Tom Hart

Spatial and temporal extinction dynamics in a freshwater cetacean

Geographical range contraction is a fundamental ecological characteristic of species population decline, but relatively little investigation has been conducted into general trends in the dynamic properties of range collapse. The Yangtze River dolphin or baiji (Lipotes vexillifer), probably the first large mammal species to have become extinct in over 50 years, was believed to have experienced major range collapse during its decline through progressive large-scale range contraction and fragmentation. This range-collapse model is challenged by a new dataset of 406 baiji last-sighting records collected from across the baijis historical range during an interview survey of Yangtze fishing communities. Although baiji regional abundance may have varied across its range, analyses of the extensive new sighting series provide comprehensive evidence that baiji population decline was not associated with any major contraction in geographical range across the middle–lower Yangtze drainage, even in the decade immediately before probable global extinction of the species. Extinction risk in baiji was therefore seemingly not related to evidence of range collapse. Baiji apparently underwent large-scale periodic and seasonal movements across their range, and we propose that range contraction and fragmentation may not be general biogeographic characteristics for declining populations of mobile species in connected landscapes.

A reversal of fortunes: climate change ‘winners’ and ‘losers’ in Antarctic Peninsula penguins

Climate change is a major threat to global biodiversity. Antarctic ecosystems are no exception. Investigating past species responses to climatic events can distinguish natural from anthropogenic impacts. Climate change produces ‘winners’, species that benefit from these events and ‘losers’, species that decline or become extinct. Using molecular techniques, we assess the demographic history and population structure of Pygoscelis penguins in the Scotia Arc related to climate warming after the Last Glacial Maximum (LGM). All three pygoscelid penguins responded positively to post-LGM warming by expanding from glacial refugia, with those breeding at higher latitudes expanding most. Northern (Pygoscelis papua papua) and Southern (Pygoscelis papua ellsworthii) gentoo sub-species likely diverged during the LGM. Comparing historical responses with the literature on current trends, we see Southern gentoo penguins are responding to current warming as they did during post-LGM warming, expanding their range southwards. Conversely, Adélie and chinstrap penguins are experiencing a ‘reversal of fortunes’ as they are now declining in the Antarctic Peninsula, the opposite of their response to post-LGM warming. This suggests current climate warming has decoupled historic population responses in the Antarctic Peninsula, favoring generalist gentoo penguins as climate change ‘winners’, while Adélie and chinstrap penguins have become climate change ‘losers’.

Too much of a good thing: sea ice extent may have forced emperor penguins into refugia during the last glacial maximum

The relationship between population structure and demographic history is critical to understanding microevolution and for predicting the resilience of species to environmental change. Using mitochondrial DNA from extant colonies and radiocarbon-dated subfossils, we present the first microevolutionary analysis of emperor penguins (Aptenodytes forsteri) and show their population trends throughout the last glacial maximum (LGM, 19.5-16 kya) and during the subsequent period of warming and sea ice retreat. We found evidence for three mitochondrial clades within emperor penguins, suggesting that they were isolated within three glacial refugia during the LGM. One of these clades has remained largely isolated within the Ross Sea, while the two other clades have intermixed around the coast of Antarctica from Adélie Land to the Weddell Sea. The differentiation of the Ross Sea population has been preserved despite rapid population growth and opportunities for migration. Low effective population sizes during the LGM, followed by a rapid expansion around the beginning of the Holocene, suggest that an optimum set of sea ice conditions exist for emperor penguins, corresponding to available foraging area.

Time series analysis of biologging data: autocorrelation reveals periodicity of diving behaviour in macaroni penguins

The nature of how behaviour at one time step influences the next is of great interest to behavioural ecologists, but rarely used for comparisons between animals. Time depth recorders (TDR) and other archival tags have been widely used to infer patterns of diving and foraging. However, while we can extract variables that describe individual dives, how runs of dives may indicate behaviours and how one dive influences the next are not fully understood. Treating TDR data as time series, we examined patterns of autocorrelation to investigate structure in the timing of behaviour. We fitted an oscillating best-fit curve to the autocorrelation and used the parameters of this curve to investigate differences in foraging strategy of 129 macaroni penguins, Eudyptes chrysolophus, of both sexes. We found interannual differences in autocorrelation parameters as well as differences between reproductive stages. In contrast to other studies of macaroni penguin diving based on depth analysis, we found no differences between the sexes. We mimicked changes in the various parameters by simulation of dive profiles, and used these to infer biological meaning from the parameters. As this technique makes very few assumptions about how to identify a dive or cluster of dives, we suggest that it is a useful first characterization of diving or cyclical behaviour in a wide range of animals.

