Use of bird-borne radar to examine shearwater interactions with legal and illegal fisheries.

Seabirds interact with fishing vessels to consume fishing discards and baits, sometimes resulting in incidental capture (bycatch) and the death of the bird, which has clear conservation implications. To understand seabird-fishery interactions at large spatiotemporal scales, researchers are increasing their use of simultaneous seabird and fishing vessel tracking. However, vessel tracking data can contain gaps due to technical problems, illicit manipulation, or lack of adoption of tracking monitoring systems. These gaps might lead to underestimating the fishing effort and bycatch rates and jeopardize the effectiveness of marine conservation. We deployed bird-borne radar detector tags capable of recording radar signals from vessels. We placed tags on 88 shearwaters (Calonectris diomedea, Calonectris borealis, and Calonectris edwardsii) that forage in the northwestern Mediterranean Sea and the Canary Current Large Marine Ecosystem. We modeled vessel radar detections registered by the tags in relation to gridded automatic identification system (AIS) vessel tracking data to examine the spatiotemporal dynamics of seabird-vessel interactions and identify unreported fishing activity areas. Our models showed a moderate fit (area under the curve >0.7) to vessel tracking data, indicating a strong association of shearwaters to fishing vessels in major fishing grounds. Although in high-marine-traffic regions, radar detections were also driven by nonfishing vessels. The tags registered the presence of potential unregulated and unreported fishing vessels in West African waters, where merchant shipping is unusual but fishing activity is intense. Overall, bird-borne radar detectors showed areas and periods when the association of seabirds with legal and illegal fishing vessels was high. Bird-borne radar detectors could improve the focus of conservation efforts.

Uso de radares en aves para analizar las interacciones de las pardelas con las pesquerías legales e ilegales Resumen Las aves marinas interactúan con los barcos pesqueros para consumir los cebos y lo que descartan, lo que a veces resulta en la captura accesoria y la muerte del ave, por lo que esto tiene implicaciones claras para la conservación. Los investigadores cada vez usan más el rastreo simultáneo de las aves marinas y los barcos pesqueros para comprender las interacciones aves marinas ­ pesquerías a gran escala espaciotemporal. Sin embargo, los datos del rastreo de barcos pueden incluir vacíos por problemas técnicos, manipulación ilícita o porque no adoptan sistemas para monitorear el rastreo. Estos vacíos pueden llevar a subestimar el esfuerzo de pesca y las tasas de captura accesoria y a comprometer la efectividad de la conservación marina. Desplegamos marcas detectoras de radar encima de aves capaces de registrar las señales de radar de los barcos. Colocamos estas marcas en 88 pardelas (Calonectris diomedea, C. borealis, y C. edwardsii) que forrajean en el noroeste del Mar Mediterráneo y el Gran Ecosistema Marino de Canarias. Modelamos las detecciones del radar de los barcos registradas por las marcas en relación con los datos reticulados de rastreo de barcos del sistema de identificación automático (AIS) para analizar las dinámicas espaciotemporales de las interacciones aves marinas­barcos e identificar áreas con actividad pesquera no reportada. Nuestros modelos mostraron un ajuste moderado (área bajo la curva > 0.7) a los datos de rastreo de barcos, lo que indica una fuerte asociación entre las pardelas y los barcos en los principales sitios de pesca, aunque en las regiones con alto tránsito de barcos las detecciones por radar también fueron causadas por barcos no pesqueros. Las marcas registraron la presencia del potencial de barcos pesqueros sin regular y sin reportar en aguas del oeste de África, en donde los buques mercantes son poco comunes pero la actividad pesquera es intensa. En general, los detectores por radar en las aves mostraron áreas y periodos en donde la asociación entre las aves marinas y los barcos pesqueros legales e ilegales es alta. Estos detectores por radar podrían mejorar el enfoque de los esfuerzos de conservación.

Animal behaviour on the move: the use of auxiliary information and semi-supervision to improve behavioural inferences from Hidden Markov Models applied to GPS tracking datasets.

