Abstract
. Historic limitations have resulted in marine ecological studies usually overlooking some well-established concepts from behavioural ecology. This is likely because the methods available were easily overwhelmed by the scale of ecological processes and sampling resolution in the marine environment. Innovations in technology, data management, and statistical modelling now provide the capacity to fully embrace behavioural ecology concepts and study marine ecological interactions from a more holistic perspective. To facilitate this vision, we propose a novel perspective and workflow for marine ecology: the Seascape of Ecological Energy, or SEE-scapes. SEE-scapes contextualizes the accumulated knowledge from marine biology and behavioural ecology research and provides a guide for marine scientists interested in grounding their research in behavioural ecology’s first principles. SEE-scapes specifies relevant considerations for contemporary seascapes, with special attention to relationships between individuals and their conspecifics, competitors, predators, and the abiotic environment. It is formulated to account for how the unique features of marine vertebrates require revisions to standard, terrestrially focused methodological approaches, and offers solutions for how to sample and model the eco-evolutionary drivers of behaviour across the dynamic and hierarchical seascape. Simply put, SEE-scapes provides the guidance to translate a dynamic system and reveal opportunities for collaborative and integrative research.

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.

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.

AbstractEffective marine ecosystem monitoring is critical for sustainable management. Monitoring seabird diets can convey important information on ecosystem health and seabird–fishery interactions. The diet of breeding black-browed albatross (Thalassarche melanophris) has previously been assessed using stomach content analysis (SCA) or stable isotope analysis (SIA), but not both methods together. Combining dietary sampling approaches reduces biases associated with using single methods. This study combines SCA and SIA to study the diet of black-browed albatross chicks, with a specific focus on fishery discard consumption, at two Falkland Islands colonies (New Island 51°43′S, 61°18′W and Steeple Jason Island 51°01′S, 61°13′W) during two consecutive breeding seasons (2019 and 2020). SCA provided high taxonomic resolution of short-term diet and priors for stable isotope mixing models, with multiple measures of dietary items (e.g. numeric frequency N%, frequency of occurrence FO%). By contrast, SIA of down feathers provided a single and more integrated dietary signal from throughout chick development. Although the two methods disagreed on the dominant prey group (SCA—crustacean; SIA—pelagic fish), the complementary information suggested a chick diet dominated by natural prey (SCA: 74%–93% [FO], 44%–98% [N]; SIA: minimum 87%–95% contribution). Nonetheless, SCA revealed that a high proportion of breeding adults do take discards. We detected consistent colony-specific diets in relation to prey species, but not in relation to higher discard use. Overall, discard consumption was highest in 2020, the year characterised by the poorest foraging conditions. Our results have implications for fisheries management and future dietary studies assessing discard use.

AbstractIn 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.

Summary
. Seabirds are declining globally and are one of the most threatened groups of birds. To halt or reverse this decline they need protection both on land and at sea, requiring site-based conservation initiatives based on seabird abundance and diversity. The Important Bird and Biodiversity Area (IBA) programme is a method of identifying the most important places for birds based on globally agreed standardised criteria and thresholds. However, while great strides have been made identifying terrestrial sites, at-sea identification is lacking. The Chagos Archipelago, central Indian Ocean, supports four terrestrial IBAs (tIBAs) and two proposed marine IBAs (mIBAs). The mIBAs are seaward extensions to breeding colonies based on outdated information and, other types of mIBA have not been explored. Here, we review the proposed seaward extension mIBAs using up-to-date seabird status and distribution information and, use global positioning system (GPS) tracking from Red-footed Booby Sula sula – one of the most widely distributed breeding seabirds on the archipelago – to identify any pelagic mIBAs. We demonstrate that due to overlapping boundaries of seaward extension to breeding colony and pelagic areas of importance there is a single mIBA in the central Indian Ocean that lays entirely within the Chagos Archipelago Marine Protected Area (MPA). Covering 62,379 km2 it constitutes ~10% of the MPA and if designated, would become the 11th largest mIBA in the world and 4th largest in the Indian Ocean. Our research strengthens the evidence of the benefits of large-scale MPAs for the protection of marine predators and provides a scientific foundation stone for marine biodiversity hotspot research in the central Indian Ocean.

