Harem choice and breeding experience of female southern elephant seals influence offspring survival

Female mammals can increase their lifetime fitness through modification of investment potential and by providing better rearing environments with improved breeding experience. We examined the relationships between reproductive fitness and the behavioural decisions that female southern elephant seals (Mirounga leonina) made during the breeding season. We examined whether mother age and breeding experience influenced reproductive success (measured as 1st-year survival probability), and whether there was a change in the choice of harem size with increasing age. Pups produced by young mothers had lower 1st-year survival probability than pups produced by older mothers. A significant increase in mean female mass with age required an analysis of both these effects on offspring survival. There was a significant positive effect of both female age and mass, and the interaction between the two, on 1st-year pup survival. The proportion of young mothers (<5 years old) decreased and the proportion of older mothers (>6 years old) increased with increasing harem size (harems surveyed from 1997 to 2001). Females chose larger harems in which to breed as they aged. Females demonstrated fidelity to breeding areas among successive breeding seasons, with older females displaying greater breeding-site fidelity than younger females. The mean number of previous breeding attempts per female within a harem (breeding experience) increased significantly with increasing harem size. Breeding females returned to breed later in the breeding season as they aged—we hypothesize that young, subordinate females gain a priority advantage by returning earlier. These results lend support to the hypothesis that there are fitness advantages, in terms of offspring survival, that are conferred to females that breed in successively larger harems with age. Potential mechanisms that select for females to improve their breeding conditions include improved mate selection and the avoidance of conspecific harassment in harems.Communicated by F. Trillmich     

Offspring sex ratio in relation to female size in southern elephant seals,Mirounga leonina

Southern elephant seals Mirounga leonina display extreme sexual dimorphism. In addition females show great variation in size and stored resources at parturition. Therefore they present an excellent opportunity for examination of responses of sex ratio to resource availability. We studied the relationships between the size of southern elephant seal females at parturition and the size and sex of their pups at South Georgia over four breeding seasons. We found a large individual variation in maternal post-partum mass (range 296–977 kg, n=151). Larger mothers gave birth to larger pups, irrespective of the sex of their pup. Male pups were on average 14% larger than females at birth and consequently more costly to bring to parturition. Our results suggest that female southern elephant seals must weigh more than 300 kg if they are to breed at all, and more than 380 kg if they are to give birth to a male pup. Above this threshold the proportion of males among offspring rapidly increases with maternal mass, and stabilizes at a level not significantly different from parity. These results show that smaller females of southern elephant seals vary offspring sex ratio in a way that is consistent with theories on adaptive offspring sex ratio. A smaller mother with a male foetus may benefit from terminating her pregnancy and allocating the resources she saves to her own growth. She could then give birth to and raise a larger pup in the subsequent season.     

Fish prey of fur seals Arctocephalus spp. at subantarctic Marion Island

The analysis of scats collected between 1989 and 1995 from the two fur seal species resident on subantarctic Marion Island, Arctocephalus gazella and A. tropicalis, showed that they fed predominantly on fish of the family Myctophidae (lanternfishes). Scat composition (prey species, abundance) was very similar for the two species. The seven species of myctophids that formed numerically 90 and 86% of the scat composition for A. gazella and A. tropicalis, respectively, all showed seasonal fluctuations in their contribution to seal diets. Electrona carlsbergi, E. subaspera, Metelectrona ventralis and Gymnoscopelus fraseri increased in winter in both species of fur seals, whereas Gymnoscopelus piabilis, Protomyctophum choriodon and P. tenisoni showed the opposite trend. Seal diets overlapped substantially with those of the king penguins (Aptenodytes patagonicus) resident on Marion Island, but no evidence for competitive exclusion could be found between these two major warmblooded consumers of marine resources at the Prince Edward Islands. Received: 16 July 1997 / Accepted: 2 March 1998     

Shift in foraging grounds and diet broadening during ontogeny in southern elephant seals from Kerguelen Islands

Highly dimorphic species like southern elephant seals (Mirounga leonina, SES hereafter) frequently exhibit resource partitioning according to sex and/or age classes. We measured carbon and nitrogen stable isotopes (δ¹³C and δ¹⁵N) of 404 blood samples (136 males and 268 females from Kerguelen Islands, 49°21′S, 70°18′E) from 2004 to 2011. Assuming that the distribution of carbon isotopes (δ¹³C value) reflects the two main foraging grounds (Polar Frontal and Antarctic Zones), we quantified the proportion of SES foraging within each zone in relation with size, a proxy for their age. We found a clear shift from Polar Frontal to Antarctic waters as male SES aged, but no relation as far as females is concerned. We also observed a widening range of nitrogen isotopic (δ¹⁵N) values, suggesting that both males and females expanded their diet spectra with age. Whereas males increased their trophic level, females remained constant on average, with some adult females feeding both at lower and at higher trophic levels than juveniles.     

