Jaume Forcada | British Antarctic Survey (original) (raw)

Papers by Jaume Forcada

Published studies of wild vertebrate populations have almost universally reported positive associ... more Published studies of wild vertebrate populations have almost universally reported positive associations between genetic variation measured at microsatellite loci and fitness, creating the impression of ubiquity both in terms of the species and the traits involved. However, there is concern that this picture may be misleading because negative results frequently go unpublished. Here, we analyse the relationship between genotypic variation at nine highly variable microsatellite loci and neonatal fitness in 1070 Antarctic fur seal pups born at Bird Island, South Georgia. Despite our relatively large sample size, we find no significant association between three different measures of heterozygosity and two fitness traits, birth weight and survival. Furthermore, increasing genetic resolution by calculating parental relatedness also yields no association between genetic variation and fitness. Our findings are consistent with necropsy data showing that most pups die from starvation or trauma, conditions that are unlikely to be influenced strongly by genetic factors, particularly if the benefits of high heterozygosity are linked to immune-related genes.

Journal of Zoology, 2004

The dynamics of leopard seal Hydrurga leptonyx abundance and residency were investigated both wit... more The dynamics of leopard seal Hydrurga leptonyx abundance and residency were investigated both within and between years at Bird Island, South Georgia, using observations of tagged and untagged individuals. Seals belonged to an open population, and were sampled outside the breeding season and during their northward winter dispersal (April-October). Seals at Bird Island were highly transient (41-63%) with only a limited number of individuals making repeat visits across years. A minimum yearly survival for resident seals was estimated to be between 0.61 and 0.85. The physical environment played an important role in leopard seal abundance, with observations following an apparent 4-year periodicity consistent with the frequency and timing of the Antarctic Circumpolar Wave. Years of low sea-surface temperature (SST) and extensive pack-ice corresponded to a higher number of seals present, earlier arrivals, and longer periods of residency. Peak leopard seal arrivals at Bird Island corresponded to periods of low SST, which was consistent with the pagophilic nature of this species.

Ecology, 2005

Climate variability has strong effects on marine ecosystems, with repercussions that range in sca... more Climate variability has strong effects on marine ecosystems, with repercussions that range in scale from those that impact individuals to those that impact the entire food web. Climate-induced changes in the abundance of species in lower trophic levels can cascade up to apex predators by depressing vital rates. However, the characteristics and predictability of predator demographic responses remain largely unexplored. We investigated the detectability, limits, and nonlinearity of changes in Antarctic fur seal pup production at South Georgia over a 20-year period in response to environmental autocorrelation created by global climate perturbations; these were identified in time series of monthly averaged sea surface temperature (SST). Environmental autocorrelation at South Georgia was evident with frequent SST anomalies between 1990 and 1999, during a decade of warm background (time-averaged) conditions. SST anomalies were preceded by, and cross-correlated with, frequent El Niñ o-La Niñ a events between 1987 and 1998, which was also a decade of warm background conditions in the tropical Pacific Ocean. Nonlinear mixedeffects models indicated that positive anomalies at South Georgia explained extreme reductions in Antarctic fur seal pup production over 20 years of study. Simulated environmental time series suggested that the effect of anomalies on Antarctic fur seals was only detectable within a narrow range of positive SST, regardless of the distribution, variance, and autocorrelation structure in SST; this explained the observed nonlinearity in responses in pup production, which were observed only under persistent high SST levels. Such anomalies at South Georgia were likely associated with low availability of prey, largely krill, which affected Antarctic fur seal females over time scales longer than their breeding cycle. Reductions in Antarctic fur seal pup production could thus be predicted in advance by the detection of large-scale anomalies, which appeared to be driven by trends in global climate perturbation.

Proceedings of The Royal Society B: Biological Sciences, 2007

Determining how climate fluctuations affect ocean ecosystems requires an understanding of how bio... more Determining how climate fluctuations affect ocean ecosystems requires an understanding of how biological and physical processes interact across a wide range of scales. Here we examine the role of physical and biological processes in generating fluctuations in the ecosystem around South Georgia in the South Atlantic sector of the Southern Ocean. Anomalies in sea surface temperature (SST) in the South Pacific sector of the Southern Ocean have previously been shown to be generated through atmospheric teleconnections with El Niñ o Southern Oscillation (ENSO)-related processes. These SST anomalies are propagated via the Antarctic Circumpolar Current into the South Atlantic (on time scales of more than 1 year), where ENSO and Southern Annular Mode-related atmospheric processes have a direct influence on short (less than six months) time scales. We find that across the South Atlantic sector, these changes in SST, and related fluctuations in winter sea ice extent, affect the recruitment and dispersal of Antarctic krill. This oceanographically driven variation in krill population dynamics and abundance in turn affects the breeding success of seabird and marine mammal predators that depend on krill as food. Such propagating anomalies, mediated through physical and trophic interactions, are likely to be an important component of variation in ocean ecosystems and affect responses to longer term change. Population models derived on the basis of these oceanic fluctuations indicate that plausible rates of regional warming of 1 o C over the next 100 years could lead to more than a 95% reduction in the biomass and abundance of krill across the Scotia Sea by the end of the century.

Canadian Journal of Zoology-revue Canadienne De Zoologie, 2005

... We are also grateful to Phil Trathan, Tim Coulson, and Peter Rothery, who provided use-ful ad... more ... We are also grateful to Phil Trathan, Tim Coulson, and Peter Rothery, who provided use-ful advice throughout the development of this manuscript, and to Rodrigo Hucke-Gaete and one anonymous reviewer, who provided helpful comments and suggestions. References ...

