Wollebaeki Research Articles

OCURRENCIA Y TASAS DE ENCUENTRO DE MAMÍFEROS MARINOS EN LAS AGUAS DE LA ISLA MALPELO Y HACIA EL CONTINENTE

This work compiles a decade (2001-2010) of marine mammal sightings in the Malpelo Fauna and Flora Sanctuary (FFS) and the area between the island and mainland coast. Four separate sources of data were consulted, which used visual searching during cruising efforts while Malpelo’s surrounding waters were surveyed from a vantage point. Seven species were identified in the FFS: October and November were the months with higher species richness. Tursiops truncatus had the highest encounter rates (17.78 groups/100 h), followed by Megaptera novaeangliae (1.62) and Stenella attenuata (0.88). These species were usually within 6 km from the island. Other species seen around the island include Stenella coeruleoalba, Delphinus delphis, Stenella longirostris and Zalophus wollebaeki. On the other hand, thirteen species were identified during cruises, and March and April were the months with the highest species richness. Megaptera novaeangliae had the highest encounter rate (5.94), followed by T. truncatus (3.30), S. attenuata (3.08), D. delphis (3.08) and S. coeruleoalba, Globicephala macrorhynchus and Orcinus orca, each one with 0.66. Other species seen during the cruises include Steno bredanensis, Pseudorca crassidens, Grampus griseus, Peponocephala electra, Physeter macrocephalus and Ziphius cavirostris. Megaptera novaeangliae were associated to the continental shelf, T. trunctaus and D. delphis to oceanic waters, and S. attenuata to both the continental shelf and oceanic waters. Delphinus delphis was more abundant in intermediate waters, during the first trimester (January-March), while T. truncatus was the most abundant species around the Sanctuary during all seasons, suggesting that is the same population. The latter was also the only species found all year round in both zones (around the island and in oceanic waters), and encounter rates did not change across years. Megaptera novaeangliae had a seasonal presence (mostly June to November) with a higher abundance during the third trimester (July-September), both around Malpelo and the transect. The presence of humpback whales calves suggests that Malpelo is used for reproductive purposes. This new information about the marine mammals found in Malpelo FFS and the area between the island and the continent contributes to the understanding of these species in Colombian waters.

Context-dependent associations between heterozygosity and immune variation in a wild carnivore.

BackgroundA multitude of correlations between heterozygosity and fitness proxies associated with disease have been reported from wild populations, but the genetic basis of these associations is unresolved. We used a longitudinal dataset on wild Galapagos sea lions (Zalophus wollebaeki) to develop a relatively new perspective on this problem, by testing for associations between heterozygosity and immune variation across age classes and between ecological contexts.ResultsHomozygosity by locus was negatively correlated with serum immunoglobulin G production in pups (0–3 months of age), suggesting that reduced genetic diversity has a detrimental influence on the early development of immune defence in the Galapagos sea lion. In addition, homozygosity by locus was positively correlated with total circulating leukocyte concentration in juveniles (6–24 months of age), but only in a colony subject to the anthropogenic environmental impacts of development, pollution and introduced species, which suggests that reduced genetic diversity influences mature immune system activity in circumstances of high antigen exposure.ConclusionsThese findings demonstrate the environmental context-dependency of the phenotypic expression of immune variation, which is implicit in the theory of ecoimmunology, but which has been rarely demonstrated in the wild. They also indicate that heterozygosity may be linked to the maintenance of heterogeneity in mammalian immune system development and response to infection, adding to the body of evidence on the nature of the mechanistic link between heterozygosity and fitness.Electronic supplementary materialThe online version of this article (doi:10.1186/s12862-015-0519-6) contains supplementary material, which is available to authorized users.

Health Status of Galápagos Sea Lions (Zalophus wollebaeki) on San Cristóbal Island Rookeries Determined by Hematology, Biochemistry, Blood Gases, and Physical Examination.

The Galápagos sea lion, Zalophus wollebaeki, is an endemic and endangered species subject to population decline associated with environmental variability, such as El Niño events, constant feeding stress, and exposure to diseases through contact with introduced species. Reference blood parameter intervals have been published for some pinniped species, but baseline biochemical and blood gas values are lacking from Z. wollebaeki. We analyzed blood samples from 30 juvenile Galápagos sea lions (19 females, 11 males) captured in two rookeries on San Cristóbal Island. A portable blood analyzer (iSTAT) was used to obtain near-immediate field results for pH, partial pressure of O2, partial pressure of CO2, bicarbonate (HCO3(-)), hematocrit (Hct), hemoglobin, Na, K, ionized Ca, and glucose, and blood lactate was measured using a portable Lactate Plus(TM) analyzer. Average heart rate, biochemistry, and hematology parameters were comparable with healthy individuals of other pinniped species. Hemoglobin was significantly correlated with body condition of juvenile Galápagos sea lions. When compared with available blood values of clinically healthy California sea lions, Galápagos sea lions had higher total protein and Hct and lower Ca and K levels. Our results provide baseline data that may be useful in comparisons among populations and in detecting changes in health status among Galápagos sea lions.

