Demographic rates reveal the benefits of protected areas in a long-lived migratory bird.

Recent studies have suggested that protected areas often fail to conserve target species. However, the efficacy of terrestrial protected areas is difficult to measure, especially for highly vagile species like migratory birds that may move between protected and unprotected areas throughout their lives. Here, we use a 30-y dataset of detailed demographic data from a migratory waterbird, the Whooper swan (Cygnus cygnus), to assess the value of nature reserves (NRs). We assess how demographic rates vary at sites with varying levels of protection and how they are influenced by movements between sites. Swans had a lower breeding probability when wintering inside NRs than outside but better survival for all age classes, generating a 30-fold higher annual growth rate within NRs. There was also a net movement of individuals from NRs to non-NRs. By combining these demographic rates and estimates of movement (into and out of NRs) into population projection models, we show that the NRs should help to double the population of swans wintering in the United Kingdom by 2030. These results highlight the major effect that spatial management can have on species conservation, even when the areas protected are relatively small and only used during short periods of the life cycle.

Geolocator-tracking seabird migration and moult reveal large-scale, temperature-driven isoscapes in the NE Atlantic.

RATIONALE: By combining precision satellite-tracking with blood sampling, seabirds can be used to validate marine carbon and nitrogen isoscapes, but it is unclear whether a comparable approach using low-precision light-level geolocators (GLS) and feather sampling can be similarly effective. METHODS: Here we used GLS to identify wintering areas of northern gannets (Morus bassanus) and sampled winter grown feathers (confirmed from image analysis of non-breeding birds) to test for spatial gradients in δ13 C and δ15 N in the NE Atlantic. RESULTS: By matching winter-grown feathers with the non-breeding location of tracked birds we found latitudinal gradients in δ13 C and δ15 N in neritic waters. Moreover, isotopic patterns were best explained by sea surface temperature. Similar isotope gradients were found in fish muscle sampled at local ports. CONCLUSIONS: Our study reveals the potential of using seabird GLS and feathers to reconstruct large-scale isotopic patterns.

Factors determining the number of seabirds impacted by oil spills and the success of their rehabilitation: Lessons learned from Namibia and South Africa.

The coastal waters of Namibia and South Africa have an extensive history of oil spills, with 71 recorded up to 2021. Thirty-nine spills reportedly affected 83,224 seabirds, with African penguins (Spheniscus demersus; 91.0 %) and Cape gannets (Morus capensis; 8.5 %) most affected. Spills affecting seabirds were caused by unknown sources (46 %), bulk/cargo carriers (43 %), tankers (38 %) and ship-to-ship transfers (14 %). The number of penguins oiled was predicted by the breeding population size within 25 to 75 km, but not the volume of oiled spilled, the month or the year. Rehabilitation records from penguins oiled in spills since 2001 reveal that the day of admission (relative to the start of the spill) was predictive of packed cell volume, body mass, and plasma total solids, with the latter two being predictive of rehabilitation success. Our results highlight the importance of rapid monitoring at colonies to locate oiled birds in the event of spills.

Intercolony variation in reproductive skipping in the African penguin.

In long-lived species, reproductive skipping is a common strategy whereby sexually mature animals skip a breeding season, potentially reducing population growth. This may be an adaptive decision to protect survival, or a non-adaptive decision driven by individual-specific constraints. Understanding the presence and drivers of reproductive skipping behavior can be important for effective population management, yet in many species such as the endangered African penguin (Spheniscus demersus), these factors remain unknown. This study uses multistate mark-recapture methods to estimate African penguin survival and breeding probabilities at two colonies between 2013 and 2020. Overall, survival (mean ± SE) was higher at Stony Point (0.82 ± 0.01) than at Robben Island (0.77 ± 0.02). Inter-colony differences were linked to food availability; under decreasing sardine (Sardinops sagax) abundance, survival decreased at Robben Island and increased at Stony Point. Additionally, reproductive skipping was evident across both colonies; at Robben Island the probability of a breeder becoming a nonbreeder was ~0.22, versus ~0.1 at Stony Point. Penguins skipping reproduction had a lower probability of future breeding than breeding individuals; this lack of adaptive benefit suggests reproductive skipping is driven by individual-specific constraints. Lower survival and breeding propensity at Robben Island places this colony in greater need of conservation action. However, further research on the drivers of inter-colony differences is needed.

