Satellite imagery can be used to detect variation in abundance of Weddell seals (Leptonychotes weddellii) in Erebus Bay, Antarctica

The Weddell seal population in Erebus Bay, Antarctica, represents one of the best-studied marine mammal populations in the world, providing an ideal test for the efficacy of satellite imagery to inform about seal abundance and population trends. Using high-resolution (0.6 m) satellite imagery, we compared counts from imagery to ground counts of adult Weddell seals and determined temporal trends in Erebus Bay during November 2004–2006 and 2009, and December 2007. Seals were counted from QuickBird-2 and WorldView-1 images, and these counts were compared with ground counts at overlapping locations within Erebus Bay during the same time. Counts were compared across years and within individual haul-out locations. We counted a total of 1,000 adult Weddell seals from five images across all years (for a total of 21 satellite-to-ground count comparisons), approximately 72% of the total counted on the ground at overlapping locations. We accurately detected an increase in abundance during 2004–2009. There was a strong, positive correlation (r = 0.98, df = 3, P < 0.003) between ground counts and counts derived from the imagery. The correlation between counts at individual haul-out locations was also strong (r = 0.80, df = 19, P < 0.001). Detection rates ranged from 30 to 88%. Overall, our results showed the utility of high-resolution imagery to provide an accurate way to detect the presence and variation in abundance of Weddell seals. Our methods may be applied to other species in polar regions, such as walruses or polar bears, particularly in areas where little is known about population status.     

Isolated faecal bacterial communities found for Weddell seals,Leptonychotes weddellii,at White Island,McMurdo Sound,Antarctica

The microbial diversity of faecal communities co-existing with mega fauna is not well understood even though these faecal communities are critical for health and development. Additionally, the transfer of microbial taxa among host animals is little studied. Here, we used 16S sequences obtained from clone libraries to characterise the faecal microbiota of Weddell seals breeding in McMurdo Sound and at White Island, Antarctica. Faecal bacterial communities were dominated by four phyla; Actinobacteria (20 %), Bacteroidetes (13 %), Firmicutes (23 %), and Proteobacteria (13 %). We also used automated ribosomal intergenic spacer analysis to examine the dispersal of bacteria between populations of Weddell seals breeding at White Island and in McMurdo Sound. The Weddell seals at White Island are isolated by the Ross Ice Shelf from the larger population of Weddell seals breeding in McMurdo Sound. We found that the faecal bacteria communities of the seals at White Island had lower diversity and that the community composition was significantly different compared with the seals in the McMurdo Sound area.     

DEVELOPMENT AND APPLICATION OF A MASS-ESTIMATION METHOD FOR WEDDELL SEALS

Traditional methods of acquiring mass data limit the ability to collect large samples from across populations of some pinnipeds, or to sample without great disturbance to the animals. In order to collect substantial samples of mass data from the Weddell seal (Leptonychotes weddellii) population in Erebus Bay, Antarctica, we developed the equipment and methods for estimating the mass of Weddell seals using digital photographs. Resulting regression models predict the mass of adult female seals to within ±13.8% of estimated mass, and ±25.9% of estimated mass for pups. We show the protocols developed are repeatable and efficient enough to be applied to a large number of animals in a relatively short period of time and may be useful for studies of other marine mammals. We caution that prediction intervals exist around mass estimates and must be accounted for when estimates are applied to biological questions. In a limited application of the method, differences in mass transfer between experienced and inexperienced maternal females and their pups were detected when prediction error variance around mass estimates was explicitly included. Similar mass‐estimation methods may therefore be useful in consideration of biological questions requiring large samples of mass previously unattainable.     