Behavioural and environmental correlates of Philophthalmus zalophi infections and their impact on survival in juvenile Galapagos sea lions

Parasitic infections can play a major role in the dynamics of animal populations by influencing the development and survival of juveniles. Understanding how intrinsic and extrinsic factors shape individual parasitisation and affect long-term consequences of infections is therefore of prime importance for conservation management. Galapagos sea lions (Zalophus wollebaeki) provide a good model system to assess the impact of environmental and behavioural differences on parasitisation. Their environment is characterised by seasonal changes in sea surface temperatures (SST), and the slow transition to independence in juveniles goes along with major behavioural changes. The present study focused on infections with Philophthalmus zalophi, a digenetic trematode, which is regularly found in the ocular cavity of young sea lions. Long-term data on P. zalophi infections were collected from a sea lion colony located in the centre of the Galapagos archipelago (0°45′S, 90°160′W, 2007–2014). The probability to become infected differed between age classes and was influenced by the SST at the time of capture. About 16 % of the animals infected during the peak of infections developed severe clinical signs which had long-term consequences for juvenile survival. The transition to independence appeared to be a particularly sensitive period when negative consequences of previous parasitic infections exerted the greatest influence on juvenile survival. Our results suggest that the prevalence of infections and thus the negative implications for juvenile survival may increase with broad-scale climatic changes.

Differences in foraging ecology align with genetically divergent ecotypes of a highly mobile marine top predator.

Foraging differentiation within a species can contribute to restricted gene flow between ecologically different groups, promoting ecological speciation. Galapagos sea lions (Zalophus wollebaeki) show genetic and morphological divergence between the western and central archipelago, possibly as a result of an ecologically mediated contrast in the marine habitat. We use global positioning system (GPS) data, time-depth recordings (TDR), stable isotope and scat data to compare foraging habitat characteristics, diving behaviour and diet composition of Galapagos sea lions from a western and a central colony. We consider both juvenile and adult life stages to assess the potential role of ontogenetic shifts that can be crucial in shaping foraging behaviour and habitat choice for life. We found differences in foraging habitat use, foraging style and diet composition that aligned with genetic differentiation. These differences were consistent between juvenile and adult sea lions from the same colony, overriding age-specific behavioural differences. Our study contributes to an understanding of the complex interaction of ecological condition, plastic behavioural response and genetic make-up of interconnected populations.

Urban life of Galapagos sea lions (Zalophus wollebaeki) on San Cristobal Island, Ecuador: colony trends and threats

Worldwide, pristine environments are influenced by human societies. In the Galapagos Islands, the endangered, endemic Galapagos sea lion (Zalophus wollebaeki) has formed one of the biggest colonies within the town center of Puerto Baquerizo Moreno. About 8,000 people live there and human wildlife interactions occur daily.With colony counts and direct observations from 2008 to 2012, we analyze cause of death, injuries and disease of urban sea lion colonies at Wreck Bay. Population increase since 2008 can be attributed to an immigration of adult sea lions in 2010, resulting in an increase in the pup and juvenile production in 2011 and 2012. Pup mortality increased drastically to 2009 and again in 2011 and 2012. Besides pup mortality, most of the deaths are caused by increased disease incidences and human activity. Our observations suggest that overall 65% of the injuries observed are produced by human interaction.The increase in threats leading to death, injuries or disease can have long-term effects on the population. Although threats that cause physical injuries can be managed locally, sea lions range movements contributes to the spread of infectious pathogens, which may affect neighbor colonies and potentially have an impact on the survival of the species. Our study reveals the need of stronger efforts towards a more complete understanding of threats and especially disease spread among Galapagos Sea lions in urban environments and the establishment of more effective management measures.

Lazy sons, self-sufficient daughters: are sons more demanding?