Abstract
. Fishery discards supplement food for many seabirds, but the impacts of declining discards are poorly understood. Discards may be beneficial for some populations but have negative impacts by increasing bycatch risk or because they are junk-food. The Falkland Islands support > 70% of global black-browed albatross Thalassarche melanophris populations, which feed on discards. However, the effect of discards on population demographics, and implications of fishery management changes, are unknown. We analysed stomach contents of black-browed albatross chicks across eight breeding seasons (2004–2020) from New Island, Falkland Islands, to assess variation in discard consumption and how this relates to foraging conditions and breeding success. Across years, 68%–98% of samples contained natural prey, whilst 23%–88% of samples contained fishery discards. Discard consumption was positively related to fishery catches of hoki Macruronus magellanicus and sea surface temperature anomalies SSTA (°C), and negatively related to breeding success. These results suggest a diet-switching behaviour for Falkland Islands albatrosses, whereby birds switch from preferred natural prey to suboptimal discards when environmental conditions, and hence natural feeding opportunities, are unfavourable. Crucially, this study highlights that fishery discards do not compensate for poor natural foraging conditions for breeding albatrosses in the long term.

Climate change and other human activities are causing profound effects on marine ecosystem productivity. We show that the breeding success of seabirds is tracking hemispheric differences in ocean warming and human impacts, with the strongest effects on fish-eating, surface-foraging species in the north. Hemispheric asymmetry suggests the need for ocean management at hemispheric scales. For the north, tactical, climate-based recovery plans for forage fish resources are needed to recover seabird breeding productivity. In the south, lower-magnitude change in seabird productivity presents opportunities for strategic management approaches such as large marine protected areas to sustain food webs and maintain predator productivity. Global monitoring of seabird productivity enables the detection of ecosystem change in remote regions and contributes to our understanding of marine climate impacts on ecosystems.

AbstractEstimating life‐history traits and understanding their variation underpins the management of long‐lived, migratory animals, while knowledge of recovery dynamics can inform the management of conservation‐dependent species. Using a combination of nest counts and individual‐based life‐history data collected since 1993, we explore the drivers underlying contrasting population recovery rates of sympatrically nesting loggerhead (Caretta caretta) and green (Chelonia mydas) turtles in North Cyprus. We found that nest counts of loggerhead and green turtles from 28 beaches across the island increased by 46% and 162%, respectively over the past 27 years. A Bayesian state‐space model revealed that, at our individual‐based monitoring site, nesting of green turtles increased annually at four times the rate of that of loggerhead turtles. Furthermore, we found that loggerhead turtles nesting at the individual‐based monitoring site had stable reproductive parameters and average adult survival for the species and are the smallest breeding adults globally. Based on results from multiple matrix model scenarios, we propose that higher mortality rates of individuals in all age classes (likely driven by differences in life history and interaction with fisheries), rather than low reproductive output, are impeding the recovery of this species. While the increase in green turtles is encouraging, the Mediterranean population is estimated to have around 3,400 adults and is restricted to the Eastern Basin. The recovery of loggerhead turtles is likely to be compromised until mortality rates in the region are adequately quantified and mitigated. As survival of immature individuals is a powerful driver for sea turtle population numbers, additional efforts should target management at pelagic and neritic foraging areas. Understanding threats faced by immature life stages is crucial to accurately parameterise population models and to target conservation actions for long‐lived marine vertebrates.