Variation of annual apparent survival and detection rates with age,year and individual identity in male Weddell seals (Leptonychotes weddellii) from long-term mark-recapture data

Exploring age- and sex-specific survival rates provides insight regarding population behavior and life-history trait evolution. However, our understanding of how age-specific patterns of survival, including actuarial senescence, compare between the sexes remains inadequate. Using 36 years of mark-recapture data for 7,516 male Weddell seals (Leptonychotes weddellii) born in Erebus Bay, Antarctica, we estimated age-specific annual survival rates using a hierarchical model for mark-recapture data in a Bayesian framework. Our male survival estimates were moderate for pups and yearlings, highest for 2-year-olds, and gradually declined with age thereafter such that the oldest animals observed had the lowest rates of any age. Reports of senescence in other wildlife populations of species with similar longevity occurred at older ages than those presented here. When compared to recently published estimates for reproductive Weddell seal females, we found that peak survival rates were similar (males: 0.94, 95% CI = 0.92–0.96; females: 0.92, 95% CI = 0.93–0.95), but survival rates at older ages were lower in males. Age-specific male Weddell seal survival rates varied across years and individuals, with greater variation occurring across years. Similar studies on a broad range of species are needed to contextualize these results for a better understanding of the variation in senescence patterns between the sexes of the same species, but our study adds information for a marine mammal species to a research topic dominated by avian and ungulate species.     

Cryptic inbreeding depression in a growing population of a long‐lived species

Genetic effects are often overlooked in endangered species monitoring, and populations showing positive growth are often assumed to be secure. However, the continued reproductive success of a few individuals may mask issues such as inbreeding depression, especially in long‐lived species. Here, we test for inbreeding depression in little spotted kiwi (Apteryx owenii) by comparing a population founded with two birds to one founded with 40 birds, both from the same source population and both showing positive population growth. We used a combination of microsatellite genotypes, nest observations and modelling to examine the consequences of assessing population viability exclusively via population growth. We demonstrate (i) significantly lower hatching success despite significantly higher reproductive effort in the population with two founders; (ii) positive growth in the population with two founders is mainly driven by ongoing chick production of the founding pair; and (iii) a substantial genetic load in the population founded with two birds (10–15 diploid lethal equivalents). Our results illustrate that substantial, cryptic inbreeding depression may still be present when a population is growing, especially in long‐lived species with overlapping generations.     

Variability in temporary emigration rates of individually marked female Weddell seals prior to first reproduction

In many species, temporary emigration (TE) is a phenomenon, often indicative of life-history characteristics such as dormancy, skipped reproduction, or partial migration, whereby certain individuals in a population are temporarily unobservable at a particular site. TE may be a flexible condition-dependent strategy that allows individuals to mitigate effects of adverse conditions. Consequently, TE rates ought to be highly variable, but sources of variations are poorly understood for most species. We used data from known-aged female Weddell seals (Leptonychotes weddellii) tagged in Erebus Bay, Antarctica, to investigate sources of variation in TE rates prior to reproduction and to evaluate possible implications for age-specific probability of first reproduction. TE rates were near 1 the year after birth, decreased to an average of 0.15 ( $ \widehat{\text{SE}} $  = 0.01) by age 8, and were similar thereafter. TE rates varied substantially from year-to-year and were lower for seals that attended reproductive colonies the previous year than for seals that did not attend (e.g., $ \overline{{\hat{\psi }_{{i,{\text{age}}\,8}}^{\text{UU}} - \hat{\psi }_{{i,\,{\text{age}}\,8}}^{\text{PU}} }} $  = 0.22). Recruitment rates were marginally greater for seals that did attend than for seals that did not attend colonies the previous year. For Weddell seals specifically, our results suggest that (1) motivation to attend colonies varied temporally, (2) as seals grew older they had increased motivation to attend even before reproductive maturity, and (3) seals appear to follow various attendance strategies. More broadly, our results support the idea of TE as a variable, condition-dependent strategy, and highlight the utility of TE models for providing population and life-history insights for diverse taxa.     