Urofaecal glucocorticoid metabolite concentrations in African penguin (Spheniscus demersus) chick populations experiencing different levels of human disturbance.

Despite the importance of ecotourism in species conservation, little is known about the industry's effects on wildlife. In South Africa, some African penguin (Spheniscus demersus) colonies have become tourist attractions. The species is globally endangered, with population sizes decreasing over the past 40 years. As African penguin chicks are altricial and unable to move away from anthropogenic stressors, it is important to evaluate the effect of tourist activities on baseline glucocorticoid levels as a measure of potential disturbance. Chicks at three study sites within two breeding colonies (Robben Island, Stony Point), with varying levels of exposure to tourism (low/moderate/high) were monitored. Urofaecal samples were collected to determine urofaecal glucocorticoid metabolite (ufGCM) concentrations as an indication of baseline stress physiology. Morphometric measurements were taken to compare body condition between sites. Penguin chicks experiencing low, infrequent human presence had significantly higher mean (± standard deviation) ufGCM levels [1.34 ± 1.70 µg/g dry weight (DW)] compared to chicks experiencing both medium (0.50 ± 0.40 µg/g DW, P = 0.001) and high levels of human presence (0.57 ± 0.47 µg/g DW, P = 0.003). There was no difference in chick body condition across sites. These results suggest that exposure to frequent human activity may induce habituation/desensitization in African penguin chicks. Acute, infrequent human presence was likely an important driver for comparatively higher ufGCM levels in chicks, though several other environmental stressors may also play an important role in driving adrenocortical activity. Nevertheless, as unhabituated chicks experiencing infrequent anthropogenic presence showed significantly higher ufGCM levels, managers and legislation should attempt to minimize all forms of activity around important breeding colonies that are not already exposed to regular tourism. Although the results of this study are crucial for developing enhanced conservation and management protocols, additional research on the long-term effect of anthropogenic activities on African penguin physiology is required.

Half a century of global decline in oceanic sharks and rays.

Overfishing is the primary cause of marine defaunation, yet declines in and increasing extinction risks of individual species are difficult to measure, particularly for the largest predators found in the high seas1-3. Here we calculate two well-established indicators to track progress towards Aichi Biodiversity Targets and Sustainable Development Goals4,5: the Living Planet Index (a measure of changes in abundance aggregated from 57 abundance time-series datasets for 18 oceanic shark and ray species) and the Red List Index (a measure of change in extinction risk calculated for all 31 oceanic species of sharks and rays). We find that, since 1970, the global abundance of oceanic sharks and rays has declined by 71% owing to an 18-fold increase in relative fishing pressure. This depletion has increased the global extinction risk to the point at which three-quarters of the species comprising this functionally important assemblage are threatened with extinction. Strict prohibitions and precautionary science-based catch limits are urgently needed to avert population collapse6,7, avoid the disruption of ecological functions and promote species recovery8,9.

Vulnerability of northern gannets to offshore wind farms; seasonal and sex-specific collision risk and demographic consequences.