Habitat-mediated dive behavior in free-ranging grey seals.

Understanding the links between foraging behaviour and habitat use of key species is essential to addressing fundamental questions about trophic interactions and ecosystem functioning. Eight female grey seals (Halichoerus grypus) were equipped with time-depth recorders linked to Fastloc GPS tags following the annual moult in southwest Ireland. Individual dives were coupled with environmental correlates to investigate the habitat use and dive behaviour of free-ranging seals. Dives were characterised as either pelagic, benthic, or shallow (where errors in location and charted water depth made differentiating between pelagic and benthic dives unreliable). Sixty-nine percent of dives occurring in water >50 m were benthic. Pelagic dives were more common at night than during the day. Seals performed more pelagic dives over fine sediments (mud/sand), and more benthic dives when foraging over more three-dimensionally complex rock substrates. We used Markov chain analysis to determine the probability of transiting between dive states. A low probability of repeat pelagic dives suggests that pelagic prey were encountered en route to the seabed. This approach could be applied to make more accurate predictions of habitat use in data-poor areas, and investigate contentious issues such as resource overlap and competition between top predators and fisheries, essential for the effective conservation of these key marine species.

Testing and Improving the Accuracy of Discriminant Function Tests: A Comparison between Morphometric and Molecular Sexing in Macaroni Penguins

Abstract. Dimorphism in various morphological traits is widely used to sex birds. However, when a threshold is set and there is overlap in the distribution of traits between sexes, there will be an error rate. Macaroni Penguins show limited sexual dimorphism in size, with the distributions of most phenotypic traits overlapping between sexes, making using morphological characters for sexing difficult, although bill depth is used in one discrimination test. Two existing DNA-based PCR tests that use different primer pairs (P2/P8 and 2550/2781 ) were adapted for use with fluorescently labelled primers to enable the size of fragments to be accurately measured via genotyping protocols and sex to be determined. The two molecular tests were applied to the same samples to estimate the error rate within and between tests. Whilst both primer pairs reproducibly identified the sex of the penguins under study with no inconsistencies between tests, the P2/P8 primer pair had a higher success rate of amplification. To estimate the error rates of morphometric tests, they were compared with molecular tests, revealing areas of disagreement. Structure in the error indicated a bias near the morphometric discrimination threshold. We suggest that larger sample sizes in discriminant function analyses and calibration with molecular data can be used to estimate the likelihood of misidentification for any particular value of the trait. A description of where error lies can then be used to target the more expensive and time-consuming DNA tests to check individuals that are most likely to be mis-assigned.

Dispersal in the sub-Antarctic: king penguins show remarkably little population genetic differentiation across their range

BackgroundSeabirds are important components of marine ecosystems, both as predators and as indicators of ecological change, being conspicuous and sensitive to changes in prey abundance. To determine whether fluctuations in population sizes are localised or indicative of large-scale ecosystem change, we must first understand population structure and dispersal. King penguins are long-lived seabirds that occupy a niche across the sub-Antarctic zone close to the Polar Front. Colonies have very different histories of exploitation, population recovery, and expansion.ResultsWe investigated the genetic population structure and patterns of colonisation of king penguins across their current range using a dataset of 5154 unlinked, high-coverage single nucleotide polymorphisms generated via restriction site associated DNA sequencing (RADSeq). Despite breeding at a small number of discrete, geographically separate sites, we find only very slight genetic differentiation among colonies separated by thousands of kilometers of open-ocean, suggesting migration among islands and archipelagos may be common. Our results show that the South Georgia population is slightly differentiated from all other colonies and suggest that the recently founded Falkland Island colony is likely to have been established by migrants from the distant Crozet Islands rather than nearby colonies on South Georgia, possibly as a result of density-dependent processes.ConclusionsThe observed subtle differentiation among king penguin colonies must be considered in future conservation planning and monitoring of the species, and demographic models that attempt to forecast extinction risk in response to large-scale climate change must take into account migration. It is possible that migration could buffer king penguins against some of the impacts of climate change where colonies appear panmictic, although it is unlikely to protect them completely given the widespread physical changes projected for their Southern Ocean foraging grounds. Overall, large-scale population genetic studies of marine predators across the Southern Ocean are revealing more interconnection and migration than previously supposed.