Abstract
. In a scientifically-transformative project, South Africa implemented a decade-long field experiment to understand how fisheries may be affecting its most iconic seabird, the African penguin Spheniscus demersus. This unique effort prohibits the take of anchovy and sardine within relatively small areas around four African penguin breeding colonies, two in the Benguela upwelling ecosystem and two in the adjacent Agulhas region. For the Benguela, fisheries closures within the birds’ primary foraging range increased their breeding productivity and perhaps reduced parental foraging efforts, indicating that the fisheries are competing with the birds for food. Results were less clear for foraging behaviour in the Agulhas, but no data on breeding success were collected there. The African penguin is endangered, its population continues to decline, and fisheries closures have been demonstrated to improve demographic traits that contribute to population growth. Therefore, given the critical status of the species, fisheries closures should be maintained, at least at Dassen Island where the population has great capacity to expand and support other nearby colonies. Continuing or implementing corresponding fisheries closures in the Agulhas region is also warranted, as well as creating and testing the value of pelagic closed areas during the non-breeding season when the penguins disperse widely across these ecosystems. These management actions would increase penguin food supplies and may help to meet societal goals of halting the decline of the penguin population, as well as maintaining the economic and cultural services provided by fisheries and ecotourism.

Abstract
. 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.

King penguins (Aptenodytes patagonicus) are an iconic Southern Ocean species, but the prey distributions that underpin their at-sea foraging tracks and diving behaviour remain unclear. We conducted simultaneous acoustic surveys off South Georgia and tracking of king penguins breeding ashore there in Austral summer 2017 to gain insight into habitat use and foraging behaviour. Acoustic surveys revealed ubiquitous deep scattering layers (DSLs; acoustically detected layers of fish and other micronekton that inhabit the mesopelagic zone) at c. 500 m and shallower ephemeral fish schools. Based on DNA extracted from penguin faecal samples, these schools were likely comprised of lanternfish (an important component of king penguin diets), icefish (Channichthyidae spp.) and painted noties (Lepidonotothen larseni). Penguins did not dive as deep as DSLs, but their prey-encounter depth-distributions, as revealed by biologging, overlapped at fine scale (10s of m) with depths of acoustically detected fish schools. We used neural networks to predict local scale (10 km) fish echo intensity and depth distribution at penguin dive locations based on environmental correlates, and developed models of habitat use. Habitat modelling revealed that king penguins preferentially foraged at locations predicted to have shallow and dense (high acoustic energy) fish schools associated with shallow and dense DSLs. These associations could be used to predict the distribution of king penguins from other colonies at South Georgia for which no tracking data are available, and to identify areas of potential ecological significance within the South Georgia and the South Sandwich Islands marine protected area.

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 ± 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.

AbstractThe 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.

Every year fisheries discard >10 million tonnes of fish. This provides a bounty for scavengers, yet the ecological impact of discarding is understudied. Seabirds are the best-studied discard scavengers and fisheries have shaped their movement ecology, demography and community structure. However, we know little about the number of scavenging seabirds that discards support, how this varies over time or might change as stocks and policy change. Here, we use a Bayesian bioenergetics model to estimate the number of scavenging birds potentially supported by discards in the North Sea (one of the highest discard-producing regions) in 1990, around the peak of production, and again after discard declines in 2010. We estimate that North Sea discards declined by 48% from 509,840 tonnes in 1990 to 267,549 tonnes in 2010. This waste had the potential to support 5.66 (95% credible intervals: 3.33–9.74) million seabirds in the 1990s, declining by 39% to 3.45 (1.98–5.78) million birds by 2010. Our study reveals the potential for fishery discards to support very large scavenging seabird communities but also shows how this has declined over recent decades. Discard bans, like the European Union's Landing Obligation, may reduce inflated scavenger communities, but come against a backdrop of gradual declines potentially buffering deleterious impacts. More work is required to reduce uncertainty and to generate global estimates, but our study highlights the magnitude of scavenger communities potentially supported by discards and thus the importance of understanding the wider ecological consequences of dumping fisheries waste.

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.

AbstractThe International Union for Conservation of Nature's (IUCN) Red List is the global standard for quantifying extinction risk but assessing population reduction (criterion A) of wide‐ranging, long‐lived marine taxa remains difficult and controversial. We show how Bayesian state–space models (BSSM), coupled with expert knowledge at IUCN Red List workshops, can combine regional abundance data into indices of global population change. To illustrate our approach, we provide examples of the process to assess four circumglobal sharks with differing temporal and spatial data‐deficiency: Blue Shark (Prionace glauca), Shortfin Mako (Isurus oxyrinchus), Dusky Shark (Carcharhinus obscurus), and Great Hammerhead (Sphyrna mokarran). For each species, the BSSM provided global population change estimates over three generation lengths bounded by uncertainty levels in intuitive outputs, enabling informed decisions on the status of each species. Integrating similar analyses into future workshops would help conservation practitioners ensure robust, consistent, and transparent Red List assessments for other long‐lived, wide‐ranging species.