BACKGROUND: State-space models, such as Hidden Markov Models (HMMs), are increasingly used to classify animal tracks into behavioural states. Typically, step length and turning angles of successive locations are used to infer where and when an animal is resting, foraging, or travelling. However, the accuracy of behavioural classifications is seldom validated, which may badly contaminate posterior analyses. In general, models appear to efficiently infer behaviour in species with discrete foraging and travelling areas, but classification is challenging for species foraging opportunistically across homogenous environments, such as tropical seas. Here, we use a subset of GPS loggers deployed simultaneously with wet-dry data from geolocators, activity measurements from accelerometers, and dive events from Time Depth Recorders (TDR), to improve the classification of HMMs of a large GPS tracking dataset (478 deployments) of red-billed tropicbirds (Phaethon aethereus), a poorly studied pantropical seabird. METHODS: We classified a subset of fixes as either resting, foraging or travelling based on the three auxiliary sensors and evaluated the increase in overall accuracy, sensitivity (true positive rate), specificity (true negative rate) and precision (positive predictive value) of the models in relation to the increasing inclusion of fixes with known behaviours. RESULTS: We demonstrate that even with a small informed sub-dataset (representing only 9% of the full dataset), we can significantly improve the overall behavioural classification of these models, increasing model accuracy from 0.77 ± 0.01 to 0.85 ± 0.01 (mean ± sd). Despite overall improvements, the sensitivity and precision of foraging behaviour remained low (reaching 0.37 ± 0.06, and 0.06 ± 0.01, respectively). CONCLUSIONS: This study demonstrates that the use of a small subset of auxiliary data with known behaviours can both validate and notably improve behavioural classifications of state space models of opportunistic foragers. However, the improvement is state-dependant and caution should be taken when interpreting inferences of foraging behaviour from GPS data in species foraging on the go across homogenous environments.

DNA metabarcoding to assess prey overlap between tuna and seabirds in the Eastern tropical Atlantic: Implications for an ecosystem-based management.

Overfishing has been drastically changing food webs in marine ecosystems, and it is pivotal to quantify these changes at the ecosystem level. This is especially important for ecosystems with a high diversity of top predators such as the Eastern Atlantic marine region. In this work we used high-throughput sequencing methods to describe the diet of the two most abundant tuna species, the Skipjack tuna (Katsuwonus pelamis) and the Yellowfin tuna (Thunnus albacares), highly targeted by fisheries off west Africa. We also explored prey diversity overlap between these tuna species and the seabird species breeding in Cabo Verde that are most likely to share prey preferences and suffer from bycatch, the Brown booby (Sula leucogaster) and Cape Verde shearwater (Calonectris edwardsii). Overall, the diet of both tuna species was more diverse than that of seabirds. Skipjack tuna diet was dominated by prey from lower trophic levels, such as krill, anchovies, and siphonophores, while the Yellowfin tuna diet was mainly based on epipelagic fish such as flying and halfbeak fishes. Some of the most abundant prey families detected in the Yellowfin tuna diet were shared with both seabird species, resulting in a high prey diversity overlap between this tuna species and seabirds These results have implications for the management of tuna fisheries in the Eastern Tropical Atlantic, because a large decrease of both tuna species might have cascading effects on both primary and secondary consumer levels, and the decrease of these underwater predators may have implications on the viability of tropical seabird populations.

Will climate change affect the survival of tropical and subtropical species? Predictions based on Bulwer's petrel populations in the NE Atlantic Ocean.

Climate change has repeatedly been shown to impact the demography and survival of marine top predators. However, most evidence comes from single populations of widely distributed species, limited mainly to polar and subpolar environments. Here, we aimed to evaluate the influence of environmental conditions on the survival of a tropical and migratory seabird over the course of its annual cycle. We used capture-mark-recapture data from three populations of Bulwer's petrel (Bulweria bulwerii) spread across the NE Atlantic Ocean, from the Azores, Canary, and Cabo Verde Islands (including temperate to tropical zones). We also inferred how the survival of this seabird might be affected under different climatic scenarios, defined by the Intergovernmental Panel on Climate Change. Among the environmental variables whose effect we evaluated (North Atlantic Oscillation index, Southern Oscillation Index, Sea Surface Temperature [SST] and wind speed), SST estimated for the breeding area and season was the variable with the greatest influence on adult survival. Negative effects of SST increase emerged across the three populations, most likely through indirect trophic web interactions. Unfortunately, our study also shows that the survival of Bulwer's petrel will be profoundly affected by the different scenarios of climate change, even with the most optimistic trajectory involving the lowest greenhouse gas emission. Furthermore, for the first time, our study predicts stronger impacts of climate change on tropical populations than on subtropical and temperate ones. This result highlights the devastating effect that climate change may also have on tropical areas, and the importance of considering multi-population approaches when evaluating its impacts which may differ across species distributions.