Age-related shifts in the diet composition of southern elephant seals expand overall foraging niche

Southern elephant seals are important apex predators in a highly variable and unpredictable marine environment. In the presence of resource limitation, foraging behaviours evolve to reduce intra-specific competition increasing a species’ overall probability of successful foraging. We examined the diet of 141 (aged 1–3 years) juvenile southern elephant seals to test the hypotheses that differences between ages, sexes and seasons in diet structure occur. We described prey species composition for common squid and fish species and the mean size of cephalopod prey items for these age groups. Three cephalopod species dominated the stomach samples, Alluroteuthis antarcticus, Histioteuthis eltaninae and Slosarczykovia circumantarcticus. We found age-related differences in both species composition and size of larger prey species that probably relate to ontogenetic changes in diving ability and haul-out behaviour and prey availability. These changes in foraging behaviour and diet are hypothesised to reduce intra-specific food competition concomitant with the increase in foraging niche of growing juveniles.     

Winter distribution and haul-out behaviour of female Antarctic fur seals from South Georgia

Telemetry-based techniques have revealed the foraging patterns of many land breeding marine predators, especially during the summer breeding season. However, during the winter, when freed from the constraints of provisioning their young, such animals are more difficult to track. Using geolocation (Global Location Sensing, GLS) loggers and satellite tags (Platform Terminal Transmitters, PTTs) we successfully tracked 16 female Antarctic fur seals from South Georgia during the austral winter. The majority of females concentrated their winter foraging in the waters around the breeding beaches (90% of locations were within 510 km). However, as the winter progressed, two of the seals spent a number of months to the south, in and around the seasonal ice edge, and five seals migrated north and northwest from South Georgia. Four of these seals clearly crossed the Polar Front and two reached the Patagonian Shelf, apparently exploiting the continental shelf edge and the Subantarctic Front. Activity (saltwater immersion) data suggested that seals spent the majority of the winter months at sea but there were rare occasions when seals hauled out, either on land or on ice floes. We obtained data from two individuals that enabled us to compare the performance of PTT and GLS devices. For these seals the mean distance between GLS and PTT locations was 122 and 132 km. Although the recovery rates were low in this study, given improvements in attachment techniques, we have demonstrated that these micro-geolocation loggers provide an ideal tool with which to study the long-term dispersal of diving marine predators at larger scales. This is the first study to show that female fur seals from South Georgia remain at sea for almost the entirety of the non-breeding winter period. Using land-based observations it has been assumed that the fur seal population at South Georgia has little temporal overlap with the krill fishery that operates mostly during the winter months in this region. We have shown that a large proportion of the female fur seals that breed on South Georgia potentially remain in the vicinity of the island and are thus in direct competition with the region’s fisheries activities.     

Trophic ecology of Grey-headed albatrosses from Marion Island, Southern Ocean: insights from stomach contents and diet tracers

During chick-rearing, albatrosses can alternate between long foraging trips that provide the main source of food for the adults and short foraging trips that they use to feed their young. This flexibility in foraging behaviour can lead to differences in diet composition between adults and chicks and implies that they may be vulnerable in different ways to food shortages. The trophic ecology of the Grey-headed albatross Thalassarche chrysostoma was investigated at the sub-Antarctic Prince Edward Islands during the chick-rearing period in April 2006 using a combination of approaches. Diets of adults and chicks were assessed using stable isotope ratios and fatty acid (FA) profiles of blood and/or stomach oils, in addition to stomach contents analysis. Fish from the family Macrouridae and cephalopods (particularly the onychoteuthid Kondakovia longimana) were the primary prey, whereas crustaceans (krill Euphausia superba) represented a smaller proportion of the stomach contents. Stomach oil FA profiles contained more monounsaturated FA than the profiles of plasma, which were richer in saturated FA and arachidonic acid (20:4n-6). There was also a distinct separation of adults from chicks, with higher levels of monounsaturates in chick plasma, and higher saturated FA levels (particularly 16:0) in the adult plasma. Stable carbon isotope ratios of whole blood were similar in adults and chicks, whereas stable nitrogen isotope ratios showed significant enrichment by >1‰ in chicks. The combined FA, stable isotopes and stomach contents analyses suggest clear differences in diet quality between adults and chicks, with chicks feeding at a higher trophic position through feeding more on highly nutritious fish and adults keeping much of the less nutritious zooplankton for themselves.     