The Bank Cormorant Phalacrocorax neglectus is endemic to the Benguela upwelling ecosystem off southwest Africa and is classified as Endangered owing to a recent large reduction in its number. It is thought that food scarcity, including a decreased abundance of West Coast rock lobster Jasus lalandii, has been a major driver of the decrease, yet its diet in South Africa is poorly known. We collected 941 pellets regurgitated by Bank Cormorants, at 18 South African breeding colonies during 1975–1985, and 1 523 pellets at 17 colonies during 1995–2002. The species composition of the diet (% numbers) was significantly different between the two periods, with widespread decreases in proportions of rock lobster in the west and of octopus and cuttlefish Sepia spp. at most localities. These taxa were replaced in the diet by fish, including Gobiidae and Clinidae. The pelagic goby Sufflogobius bibarbatus, an important prey of Bank Cormorants in Namibia, was absent from pellets collected in 1975–1985 but common at northern localities from 1995–2002. Composition of the diet by frequency of occurrence was only determined for 1995–2002, when rock lobster was present in 67% of all samples collected, cuttlefish in 39%, and Clinidae in 32%. Data for 1975–1985 and 1995–2002 showed that carapace lengths of rock lobsters eaten by Bank Cormorants averaged 56 mm (range 22–82 mm) and 50 mm (range 22–75 mm), respectively, which compares to the minimum legal size of 75 mm for fisheries in South Africa. This energy- rich prey item was an important constituent of the diet in the winter breeding period.

Penguins face a wide range of threats. Most observed population changes have been negative and have happened over the last 60 years. Today, populations of 11 penguin species are decreasing. Here we present a review that synthesizes details of threats faced by the world's 18 species of penguins. We discuss alterations to their environment at both breeding sites on land and at sea where they forage. The major drivers of change appear to be climate, and food web alterations by marine fisheries. In addition, we also consider other critical and/or emerging threats, namely human disturbance near nesting sites, pollution due to oil, plastics and chemicals such as mercury and persistent organic compounds. Finally, we assess the importance of emerging pathogens and diseases on the health of penguins. We suggest that in the context of climate change, habitat degradation, introduced exotic species and resource competition with fisheries, successful conservation outcomes will require new and unprecedented levels of science and advocacy. Successful conservation stories of penguin species across their geographical range have occurred where there has been concerted effort across local, national and international boundaries to implement effective conservation planning.

Abstract


Understanding the functional relationship between marine predators and their prey is vital to inform ecosystem‐based management. However, collecting concurrent data on predator behaviour and their prey at relevant scales is challenging. Moreover, opportunities to study these relationships in the absence of industrial fishing are extremely rare.

We took advantage of an experimental fisheries closure to study how local prey abundance influences foraging success and chick condition of Endangered African penguins Spheniscus demersus in the Benguela ecosystem.

We tracked 75 chick‐provisioning penguins with GPS–time–depth devices, measured body condition of 569 chicks, quantified the diet of 83 breeding penguins and conducted 12 forage fish hydro‐acoustic surveys within a 20 km radius of Robben Island, South Africa, over three years (2011–2013). Commercial fishing for the penguins' main prey, sardine Sardinops sagax and anchovy Engraulis encrasicolus, was prohibited within this 20 km radius during the study period.

Local forage fish abundance explained 60% of the variation in time spent diving for 14 penguins at sea within 2 days of a hydro‐acoustic survey. Penguin foraging effort (time spent diving, number of wiggles per trip, number of foraging dives and the maximum distance travelled) increased and offspring body condition decreased as forage fish abundance declined. In addition, quantile regression revealed that variation in foraging effort increased as prey abundance around the colony declined.