Parent–offspring conflict exists because the optima of parental care differ between parent and offspring. In many polygynous species, males are larger than females and may require additional allocation of maternal resources. Differential allocation to sons and daughters requires either spontaneously increased supply of maternal resources in the more expensive sex or that sex to induce additional supply by its greater demand. In the Galapagos sea lion, Zalophus wollebaeki, males are larger than females from birth to weaning. However, offspring begin to forage by diving before weaning. Therefore, we investigated whether sons demand more supply of maternal resources than daughters or contribute more by independent foraging. Our results show that during the first 2 years of life, sons were much less likely than daughters to forage independently, despite very similar diving abilities. During this time, females apparently sustained sons by providing milk whereas daughters contributed substantially to their energetic needs by independent foraging. The observed difference in reliance on self-feeding between male and female offspring is consistent with the idea that males demand more, but could also be due to benefits to the mother when supplying more milk in sons. Sex differences in offspring contribution to their own maintenance during the transition to independence offer a new perspective on mother–offspring conflict worth further investigation.

Competition influence in the segregation of the trophic niche of otariids: a case study using isotopic Bayesian mixing models in Galapagos pinnipeds.

The feeding success of predators is associated with the competition level for resources, and, thus, sympatric species are exposed to a potential trophic overlap. Isotopic Bayesian mixing models should provide a better understanding of the contribution of preys to the diet of predators and the feeding behavior of a species over time. The carbon and nitrogen isotopic signatures from pup hair samples of 93 Galapagos sea lions and 48 Galapagos fur seals collected between 2003 and 2009 in different regions (east and west) of the archipelago were analyzed. A PDZ Europa ANCA-GSL elemental analyzer interfaced with a PDZ Europa 20-20 continuous flow gas source mass spectrometer was employed. Bayesian models, SIAR and SIBER, were used to estimate the contribution of prey to the diet of predators, the niche breadth, and the trophic overlap level between the populations. Statistical differences in the isotopic values of both predators were observed over the time. The mixing model determined that Galapagos fur seals had a primarily teutophagous diet, whereas the Galapagos sea lions fed exclusively on fish in both regions of the archipelago. The SIBER analysis showed differences in the trophic niche between the two sea lion populations, with the western rookery of the Galapagos sea lion being the population with the largest trophic niche area. A trophic niche partitioning between Galapagos fur seals and Galapagos sea lions in the west of the archipelago is suggested by our results. At intraspecific level, the western population of the Galapagos sea lion (ZwW) showed higher trophic breadth than the eastern population, a strategy adopted by the ZwW to decrease the interspecific competition levels in the western region.

Applicability of single-camera photogrammetry to determine body dimensions of pinnipeds: Galapagos sea lions as an example.

Morphological features correlate with many life history traits and are therefore of high interest to behavioral and evolutionary biologists. Photogrammetry provides a useful tool to collect morphological data from species for which measurements are otherwise difficult to obtain. This method reduces disturbance and avoids capture stress. Using the Galapagos sea lion (Zalophus wollebaeki) as a model system, we tested the applicability of single-camera photogrammetry in combination with laser distance measurement to estimate morphological traits which may vary with an animal’s body position. We assessed whether linear morphological traits estimated by photogrammetry can be used to estimate body length and mass. We show that accurate estimates of body length (males: ±2.0%, females: ±2.6%) and reliable estimates of body mass are possible (males: ±6.8%, females: 14.5%). Furthermore, we developed correction factors that allow the use of animal photos that diverge somewhat from a flat-out position. The product of estimated body length and girth produced sufficiently reliable estimates of mass to categorize individuals into 10 kg-classes of body mass. Data of individuals repeatedly photographed within one season suggested relatively low measurement errors (body length: 2.9%, body mass: 8.1%). In order to develop accurate sex- and age-specific correction factors, a sufficient number of individuals from both sexes and from all desired age classes have to be captured for baseline measurements. Given proper validation, this method provides an excellent opportunity to collect morphological data for large numbers of individuals with minimal disturbance.