Philosophical Transactions of The Royal Society B: Biological Sciences, 2007

The Southern Ocean is a major component within the global ocean and climate system and potentiall... more The Southern Ocean is a major component within the global ocean and climate system and potentially the location where the most rapid climate change is most likely to happen, particularly in the high-latitude polar regions. In these regions, even small temperature changes can potentially lead to major environmental perturbations. Climate change is likely to be regional and may be expressed in various ways, including alterations to climate and weather patterns across a variety of time-scales that include changes to the long interdecadal background signals such as the development of the El Niño-Southern Oscillation (ENSO). Oscillating climate signals such as ENSO potentially provide a unique opportunity to explore how biological communities respond to change. This approach is based on the premise that biological responses to shorter-term sub-decadal climate variability signals are potentially the best predictor of biological responses over longer time-scales. Around the Southern Ocean, marine predator populations show periodicity in breeding performance and productivity, with relationships with the environment driven by physical forcing from the ENSO region in the Pacific. Wherever examined, these relationships are congruent with mid-trophic-level processes that are also correlated with environmental variability. The short-term changes to ecosystem structure and function observed during ENSO events herald potential long-term changes that may ensue following regional climate change. For example, in the South Atlantic, failure of Antarctic krill recruitment will inevitably foreshadow recruitment failures in a range of higher trophic-level marine predators. Where predator species are not able to accommodate by switching to other prey species, population-level changes will follow. The Southern Ocean, though oceanographically interconnected, is not a single ecosystem and different areas are dominated by different food webs. Where species occupy different positions in different regional food webs, there is the potential to make predictions about future change scenarios.

Proceedings of The National Academy of Sciences, 2008

Global Change Biology, 2006

Climate warming and associated sea ice reductions in Antarctica have modified habitat conditions ... more Climate warming and associated sea ice reductions in Antarctica have modified habitat conditions for some species. These include the congeneric Adélie, chinstrap and gentoo penguins, which now demonstrate remarkable population responses to regional warming. However, inconsistencies in the direction of population changes between species at different study sites complicate the understanding of causal processes. Here, we show that at the South Orkney Islands where the three species breed sympatrically, the less ice-adapted gentoo penguins increased significantly in numbers over the last 26 years, whereas chinstrap and Adélie penguins both declined. These trends occurred in parallel with regional long-term warming and significant reduction in sea ice extent. Periodical warm events, with teleconnections to the tropical Pacific, caused cycles in sea ice leading to reduced prey biomass, and simultaneous interannual population decreases in the three penguin species. With the loss of sea ice, Adélie penguins were less buffered against the environment, their numbers fluctuated greatly and their population response was strong and linear. Chinstrap penguins, considered to be better adapted to ice-free conditions, were affected by discrete events of locally increased ice cover, but showed less variable, nonlinear responses to sea ice loss. Gentoo penguins were temporarily affected by negative anomalies in regional sea ice, but persistent sea ice reductions were likely to increase their available niche, which is likely to be substantially segregated from that of their more abundant congeners. Thus, the regional consequences of global climate perturbations on the sea ice phenology affect the marine ecosystem, with repercussions for penguin food supply and competition for resources. Ultimately, variability in penguin populations with warming reflects the local balance between penguin adaptation to ice conditions and trophic-mediated changes cascading from global climate forcing.

Global Change Biology, 2008

The consequences of warming for Antarctic long-lived organisms depend on their ability to survive... more The consequences of warming for Antarctic long-lived organisms depend on their ability to survive changing patterns of climate and environmental variation. Among birds and mammals of different Antarctic regions, including emperor penguins, snow petrels, southern fulmars, Antarctic fur seals and Weddell seals, we found strong support for selection of life history traits that reduce interannual variation in fitness. These species maximize fitness by keeping a low interannual variance in the survival of adults and in their propensity to breed annually, which are the vital rates that influence most the variability in population growth rate (λ). All these species have been able to buffer these rates against the effects of recent climate-driven habitat changes except for Antarctic fur seals, in the Southwest Atlantic. In this region of the Southern Ocean, the rapid increase in ecosystem fluctuation, associated with increasing climate variability observed since 1990, has limited and rendered less predictable the main fur seal food supply, Antarctic krill. This has increased the fitness costs of breeding for females, causing significant short-term changes in population structure through mortality and low breeding output. Changes occur now with a frequency higher than the mean female fur seal generation time, and therefore are likely to limit their adaptive response. Fur seals are more likely to rely on phenotypic plasticity to cope with short-term changes in order to maximize individual fitness. With more frequent extreme climatic events driving more frequent ecosystem fluctuation, the repercussions for life histories in many Antarctic birds and mammals are likely to increase, particularly at regional scales. In species with less flexible life histories that are more constrained by fluctuation in their critical habitats, like sea-ice, this may cause demographic changes, population compensation and changes in distribution, as already observed in penguin species living in the Antarctic Peninsula and adjacent islands.