There is a pressing need to quantify the risks of renewable energy developments such as offshore wind farms for protected populations. However, assessments are often based on incomplete data, or fail to consider variation in risk between sexes and at different times of year. We tracked northern gannets foraging from the world's largest colony (Bass Rock, Scotland) across five consecutive breeding seasons. We examine how seasonal and sex differences in behaviour affect the collision risk from planned and operational wind farms within their foraging range and assess the likely consequences for long-term population viability. Both sexes made shorter trips during chick-rearing than prior to chick-hatching, spent a greater proportion of time within wind farm sites and had an eight times greater potential collision risk during chick-rearing. Females made longer trips than males at both these times of year, flew higher and spent more time within wind farm sites, leading to three times greater collision risk for females. After accounting for the potential additional mortality from collisions, and assuming that the death of a parent also led to the loss of its offspring, the breeding population was projected to increase by 3.57% (95% CI: 2.16-5.15%) per year, compared with 6.56% (95% CI: 4.59-8.73%) in the absence of turbines, suggesting a negligible effect on population viability. However, additional mortality could result in greater immigration from neighbouring colonies, potentially affecting their viability and highlighting a need for research within a metapopulation framework to assess the impacts of offshore wind developments on vulnerable species across multiple connected sites.

The conservation status and population decline of the African penguin deconstructed in space and time.

Understanding changes in abundance is crucial for conservation, but population growth rates often vary over space and time. We use 40 years of count data (1979-2019) and Bayesian state-space models to assess the African penguin Spheniscus demersus population under IUCN Red List Criterion A. We deconstruct the overall decline in time and space to identify where urgent conservation action is needed. The global African penguin population met the threshold for Endangered with a high probability (97%), having declined by almost 65% since 1989. An historical low of ~17,700 pairs bred in 2019. Annual changes were faster in the South African population (-4.2%, highest posterior density interval, HPDI: -7.8 to -0.6%) than the Namibian one (-0.3%, HPDI: -3.3 to +2.6%), and since 1999 were almost -10% at South African colonies north of Cape Town. Over the 40-year period, the Eastern Cape colonies went from holding ~25% of the total penguin population to ~40% as numbers decreased more rapidly elsewhere. These changes coincided with an altered abundance and availability of the main prey of African penguins. Our results underline the dynamic nature of population declines in space as well as time and highlight which penguin colonies require urgent conservation attention.

Parenting in a warming world: thermoregulatory responses to heat stress in an endangered seabird.

The frequency of extreme weather events, including heat waves, is increasing with climate change. The thermoregulatory demands resulting from hotter weather can have catastrophic impacts on animals, leading to mass mortalities. Although less dramatic, animals also experience physiological costs below, but approaching, critical temperature thresholds. These costs may be particularly constraining during reproduction, when parents must balance thermoregulation against breeding activities. Such challenges should be acute among seabirds, which often nest in locations exposed to high solar radiation and predation risk. The globally endangered bank cormorant Phalacrocorax neglectus breeds in southern Africa in the winter, giving little scope for poleward or phenological shifts in the face of increasing temperatures. Physiological studies of endangered species sensitive to human disturbance, like the bank cormorant, are challenging, because individuals cannot be captured for experimental research. Using a novel, non-invasive, videographic approach, we investigated the thermoregulatory responses of this seabird across a range of environmental temperatures at three nesting colonies. The time birds spent gular fluttering, a behaviour enhancing evaporative heat loss, increased with temperature. Crouching or standing birds spent considerably less time gular fluttering than birds sitting on nests (ca 30% less at 22°C), showing that postural adjustments mediate exposure to heat stress and enhance water conservation. Crouching or standing, however, increases the vulnerability of eggs and chicks to suboptimal temperatures and/or expose nest contents to predation, suggesting that parents may trade-off thermoregulatory demands against offspring survival. We modelled thermoregulatory responses under future climate scenarios and found that nest-bound bank cormorants will gular flutter almost continuously for several hours a day by 2100. The associated increase in water loss may lead to dehydration, forcing birds to prioritize survival over breeding, a trade-off that would ultimately deteriorate the conservation status of this species.

Microplastic study reveals the presence of natural and synthetic fibres in the diet of King Penguins (Aptenodytes patagonicus) foraging from South Georgia.