Being on time: size-dependent attendance patterns affect male reproductive success

When reproduction occurs seasonally, males are expected to concentrate their reproductive effort when the abundance of females in oestrus is highest. In colonial species, large body size and prior residency correlate with the ability to establish dominance and ultimately to achieve matings. However, in addition to dominance, attendance in proximity to females may also enhance male mating success. If females come into oestrus highly asynchronously, males are unable to attend females for the entire breeding season. We expect the attendance of the most competitive males to increase with the availability of receptive females while less competitive males may primarily attend colonies when more competitive males are rare. In Galapagos sea lions, Zalophus wollebaeki, large males defend semiaquatic territories, but are unable to hold these over the entire reproductive season. They are sighted most often during the peak reproductive season. In this study, we found that nonterritorial males frequented the colony less during this time. However, the percentage of days intermediate-sized and resident males were sighted increased towards the peak of the reproductive season. Reproductive success was positively correlated with male attendance and therefore highest among territorial males. However, the correlation between attendance and reproductive success also existed for nonterritorial males. This suggests that male Galapagos sea lions adjust their attendance to the competitive situation in the colony. By increasing attendance during the peak of female oestrus, territorial males increase their chance of reproducing successfully. By adopting size-dependent strategies, nonterritorial males are able to achieve fitness while still growing.

Environment-driven changes in terrestrial habitat use and distribution of the Galapagos sea lion

Understanding habitat use patterns of marine apex predators is of paramount impor- tance for the effective design of conservation plans. Due to their amphibious life cycle, pinnipeds are one of the most vulnerable groups to habitat loss and degradation. Reducing knowledge gaps that still exist regarding temporal changes in tropical pinniped habitat use is therefore of particu- lar importance for conservation efforts. We analyzed terrestrial habitat use phenology of a tropical otariid, the Galapagos sea lion Zalophus wollebaeki. Regular land-based censuses were carried out in Wreck Bay of San Cristobal Island, Galapagos, between 2008 and 2012 to explore seasonal changes in sea lion abundance, distribution, and preferences among different habitat types. A daily cycle of haul-out patterns was observed. Animals aggregated on sandy beaches during the coldest periods of the day and were more abundant on rocky and artificial structures when air temperature was warmer. The use of artificial floating platforms as resting places by Z. wollebaeki was described for the first time and linked to environmental variability, suggesting that this may be a successful management tool for pinnipeds inhabiting low latitudes. Further, this species demonstrated seasonal distribution changes, aggregating in high densities during warm seasons and maintaining more even distributions during cold seasons. Sea surface temperature, air tem- perature, and the onset of the breeding season were the variables that most significantly influ- enced haul-out patterns of Z. wollebaeki, indicating that this species can adapt to changing envi- ronmental conditions by displaying a high degree of plasticity regarding its distribution and terrestrial microhabitat preferences.

Spatial variation in the foraging behaviour of the Galapagos sea lions (Zalophus wollebaeki) assessed using scat collections and stable isotope analysis

This study focuses on the comparative analysis of variables related to the trophic niche plasticity in the Galapagos sea lion (Zalophus wollebaeki). There is great concern regarding the future of this species, so study of the diet and the way these animals obtain their food is useful to understand and predict their survival possibilities in the long term. The combined use of scat analysis and stable isotopes helps to determine foraging habits of this species in greater detail. The objective of the study was to assess the feeding habits of Z. wollebaeki and its space–time variation. The scat analysis (n = 200) gave as a result space–time changes in the foraging strategies of this species. The isotopic values (n = 80) showed differences in relation to foraging grounds (δ13C: P = 0.001), but also suggested an apparent stability in the trophic level of their diet (δ15N: P = 0.084). These results constitute a relevant finding in the evolutionary behaviour of the species, showing that Z. wollebaeki has developed a high degree of plasticity in its foraging habits that may improve its survival in a highly demanding ecosystem in terms of limited and fluctuating resources.

Individual Foraging Strategies Reveal Niche Overlap between Endangered Galapagos Pinnipeds

Most competition studies between species are conducted from a population-level approach. Few studies have examined inter-specific competition in conjunction with intra-specific competition, with an individual-based approach. To our knowledge, none has been conducted on marine top predators. Sympatric Galapagos fur seals (Arctocephalus galapagoensis) and sea lions (Zalophus wollebaeki) share similar geographic habitats and potentially compete. We studied their foraging niche overlap at Cabo Douglas, Fernandina Island from simultaneously collected dive and movement data to examine spatial and temporal inter- and intra-specific competition. Sea lions exhibited 3 foraging strategies (shallow, intermediate and deep) indicating intra-specific competition. Fur seals exhibited one foraging strategy, diving predominantly at night, between 0–80 m depth and mostly at 19–22 h. Most sea lion dives also occurred at night (63%), between 0–40 m, within fur seals' diving depth range. 34% of sea lions night dives occurred at 19–22 h, when fur seals dived the most, but most of them occurred at dawn and dusk, when fur seals exhibited the least amount of dives. Fur seals and sea lions foraging behavior overlapped at 19 and 21 h between 0–30 m depths. Sea lions from the deep diving strategy exhibited the greatest foraging overlap with fur seals, in time (19 h), depth during overlapping time (21–24 m), and foraging range (37.7%). Fur seals foraging range was larger. Cabo Douglas northwest coastal area, region of highest diving density, is a foraging “hot spot” for both species. Fur seals and sea lions foraging niche overlap occurred, but segregation also occurred; fur seals primarily dived at night, while sea lions exhibited night and day diving. Both species exploited depths and areas exclusive to their species. Niche breadth generally increases with environmental uncertainty and decreased productivity. Potential competition between these species could be greater during warmer periods when prey availability is reduced.