Global Change Biology, 2009

Penguins are adapted to live in extreme environments, but they can be highly sensitive to climate... more Penguins are adapted to live in extreme environments, but they can be highly sensitive to climate change, which disrupts penguin life history strategies when it alters the weather, oceanography and critical habitats. For example, in the southwest Atlantic, the distributional range of the ice-obligate emperor and Adélie penguins has shifted poleward and contracted, while the ice-intolerant gentoo and chinstrap penguins have expanded their range southward. In the Southern Ocean, the El Niño-Southern Oscillation and the Southern Annular Mode are the main modes of climate variability that drive changes in the marine ecosystem, ultimately affecting penguins. The interaction between these modes is complex and changes over time, so that penguin responses to climate change are expected to vary accordingly, complicating our understanding of their future population processes. Penguins have long life spans, which slow microevolution, and which is unlikely to increase their tolerance to rapid warming. Therefore, in order that penguins may continue to exploit their transformed ecological niche and maintain their current distributional ranges, they must possess adequate phenotypic plasticity. However, past species-specific adaptations also constrain potential changes in phenology, and are unlikely to be adaptive for altered climatic conditions. Thus, the paleoecological record suggests that penguins are more likely to respond by dispersal rather than adaptation. Ecosystem changes are potentially most important at the borders of current geographic distributions, where penguins operate at the limits of their tolerance; species with low adaptability, particularly the ice-obligates, may therefore be more affected by their need to disperse in response to climate and may struggle to colonize new habitats. While future sea-ice contraction around Antarctica is likely to continue affecting the ice-obligate penguins, understanding the responses of the ice-intolerant penguins also depends on changes in climate mode periodicities and interactions, which to date remain difficult to reproduce in general circulation models.

Biological Conservation, 2008

Goudier Island is located in the Palmer Archipelago, to the west of the Antarctic Peninsula; it i... more Goudier Island is located in the Palmer Archipelago, to the west of the Antarctic Peninsula; it is one of the most frequently visited tourist sites in Antarctica. A number of gentoo penguin (Pygoscelis papua) breeding colonies are located on the island and these have been the focus of one of the longest running experiments to examine the impacts of tourist

Journal of Sea Research, 1998

Line-transect data from sighting surveys conducted in the western Mediterranean (in 1991) and the... more Line-transect data from sighting surveys conducted in the western Mediterranean (in 1991) and the Alboran Sea (in 1992) were analysed to estimate densities and numbers of striped and common dolphins in various areas of the western Mediterranean. Density of striped dolphins in the northwestern Mediterranean was estimated as 0.20 dolphins km 2 (CV D 0.24; 95% CI D 0.12 and 0.32) and was 41% higher than in the southwestern Mediterranean, where it was estimated as 0.12 dolphins km 2 (CV D 0.38; 95% CI D 0.05 and 0.25). The highest densities were observed in the Liguro-Provençal basin, with 0.24 dolphins km 2 (CV D 0.26; 95% CI D 0.14 and 0.40), and the Alboran Sea, with 0.20 dolphins km 2 (CV D 0.33; 95% CI D 0.10 and 0.36). These areas, and especially the Ligurian Sea, appear to be the most productive in terms of the food consumed by striped dolphins. Common dolphins were abundant only in the Alboran Sea with an estimated density of 0.16 dolphins km 2 (CV D 0.40; 95% CI D 0.08 and 0.35), scarce in the south Balearic area and almost absent in the northwestern Mediterranean. The magnitude of the dolphin by-catch in fishing operations in the Alboran Sea and other areas stresses the need for further assessment of densities and numbers, notably in the Alboran Sea and the North African Mediterranean waters.

Marine Ecology-progress Series, 1999

The status of the Mediterranean Monk Seal, Monachus monachus has been studied during a five-year ... more The status of the Mediterranean Monk Seal, Monachus monachus has been studied during a five-year period in the Foça Pilot Monk Seal Conservation Area between 1994 and 1998. The identification of individual seals occurring in the area, their spatial distribution and biology were studied using the results of direct cave surveys, land based seal observations and interviews. The population of Monk Seals was estimated to consist of 9 individuals, 6 of which were adult females and 3 were juveniles, one female, one male and one of unknown sex.. Islands off the Foça town provide suitable habitats and enables the survival of the species. A total of 11 caves were discovered, 2 of which were found to be used for reproduction.

Journal of The Marine Biological Association of The United Kingdom, 2007

Monachus monachus: an unusual pattern for a phocid This paper presents baseline information on ma... more Monachus monachus: an unusual pattern for a phocid This paper presents baseline information on maternal behaviour and lactation in the Mediterranean monk seal, with particular focus on the age at which pups are weaned. The study was conducted in the western Saharan population, the only surviving colony of the species. The first moult finished at a mean pup age of 72.3±17 d (N=17) and, in contrast to other taxonomically-related phocids, this process was not associated with weaning. Lactation lasted a mean of 119.4 d (N=9; range: 103-149 d) in the pups that could be monitored until full weaning had taken place. This period almost doubles the maximum lactation length reported in other phocid species. During the first week after birth the mother-pup bond was well developed and mothers always remained with their pups. The time invested in nursing (17%, SD: ±36) and in mother-pup interactions (14%, SD: ±32) was higher during this period than afterwards (8%, SD: ±23 and 4%, SD: ±19, respectively). After the first week, nursing continued but mothers started to leave their pups in order to feed at sea. Weaning occurred gradually. Already since birth, pups were active and mobile, and swam frequently before moulting or weaning occurred. Fostering and milk stealing were common patterns of behaviour for both lactating females and pups. In 26.6% of the suckling episodes observed in mother-pup pairs of known identity, pups suckled from females other than their mothers. Some females nursed more than one pup, at least occasionally, and in some cases a pup was fostered long-term by an alien female. The Mediterranean monk seal exhibits maternalcare characteristics that are more like otarids than phocids. This observation contradicts previous proposals that a short lactation period is a phylogenetic characteristic of phocids. Several of the unusual maternal traits observed may be favoured by year-round access to abundant food supply, availability of breeding sites, and mild climatic conditions. This information should be taken into account when designing conservation strategies for the species and, very particularly, in the implementation of pup rehabilitation programmes.