Sea surface temperature, rather than land mass or geographic distance, may drive genetic differentiation in a species complex of highly dispersive seabirds.

Seabirds, particularly Procellariiformes, are highly mobile organisms with a great capacity for long dispersal, though simultaneously showing high philopatry, two conflicting life-history traits that may lead to contrasted patterns of genetic population structure. Landmasses were suggested to explain differentiation patterns observed in seabirds, but philopatry, isolation by distance, segregation between breeding and nonbreeding zones, and oceanographic conditions (sea surface temperatures) may also contribute to differentiation patterns. To our knowledge, no study has simultaneously contrasted the multiple factors contributing to the diversification of seabird species, especially in the gray zone of speciation. We conducted a multilocus phylogeographic study on a widespread seabird species complex, the little shearwater complex, showing highly homogeneous morphology, which led to considerable taxonomic debate. We sequenced three mitochondrial and six nuclear markers on all extant populations from the Atlantic (lherminieri) and Indian Oceans (bailloni), that is, five nominal lineages from 13 populations, along with one population from the eastern Pacific Ocean (representing the dichrous lineage). We found sharp differentiation among populations separated by the African continent with both mitochondrial and nuclear markers, while only mitochondrial markers allowed characterizing the five nominal lineages. No differentiation could be detected within these five lineages, questioning the strong level of philopatry showed by these shearwaters. Finally, we propose that Atlantic populations likely originated from the Indian Ocean. Within the Atlantic, a stepping-stone process accounts for the current distribution. Based on our divergence time estimates, we suggest that the observed pattern of differentiation mostly resulted from historical and current variation in sea surface temperatures.

Global political responsibility for the conservation of albatrosses and large petrels.

Migratory marine species cross political borders and enter the high seas, where the lack of an effective global management framework for biodiversity leaves them vulnerable to threats. Here, we combine 10,108 tracks from 5775 individual birds at 87 sites with data on breeding population sizes to estimate the relative year-round importance of national jurisdictions and high seas areas for 39 species of albatrosses and large petrels. Populations from every country made extensive use of the high seas, indicating the stake each country has in the management of biodiversity in international waters. We quantified the links among national populations of these threatened seabirds and the regional fisheries management organizations (RFMOs) which regulate fishing in the high seas. This work makes explicit the relative responsibilities that each country and RFMO has for the management of shared biodiversity, providing invaluable information for the conservation and management of migratory species in the marine realm.

Year-round individual specialization in the feeding ecology of a long-lived seabird.

Many generalist species are composed of individuals varying in the size of their realized niches within a population. To understand the underlying causes and implications of this phenomenon, repeated samplings on the same individuals subjected to different environmental conditions are needed. Here, we studied individual specialization of feeding strategies in breeding and non-breeding grounds of Cory's shearwaters (Calonectris borealis) for 2-8 years, and its relationship with fitness. Individuals were relatively flexible in non-breeding destinations, but specialized in diet, habitat use and daily activity across years. Daily activity was also consistent throughout the year for the same individual, suggesting that it is driven by individual constraints, whereas individual diet and habitat use changed between breeding and non-breeding grounds, indicating that these specializations may be learned at each area. Moreover, individuals that were intermediate specialized in their diet tended to show higher breeding success than those with weakly and highly specialized diets, suggesting stabilizing selection. Overall, this study suggests that the development of individual specialization is more flexible than previously thought, i.e. it emerges under specific environmental conditions and can develop differently when environmental conditions vary. However, once established, individual specialization may compromise the ability of individuals to cope with environmental stochasticity.

Does sexual segregation occur during the nonbreeding period? A comparative analysis in spatial and feeding ecology of three Calonectris shearwaters.