Feeding and predation impact of two chaetognath species, Eukrohnia hamata and Sagitta gazellae, in the vicinity of Marion Island (southern ocean)

The predation impact of the two chaetognaths Eukrohnia hamata and Sagitta gazellae on mesozooplankton standing stock were investigated in three depth layers during two 24 h stations occupied in the vicinity of Marion Island in late austral summer (April/May) 1986. The zooplankton community at both stations was dominated by small copepods (Oithona spp., Microcalanus spp.), which accounted for >95% of total zooplankton abundance. Chaetognaths comprised <2% of total zooplankton abundance. E. hamata constituted >95% of the total chaetognath stock. The general trend in both species was decreasing abundance with increasing depth, which appeared to be correlated to the distribution of copepods (r ² = 0.45; P <0.05). Gut-content analysis showed that copepods (mainly Oithona spp., Calanus spp. and Rhincalanus gigas) and ostracods were the main prey of both species, accounting for 87 and 61% of the total number of prey in E. hamata and S.␣gazellae stomachs, respectively. In the guts of S.␣gazellae, pteropods (Limacina spp.) and chaetognaths were also well represented. The mean number of prey items (NPC) for E. hamata ranged from 0.02 to 0.06 prey individual⁻¹ which corresponds to an individual feeding rate (Fr) of between 0.05 and 0.12 prey d⁻¹. For S.␣gazellae, the NPC values were higher, varying between 0.04␣and 0.20 prey individual⁻¹, or between 0.15 and 0.76 prey d⁻¹. The daily predation impact of the two chaetognaths was estimated at between 0.3 and 1.2% of the copepod standing stock or between 7 and 16% of the daily copepod production. Predation by S. gazellae on chaetognaths accounted for up to 1.6% of the chaetognath standing stock per day. Received: 26 November 1996 / Accepted: 31 October 1997     

Stable isotopes document winter trophic ecology and maternal investment of adult female southern elephant seals (<Emphasis Type="Italic">Mirounga leonina</Emphasis>) breeding at the Kerguelen Islands

Individual specialisation is widespread and can affect a population’s ecological and evolutionary dynamics. Whether intra-specific niche differences can influence reproductive investment was examined in a marine mammal, the southern elephant seal (Mirounga leonina), whose females were known to forage in two different areas during the austral winter. The study was conducted at Kerguelen Islands (49°21′S, 70°18′E), southern Indian Ocean, in late winter–early spring 2006. Pups were used as proxies of their mothers’ biology and combined information on their weaning mass (a proxy of females’ foraging success and short-term fitness) together with their blood δ¹³C value (a proxy of female foraging zone). First, the use of isotopic signature of pups was validated to study the female foraging ecology during their pre-breeding trip by demonstrating that δ¹³C and δ¹⁵N values of pups and their mothers were positively and linearly correlated. Then, blood samples were taken from a large number of newly-weaned pups, which were also weighed, to provide information at the population level. Estimated δ¹³C values of female seals encompassed a large range of values (from −23.7 to −19.1‰) with an unimodal frequency distribution, suggesting no contrasted foraging areas within the population. No significant relationship was found between pup weaning mass and their carbon signature, indicating no link between female foraging areas and maternal foraging success and investment. Finally, blood δ¹³C and δ¹⁵N values gave new insights into southern elephant seal ecology, suggesting that females mainly foraged north of the Polar Front where they preyed upon myctophid fish in late winter.     

The cost of success: reproductive effort in male southern elephant seals (<Emphasis Type="Italic">Mirounga leonina</Emphasis>)

Reproductive effort is a key parameter of life history because it measures the resources allocated to reproduction at the expense of growth and maintenance. Male reproductive effort always had a minor role with respect to female effort both in the development of theories and in field research. Elephant seals are an ideal subject for reproductive effort studies because they fast during the breeding season, splitting the phase of energy acquisition from the phase of energy use for breeding. In this paper, we present results on male reproductive effort (weight loss estimated by photogrammetry) in southern elephant seals (Mirounga leonina), the most dimorphic and polygynous of all mammal species. We show that total reproductive effort increases with age, with no sign of late decrease or senescence. Male reproductive effort in this species depends mostly on behavioral factors, i.e., the success in competition with other males, and the intensity of interaction with females. A large effort results in large gains in both mating success and fertilizations. The large reproductive success that a few males are able to achieve come at a big cost in terms of energy expenditure, but this cost does not seem to affect the likelihood of survival to the following breeding season.     