Policy implications. Our results demonstrate that local forage fish abundance influences seabird foraging and offspring fitness. They also highlight the potential for offspring condition and the mean–variance relationship in foraging behaviour to act as leading indicators of poor prey abundance. By rapidly indicating periods where forage resources are scarce, these metrics could help limit seabird fisheries competition and aid the implementation of dynamic ocean management.

Marine vertebrates show a diversity of migration strategies, including sex differences. This may lead to differential demography, but the consequences of such between-sex variation are little understood. Here, we studied the migration of known-sex northern gannets Morus bassanus — a partial migrant with females ~8 % heavier than males. We used geolocators to determine wintering areas of 49 breeding adults (19 females and 30 males during 2010 to 2014) from 2 colonies in the northeast Atlantic (Bass Rock and Grassholm, UK). We also tested for sex-specific survival probabilities using capture−mark−recapture methods (n = 72 individuals Bass Rock, n = 229 individuals Grassholm; 2010−2018) and applied sex-specific population projection matrices (PPMs) to quantify population-level effects. Tracked gannets wintered in a range of large marine ecosystems (LMEs): Canary Current LME (CCLME; 69 %), Celtic-Biscay Shelf LME (16 %), Iberian Coastal LME (8 %), North Sea LME (4 %) or Mediterranean LME (2 %). Migratory destination differed between the sexes: 90 % of females vs. 57 % of males wintered in the CCLME. Survival was similar between the sexes at Bass Rock (mean ± 95 % CI = 0.951 ± 0.053 and 0.956 ± 0.047 for females and males, respectively). At Grassholm, there was evidence of slight sex differences in breeder survival: females had lower annual survival (0.882 ± 0.040) than males (0.946 ± 0.026). At Bass Rock, PPMs with no sex effect best fitted the observed population increase (1994−2014). Sex-specific PPMs fitted the population estimates for Grassholm (1995−2015). Our results reveal that female gannets are more likely to travel further than males to winter in the CCLME. This difference is unlikely due to morphological differences, unlike in other bird species. However, the reason for slightly higher over-winter female mortality at Grassholm is unclear.

Abstract


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.

The Cape Gannet Morus capensis is one of several seabird species endemic to the Benguela upwelling ecosystem (BUS) but whose population has recently decreased, leading to an unfavourable IUCN Red List assessment. Application of ‘JARA’ (‘Just Another Red-List Assessment,’ a Bayesian state-space tool used for IUCN Red List assessments) to updated information on the areas occupied by Cape Gannets and the nest densities of breeding birds at their six colonies, suggested that the species should be classified as Vulnerable. However, the rate of decrease of Cape Gannets in their most-recent generation exceeded that of the previous generation, primarily as a result of large decreases at Bird Island, Lambert’s Bay, and Malgas Island, off South Africa’s west coast (the western part of their range). Since the 1960s, there has been an ongoing redistribution of the species from northwest to southeast around southern Africa, and ∼70% of the population now occurs on the south coast of South Africa, at Bird Island in Algoa Bay, on the eastern border of the BUS. Recruitment rather than adult survival may be limiting the present population; however, information on the seabird’s demographic parameters and mortality in fisheries is lacking for colonies in the northern part of the BUS. Presently, major threats to Cape Gannet include: substantially decreased availability of their preferred prey in the west; heavy mortalities of eggs, chicks and fledglings at and around colonies, inflicted by Cape Fur Seals Arctocephalus pusillus and other seabirds; substantial disturbance at colonies caused by Cape Fur Seals attacking adult gannets ashore; oiling; and disease.

AbstractThere 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.

. 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.
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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.

The nominate race of the greater crested tern Thalasseus bergii breeds only along the coast of the Benguela region (west coast) of southern Africa, where its population is increasing, in contrast to other species of breeding seabirds in the region which similarly depend on forage fish. Although this population’s trends are well known, its demographic parameters remain obscure. We used multi-event capture–recapture–recovery modelling to provide the first estimates of survival. The survival rate of birds older than two years was 0.93 (0.91–0.95); second-year survival was 0.93 (0.90–0.95) and first-year survival was 0.80 (0.73–0.85). A resighting of a 34-year-old bird is a new longevity record for the species. Population growth rates projected with a Leslie matrix model that included our survival estimates were very similar to estimates from annual counts of the breeding population (7–8% y−1), suggesting that these survival estimates adequately describe the species’ demography. High survival rates have likely contributed to the species’ recent population increase in the region.