Sexual segregation (SS) is widespread among animal taxa, with males and females segregated in distribution, behavior, or feeding ecology but so far, most studies on birds have focused on the breeding period. Outside this period, the relevance of segregation and the potential drivers of its persistence remain elusive, especially in the marine environment, where animals can disperse over vast areas and are not easily observed. We evaluated the degree of SS in spatio-temporal distribution and phenology, at-sea behavior, and feeding ecology during the nonbreeding period among three closely related shearwaters: Scopoli's, Cory's, and Cape Verde shearwaters (Calonectris diomedea, C. borealis, and C. edwardsii, respectively). We tracked 179 birds (92 males and 87 females) from 2008 to 2013 using geolocation-immersion loggers and collected the 13th secondary remige (molted in winter) for stable isotope analyses as a proxy of trophic level and diet. The global nonbreeding distribution did not differ between sexes for the three species, but one specific nonbreeding area was visited only by males. Cory's shearwater males remained in areas closer to the colony in a larger proportion compared to females and returned earlier to the colony, probably to defend their nests. Males presented a slightly lower nocturnal flying activity and slightly (but consistently) higher isotopic values of δ13C and δ15N compared to females. These differences suggest subtle sexual differences in diet and a slightly higher trophic level in males, but the extent to which sexual dimorphism in bill size can determine them remains unclear. Our study showed that SS in ecological niche in seabirds can persist year-round consistently but at a different extent when comparing the breeding and nonbreeding periods. Based on our findings, we propose that SS in these seabird species might have its origin in an ecological specialization derived from the different roles of males and females during reproduction, rather than from social dominance during the nonbreeding period.

Using marine isoscapes to infer movements of oceanic migrants: The case of Bulwer's petrel, Bulweria bulwerii, in the Atlantic Ocean.

Studying the movements of oceanic migrants has been elusive until the advent of several tracking devices, such as the light-level geolocators. Stable isotope analysis (SIA) offers a complementary approach to infer areas used year-round, but its suitability in oceanic environments remains almost unexplored. To evaluate SIA as a tool for inferring movements of oceanic migrants, we sampled an oceanic seabird, the Bulwer's petrel, Bulweria bulwerii, in four breeding colonies spread along its Atlantic distribution. We first studied the species moulting pattern from 29 corpses collected in the colonies. Secondly, based on this moult knowledge, we selected three feathers from tracked birds to infer their breeding and non-breeding grounds using SIA: the 1st primary (P1), the 8th secondary (S8) and the 6th rectrix (R6) feathers. Birds migrated to two main non-breeding areas, the Central or the South Atlantic Ocean. P1 showed similar isotopic values among petrels from different breeding colonies, suggesting this feather is replaced early in the non-breeding period in a common area used by most birds, the Central Atlantic. S8 and R6 feathers correctly assigned 92% and 81%, respectively, of the birds to their non-breeding areas, suggesting they were replaced late in season, when birds were settled in their main non-breeding grounds. Our results showed that the isotopic baseline levels of the Central and South Atlantic are propagated through the food web until reaching top predators, suggesting these ratios can be used to infer the movement of long-distance migrants among oceanic water masses.

Population size, breeding biology and on-land threats of Cape Verde petrel (Pterodroma feae) in Fogo Island, Cape Verde.

Cape Verde petrel (Pterodroma feae) is currently considered near threatened, but little is known about its population size, breeding biology and on land threats, jeopardizing its management and conservation. To improve this situation, we captured, marked and recaptured (CMR) birds using mist-nets over 10 years; measured and sexed them; monitored up to 14 burrows, deployed GPS devices on breeders and analyzed activity data of geolocators retrieved from breeders in Fogo (Cape Verde). We set cat traps over the colony and investigated their domestic/feral origin by marking domestic cats from a nearby village with transponders, by deploying GPS devices on domestic cats and by performing stable isotope analyses of fur of the trapped and domestic cats. The population of Fogo was estimated to be 293 birds, including immatures (95% CI: 233-254, CMR modelling). Based on geolocator activity data and nest monitoring we determined the breeding phenology of this species and we found biometric differences between sexes. While monitoring breeding performance, we verified a still ongoing cat predation and human harvesting. Overall, data gathered from trapped cats without transponder, cats GPS trips and the distinct isotopic values between domestic and trapped cats suggest cats visiting the colony are of feral origin. GPS tracks from breeders showed birds left and returned to the colony using the sector NE of the islands, where high level of public lights should be avoided specially during the fledging period. Main threats for the Cape Verde petrel in the remaining breeding islands are currently unknown but likely to be similar to Fogo, calling for an urgent assessment of population trends and the control of main threats in all Cape Verde Islands and uplisting its conservation status.

