Species‐specific foraging strategies and segregation mechanisms of sympatric Antarctic fulmarine petrels throughout the annual cycle

Determining the year‐round distribution and behaviour of birds is necessary for a better understanding of their ecology and foraging strategies. Petrels form an important component of the high‐latitude seabird assemblages in terms of species and individuals. The distribution and foraging ecology of three sympatric fulmarine petrels (Southern Fulmar Fulmarus glacialoides, Cape Petrel Daption capense and Snow Petrel Pagodroma nivea) were studied at Adélie Land, East Antarctica, by combining information from miniaturized saltwater immersion geolocators and stable isotopes from feathers. During the breeding season at a large spatial scale (c. 200 km), the three species overlapped in their foraging areas located in the vicinity of the colonies but were segregated by their diet and trophic level, as indicated by the different chick δ¹⁵N values that increased in the order Cape Petrel < Southern Fulmar < Snow Petrel. During the non‐breeding season, the three fulmarines showed species‐specific migration strategies along a wide latitudinal gradient. Snow Petrels largely remained in ice‐associated Antarctic waters, Southern Fulmars targeted primarily the sub‐Antarctic zone and Cape Petrels migrated further north. Overall, birds spent less time in flight during the non‐breeding period than during the breeding season, with the highest percentage of time spent sitting on the water occurring during the breeding season and at the beginning of the non‐breeding period before migration. This activity pattern, together with the δ¹³C values of most feathers, strongly suggests that moult of the three fulmarine petrels occurred at that time in the very productive high Antarctic waters, where birds fed on a combination of crustaceans and fish. The study highlights different segregating mechanisms that allow the coexistence of closely related species, specifically, prey partitioning during the breeding season and spatial segregation at sea during the non‐breeding season.     

Life history buffering in Antarctic mammals and birds against changing patterns of climate and environmental variation

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

The separation of Pterodroma madeira (Zino's Petrel) from Pterodroma feae (Fea's Petrel) (Aves: Procellariidae)

The taxonomic status of petrels from the North East Atlantic has long been a matter of debate. Breeding colonies of petrels occurring on the islands of Madeira, Bugio and Cape Verde were originally thought to be outlying populations of the polytypic species Pterodroma mollis . Subsequent taxonomic treatments have varied considerably in their classification of birds from these islands. The petrel populations on Madeira and Bugio represent some of Europe's rarest breeding birds and their exact species designation, and hence relation to conservation mandates, is a question of considerable importance. In this study we use molecular techniques alongside more traditional taxonomic characters to confirm the existence of two species of the genus Pterodroma in the Archipelago of Madeira. We also discuss identification of these species in the field and the implications for their conservation management.     

Skuas (Stercorarius spp.) moult body feathers during both the breeding and inter‐breeding periods: implications for stable isotope investigations in seabirds

Seabirds are mostly thought to moult during the inter‐breeding period and the isotopic values of their feathers are often therefore assumed to relate to their assimilated diet during such periods. We observed Brown Skuas Stercorarius antarcticus lonnbergi and South Polar Skuas Stercorarius maccormicki moulting on a breeding site at King George Island, Antarctica. This raises concerns about the reliability of using stable isotopes in feathers to infer feeding localities of birds during the inter‐breeding period. We analysed the δ¹³C and δ¹⁵N values of growing and fully grown body feathers collected from the same individuals. For both species, δ¹³C values of growing feathers indicated feeding areas in the Antarctic zone (breeding grounds), whereas most fully grown feathers (100% for South Polar Skuas and 93.3% for Brown Skuas) could be assigned to northern latitudes (non‐breeding grounds). However, a few fully grown body feathers of Brown Skuas (6.7% of the feathers, belonging to two birds) showed isotopic values that indicated moult in the Antarctic zone. As the growth period of those feathers was unknown, they could not be used with confidence to depict the foraging behaviour of the birds during the non‐breeding period. Although precautions must be taken when inferring dietary information from feathers in seabirds where the moulting pattern is unknown, this study shows that if the development stage of a feather (growing/fully grown) is identified, then dietary information from both breeding and non‐breeding seasons can be obtained on the same individual birds.     

Cool,cold or colder? Spatial segregation of prions and blue petrels is explained by differences in preferred sea surface temperatures

The Southern Ocean provides one of the largest environmental gradients on Earth that lacks geographical barriers, and small but highly mobile petrels living there may offer fine models of evolution of diversity along environmental gradients. Using geolocation devices, we investigated the winter distribution of closely related petrel species breeding sympatrically in the southern Indian Ocean, and applied ecological niche models to compare environmental conditions in the habitat used. We show that thin-billed prions (Pachyptila belcheri), Antarctic prions (Pachyptila desolata) and blue petrels (Halobaena caerulea) from the Kerguelen archipelago in the southern Indian Ocean segregate latitudinally, sea surface temperature being the most important variable separating the distribution of the species. Antarctic prions spent the winter north of the Polar Front in temperate waters, whereas blue petrels were found south of the Polar Front in Antarctic waters. Thin-billed prions preferred intermediate latitudes and temperatures. Stable isotope values of feathers reflected this near complete niche separation across an ecological gradient that spans large scales, and suggest evolutionary isolation by environment. In pelagic seabirds that exploit large areas of ocean, spatial niche partitioning may not only facilitate coexistence among ecologically similar species, but may also have driven their evolution in the absence of geographical barriers.     