Variation in probability of first reproduction of Weddell seals

1. For many species, when to begin reproduction is an important life-history decision that varies by individual and can have substantial implications for lifetime reproductive success and fitness. 2. We estimated age-specific probabilities of first-time breeding and modelled variation in these rates to determine age at first reproduction and understand why it varies in a population of Weddell seals in Erebus Bay, Antarctica. We used multistate mark-recapture modelling methods and encounter histories of 4965 known-age female seals to test predictions about age-related variation in probability of first reproduction and the effects of annual variation, cohort and population density. 3. Mean age at first reproduction in this southerly located study population (7.62 years of age, SD=1.71) was greater than age at first reproduction for a Weddell seal population at a more northerly and typical latitude for breeding Weddell seals (mean=4-5 years of age). This difference suggests that age at first reproduction may be influenced by whether a population inhabits the core or periphery of its range. 4. Age at first reproduction varied from 4 to 14 years, but there was no age by which all seals recruited to the breeding population, suggesting that individual heterogeneity exists among females in this population. 5. In the best model, the probability of breeding for the first time varied by age and year, and the amount of annual variation varied with age (average variance ratio for age-specific rates=4.3%). 6. Our results affirmed the predictions of life-history theory that age at first reproduction in long-lived mammals will be sensitive to environmental variation. In terms of life-history evolution, this variability suggests that Weddell seals display flexibility in age at first reproduction in order to maximize reproductive output under varying environmental conditions. Future analyses will attempt to test predictions regarding relationships between environmental covariates and annual variation in age at first reproduction and evaluate the relationship between age at first reproduction and lifetime reproductive success.     

Long-term monitoring of Antarctic pinnipeds in Admiralty Bay (South Shetlands, Antarctica)

Year-round monitoring of five Antarctic pinnipeds was conducted in Admiralty Bay from 1988 up to 2000. Two breeding species: southern elephant sealsMirounga leonina (Linnaeus, 1758) and Weddell sealsLeptonychotes weddellii (Lesson, 1826), were present throughout the year. Three other species: crabeater seals Lobodon carcinophagus (Hobron and Jacquinot, 1842), leopard sealsHydrurga leptonyx (Blainville, 1820), and Antarctic fur sealsArctocephalus gazella (Peters, 1875) visited the area only for short periods. During this study, the abundance of elephant seals was stable, whereas those of Weddell and crabeater seals declined. Leopard seals numbers fluctuated irregularly. We detected a possible immigration from South Georgia: of a stable magnitude for elephant seals, and of variable magnitude, depending on food accessibility, for Antarctic fur seals. We found a strong recurrence of the spatial distributions of elephant, Weddell, and Antarctic fur seals in the 13 oases on the shore of Admiralty Bay. Annual distribution patterns were characteristic for each species. The innermost beaches were used predominantly by the animals during their annual fasts: the breeding and the moulting seasons.     

The roles of demography and genetics in the early stages of colonization

Colonization success increases with the size of the founding group. Both demographic and genetic factors underlie this relationship, yet because genetic diversity normally increases with numbers of individuals, their relative importance remains unclear. Furthermore, their influence may depend on the environment and may change as colonization progresses from establishment through population growth and then dispersal. We tested the roles of genetics, demography and environment in the founding of Tribolium castaneum populations. Using three genetic backgrounds (inbred to outbred), we released individuals of four founding sizes (2–32) into two environments (natal and novel), and measured establishment success, initial population growth and dispersal. Establishment increased with founding size, whereas population growth was shaped by founding size, genetic background and environment. Population growth was depressed by inbreeding at small founding sizes, but growth rates were similar across genetic backgrounds at large founding size, an interaction indicating that the magnitude of the genetic effects depends upon founding population size. Dispersal rates increased with genetic diversity. These results suggest that numbers of individuals may drive initial establishment, but that subsequent population growth and spread, even in the first generation of colonization, can be driven by genetic processes, including both reduced growth owing to inbreeding depression, and increased dispersal with increased genetic diversity.     

Influence of age and individual identity in the use of breeding colony habitat by male Weddell seals in Erebus Bay,Antarctica

Male–male contest behavior can contribute to spatial distributions of male pinnipeds during breeding seasons. To maximize breeding opportunities, the most competitive males would be expected to be surrounded by the highest numbers of reproductive-age females. As information regarding fine-scale spatial ecology of Weddell seals is lacking, we performed an exploratory study using kernel density analyses to evaluate age-specific habitat use of male Weddell seals in Erebus Bay, Antarctica. Additionally, we investigated the relationship between age and number of surrounding reproductive-age females using a competing set of regression models in a Bayesian framework that considered different functional forms of age while incorporating individual heterogeneity. As male adult Weddell seals aged, to at least 20 years, they were more likely to be found in areas associated with the greatest densities of reproductive-age females, but individual heterogeneity also influenced the number of reproductive-age female neighbors. The youngest males tended to haul out in offshore areas associated with better hunting, and older males tended to settle in more nearshore areas associated with more pup production. Our findings from this preliminary investigation indicate that male Weddell seal spatial behavior during the breeding season varies with age and individual and might be related to reproductive activity.     