Marine Mammal Science, 2000

The use of photo-identification and its reliability in capture-recapture studies of Mediterranean... more The use of photo-identification and its reliability in capture-recapture studies of Mediterranean monk seals were assessed using slides collected in the colony at Cap Blanc, western Sahara, from 1993 to 1996. Five tests indicated that researchers involved in photo-identification were proficient in matching slides of identified seals, consistent in classifying the side of the seal shown in slides and in assigning the morphological stage of the seal, and that changes of markings over a period of three years were insufficient to affect matching success. The certainty of identifying a seal was not dependent on the number of slides used but on distinctiveness of the markings and the quality of the slides taken. Capture-recapture abundance estimates were biased upwards when including poor quality slides. The exclusive use of excellent- and good quality slides provided the best estimates. The proportion of distinctive seals varied between morphological stages and was significantly lower in juveniles. When including the identification histories of juveniles, the heterogeneity of capture probabilities was higher. Therefore, abundance estimates were less biased when all juveniles were considered as non-distinctive seals. Reliable abundance estimates required a balance between duration of capture occasions and time interval between these.

Journal of Zoology, 1996

A sightings survey was conducted in the summer of 1991 in the western Mediterranean to describe t... more A sightings survey was conducted in the summer of 1991 in the western Mediterranean to describe the distribution of fin whales and to estimate their absolute abundance during the summer feeding season. Fin whales were only found in the Liguro-Provençal basin, in deep waters (mean depth 2,360, S.E.: 46.8 m), beyond the continental shelf. The highest densities of fin whales were found in relatively cool waters (mean = 23.9°C), and water in areas in which fin whales were found was significantly cooler than that in areas without fin whales. Patterns of distribution of whales, with remarkably high densities in the feeding grounds, and composition of schools, which were found to be smaller than in other regions, suggest that food resources for the species in the north-western Mediterranean feeding grounds occur at higher densities but are more patchily distributed than in other fin whale feeding grounds so far studied. Abundance was estimated as 3,583 individuals (S.E.: 967; 95% CI: 2,130-6,027).

Marine Ecology-progress Series, 2004

Line-transect estimators were developed to assess abundance of coastal dolphins Tursiops truncatu... more Line-transect estimators were developed to assess abundance of coastal dolphins Tursiops truncatus and Stenella coeruleoalba encountered in low densities during aerial sighting surveys. The analysis improved on conventional approaches by objectively combining data from different species, survey areas and other covariates affecting dolphin detectability. Model selection and multimodel inference allowed robust estimates of precision in accounting for covariate selection uncertainty. These methods were used to estimate bottlenose dolphin abundance in NE Mediterranean waters that included a putative subpopulation in the Balearic Islands. Total abundance was estimated as 7654 (coefficient of variation, CV = 0.47; 95% CI = 1608 to 15 766) and the abundance in inshore waters of the Balearic Islands varied from 727 (CV = 0.47; 95% CI = 149 to 1481) dolphins in spring 2002 to 1333 (CV = 0.44; 95% CI = 419 to 2617) dolphins in autumn 2002, with an average estimate of 1030 (CV = 0.35; 95% CI = 415 to 1849). The results do not support an exclusively coastal Balearic Island subpopulation, but they strongly indicate that the islands contain critical habitats required for the conservation of the species. Given the observed decline of the species during the last few decades, conservation-oriented management should focus on reducing or eliminating adverse fishing interactions while key areas are protected from encroachment produced by human development.

Marine Ecology-progress Series, 2004

Exposure to organochlorines induces retinoid deficiency in mammals; hence, retinoids are potentia... more Exposure to organochlorines induces retinoid deficiency in mammals; hence, retinoids are potential biomarkers of the impact of these pollutants. Appropriate target tissues to monitor retinoids in cetaceans have not been properly identified because of a lack of information on the contribution of each tissue to total body retinoids. Therefore, we have addressed this issue by studying the contribution of the main body tissues to retinoids in 21 common dolphins obtained from incidental catches and in apparent good health and nutritive condition. Although concentrations in the liver were highest, those in blubber were also high and accounted for 43% of the total retinoid load of the compartments examined. As blubber can be obtained using non-invasive biopsy techniques, this tissue is proposed as a reliable indicator of retinoid status in cetaceans. However, blubber topographical variation in structure and composition requires standardization of sampling sites. Retinoid concentrations did not differ significantly between sexes or with body size for any of the tissues, but the lipid content of blubber strongly influenced these concentrations. Biopsies from healthy, freeranging individuals are preferred to samples from stranded animals. Further research on the influence of factors (age, sex, reproductive condition, diet) that potentially affect retinoid levels is required to implement the use of retinoids as biomarkers of pollutant exposure in cetaceans.

Marine Mammal Science, 1994

A survey was conducted in the summer of 1991 in the western Mediterranean to map the distribution... more A survey was conducted in the summer of 1991 in the western Mediterranean to map the distribution and to estimate the number of striped dolphins surviving the 1990 outbreak of the morbillivirus epizootic. The highest densities of striped dolphins were found in the Alboran Sea and the Ligurian Sea. Total numbers were estimated at 117,880 (SE: 38,962; 95% CI: 68,379–214,800). The mortality resulting from the epizootic is unknown but may have been severe because the mean size of dolphin schools observed during the epizootic outbreak was significantly smaller than that before the event. A return to larger school sizes after the 1990 outbreak is not evidence of a recovery but can be viewed as a regrouping of individuals into schools of a preferred size. Calves were observed in 24.6% of the schools. They were not seen in schools of ten or less but were present in about half the schools of 30 or more dolphins, a result which suggests segregation similar to that seen in populations of this species elsewhere in the world.