Marine ecosystems are experiencing substantial disturbances due to climate change and overfishing, and plastic pollution is an additional growing threat. Microfibres are among the most pervasive pollutants in the marine environment, including in the Southern Ocean. However, evidence for microfibre contamination in the diet of top predators in the Southern Ocean is rare. King Penguins (Aptenodytes patagonicus) feed on mesopelagic fish, which undergo diel vertical migrations towards the surface at night. Microfibres are concentrated in surface waters and sediments but can also be concentrated in fish, therefore acting as contamination vectors for diving predators feeding at depth. In this study, we investigate microfibre contamination of King Penguin faecal samples collected in February and March 2017 at South Georgia across three groups: incubating, chick-rearing and non-breeding birds. After a KOH digestion to dissolve the organic matter and a density separation step using a NaCl solution, the samples were filtered to collect microfibres. A total of 77% of the penguin faecal samples (36 of 47) contained microfibres. Fibres were measured and characterized using Fourier-Transform Infrared spectroscopy to determine their polymeric identity. Most fibres (88%) were made of natural cellulosic materials (e.g. cotton, linen), with only 12% synthetic (e.g. polyester, nylon) or semi-synthetic (e.g. rayon). An average of 21.9 ±â€¯5.8 microfibres g-1 of faeces (lab dried mass) was found, with concentrations more than twice as high in incubating penguins than in penguins rearing chicks. Incubating birds forage further north at the Antarctic Polar Front and travel longer distances from South Georgia than chick-rearing birds. This suggests that long-distance travelling penguins are probably more exposed to the risk of ingesting microfibres when feeding north of the Antarctic Polar Front, which might act as a semi-permeable barrier for microfibres. Microfibres could therefore provide a signature for foraging location in King Penguins.

Sex-specific effects of fisheries and climate on the demography of sexually dimorphic seabirds.

Many animal taxa exhibit sex-specific variation in ecological traits, such as foraging and distribution. These differences could result in sex-specific responses to change, but such demographic effects are poorly understood. Here, we test for sex-specific differences in the demography of northern (NGP, Macronectes halli) and southern (SGP, M. giganteus) giant petrels - strongly sexually size-dimorphic birds that breed sympatrically at South Georgia, South Atlantic Ocean. Both species feed at sea or on carrion on land, but larger males (30% heavier) are more reliant on terrestrial foraging than the more pelagic females. Using multi-event mark-recapture models, we examine the impacts of long-term changes in environmental conditions and commercial fishing on annual adult survival and use two-sex matrix population models to forecast future trends. As expected, survival of male NGP was positively affected by carrion availability, but negatively affected by zonal winds. Female survival was positively affected by meridional winds and El Niño-Southern Oscillation (ENSO), and negatively affected by sea ice concentration and pelagic longline effort. Survival of SGPs did not differ between sexes; however, survival of males only was positively correlated with the Southern Annular Mode (SAM). Two-sex population projections indicate that future environmental conditions are likely to benefit giant petrels. However, any potential increase in pelagic longline fisheries could reduce female survival and population growth. Our study reveals that sex-specific ecological differences can lead to divergent responses to environmental drivers (i.e. climate and fisheries). Moreover, because such effects may not be apparent when all individuals are considered together, ignoring sex differences could underestimate the relative influence of a changing environment on demography.

A non-invasive approach to estimate the energetic requirements of an increasing seabird population in a perturbed marine ecosystem.

There is a growing desire to integrate the food requirements of predators living in marine ecosystems impacted by humans into sustainable fisheries management. We used non-invasive video-recording, photography and focal observations to build time-energy budget models and to directly estimate the fish mass delivered to chicks by adult greater crested terns Thalasseus bergii breeding in the Benguela ecosystem. Mean modelled adult daily food intake increased from 140.9 g·d-1 of anchovy Engraulis capensis during incubation to 171.7 g·d-1 and 189.2 g·d-1 when provisioning small and large chicks, respectively. Modelled prey intake expected to be returned to chicks was 58.3 g·d-1 (95% credible intervals: 44.9-75.8 g·d-1) over the entire growth period. Based on our observations, chicks were fed 19.9 g·d-1 (17.2-23.0 g·d-1) to 45.1 g·d-1 (34.6-58.7 g·d-1) of anchovy during early and late provisioning, respectively. Greater crested terns have lower energetic requirements at the individual (range: 15-34%) and population level (range: 1-7%) than the other Benguela endemic seabirds that feed on forage fish. These modest requirements - based on a small body size and low flight costs - coupled with foraging plasticity have allowed greater crested terns to cope with changing prey availability, unlike the other seabirds species using the same exploited prey base.