Dinucleotide microsatellite markers from the Antarctic seals and their use in other Pinnipeds

Twenty‐four microsatellite loci were isolated from three species of Antarctic seals (Subfamily Monachinae, Tribe Lobodontini). Eleven loci were cloned from Weddell seal, Leptonychotes weddellii, seven from leopard seal, Hydrurga leptonyx, and six from crabeater seal, Lobodon carcinophagus. Variability was assessed in Weddell seals collected in McMurdo Sound, leopard seals from Bird Island, South Georgia, and crabeater seals sampled in the eastern Ross Sea. All loci were variable in the three species used for cloning and 22 of these loci amplified variable products in the Ross seal, Ommatophoca rossii. Cross‐species amplification was largely successful, with an average of 19 loci amplifying products in other phocids.     

Genetically effective population sizes of Antarctic seals estimated from nuclear genes

We analyzed eight nuclear microsatellite loci in three species of Antarctic seals; Weddell seal (Leptonychotes weddellii; mean N = 163), crabeater seal (Lobodon carcinophaga; 138) and Ross seal (Ommatophoca rossii; 35). We estimated genetic diversity (Θ) and effective population size (N _E) for each species. Autosomal microsatellite based N _E estimates were 151,200 for Weddell seals, 880,200 for crabeater seals, and 254,500 for Ross seals. We screened one X-linked microsatellite (Lw18), which yielded similar N _E estimates to the autosomal loci for all species except the Ross seals, where it was considerably larger (~103 times). Microsatellite N _E estimates were comparable with previously published N _E estimates from mitochondrial DNA, but both are substantially lower than direct estimates of population size in all species except the Ross seals. The ratio of maternally versus biparentally derived estimates of N _E for Ross seals was not consistent with the hypothesis that they are a polygynous species. We found no sign of a recent, sustained genetic bottleneck in any of the species.     

Inbreeding depression in a critically endangered carnivore

Harmful effects arising from matings between relatives (inbreeding) is a long‐standing observation that is well founded in theory. Empirical evidence for inbreeding depression in natural populations is however rare because of the challenges of assembling pedigrees supplemented with fitness traits. We examined the occurrence of inbreeding and subsequent inbreeding depression using a unique data set containing a genetically verified pedigree with individual fitness traits for a critically endangered arctic fox (Vulpes lagopus) population. The study covered nine years and was comprised of 33 litters with a total of 205 individuals. We recorded that the present population was founded by only five individuals. Over the study period, the population exhibited a tenfold increase in average inbreeding coefficient with a final level corresponding to half‐sib matings. Inbreeding mainly occurred between cousins, but we also observed two cases of full‐sib matings. The pedigree data demonstrated clear evidence of inbreeding depression on traditional fitness traits where inbred individuals displayed reduced survival and reproduction. Fitness traits were however differently affected by the fluctuating resource abundande. Inbred individuals born at low‐quality years displayed reduced first‐year survival, while inbred individuals born at high‐quality years were less likely to reproduce. The documentation of inbreeding depression in fundamental fitness traits suggests that inbreeding depression can limit population recovery. Introducing new genetic material to promote a genetic rescue effect may thus be necessary for population long‐term persistence.     

An apparent population decrease,or change in distribution,of Weddell seals along the Victoria Land coast

Ground counts during 1959–1968 compared with counts using high resolution (0.6 m²) satellite imagery during 2008–2012 indicated many fewer Weddell seals (Leptonychotes weddellii) at two major molting areas in the western Ross Sea: Edisto Inlet‐Moubray Bay, northern Victoria Land, and McMurdo Sound, southern Victoria Land. Breeding seals have largely disappeared from Edisto‐Moubray, though the breeding population in McMurdo Sound appears to have recovered from harvest in the 1960s. The timing of decline, or perhaps spreading (lower numbers of seals in more places), is unknown but appears unrelated to changes in sea ice conditions. We analyzed both historic and satellite‐derived ice data confirming a large expansion of pack ice mostly offshore of the Ross Sea, and not over the continental shelf (main Weddell seal habitat), and a thinning of fast ice along Victoria Land (conceivably beneficial to seals). Timing of fast ice presence and extent in coves and bays along Victoria Land, remains the same. The reduction in numbers is consistent with an altered food web, the reasons for which are complex. In the context of a recent industrial fishery targeting a seal prey species, a large‐scale seal monitoring program is required to increase understanding of seal population changes.     