Published studies of wild vertebrate populations have almost universally reported positive associ... more Published studies of wild vertebrate populations have almost universally reported positive associations between genetic variation measured at microsatellite loci and fitness, creating the impression of ubiquity both in terms of the species and the traits involved. However, there is concern that this picture may be misleading because negative results frequently go unpublished. Here, we analyse the relationship between genotypic variation at nine highly variable microsatellite loci and neonatal fitness in 1070 Antarctic fur seal pups born at Bird Island, South Georgia. Despite our relatively large sample size, we find no significant association between three different measures of heterozygosity and two fitness traits, birth weight and survival. Furthermore, increasing genetic resolution by calculating parental relatedness also yields no association between genetic variation and fitness. Our findings are consistent with necropsy data showing that most pups die from starvation or trauma, conditions that are unlikely to be influenced strongly by genetic factors, particularly if the benefits of high heterozygosity are linked to immune-related genes.

Journal of Zoology, 2004

The dynamics of leopard seal Hydrurga leptonyx abundance and residency were investigated both wit... more The dynamics of leopard seal Hydrurga leptonyx abundance and residency were investigated both within and between years at Bird Island, South Georgia, using observations of tagged and untagged individuals. Seals belonged to an open population, and were sampled outside the breeding season and during their northward winter dispersal (April-October). Seals at Bird Island were highly transient (41-63%) with only a limited number of individuals making repeat visits across years. A minimum yearly survival for resident seals was estimated to be between 0.61 and 0.85. The physical environment played an important role in leopard seal abundance, with observations following an apparent 4-year periodicity consistent with the frequency and timing of the Antarctic Circumpolar Wave. Years of low sea-surface temperature (SST) and extensive pack-ice corresponded to a higher number of seals present, earlier arrivals, and longer periods of residency. Peak leopard seal arrivals at Bird Island corresponded to periods of low SST, which was consistent with the pagophilic nature of this species.

Ecology, 2005

Climate variability has strong effects on marine ecosystems, with repercussions that range in sca... more Climate variability has strong effects on marine ecosystems, with repercussions that range in scale from those that impact individuals to those that impact the entire food web. Climate-induced changes in the abundance of species in lower trophic levels can cascade up to apex predators by depressing vital rates. However, the characteristics and predictability of predator demographic responses remain largely unexplored. We investigated the detectability, limits, and nonlinearity of changes in Antarctic fur seal pup production at South Georgia over a 20-year period in response to environmental autocorrelation created by global climate perturbations; these were identified in time series of monthly averaged sea surface temperature (SST). Environmental autocorrelation at South Georgia was evident with frequent SST anomalies between 1990 and 1999, during a decade of warm background (time-averaged) conditions. SST anomalies were preceded by, and cross-correlated with, frequent El Niñ o-La Niñ a events between 1987 and 1998, which was also a decade of warm background conditions in the tropical Pacific Ocean. Nonlinear mixedeffects models indicated that positive anomalies at South Georgia explained extreme reductions in Antarctic fur seal pup production over 20 years of study. Simulated environmental time series suggested that the effect of anomalies on Antarctic fur seals was only detectable within a narrow range of positive SST, regardless of the distribution, variance, and autocorrelation structure in SST; this explained the observed nonlinearity in responses in pup production, which were observed only under persistent high SST levels. Such anomalies at South Georgia were likely associated with low availability of prey, largely krill, which affected Antarctic fur seal females over time scales longer than their breeding cycle. Reductions in Antarctic fur seal pup production could thus be predicted in advance by the detection of large-scale anomalies, which appeared to be driven by trends in global climate perturbation.

Proceedings of The Royal Society B: Biological Sciences, 2007

Determining how climate fluctuations affect ocean ecosystems requires an understanding of how bio... more Determining how climate fluctuations affect ocean ecosystems requires an understanding of how biological and physical processes interact across a wide range of scales. Here we examine the role of physical and biological processes in generating fluctuations in the ecosystem around South Georgia in the South Atlantic sector of the Southern Ocean. Anomalies in sea surface temperature (SST) in the South Pacific sector of the Southern Ocean have previously been shown to be generated through atmospheric teleconnections with El Niñ o Southern Oscillation (ENSO)-related processes. These SST anomalies are propagated via the Antarctic Circumpolar Current into the South Atlantic (on time scales of more than 1 year), where ENSO and Southern Annular Mode-related atmospheric processes have a direct influence on short (less than six months) time scales. We find that across the South Atlantic sector, these changes in SST, and related fluctuations in winter sea ice extent, affect the recruitment and dispersal of Antarctic krill. This oceanographically driven variation in krill population dynamics and abundance in turn affects the breeding success of seabird and marine mammal predators that depend on krill as food. Such propagating anomalies, mediated through physical and trophic interactions, are likely to be an important component of variation in ocean ecosystems and affect responses to longer term change. Population models derived on the basis of these oceanic fluctuations indicate that plausible rates of regional warming of 1 o C over the next 100 years could lead to more than a 95% reduction in the biomass and abundance of krill across the Scotia Sea by the end of the century.

Canadian Journal of Zoology-revue Canadienne De Zoologie, 2005

... We are also grateful to Phil Trathan, Tim Coulson, and Peter Rothery, who provided use-ful ad... more ... We are also grateful to Phil Trathan, Tim Coulson, and Peter Rothery, who provided use-ful advice throughout the development of this manuscript, and to Rodrigo Hucke-Gaete and one anonymous reviewer, who provided helpful comments and suggestions. References ...