Foraging plasticity in seabirds: A non-invasive study of the diet of greater crested terns breeding in the Benguela region.

Marine predators, such as seabirds, are useful indicators of marine ecosystem functioning. In particular, seabird diet may reflect variability in food-web composition due to natural or human-induced environmental change. Diet monitoring programmes, which sample diet non-invasively, are valuable aids to conservation and management decision-making. We investigated the diet of an increasing population of greater crested terns Thalasseus bergii in the Western Cape, South Africa, during three successive breeding seasons (2013 to 2015), when populations of other seabirds feeding on small pelagic schooling fish in the region were decreasing. Breeding greater crested terns carry prey in their bills, so we used an intensive photo-sampling method to record their diet with little disturbance. We identified 24,607 prey items from at least 47 different families, with 34 new prey species recorded. Fish dominated the diet, constituting 94% of prey by number, followed by cephalopods (3%), crustaceans (2%) and insects (1%). The terns mainly targeted surface-schooling Clupeiformes, with anchovy Engraulis encrasicolus the most abundant prey in all three breeding seasons (65% overall). Prey composition differed significantly between breeding stages and years, with anchovy most abundant at the start of the breeding season, becoming less frequent as the season progressed. The proportion of anchovy in the diet also was influenced by environmental factors; anchovy occurred more frequently with increasing wind speeds and was scarce on foggy days, presumably because terns rely in part on social facilitation to locate anchovy schools. The application of this intensive and non-invasive photo-sampling method revealed an important degree of foraging plasticity for this seabird within a context of locally reduced food availability, suggesting that, unlike species that specialise on a few high-quality prey, opportunistic seabirds may be better able to cope with reductions in the abundance of their preferred prey.

Bayesian inference reveals positive but subtle effects of experimental fishery closures on marine predator demographics.

Global forage-fish landings are increasing, with potentially grave consequences for marine ecosystems. Predators of forage fish may be influenced by this harvest, but the nature of these effects is contentious. Experimental fishery manipulations offer the best solution to quantify population-level impacts, but are rare. We used Bayesian inference to examine changes in chick survival, body condition and population growth rate of endangered African penguins Spheniscus demersus in response to 8 years of alternating time-area closures around two pairs of colonies. Our results demonstrate that fishing closures improved chick survival and condition, after controlling for changing prey availability. However, this effect was inconsistent across sites and years, highlighting the difficultly of assessing management interventions in marine ecosystems. Nevertheless, modelled increases in population growth rates exceeded 1% at one colony; i.e. the threshold considered biologically meaningful by fisheries management in South Africa. Fishing closures evidently can improve the population trend of a forage-fish-dependent predator-we therefore recommend they continue in South Africa and support their application elsewhere. However, detecting demographic gains for mobile marine predators from small no-take zones requires experimental time frames and scales that will often exceed those desired by decision makers.

Seabird species vary in behavioural response to drone census.

Unmanned aerial vehicles (UAVs) provide an opportunity to rapidly census wildlife in remote areas while removing some of the hazards. However, wildlife may respond negatively to the UAVs, thereby skewing counts. We surveyed four species of Arctic cliff-nesting seabirds (glaucous gull Larus hyperboreus, Iceland gull Larus glaucoides, common murre Uria aalge and thick-billed murre Uria lomvia) using a UAV and compared censusing techniques to ground photography. An average of 8.5% of murres flew off in response to the UAV, but >99% of those birds were non-breeders. We were unable to detect any impact of the UAV on breeding success of murres, except at a site where aerial predators were abundant and several birds lost their eggs to predators following UAV flights. Furthermore, we found little evidence for habituation by murres to the UAV. Most gulls flew off in response to the UAV, but returned to the nest within five minutes. Counts of gull nests and adults were similar between UAV and ground photography, however the UAV detected up to 52.4% more chicks because chicks were camouflaged and invisible to ground observers. UAVs provide a less hazardous and potentially more accurate method for surveying wildlife. We provide some simple recommendations for their use.