Size and seasonal influences on the foraging range of female grey seals in the northeast Atlantic

Evidence of segregation in foraging habitat has been demonstrated in some top marine predators, including cetaceans, pinnipeds and seabirds. However, most data are not adequate to assess differences relating to body size or seasonal influences. This has implications for quantitative modelling of population-level predator–prey interactions and ecosystem structure. We examined potential influence of body size and ‘fatness’ on the foraging trip characteristics of a top marine predator, the Atlantic grey seal (Halichoerus grypus), in southwest Ireland within the framework of optimal foraging theory to examine how female grey seals foraging behaviour varied with size, and across the period between moult and breeding. Larger seals undertook trips of greater duration and travelled further from haul-out sites than smaller seals. However, body fat was negatively associated with trip duration and extent. Seals spent more time at sea during the summer, but trips were shorter in extent, suggesting more localized foraging during this season.     

Quantitative and qualitative observations on Laminaria dentigera and other subtidal kelps of southern Kodiak Island,Alaska

Observations were made and quantitative samples of non-canopy kelps were collected, using SCUBA, at 9 subtidal sites off southern Kodiak Island in May, 1976. At a 10th site, only observations were made. The most abundant species in the quadrats were Laminaria dentigera, L. yezoensis, Pleurophycus gardneri, Agarum cribrosum, and Alaria marginata. We found fertile plants of all these species. L. dentigera dominated at all sampled sites except one within a bed of Nereocystis luetkeana, where P. gardneri was dominant. L. yezoensis was dominant at the site which was visited but not sampled. The average wet weight of non-canopy kelps in the 55 samples was 12 kg m^-2, and the quadrat range was 2 to 35 kg m^-2. The reported ranges of P. gardneri and Alaria marginata are extended westward from Montague Island (147°22W) to Bumble Bay, Kodiak Island (154°43W), where A. marginata was found at a depth of 8.5 m.     

How dynamics and drivers of land use/land cover change impact elephant conservation and agricultural livelihood development in Rombo,Tanzania

Land use/land cover (LULC) change affects the provision of ecosystem services for humans and habitat for wildlife. Hence, it is crucial to monitor LULC particularly adjacent to protected areas. In this study, we measured LULC change in Rombo, Tanzania, an area with high-potential agro-ecological zones that is dominated by human–elephant conflicts (HECs). We used remote sensing and geographical information system techniques, questionnaires and village meetings to assess spatio-temporal patterns of the LULC changes in the study area. Using Landsat imagery, digital elevation model (DEM) and ground truthing, we classified and monitored changes in LULC from the years 1987 to 2015. We found that within Rombo, settlements were increasing, while agricultural and agroforestry lands were decreasing and respondents’ perceptions varied along the altitudinal gradient. Patterns of HEC and LULC were observed to change along the gradient and the later threatened the agricultural land and ecological integrity for elephant habitat, leading to high tension and competition between elephants and people. This research offers baseline information for land use planning to balance wildlife conservation with livelihood development in Rombo and highlights that managing the impacts of LULC changes on HEC and elephant habitat loss is a matter of urgency.     

南太湖沉积岩芯中金属和营养元素的垂向分布特征及其意义

通过太湖南部钻孔(THS)沉积物^137Cs和^210Pb、总有机碳、总氮、总磷、粒度和金属元素分析，探讨其垂向分布特征及意义。表层^137Cs比活度高值表明表层沉积物具有混合作用的特征。粒度分析表明，沉积物呈现随深度变浅而变粗的总体趋势；绝大多数金属元素含量垂向上的变化与粘土含量呈正相关。铁锰界面循环也影响着金属元素的迁移。沉积物W(C)／w(N)比值的状况显示了其有机质主要为陆源性质，但最近60年内来源于水体的有机质呈现增加的趋势。沉积物中元素的演化序列表明，元素铅受到明显的人为活动来源的污染。     

崇明岛典型土地利用方式对土壤有机碳和酶活性的影响

为了研究上海崇明岛不同土地利用方式对土壤有机碳以及酶活性的影响,对冬小麦地、蔬菜地、柑桔地和冬绿肥地4种土地利用方式的土壤进行了调查、样品测定和分析。结果表明,土地利用方式对有机碳和4种酶活性的影响极显著。冬小麦地总有机碳和活性有机碳质量分数均最高,分别为19.23g·kg^-1和2.26mg·g^-1,柑桔地土壤脲酶、碱性磷酸酶、纤维素酶和蛋白酶等4种酶活性均高于其他3种土地利用方式,表明采用秸秆还田式的冬小麦地,可以增加土壤总有机碳和活性有机碳质量分数;常年处于免耕状态的柑桔地可以使土壤酶活性增强。     