Global spatial ecology of three closely-related gadfly petrels.

The conservation status and taxonomy of the three gadfly petrels that breed in Macaronesia is still discussed partly due to the scarce information on their spatial ecology. Using geolocator and capture-mark-recapture data, we examined phenology, natal philopatry and breeding-site fidelity, year-round distribution, habitat usage and at-sea activity of the three closely-related gadfly petrels that breed in Macaronesia: Zino's petrel Pterodroma madeira, Desertas petrel P. deserta and Cape Verde petrel P. feae. All P. feae remained around the breeding area during their non-breeding season, whereas P. madeira and P. deserta dispersed far from their colony, migrating either to the Cape Verde region, further south to equatorial waters in the central Atlantic, or to the Brazil Current. The three taxa displayed a clear allochrony in timing of breeding. Habitat modelling and at-sea activity patterns highlighted similar environmental preferences and foraging behaviours of the three taxa. Finally, no chick or adult was recaptured away from its natal site and survival estimates were relatively high at all study sites, indicating strong philopatry and breeding-site fidelity for the three taxa. The combination of high philopatry, marked breeding asynchrony and substantial spatio-temporal segregation of their year-round distribution suggest very limited gene flow among the three taxa.

Comparing multiple criteria for species identification in two recently diverged seabirds.

Correct species identification is a crucial issue in systematics with key implications for prioritising conservation effort. However, it can be particularly challenging in recently diverged species due to their strong similarity and relatedness. In such cases, species identification requires multiple and integrative approaches. In this study we used multiple criteria, namely plumage colouration, biometric measurements, geometric morphometrics, stable isotopes analysis (SIA) and genetics (mtDNA), to identify the species of 107 bycatch birds from two closely related seabird species, the Balearic (Puffinus mauretanicus) and Yelkouan (P. yelkouan) shearwaters. Biometric measurements, stable isotopes and genetic data produced two stable clusters of bycatch birds matching the two study species, as indicated by reference birds of known origin. Geometric morphometrics was excluded as a species identification criterion since the two clusters were not stable. The combination of plumage colouration, linear biometrics, stable isotope and genetic criteria was crucial to infer the species of 103 of the bycatch specimens. In the present study, particularly SIA emerged as a powerful criterion for species identification, but temporal stability of the isotopic values is critical for this purpose. Indeed, we found some variability in stable isotope values over the years within each species, but species differences explained most of the variance in the isotopic data. Yet this result pinpoints the importance of examining sources of variability in the isotopic data in a case-by-case basis prior to the cross-application of the SIA approach to other species. Our findings illustrate how the integration of several methodological approaches can help to correctly identify individuals from recently diverged species, as each criterion measures different biological phenomena and species divergence is not expressed simultaneously in all biological traits.

Plastic debris in Mediterranean seabirds.

Plastic debris is often ingested by marine predators and can cause health disorders and even death. We present the first assessment of plastic ingestion in Mediterranean seabirds. We quantified and measured plastics accumulated in the stomach of 171 birds from 9 species accidentally caught by longliners in the western Mediterranean from 2003 to 2010. Cory's shearwaters (Calonectris diomedea) showed the highest occurrence (94%) and large numbers of small plastic particles per affected bird (on average N = 15.3 ± 24.4 plastics and mass = 23.4 ± 49.6 mg), followed by Yelkouan shearwaters (Puffinus yelkouan, 70%, N = 7.0 ± 7.9, 42.1 ± 100.0 mg), Balearic shearwaters (Puffinus mauretanicus, 70%, N = 3.6 ± 2.9, 5.5 ± 9.7 mg) and the rest of species (below 33%, N = 2.7, 113.6 ± 128.4 mg). Plastic characteristics did not differ between sexes and were not related to the physical condition of the birds. Our results point out the three endemic and threatened shearwater species as being particularly exposed to plastic accumulation.