Philosophical Transactions of The Royal Society B: Biological Sciences, 2007

The Southern Ocean is a major component within the global ocean and climate system and potentiall... more The Southern Ocean is a major component within the global ocean and climate system and potentially the location where the most rapid climate change is most likely to happen, particularly in the high-latitude polar regions. In these regions, even small temperature changes can potentially lead to major environmental perturbations. Climate change is likely to be regional and may be expressed in various ways, including alterations to climate and weather patterns across a variety of time-scales that include changes to the long interdecadal background signals such as the development of the El Niño-Southern Oscillation (ENSO). Oscillating climate signals such as ENSO potentially provide a unique opportunity to explore how biological communities respond to change. This approach is based on the premise that biological responses to shorter-term sub-decadal climate variability signals are potentially the best predictor of biological responses over longer time-scales. Around the Southern Ocean, marine predator populations show periodicity in breeding performance and productivity, with relationships with the environment driven by physical forcing from the ENSO region in the Pacific. Wherever examined, these relationships are congruent with mid-trophic-level processes that are also correlated with environmental variability. The short-term changes to ecosystem structure and function observed during ENSO events herald potential long-term changes that may ensue following regional climate change. For example, in the South Atlantic, failure of Antarctic krill recruitment will inevitably foreshadow recruitment failures in a range of higher trophic-level marine predators. Where predator species are not able to accommodate by switching to other prey species, population-level changes will follow. The Southern Ocean, though oceanographically interconnected, is not a single ecosystem and different areas are dominated by different food webs. Where species occupy different positions in different regional food webs, there is the potential to make predictions about future change scenarios.

Proceedings of The National Academy of Sciences, 2008

Global Change Biology, 2006

Climate warming and associated sea ice reductions in Antarctica have modified habitat conditions ... more Climate warming and associated sea ice reductions in Antarctica have modified habitat conditions for some species. These include the congeneric Adélie, chinstrap and gentoo penguins, which now demonstrate remarkable population responses to regional warming. However, inconsistencies in the direction of population changes between species at different study sites complicate the understanding of causal processes. Here, we show that at the South Orkney Islands where the three species breed sympatrically, the less ice-adapted gentoo penguins increased significantly in numbers over the last 26 years, whereas chinstrap and Adélie penguins both declined. These trends occurred in parallel with regional long-term warming and significant reduction in sea ice extent. Periodical warm events, with teleconnections to the tropical Pacific, caused cycles in sea ice leading to reduced prey biomass, and simultaneous interannual population decreases in the three penguin species. With the loss of sea ice, Adélie penguins were less buffered against the environment, their numbers fluctuated greatly and their population response was strong and linear. Chinstrap penguins, considered to be better adapted to ice-free conditions, were affected by discrete events of locally increased ice cover, but showed less variable, nonlinear responses to sea ice loss. Gentoo penguins were temporarily affected by negative anomalies in regional sea ice, but persistent sea ice reductions were likely to increase their available niche, which is likely to be substantially segregated from that of their more abundant congeners. Thus, the regional consequences of global climate perturbations on the sea ice phenology affect the marine ecosystem, with repercussions for penguin food supply and competition for resources. Ultimately, variability in penguin populations with warming reflects the local balance between penguin adaptation to ice conditions and trophic-mediated changes cascading from global climate forcing.

Global Change Biology, 2008

The consequences of warming for Antarctic long-lived organisms depend on their ability to survive... more The consequences of warming for Antarctic long-lived organisms depend on their ability to survive changing patterns of climate and environmental variation. Among birds and mammals of different Antarctic regions, including emperor penguins, snow petrels, southern fulmars, Antarctic fur seals and Weddell seals, we found strong support for selection of life history traits that reduce interannual variation in fitness. These species maximize fitness by keeping a low interannual variance in the survival of adults and in their propensity to breed annually, which are the vital rates that influence most the variability in population growth rate (λ). All these species have been able to buffer these rates against the effects of recent climate-driven habitat changes except for Antarctic fur seals, in the Southwest Atlantic. In this region of the Southern Ocean, the rapid increase in ecosystem fluctuation, associated with increasing climate variability observed since 1990, has limited and rendered less predictable the main fur seal food supply, Antarctic krill. This has increased the fitness costs of breeding for females, causing significant short-term changes in population structure through mortality and low breeding output. Changes occur now with a frequency higher than the mean female fur seal generation time, and therefore are likely to limit their adaptive response. Fur seals are more likely to rely on phenotypic plasticity to cope with short-term changes in order to maximize individual fitness. With more frequent extreme climatic events driving more frequent ecosystem fluctuation, the repercussions for life histories in many Antarctic birds and mammals are likely to increase, particularly at regional scales. In species with less flexible life histories that are more constrained by fluctuation in their critical habitats, like sea-ice, this may cause demographic changes, population compensation and changes in distribution, as already observed in penguin species living in the Antarctic Peninsula and adjacent islands.