Early colonisation population structure of a Norway rat island invasion

Colonists undergo non-equilibrium processes such as founder effects, inbreeding and changing population size which influence the mating system and demography of a population. Understanding these processes in colonising populations informs management and helps prevent further invasions. We sampled and genotyped most individuals of a Norway rat (Rattus norvegicus) reinvasion on Moturemu island (5 ha) in New Zealand. Population size was most likely between 30 and 33 rats. Genetic methods detected a clear bottleneck signal from the founding population. Parentage assignment revealed promiscuous mating dominated by a few individuals with increasing inbreeding, both putatively a result of small island size. Combining ecological and genetic data from a single sample allowed inferences on population structure and functioning. Invading Norway rats rapidly achieve population structure similar to established island populations despite a small number of colonists and associated inbreeding. Overcoming these initial obstacles to population establishment contributes to the global success of invasive rats.     

Conservation genetics of the Critically Endangered Saint Croix ground lizard (<Emphasis Type="Italic">Ameiva polops</Emphasis> Cope 1863)

The Saint Croix ground lizard (Ameiva polops) is a Critically Endangered species endemic to Saint Croix, U.S. Virgin Islands. Although it is completely extirpated from Saint Croix Island (last seen in 1968), two small natural satellite populations survive on two islets off St. Croix: one on Protestant Cay (estimated at ~30 individuals in 2002); and one on Green Cay (estimated at ~180 individuals in 2002). Two additional small populations exist that were founded with individuals translocated from the two surviving natural populations. One is on Ruth Island, a man-made islet off St. Croix, founded in 1990 with 10 individuals from Protestant Cay. The other is on Buck Island, ~2.5 km from Saint Croix, founded in 2008 with 57 individuals from Green Cay. All populations are vulnerable to catastrophic events such as hurricanes, sea level rise, introduction of exotic species, and landscape transformation. Herein, we used mitochondrial and nuclear-microsatellite markers to examine levels of genetic diversity within extant populations of A. polops and the degree of genetic differentiation among them. We also conducted analyses to search for signatures of recent bottlenecks in these populations and to estimate their effective population size (N _e ). We found low genetic variability within populations of this lizard, comparable to that observed in other threatened vertebrates. We also found significant genetic differentiation among the three populations examined, as well as signatures of recent bottlenecks and critically low N _e values in all populations. Based on our results, we suggest two different conservation units for A. polops: (1) Green Cay and its replicate population at Buck Island; and (2) Protestant Cay and its replicate population at Ruth Island. We discuss the implications of our findings on the conservation and management of A. polops.     

Summer diet of Weddell Seals (Leptonychotes weddelli) in the eastern and southern Weddell Sea,Antarctica

Summary Stomach and intestine samples from 21 adult Weddell seals were used to study the diet of these seals from the eastern and southern Weddell Sea coast from January to February 1983 and 1985. Fish occurred in all seals, squid in five, octopods in three and Euphausia crystallorophias in one seal. Pleuragramma antarcticum was the predominant fish in the diet, constituting 61.1% of otoliths in 1983 samples and 93.8% in 1985. Aethotaxis mitopteryx, Dissostichus mawsoni, unidentified Trematomus spp. and channichthyids were also recorded. Size and wet weight of P. antarcticum were calculated from uneroded otoliths, found in 6 seal stomachs with liquid food pulp, collected during early morning hours in 1985. Size distribution of P. antarcticum from individual seals was reasonably constant, ranging between 5.0 and 22.0 cm SL; adult fish from about 14.0 to 19.0 cm SL predominated. P. antarcticum in seals from the southern area had a larger median size (16.5 cm SL), than those from further east (15.5 cm SL). Calculated wet weights of all P. antarcticum from individual seal stomachs ranged between 4.7 and 16.9 kg the mean was 12.8 kg. Comparisons with net-hauls from the southern Gould Bay suggest that Weddell seals feed mainly in deeper water layers (>400 m) where adult P. antarcticum occur at higher densities.     