Seabirds.

In this Quick Guide, Votier and Sherley explain how diverse seabirds play important roles in ecosystem functioning, global nutrient cycling and climate regulation, but are declining in the face of multiple threats.

Defining ecologically relevant scales for spatial protection with long-term data on an endangered seabird and local prey availability.

Human activities are important drivers of marine ecosystem functioning. However, separating the synergistic effects of fishing and environmental variability on the prey base of nontarget predators is difficult, often because prey availability estimates on appropriate scales are lacking. Understanding how prey abundance at different spatial scales links to population change can help integrate the needs of nontarget predators into fisheries management by defining ecologically relevant areas for spatial protection. We investigated the local population response (number of breeders) of the Bank Cormorant (Phalacrocorax neglectus), a range-restricted endangered seabird, to the availability of its prey, the heavily fished west coast rock lobster (Jasus lalandii). Using Bayesian state-space modeled cormorant counts at 3 colonies, 22 years of fisheries-independent data on local lobster abundance, and generalized additive modeling, we determined the spatial scale pertinent to these relationships in areas with different lobster availability. Cormorant numbers responded positively to lobster availability in the regions with intermediate and high abundance but not where regime shifts and fishing pressure had depleted lobster stocks. The relationships were strongest when lobsters 20-30 km offshore of the colony were considered, a distance greater than the Bank Cormorant's foraging range when breeding, and may have been influenced by prey availability for nonbreeding birds, prey switching, or prey ecology. Our results highlight the importance of considering the scale of ecological relationships in marine spatial planning and suggest that designing spatial protection around focal species can benefit marine predators across their full life cycle. We propose the precautionary implementation of small-scale marine protected areas, followed by robust assessment and adaptive-management, to confirm population-level benefits for the cormorants, their prey, and the wider ecosystem, without negative impacts on local fisheries.

Metapopulation Tracking Juvenile Penguins Reveals an Ecosystem-wide Ecological Trap.

Climate change and fisheries are transforming the oceans, but we lack a complete understanding of their ecological impact [1-3]. Environmental degradation can cause maladaptive habitat selection, inducing ecological traps with profound consequences for biodiversity [4-6]. However, whether ecological traps operate in marine systems is unclear [7]. Large marine vertebrates may be vulnerable to ecological traps [6], but their broad-scale movements and complex life histories obscure the population-level consequences of habitat selection [8, 9]. We satellite tracked postnatal dispersal in African penguins (Spheniscus demersus) from eight sites across their breeding range to test whether they have become ecologically trapped in the degraded Benguela ecosystem. Bayesian state-space and habitat models show that penguins traversed thousands of square kilometers to areas of low sea surface temperatures (14.5°C-17.5°C) and high chlorophyll-a (∼11 mg m-3). These were once reliable cues for prey-rich waters, but climate change and industrial fishing have depleted forage fish stocks in this system [10, 11]. Juvenile penguin survival is low in populations selecting degraded areas, and Bayesian projection models suggest that breeding numbers are ∼50% lower than if non-impacted habitats were used, revealing the extent and effect of a marine ecological trap for the first time. These cascading impacts of localized forage fish depletion-unobserved in studies on adults-were only elucidated via broad-scale movement and demographic data on juveniles. Our results support suspending fishing when prey biomass drops below critical thresholds [12, 13] and suggest that mitigation of marine ecological traps will require matching conservation action to the scale of ecological processes [14].