Global Change Biology, 2009

Penguins are adapted to live in extreme environments, but they can be highly sensitive to climate... more Penguins are adapted to live in extreme environments, but they can be highly sensitive to climate change, which disrupts penguin life history strategies when it alters the weather, oceanography and critical habitats. For example, in the southwest Atlantic, the distributional range of the ice-obligate emperor and Adélie penguins has shifted poleward and contracted, while the ice-intolerant gentoo and chinstrap penguins have expanded their range southward. In the Southern Ocean, the El Niño-Southern Oscillation and the Southern Annular Mode are the main modes of climate variability that drive changes in the marine ecosystem, ultimately affecting penguins. The interaction between these modes is complex and changes over time, so that penguin responses to climate change are expected to vary accordingly, complicating our understanding of their future population processes. Penguins have long life spans, which slow microevolution, and which is unlikely to increase their tolerance to rapid warming. Therefore, in order that penguins may continue to exploit their transformed ecological niche and maintain their current distributional ranges, they must possess adequate phenotypic plasticity. However, past species-specific adaptations also constrain potential changes in phenology, and are unlikely to be adaptive for altered climatic conditions. Thus, the paleoecological record suggests that penguins are more likely to respond by dispersal rather than adaptation. Ecosystem changes are potentially most important at the borders of current geographic distributions, where penguins operate at the limits of their tolerance; species with low adaptability, particularly the ice-obligates, may therefore be more affected by their need to disperse in response to climate and may struggle to colonize new habitats. While future sea-ice contraction around Antarctica is likely to continue affecting the ice-obligate penguins, understanding the responses of the ice-intolerant penguins also depends on changes in climate mode periodicities and interactions, which to date remain difficult to reproduce in general circulation models.

Biological Conservation, 2008

Goudier Island is located in the Palmer Archipelago, to the west of the Antarctic Peninsula; it i... more Goudier Island is located in the Palmer Archipelago, to the west of the Antarctic Peninsula; it is one of the most frequently visited tourist sites in Antarctica. A number of gentoo penguin (Pygoscelis papua) breeding colonies are located on the island and these have been the focus of one of the longest running experiments to examine the impacts of tourist

Journal of Sea Research, 1998

Line-transect data from sighting surveys conducted in the western Mediterranean (in 1991) and the... more Line-transect data from sighting surveys conducted in the western Mediterranean (in 1991) and the Alboran Sea (in 1992) were analysed to estimate densities and numbers of striped and common dolphins in various areas of the western Mediterranean. Density of striped dolphins in the northwestern Mediterranean was estimated as 0.20 dolphins km 2 (CV D 0.24; 95% CI D 0.12 and 0.32) and was 41% higher than in the southwestern Mediterranean, where it was estimated as 0.12 dolphins km 2 (CV D 0.38; 95% CI D 0.05 and 0.25). The highest densities were observed in the Liguro-Provençal basin, with 0.24 dolphins km 2 (CV D 0.26; 95% CI D 0.14 and 0.40), and the Alboran Sea, with 0.20 dolphins km 2 (CV D 0.33; 95% CI D 0.10 and 0.36). These areas, and especially the Ligurian Sea, appear to be the most productive in terms of the food consumed by striped dolphins. Common dolphins were abundant only in the Alboran Sea with an estimated density of 0.16 dolphins km 2 (CV D 0.40; 95% CI D 0.08 and 0.35), scarce in the south Balearic area and almost absent in the northwestern Mediterranean. The magnitude of the dolphin by-catch in fishing operations in the Alboran Sea and other areas stresses the need for further assessment of densities and numbers, notably in the Alboran Sea and the North African Mediterranean waters.

Marine Ecology-progress Series, 1999

The status of the Mediterranean Monk Seal, Monachus monachus has been studied during a five-year ... more The status of the Mediterranean Monk Seal, Monachus monachus has been studied during a five-year period in the Foça Pilot Monk Seal Conservation Area between 1994 and 1998. The identification of individual seals occurring in the area, their spatial distribution and biology were studied using the results of direct cave surveys, land based seal observations and interviews. The population of Monk Seals was estimated to consist of 9 individuals, 6 of which were adult females and 3 were juveniles, one female, one male and one of unknown sex.. Islands off the Foça town provide suitable habitats and enables the survival of the species. A total of 11 caves were discovered, 2 of which were found to be used for reproduction.

Journal of The Marine Biological Association of The United Kingdom, 2007

Monachus monachus: an unusual pattern for a phocid This paper presents baseline information on ma... more Monachus monachus: an unusual pattern for a phocid This paper presents baseline information on maternal behaviour and lactation in the Mediterranean monk seal, with particular focus on the age at which pups are weaned. The study was conducted in the western Saharan population, the only surviving colony of the species. The first moult finished at a mean pup age of 72.3±17 d (N=17) and, in contrast to other taxonomically-related phocids, this process was not associated with weaning. Lactation lasted a mean of 119.4 d (N=9; range: 103-149 d) in the pups that could be monitored until full weaning had taken place. This period almost doubles the maximum lactation length reported in other phocid species. During the first week after birth the mother-pup bond was well developed and mothers always remained with their pups. The time invested in nursing (17%, SD: ±36) and in mother-pup interactions (14%, SD: ±32) was higher during this period than afterwards (8%, SD: ±23 and 4%, SD: ±19, respectively). After the first week, nursing continued but mothers started to leave their pups in order to feed at sea. Weaning occurred gradually. Already since birth, pups were active and mobile, and swam frequently before moulting or weaning occurred. Fostering and milk stealing were common patterns of behaviour for both lactating females and pups. In 26.6% of the suckling episodes observed in mother-pup pairs of known identity, pups suckled from females other than their mothers. Some females nursed more than one pup, at least occasionally, and in some cases a pup was fostered long-term by an alien female. The Mediterranean monk seal exhibits maternalcare characteristics that are more like otarids than phocids. This observation contradicts previous proposals that a short lactation period is a phylogenetic characteristic of phocids. Several of the unusual maternal traits observed may be favoured by year-round access to abundant food supply, availability of breeding sites, and mild climatic conditions. This information should be taken into account when designing conservation strategies for the species and, very particularly, in the implementation of pup rehabilitation programmes.