Isolation and characterization of macaroni penguin (Eudyptes chrysolophus) microsatellite loci and their utility in other penguin species (Spheniscidae, AVES)

We report the characterization of 25 microsatellite loci isolated from the macaroni penguin (Eudyptes chrysolophus). Thirteen loci were arranged into four multiplex sets for future genetic studies of macaroni penguin populations. All 25 loci were tested separately in each of four other penguin species [Adélie penguin (Pygoscelis adeliae), chinstrap penguin (Pygoscelis antarctica), gentoo penguin (Pygoscelis papua) and king penguin (Aptenodytes patagonicus)]. Between eight and 12 loci were polymorphic per species. These loci are expected to be useful for studies of population genetic structure in a range of penguin species.

Why Huddle? Ecological Drivers of Chick Aggregations in Gentoo Penguins, Pygoscelis papua, across Latitudes.

Aggregations of young animals are common in a range of endothermic and ectothermic species, yet the adaptive behavior may depend on social circumstance and local conditions. In penguins, many species form aggregations (aka. crèches) for a variety of purposes, whilst others have never been observed exhibiting this behavior. Those that do form aggregations do so for three known benefits: 1) reduced thermoregulatory requirements, 2) avoidance of unrelated-adult aggression, and 3) lower predation risk. In gentoo penguins, Pygoscelis papua, chick aggregations are known to form during the post-guard period, yet the cause of these aggregations is poorly understood. Here, for the first time, we study aggregation behavior in gentoo penguins, examining four study sites along a latitudinal gradient using time-lapse cameras to examine the adaptive benefit of aggregations to chicks. Our results support the idea that aggregations of gentoo chicks decrease an individual’s energetic expenditure when wet, cold conditions are present. However, we found significant differences in aggregation behavior between the lowest latitude site, Maiviken, South Georgia, and two of the higher latitude sites on the Antarctic Peninsula, suggesting this behavior may be colony specific. We provide strong evidence that more chicks aggregate and a larger number of aggregations occur on South Georgia, while the opposite occurs at Petermann Island in Antarctica. Future studies should evaluate multiple seabird colonies within one species before generalizing behaviors based on one location, and past studies may need to be re-evaluated to determine whether chick aggregation and other behaviors are in fact exhibited species-wide.

The challenges of detecting subtle population structure and its importance for the conservation of emperor penguins

Understanding the boundaries of breeding populations is of great importance for conservation efforts and estimates of extinction risk for threatened species. However, determining these boundaries can be difficult when population structure is subtle. Emperor penguins are highly reliant on sea ice, and some populations may be in jeopardy as climate change alters sea‐ice extent and quality. An understanding of emperor penguin population structure is therefore urgently needed. Two previous studies have differed in their conclusions, particularly whether the Ross Sea, a major stronghold for the species, is isolated or not. We assessed emperor penguin population structure using 4,596 genome‐wide single nucleotide polymorphisms (SNPs), characterized in 110 individuals (10–16 per colony) from eight colonies around Antarctica. In contrast to a previous conclusion that emperor penguins are panmictic around the entire continent, we find that emperor penguins comprise at least four metapopulations, and that the Ross Sea is clearly a distinct metapopulation. Using larger sample sizes and a thorough assessment of the limitations of different analytical methods, we have shown that population structure within emperor penguins does exist and argue that its recognition is vital for the effective conservation of the species. We discuss the many difficulties that molecular ecologists and managers face in the detection and interpretation of subtle population structure using large SNP data sets, and argue that subtle structure should be taken into account when determining management strategies for threatened species, until accurate estimates of demographic connectivity among populations can be made.