risk during the breeding season

Marine megafauna, including seabirds, are critically affected by fisheries bycatch. However, bycatch risk may differ on temporal and spatial scales due to the uneven distribution and effort of fleets operating different fishing gear, and to focal species distribution and foraging behavior. Scopoli's shearwater Calonectris diomedea is a long‐lived seabird that experiences high bycatch rates in longline fisheries and strong population‐level impacts due to this type of anthropogenic mortality. Analyzing a long‐term dataset on individual monitoring, we compared adult survival (by means of multi‐event capture–recapture models) among three close predator‐free Mediterranean colonies of the species. Unexpectedly for a long‐lived organism, adult survival varied among colonies. We explored potential causes of this differential survival by (1) measuring egg volume as a proxy of food availability and parental condition; (2) building a specific longline bycatch risk map for the species; and (3) assessing the distribution patterns of breeding birds from the three study colonies via GPS tracking. Egg volume was very similar between colonies over time, suggesting that environmental variability related to habitat foraging suitability was not the main cause of differential survival. On the other hand, differences in foraging movements among individuals from the three colonies expose them to differential mortality risk, which likely influenced the observed differences in adult survival. The overlap of information obtained by the generation of specific bycatch risk maps, the quantification of population demographic parameters, and the foraging spatial analysis should inform managers about differential sensitivity to the anthropogenic impact at mesoscale level and guide decisions depending on the spatial configuration of local populations. The approach would apply and should be considered in any species where foraging distribution is colony‐specific and mortality risk varies spatially.     

Cryptic speciation and population differentiation in the yellow-nosed albatross species complex

The two species of yellow-nosed albatross, Atlantic (Thalassarche chlororhynchos) and Indian (T. carteri), are morphologically similar, but they differ in breeding behaviour and distribution. Both species are listed as endangered by the IUCN due to the limited number of breeding sites, threats from introduced predators and diseases, and impact of commercial fishing. We quantified genetic variation between and within the two species. Using nuclear (microsatellites and two nuclear sequences) and mitochondrial (control region) markers, we analysed 354 samples from four breeding islands (Atlantic: Nightingale, Inaccessible, and Gough; Indian: Amsterdam) and bycatch samples from South Africa and New Zealand. In addition to all markers separating the two species, nuclear markers showed Atlantic yellow-nosed albatrosses from Gough Island are genetically distinct from those breeding at Nightingale and Inaccessible Islands in the Tristan da Cunha archipelago. Nuclear markers confirmed that all bycatch samples were Indian yellow-nosed albatrosses, however, the bycatch birds from South Africa and New Zealand were distinct from each other and from birds breeding on Amsterdam Island, suggesting colony specific dispersal at sea. Our study supports the current recognition of two yellow-nosed albatross species and recognises genetically distinct groups of both Atlantic and Indian yellow-nosed albatross breeding on different islands, which is important for their conservation and management.

     

Small-scale fisheries bycatch jeopardizes endangered Pacific loggerhead turtles

BACKGROUND: Although bycatch of industrial-scale fisheries can cause declines in migratory megafauna including seabirds, marine mammals, and sea turtles, the impacts of small-scale fisheries have been largely overlooked. Small-scale fisheries occur in coastal waters worldwide, employing over 99% of the world's 51 million fishers. New telemetry data reveal that migratory megafauna frequent coastal habitats well within the range of small-scale fisheries, potentially producing high bycatch. These fisheries occur primarily in developing nations, and their documentation and management are limited or non-existent, precluding evaluation of their impacts on non-target megafauna. PRINCIPAL FINDINGS/METHODOLOGY: 30 North Pacific loggerhead turtles that we satellite-tracked from 1996-2005 ranged oceanwide, but juveniles spent 70% of their time at a high use area coincident with small-scale fisheries in Baja California Sur, Mexico (BCS). We assessed loggerhead bycatch mortality in this area by partnering with local fishers to 1) observe two small-scale fleets that operated closest to the high use area and 2) through shoreline surveys for discarded carcasses. Minimum annual bycatch mortality in just these two fleets at the high use area exceeded 1000 loggerheads year(-1), rivaling that of oceanwide industrial-scale fisheries, and threatening the persistence of this critically endangered population. As a result of fisher participation in this study and a bycatch awareness campaign, a consortium of local fishers and other citizens are working to eliminate their bycatch and to establish a national loggerhead refuge. CONCLUSIONS/SIGNIFICANCE: Because of the overlap of ubiquitous small-scale fisheries with newly documented high-use areas in coastal waters worldwide, our case study suggests that small-scale fisheries may be among the greatest current threats to non-target megafauna. Future research is urgently needed to quantify small-scale fisheries bycatch worldwide. Localizing coastal high use areas and mitigating bycatch in partnership with small-scale fishers may provide a crucial solution toward ensuring the persistence of vulnerable megafauna.     