Marine Mammal Science, 2000

The use of photo-identification and its reliability in capture-recapture studies of Mediterranean... more The use of photo-identification and its reliability in capture-recapture studies of Mediterranean monk seals were assessed using slides collected in the colony at Cap Blanc, western Sahara, from 1993 to 1996. Five tests indicated that researchers involved in photo-identification were proficient in matching slides of identified seals, consistent in classifying the side of the seal shown in slides and in assigning the morphological stage of the seal, and that changes of markings over a period of three years were insufficient to affect matching success. The certainty of identifying a seal was not dependent on the number of slides used but on distinctiveness of the markings and the quality of the slides taken. Capture-recapture abundance estimates were biased upwards when including poor quality slides. The exclusive use of excellent- and good quality slides provided the best estimates. The proportion of distinctive seals varied between morphological stages and was significantly lower in juveniles. When including the identification histories of juveniles, the heterogeneity of capture probabilities was higher. Therefore, abundance estimates were less biased when all juveniles were considered as non-distinctive seals. Reliable abundance estimates required a balance between duration of capture occasions and time interval between these.

Journal of Zoology, 1996

A sightings survey was conducted in the summer of 1991 in the western Mediterranean to describe t... more A sightings survey was conducted in the summer of 1991 in the western Mediterranean to describe the distribution of fin whales and to estimate their absolute abundance during the summer feeding season. Fin whales were only found in the Liguro-Provençal basin, in deep waters (mean depth 2,360, S.E.: 46.8 m), beyond the continental shelf. The highest densities of fin whales were found in relatively cool waters (mean = 23.9°C), and water in areas in which fin whales were found was significantly cooler than that in areas without fin whales. Patterns of distribution of whales, with remarkably high densities in the feeding grounds, and composition of schools, which were found to be smaller than in other regions, suggest that food resources for the species in the north-western Mediterranean feeding grounds occur at higher densities but are more patchily distributed than in other fin whale feeding grounds so far studied. Abundance was estimated as 3,583 individuals (S.E.: 967; 95% CI: 2,130-6,027).

Marine Ecology-progress Series, 2004

Line-transect estimators were developed to assess abundance of coastal dolphins Tursiops truncatu... more Line-transect estimators were developed to assess abundance of coastal dolphins Tursiops truncatus and Stenella coeruleoalba encountered in low densities during aerial sighting surveys. The analysis improved on conventional approaches by objectively combining data from different species, survey areas and other covariates affecting dolphin detectability. Model selection and multimodel inference allowed robust estimates of precision in accounting for covariate selection uncertainty. These methods were used to estimate bottlenose dolphin abundance in NE Mediterranean waters that included a putative subpopulation in the Balearic Islands. Total abundance was estimated as 7654 (coefficient of variation, CV = 0.47; 95% CI = 1608 to 15 766) and the abundance in inshore waters of the Balearic Islands varied from 727 (CV = 0.47; 95% CI = 149 to 1481) dolphins in spring 2002 to 1333 (CV = 0.44; 95% CI = 419 to 2617) dolphins in autumn 2002, with an average estimate of 1030 (CV = 0.35; 95% CI = 415 to 1849). The results do not support an exclusively coastal Balearic Island subpopulation, but they strongly indicate that the islands contain critical habitats required for the conservation of the species. Given the observed decline of the species during the last few decades, conservation-oriented management should focus on reducing or eliminating adverse fishing interactions while key areas are protected from encroachment produced by human development.

Marine Ecology-progress Series, 2004

Exposure to organochlorines induces retinoid deficiency in mammals; hence, retinoids are potentia... more Exposure to organochlorines induces retinoid deficiency in mammals; hence, retinoids are potential biomarkers of the impact of these pollutants. Appropriate target tissues to monitor retinoids in cetaceans have not been properly identified because of a lack of information on the contribution of each tissue to total body retinoids. Therefore, we have addressed this issue by studying the contribution of the main body tissues to retinoids in 21 common dolphins obtained from incidental catches and in apparent good health and nutritive condition. Although concentrations in the liver were highest, those in blubber were also high and accounted for 43% of the total retinoid load of the compartments examined. As blubber can be obtained using non-invasive biopsy techniques, this tissue is proposed as a reliable indicator of retinoid status in cetaceans. However, blubber topographical variation in structure and composition requires standardization of sampling sites. Retinoid concentrations did not differ significantly between sexes or with body size for any of the tissues, but the lipid content of blubber strongly influenced these concentrations. Biopsies from healthy, freeranging individuals are preferred to samples from stranded animals. Further research on the influence of factors (age, sex, reproductive condition, diet) that potentially affect retinoid levels is required to implement the use of retinoids as biomarkers of pollutant exposure in cetaceans.

Marine Mammal Science, 1994

A survey was conducted in the summer of 1991 in the western Mediterranean to map the distribution... more A survey was conducted in the summer of 1991 in the western Mediterranean to map the distribution and to estimate the number of striped dolphins surviving the 1990 outbreak of the morbillivirus epizootic. The highest densities of striped dolphins were found in the Alboran Sea and the Ligurian Sea. Total numbers were estimated at 117,880 (SE: 38,962; 95% CI: 68,379–214,800). The mortality resulting from the epizootic is unknown but may have been severe because the mean size of dolphin schools observed during the epizootic outbreak was significantly smaller than that before the event. A return to larger school sizes after the 1990 outbreak is not evidence of a recovery but can be viewed as a regrouping of individuals into schools of a preferred size. Calves were observed in 24.6% of the schools. They were not seen in schools of ten or less but were present in about half the schools of 30 or more dolphins, a result which suggests segregation similar to that seen in populations of this species elsewhere in the world.