Divergent allele advantage at MHC-DRB through direct and maternal genotypic effects and its consequences for allele pool composition and mating

It is still debated whether main individual fitness differences in natural populations can be attributed to genome-wide effects or to particular loci of outstanding functional importance such as the major histocompatibility complex (MHC). In a long-term monitoring project on Galápagos sea lions (Zalophus wollebaeki), we collected comprehensive fitness and mating data for a total of 506 individuals. Controlling for genome-wide inbreeding, we find strong associations between the MHC locus and nearly all fitness traits. The effect was mainly attributable to MHC sequence divergence and could be decomposed into contributions of own and maternal genotypes. In consequence, the population seems to have evolved a pool of highly divergent alleles conveying near-optimal MHC divergence even by random mating. Our results demonstrate that a single locus can significantly contribute to fitness in the wild and provide conclusive evidence for the 'divergent allele advantage' hypothesis, a special form of balancing selection with interesting evolutionary implications.

Immune Activity, Body Condition and Human-Associated Environmental Impacts in a Wild Marine Mammal

Within individuals, immunity may compete with other life history traits for resources, such as energy and protein, and the damage caused by immunopathology can sometimes outweigh the protective benefits that immune responses confer. However, our understanding of the costs of immunity in the wild and how they relate to the myriad energetic demands on free-ranging organisms is limited. The endangered Galapagos sea lion (Zalophus wollebaeki) is threatened simultaneously by disease from domestic animals and rapid changes in food availability driven by unpredictable environmental variation. We made use of this unique ecology to investigate the relationship between changes in immune activity and changes in body condition. We found that during the first three months of life, changes in antibody concentration were negatively correlated with changes in mass per unit length, skinfold thickness and serum albumin concentration, but only in a sea lion colony exposed to anthropogenic environmental impacts. It has previously been shown that changes in antibody concentration during early Galapagos sea lion development were higher in a colony exposed to anthropogenic environmental impacts than in a control colony. This study allows for the possibility that these relatively large changes in antibody concentration are associated with negative impacts on fitness through an effect on body condition. Our findings suggest that energy availability and the degree of plasticity in immune investment may influence disease risk in natural populations synergistically, through a trade-off between investment in immunity and resistance to starvation. The relative benefits of such investments may change quickly and unpredictably, which allows for the possibility that individuals fine-tune their investment strategies in response to changes in environmental conditions. In addition, our results suggest that anthropogenic environmental impacts may impose subtle energetic costs on individuals, which could contribute to population declines, especially in times of energy shortage.

Site fidelity of male Galápagos sea lions: a lifetime perspective

Knowledge about the distribution of resources can lead to the development of spatial preferences and long-term site fidelity. Individuals are expected to choose sites that best suit their needs. However, dominant individuals restrict movements of less competitive ones. Accordingly, one may expect spatial preferences to differ with regard to individual characteristics and to change over time. We investigated lifetime changes of site fidelity patterns with regard to reproductive success in male Galapagos sea lions (Zalophus wollebaeki). Showing a high degree of natal philopatry in the first 2 years of life, non-territorial males pass through a stage of fidelity to their natal colony where they develop preferences for areas outside prime breeding areas. Variation in the degree of spatial preferences was not associated with age or size, characteristics linked to an individual’s dominance status. For non-territorial males, roaming proved to be an adequate strategy to gain reproductive success. Only the most competitive males established territories in areas preferentially visited by females. They had a high probability to return to breeding areas where they successfully reproduced in previous seasons. Overall, the results reveal lifetime changes in site fidelity with regard to male status. The degree of site fidelity observed within the colony suggests familiarity and thus a high degree of tolerance among individuals using the same areas. This seems to facilitate attendance in the colony and thus the possibility to prospect for oestrus females.