Worldwide, in recent years capture fisheries targeting lower-trophic level forage fish and euphausiid crustaceans have been substantial (∼20 million metric tons [MT] annually). Landings of forage species are projected to increase in the future, and this harvest may affect marine ecosystems and predator-prey interactions by removal or redistribution of biomass central to pelagic food webs. In particular, fisheries targeting forage fish and euphausiids may be in competition with seabirds, likely the most sensitive of marine vertebrates given limitations in their foraging abilities (ambit and gape size) and high metabolic rate, for food resources. Lately, apparent competition between fisheries and seabirds has led to numerous high-profile conflicts over interpretations, as well as the approaches that could and should be used to assess the magnitude and consequences of fisheries-seabird resource competition. In this paper, we review the methods used to date to study fisheries competition with seabirds, and present “best practices” for future resource competition assessments. Documenting current fisheries competition with seabirds generally involves addressing two major issues: 1) are fisheries causing localized prey depletion that is sufficient to affect the birds? (i.e. are fisheries limiting food resources?), and 2) how are fisheries-induced changes to forage stocks affecting seabird populations given the associated functional or numerical response relationships? Previous studies have been hampered by mismatches in the scale of fisheries, fish, and seabird data, and a lack of causal understanding due to confounding by climatic and other ecosystem factors (e.g. removal of predatory fish). Best practices for fisheries-seabird competition research should include i) clear articulation of hypotheses, ii) data collection (or summation) of fisheries, fish, and seabirds on matched spatio-temporal scales, and iii) integration of observational and experimental (including numerical simulation) approaches to establish connections and causality between fisheries and seabirds. As no single technique can provide all the answers to this vexing issue, an integrated approach is most promising to obtain robust scientific results and in turn the sustainability of forage fish fisheries from an ecosystem perspective.

The use of wildlife rehabilitation for conservation is growing, but quantitative criteria are rarely used to guide whether and when to remove animals from the wild. Since 2006, large numbers of African penguin Spheniscus demersus chicks have been abandoned annually when adults enter moult with dependent young still in the nest. As part of conservation initiatives for this endangered species, these chicks were collected and hand reared to fledging age. Post-release survival has been well documented; in this study we develop models to predict survival of individuals during rehabilitation with the aim of improving hand-rearing success and guiding the use of scarce resources. For 1455 chicks abandoned between 2008 and 2013, we assessed whether a chick body condition index (BCI) could predict outcome (death or release) and time spent in rearing. In addition, for a subset of 173 chicks in 2012, we assessed whether BCI at admission influenced chick growth rates during rehabilitation and examined whether the use of additional structural measurements and sex provided additional power to predict outcome. Models predicted an 82.9% (95% confidence interval: 73.3–89.5%) release rate for chicks admitted with a BCI >0, the proposed guideline for removal from colonies. This fell below 50% for BCIs 

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.

Dietary studies give vital insights into foraging behaviour, with implications for understanding changing environmental conditions and the anthropogenic impacts on natural resources. Traditional diet sampling methods may be invasive or subject to biases, so developing non-invasive and unbiased methods applicable to a diversity of species is essential. We used digital photography to investigate the diet fed to chicks of a prey-carrying seabird and compared our approach (photo-sampling) to a traditional method (regurgitations) for the greater crested tern Thalasseus bergii. Over three breeding seasons, we identified >24 000 prey items of at least 48 different species, more than doubling the known diversity of prey taken by this population of terns. We present a method to estimate the length of the main prey species (anchovy Engraulis encrasicolus) from photographs, with an accuracy

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.