Effects of predators on sea urchin density and habitat use in a southern California kelp forest

Grazing sea urchins can reduce kelp abundance and therefore strongly affect kelp forest community structure. Despite the ecological importance of sea urchins, direct field studies on the role that urchin predators play in shaping urchin populations are rare for southern California. We conducted surveys and manipulative experiments within kelp forests near San Diego, CA, (32–51′28″N, 117–16′00″W) from 2006 to 2009 to determine whether predators such as sheephead (Semicossyphus pulcher) and spiny lobsters (Panulirus interruptus) may be linked to purple urchin (Strongylocentrotus purpuratus) and red urchin (Strongylocentrotus franciscanus) distribution and habitat use, as well as purple urchin density-dependent mortality. Purple urchins were less dense and more cryptic inside a local marine protected area (MPA) that contained high predator abundance than in nearby heavily fished areas, whereas red urchins rarely were found outside the MPA. Urchin proportional mortality was inversely density dependent during the day when sheephead were active, despite fish aggregations in plots of high urchin density, but was density independent during the night when lobsters were active. Urchin mortality was reduced under understory algal cover during the day, but not during the night. Examining whether urchin mortality from predation is density dependent and how habitat complexity influences this relationship is imperative because behavioral changes and increases in urchin populations can have vast ecological and economic consequences in kelp forest communities.     

Costs of courtship and mating in a sexually cannibalistic orb-web spider: female mating strategies and their consequences for males

The costs of courtship and mating may include increased risks of predation, the transmission of pathogens, and a loss of foraging opportunities. Thus, a female's decision to tolerate a courting male will depend upon how these costs offset the benefits of mating, which will depend on her reproductive and nutritional status. While these costs may be similar for mated and unmated females, the benefits of mating will be less for mated than virgin females. However, the cost of lost foraging opportunities may be higher for females with fewer nutritional reserves necessary for forming eggs. We examined how these costs and benefits influence the courtship and mating behaviour of male and female orb-web spiders, Argiope keyserlingi. In the field, females on webs that also contained a courting male intercepted fewer prey items per hour than females on webs without a male. In the laboratory, the presence of a courting male at the hub also attracted mantid predators to the web, increasing the risk of predation for both male and female. Staged mating experiments in the laboratory revealed that the frequency of female attacks and pre-copulatory cannibalism was greater among mated than virgin females. Feeding history did not affect aggression in virgin females but, among mated females, food-deprived spiders attacked and cannibalized males more frequently than sated females and only the latter ever remated. These differences in female behaviour influenced male mating strategies. Choice experiments demonstrated that males preferred to venture onto the silk threads of virgin rather than those of mated females. Similar patterns of mate selectivity were observed in the field; females with narrow abdomens attracted more males to the webs than females with broad abdomens, and copulations were observed more frequently among females with narrow abdomens. These smaller females are likely to be virgins that have recently molted. Males that preferentially mate with virgin females will not only avoid potentially fatal attacks but also obtain, on average, a higher fertilization success.     

Infant care,infanticide, and female reproductive strategies in polygynous groups of common marmosets (Callithrix jacchus)

Reproduction in marmoset and tamarin groups is typically restricted to a single dominant female, but it is unclear why subordinate females tolerate delayed reproduction. The presence of two breeding females in free-ranging groups of common marmosets (Callithrix jacchus) presented a unique opportunity to examine differences in the reproductive strategies of dominant and subordinate females. Three groups were monitored for 12–18 months at a forest reserve in northeastern Brazil. Data on infant care were collected during two consecutive all-day follows every 10 days until the infants were 2 months old. Carrying patterns for infants born to dominant females were similar to those observed in groups containing a single breeding female. All group members over 5 months of age participated in infant care, and dominant females allowed some group members to carry their infants from the 1 st day of life. In contrast, subordinate females were protective of their offspring and were their sole caretakers for at least a week following birth. One infant born to a subordinate was killed in an attack involving the dominant female. Overall, dominant females gave birth to more infants and had higher infant survival than did subordinate breeding females. subordinate females were successful in rearing young only when the timing of births was such that they did not overlap with the dependency period of infants born to the dominant female. These patterns suggest that subordinate females may face resource competition, especially over access to helpers. In times of severe competition, subordinate females that delay reproduction may be avoiding a wasted reproductive effort.