Proximity loggers on amphibious mammals: a new method to study social relations in their terrestrial habitat

Amphibious mammal species alternate between foraging at sea and attendance on land. Due to thermoregulatory requirements, they often haul out during the night, making social interactions difficult to observe. We tested the suitability of UHF-proximity loggers for assessing social relationships among Galapagos sea lions Zalophus wollebaeki. To survive periods at sea, proximity loggers were embedded in epoxy. Automatic downloads to receiving stations rendered logger recovery unnecessary. Encounters were logged within a range of 10 m. Logs provided information about interacting individuals as well as time and duration of encounters. 'Received signal strength indicator' values correlated with distance, but were influenced by antenna angle and environmental factors. Laboratory tests and validation in the field demonstrated that the spa- tial resolution corresponded to 2 m. Data needed to be corrected, as single encounters were recorded as multiple logs and reciprocity of loggings between animals was not always achieved. Digital data correlated with observational data, but associations were 4 times more likely to be detected using proximity loggers. Data revealed that non-territorial males frequently associated for extended periods of time, especially during the night and even outside the study colony. The modified proximity logger system proves an excellent tool to determine social structure in situations where direct observation is limited.

Same size – same niche? Foraging niche separation between sympatric juvenile Galapagos sea lions and adult Galapagos fur seals

1. In vertebrates, patterns of resource utilization change throughout development according to age- and or size-specific abilities and requirements. Thus, interspecific competition affects different age classes differently. 2. Adults of sympatric species often show distinct foraging niche segregation, but juvenile resource use might overlap with adult competitors of similar body size. Resultant negative effects on juveniles can have important consequences for population dynamics, yet such interactions have received little attention in studies of mammalian communities. 3. Using GPS tracking devices, time-depth recorders and stable isotope data, we compared diving depth, activity time, trophic position and foraging habitat characteristics to investigate foraging niche overlap between similar-sized sympatric juvenile Galapagos sea lions (Zalophus wollebaeki) and adult Galapagos fur seals (Arctocephalus galapagoensis) and compared each group with much larger-bodied adult Galapagos sea lions. 4. We found little indication for direct competition but a complex pattern of foraging niche segregation: juvenile sea lions and adult fur seals dived to shallow depths at night, but foraged in different habitats with limited spatial overlap. Conversely, juvenile and adult sea lions employed different foraging patterns, but their foraging areas overlapped almost completely. 5. Consistency of foraging habitat characteristics between juvenile and adult sea lions suggests that avoidance of competition may be important in shaping foraging habitat utilization. Resultant specialization on a limited habitat could contribute to low sea lion numbers that contrast with high fur seal abundance. Our data suggest that exploitation by multiple predators within spatially restricted foraging ranges of juveniles might negatively impact juvenile foraging success and ultimately influence population dynamics.

Consumo oportunista de placenta en la Rata arrocera endémica de Galápagos (Cricetidae, Aegiolomys galapagoensis bauri) en la isla Santa Fe, Galápagos

La rata endémica Aegiolomys galapagoensis bauri de la Isla Santa Fe en el centro del Archipiélago de Galápagos fue observada consumiendo la placenta de un lobo marino (Zalophus wollebacki). Para un mamífero pequeño que consume principalmente materia vegetal, la posibilidad de aprovechar una fuente de proteínas concentradas debe ser una oportunidad valiosa.

Mama’s boy: sex differences in juvenile survival in a highly dimorphic large mammal, the Galapagos sea lion

In many mammals, early survival differs between the sexes, with males proving the more fragile sex ["Fragile male (FM) hypothesis"], especially in sexually dimorphic species where males are the larger sex. Male-biased allocation (MBA) by females may offset this difference. Here, we evaluate support for the FM and MBA hypotheses using a dataset on Galapagos sea lions (Zalophus wollebaeki). We statistically model sex-specific survival as it depends on body mass and environmental conditions (sea surface temperature, SST, a correlate of marine productivity) at three developmental stages, the perinatal phase (1st month), the main lactation period (1st year), and the weaning period (2nd year). Supporting the FM hypothesis, we found that, early in life (1st month), at equal birth mass, males survived less well than females. During the remainder of the first year of life, male survival was actually less sensitive to harsh environmental conditions than that of females, contradicting the FM hypothesis and supporting the MBA hypothesis. During the second year of life, only male survival suffered with high SSTs as predicted by the FM hypothesis. At each developmental stage, observed survival rates were almost equal for both sexes, suggesting that mothers buffer against the inherent fragility of male offspring through increased allocation, thereby masking the differences in survival prospects between the sexes.