Global relationships amongst black-browed and grey-headed albatrosses: analysis of population structure using mitochondrial DNA and microsatellites

The population structure of black-browed (Thalassarche melanophris and T. impavida) and grey-headed (T. chrysostoma) albatrosses was examined using both mitochondrial DNA (mtDNA) and microsatellite analyses. mtDNA sequences from 73 black-browed and 50 grey-headed albatrosses were obtained from five island groups in the Southern Ocean. High levels of sequence divergence were found in both taxa (0.55-7.20% in black-browed albatrosses and 2.10-3.90% in grey-headed albatrosses). Black-browed albatrosses form three distinct groups: Falklands, Diego Ramirez/South Georgia/Kerguelen, and Campbell Island (T. impavida). T. melanophris from Campbell Island contain birds from each of the three groups, indicating high levels of mixture and hybridization. In contrast, grey-headed albatrosses form one globally panmictic population. Microsatellite analyses on a larger number of samples using seven highly variable markers found similar population structure to the mtDNA analyses in both black-browed and grey-headed albatrosses. Differences in population structure between these two very similar and closely related species could be the result of differences in foraging and dispersal patterns. Breeding black-browed albatrosses forage mainly over continental shelves and migrate to similar areas when not breeding. Grey-headed albatrosses forage mainly at frontal systems, travelling widely across oceanic habitats outside the breeding season. Genetic analyses support the current classification of T. impavida as being distinct from T. melanophris, but would also suggest splitting T. melanophris into two groups: Falkland Islands, and Diego Ramirez/South Georgia/Kerguelen.     

Capture and blood sampling do not affect foraging behaviour, breeding success and return rate of a large seabird: the black-browed albatross

During the last decades, eco-physiological studies have usually relied on the collection of blood from wild organisms in order to obtain relevant physiological measures. However, accurate estimates of the impact of capture and blood collection on performances of Polar seabird species have rarely been conducted. We investigated for the first time the effects of a blood sampling process on subsequent foraging behaviour, reproductive performance and return rate of black-browed albatrosses (Thalassarche melanophris) at Kerguelen Islands. We did not find any evidence that the blood sampling process as conducted in our study had detrimental effects on the breeding or foraging strategies or performance of black-browed albatrosses. Because blood collection can be performed in several different ways, we recommend that eco-physiologists conduct pilot studies to test whether their blood sampling process affects the performances of their study species.     

A comprehensive isotopic investigation of habitat preferences in nonbreeding albatrosses from the Southern Ocean

Albatrosses are among the world's most endangered seabirds. Threats during the nonbreeding period have major impacts on their population dynamics, but for most species, detailed information on distribution and ecology remains essentially unknown. We used stable isotope values (δ¹³C and δ¹⁵N) in feathers to infer and compare the moulting (nonbreeding) habitats of 35 populations that include all the 20 species and subspecies (444 individuals) of albatrosses breeding within the Southern Ocean and in fringing subtropical waters. Isotopic values together with a review of available information show that the 20 albatrosses can be categorized into three groups depending on their favoured moulting grounds: 12 (60%) taxa forage primarily in warm neritic waters, six (30%) in northern oceanic waters and two (10%) in oceanic waters of the Southern Ocean. Stable isotopes indicate that habitat preferences during the nonbreeding period vary much less among different breeding populations in some species (wandering, Salvin's, grey‐headed and light‐mantled sooty albatrosses), than others (black‐browed, Indian yellow‐nosed and sooty albatrosses). The major finding of our isotopic investigation is that the great majority of albatrosses spend the nonbreeding period outside the Southern Ocean, with only three species (and in the sooty albatross, just one of the breeding populations) favouring oceanic subantarctic waters at that time. Hence, the study highlights the overwhelming importance of subtropical waters for albatrosses, where the birds are known to interact with human activities and are more likely to be negatively affected by the diverse range of fisheries operating in both neritic and oceanic waters.     

Orientation in the wandering albatross: interfering with magnetic perception does not affect orientation performance

After making foraging flights of several thousands of kilometers, wandering albatrosses (Diomedea exulans) are able to pinpoint a specific remote island where their nests are located. This impressive navigation ability is highly precise but its nature is mysterious. Here we examined whether albatrosses rely on the perception of the Earth's magnetic field to accomplish this task. We disturbed the perception of the magnetic field using mobile magnets glued to the head of nine albatrosses and compared their performances with those of 11 control birds. We then used satellite telemetry to monitor their behavior. We found that the ability of birds to home specific nest sites was unimpaired by this manipulation. In particular, experimental and control birds did not show significant differences with respect to either foraging trip duration, or length, or with respect to homing straightness index. Our data suggest that wandering albatrosses do not require magnetic cues to navigate back to their nesting birds.     

Projected distributions of Southern Ocean albatrosses,petrels and fisheries as a consequence of climatic change

Given the major ongoing influence of environmental change on the oceans, there is a need to understand and predict the future distributions of marine species in order to plan appropriate mitigation to conserve vulnerable species and ecosystems. In this study we use tracking data from seven large seabird species of the Southern Ocean (black‐browed albatross Thalassarche melanophris, grey‐headed albatross T. chrysostoma, northern giant petrel Macronectes halli, southern giant petrel M. giganteus, Tristan albatross Diomedea dabbenena, wandering albatross D. exulans and white‐chinned petrel Procellaria aequinoctialis, and on fishing effort in two types of fisheries (characterised by low or high‐bycatch rates), to model the associations with environmental variables (bathymetry, chlorophyll‐a concentration, sea surface temperature and wind speed) through ensemble species distribution models. We then projected these distributions according to four climate change scenarios built by the Intergovernmental Panel for Climate Change for 2050 and 2100. The resulting projections were consistent across scenarios, indicating that there is a strong likelihood of poleward shifts in distribution of seabirds, and several range contractions (resulting from a shift in the northern, but no change in the southern limit of the range in four species). Current trends for southerly shifts in fisheries distributions are also set to continue under these climate change scenarios at least until 2100; some of these may reflect habitat loss for target species that are already over‐fished. It is of particular concern that a shift in the distribution of several highly threatened seabird species would increase their overlap with fisheries where there is a high‐bycatch risk. Under such scenarios, the associated shifts in distribution of seabirds and increases in bycatch risk will require much‐improved fisheries management in these sensitive areas to minimise impacts on populations in decline.     

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.