Inbreeding in reintroduced populations: the effects of early reintroduction history and contemporary processes

Maintaining genetic variation and minimizing inbreeding are central goals of conservation genetics. It is therefore crucial to understand the important population parameters that affect inbreeding, particularly in reintroduction programs. Using data from 41 reintroduced Alpine ibex (Capra ibex ibex) populations we estimated inbreeding since the beginning of reintroductions using population-specific Fst, and inbreeding over the last few generations with contemporary effective population sizes. Total levels of inbreeding since reintroduction of ibex were, on average, close to that from one generation of half-sib mating. Contemporary effective population sizes did not reflect total inbreeding since reintroduction, but 16% of variation in contemporary effective population sizes among populations was due to variation in current population sizes. Substantial variation in inbreeding levels among populations was explained by founder group sizes and the harmonic mean population sizes since founding. This study emphasizes that, in addition to founder group sizes, early population growth rates are important parameters determining inbreeding levels in reintroduced populations.     

Role of propagule pressure in colonization success: disentangling the relative importance of demographic,genetic and habitat effects

High propagule pressure is arguably the only consistent predictor of colonization success. More individuals enhance colonization success because they aid in overcoming demographic consequences of small population size (e.g. stochasticity and Allee effects). The number of founders can also have direct genetic effects: with fewer individuals, more inbreeding and thus inbreeding depression will occur, whereas more individuals typically harbour greater genetic variation. Thus, the demographic and genetic components of propagule pressure are interrelated, making it difficult to understand which mechanisms are most important in determining colonization success. We experimentally disentangled the demographic and genetic components of propagule pressure by manipulating the number of founders (fewer or more), and genetic background (inbred or outbred) of individuals released in a series of three complementary experiments. We used Bemisia whiteflies and released them onto either their natal host (benign) or a novel host (challenging). Our experiments revealed that having more founding individuals and those individuals being outbred both increased the number of adults produced, but that only genetic background consistently shaped net reproductive rate of experimental populations. Environment was also important and interacted with propagule size to determine the number of adults produced. Quality of the environment interacted also with genetic background to determine establishment success, with a more pronounced effect of inbreeding depression in harsh environments. This interaction did not hold for the net reproductive rate. These data show that the positive effect of propagule pressure on founding success can be driven as much by underlying genetic processes as by demographics. Genetic effects can be immediate and have sizable effects on fitness.     

Assessing polar bear (Ursus maritimus) population structure in the Hudson Bay region using SNPs

Defining subpopulations using genetics has traditionally used data from microsatellite markers to investigate population structure; however, single‐nucleotide polymorphisms (SNPs) have emerged as a tool for detection of fine‐scale structure. In Hudson Bay, Canada, three polar bear (Ursus maritimus) subpopulations (Foxe Basin (FB), Southern Hudson Bay (SH), and Western Hudson Bay (WH)) have been delineated based on mark–recapture studies, radiotelemetry and satellite telemetry, return of marked animals in the subsistence harvest, and population genetics using microsatellites. We used SNPs to detect fine‐scale population structure in polar bears from the Hudson Bay region and compared our results to the current designations using 414 individuals genotyped at 2,603 SNPs. Analyses based on discriminant analysis of principal components (DAPC) and STRUCTURE support the presence of four genetic clusters: (i) Western—including individuals sampled in WH, SH (excluding Akimiski Island in James Bay), and southern FB (south of Southampton Island); (ii) Northern—individuals sampled in northern FB (Baffin Island) and Davis Strait (DS) (Labrador coast); (iii) Southeast—individuals from SH (Akimiski Island in James Bay); and (iv) Northeast—individuals from DS (Baffin Island). Population structure differed from microsatellite studies and current management designations demonstrating the value of using SNPs for fine‐scale population delineation in polar bears.