African penguins (Spheniscus demersus) in South Africa have been subject to rapid decline in the 20th century and are currently listed as "endangered" on the IUCN Red List. A stochastic, stage-specific system dynamics model with spatial components was developed to investigate the interaction of multiple pressures on penguin population development, and originally applied to study conservation management options for the penguin colony at Robben Island, Table Bay, South Africa. We here present an extension of the model to the nearby colony at Dyer Island. The modelled population was found to be strongly dominated by the effects of Cape fur seal (Arctocephalus pusillus pusillus) predation and immature emigration, which appear to be key drivers behind current declines in population numbers at this colony. Chronic low-level oiling also had a consistent impact despite ongoing mitigation measures, while kelp gull predation had a lower effect than expected. At current low population size, the pressure from either seal predation or immature emigration seems sufficient to mask any beneficial effects to penguins from possible improvements in available food biomass (e.g. from fishery restrictions). Results suggest that conservation management at this colony should focus on regular culling of predating seals combined with improving availability of prey (thus discouraging emigration) both in the foraging range of breeding penguins and in their general foraging area. Our findings demonstrate the use of site-specific scenario tools to explore conservation strategies in data-poor management situations.

. Marine no-take zones can have positive impacts for target species and are increasingly important management tools. However, whether they indirectly benefit higher order predators remains unclear. The endangered African penguin (
. Spheniscus demersus
. ) depends on commercially exploited forage fish. We examined how chick survival responded to an experimental 3-year fishery closure around Robben Island, South Africa, controlling for variation in prey biomass and fishery catches. Chick survival increased by 18% when the closure was initiated, which alone led to a predicted 27% higher population compared with continued fishing. However, the modelled population continued to decline, probably because of high adult mortality linked to poor prey availability over larger spatial scales. Our results illustrate that small no-take zones can have bottom-up benefits for highly mobile marine predators, but are only one component of holistic, ecosystem-based management regimes.
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Reliable estimates of survival and dispersal are crucial to understanding population dynamics, but for seabirds, in which some individuals spend years away from land, mortality and emigration are often confounded. Multistate mark–recapture methods reduce bias by incorporating movement into the process of estimating survival. We used a multistate model to provide unbiased age‐specific survival and movement probabilities for the Endangered African Penguin Spheniscus demersus based on 5281 nestlings and 31 049 adults flipper‐banded and resighted in the Western Cape, South Africa, between 1994 and 2012. Adult survival was initially high (≥ 0.74) but declined after 2003–2004 coincident with a reduction in the availability of Sardine Sardinops sagax and Anchovy Engraulis encrasicolus on the west coast of South Africa. Juvenile survival was poorly estimated, but was lower and more variable than adult survival. Fidelity to the locality of origin varied over time, but was high in adults at Robben and Dassen islands (≥ 0.88) and above 0.55 for juvenile and immature Penguins at all localities. Movement occurred predominantly during 1994–2003 and was indicative of immigration to Robben and Dassen islands. Our results confirm that a prolonged period of adult mortality contributed to the observed decline in the African Penguin population and suggest a need for approaches operating over large spatial scales to ensure food security for marine top predators.

AbstractColonial breeding is characteristic of seabirds but nesting at high density has both advantages and disadvantages and may reduce survival and fecundity. African penguins (Spheniscus demersus) initiated breeding at Robben Island, South Africa in 1983. The breeding population on the island increased in the late 1990s and early 2000s before decreasing rapidly until 2010. Before the number breeding peaked, local nest density in the areas where the colony was initiated plateaued, suggesting that preferred nests sites were mostly occupied, and the area used by breeding birds expanded. However, it did not contract again as the population decreased, so that nesting density varied substantially. Breeding success was related positively to the prey available to the breeding birds and negatively to local nest density, particularly during the chick‐rearing period, suggesting a density‐dependence operating through social interactions in the colony, possibly exacerbated by poor prey availability when the breeding population was large. Although nest density at Robben Island was not high, nesting burrows, which probably reduce the incidence of aggressive encounters in the colony, are scarce and our results suggest that habitat alteration has modified the strength of density‐dependent relationships for African penguins. Gaining a better understanding of how density dependence affects fecundity and population growth rates in colonial breeders is important for informing conservation management of the African penguin and other threatened taxa.