Wilson's Storm Petrels Oceanites oceanicus Recognise the Olfactory Signature of Their Mate

Chemical signals in birds have rarely been considered as recognition cues. Nevertheless, recent experiments showed that several petrel species are able to recognize their nest by smell, and in at least one species even their mate. But the use of smell may be different across the petrel species and olfactory nest recognition appears to be dependent on species’ breeding biology. To increase our knowledge of individual olfactory recognition in petrels and the relationships between breeding biology and use of smell, we tested Wilson’s storm petrels Oceanites oceanicus in Antarctica. In previous experiments, these birds failed to home if rendered anosmic, but the method employed to obtain anosmia (potentially stressing birds) and the fact that they breed in 24‐h daylight suggest that they might use visual, rather than olfactory, cues to recognize their nest. Our birds were tested in T‐maze experiments where nest odours or partner odours were presented. Wilson’s storm petrels preferred odours of their own nest and mate. Results on olfactory nest recognition confirm and complete previous results, viz. anosmic Wilson’s storm petrels do not home. Storm petrels olfactory mate recognition suggests that this ability may be widespread in burrowing petrels and implements olfactory nest recognition.     

An effective method for trapping scavenging seabirds at sea

ABSTRACT Most studies of seabirds that involve trapping and marking birds are carried out at breeding colonies. This bias toward the breeding period and colony‐based research is partially caused by difficulties in capturing birds at sea. From 2005 to 2007, we used a cast net thrown by hand from a fishing boat to capture albatrosses and petrels at sea in the southwestern Atlantic Ocean. About 500 birds of 13 species were captured, ranging in size from the 30‐g Wilson's Storm‐petrel (Oceanites oceanicus) to the 10‐kg Wandering Albatross (Diomedea exulans). Cast nets are potentially useful for capturing any seabird that can be attracted close to fishing vessels by bait, such as sardines, squid, offal, or shark liver, thrown into the water. Our method was most effective for capturing bold species, such as Cape (Daption capense), Spectacled (Procellaria conspicillata), and White‐chinned (Procellaria aequinoctialis) petrels and Great Shearwaters (Puffinus gravis), but was not effective for capturing shy species, such as Cory's (Calonectris diomedea), Cape Verde (Calonectris edwardsii) and Manx (Puffinus puffinus) shearwaters, species that rarely sit on the water, such as Wilson's Storm‐petrels, Thin‐billed (Pachyptila belcheri) and Antarctic (Pachyptila desolata) prions and Atlantic Petrels (Pterodroma incerta), and species with excellent diving abilities, such as Sooty Shearwaters (Puffinus griseus). For many species of seabirds, cast nets would likely be more efficient for capturing large numbers of birds than other methods, such as hoop nets.     

Novel tri‐isotope ellipsoid approach reveals dietary variation in sympatric predators

Sympatric species may partition resources to reduce competition and facilitate co‐existence. While spatial variation and specialization in feeding strategies may be prevalent among large marine predators, studies have focussed on sharks, birds, and marine mammals. We consider for the first time the isotopic niche partitioning of co‐occurring, teleost reef predators spanning multiple families. Using a novel tri‐isotope ellipsoid approach, we investigate the feeding strategies of seven of these species across an atoll seascape in the Maldives. We demonstrate substantial spatial variation in resource use of all predator populations. Furthermore, within each area, there was evidence of intraspecific variation in feeding behaviors that could not wholly be attributed to individual body size. Assessing species at the population level will mask these intraspecific differences in resource use. Knowledge of resource use is important for predicting how species will respond to environmental change and spatial variation should be considered when investigating trophic diversity.     

One House Two Families: Petrel Squatters Get a Sniff of Low‐Cost Breeding Opportunities

Burrowing is a widespread nesting behaviour, found in vertebrates and invertebrates. It is particularly common in small procellariiform seabirds such as blue petrels (Halobaena caerulea) and Antarctic prions (Pachyptila desolata), two closely related petrel species. However, digging a burrow is costly and alternative strategies may evolve. Accordingly, blue petrel males can adopt two alternative nesting strategies: digging a new burrow or squatting in an empty one. Importantly, a blue petrel squatter arriving at the colony to breed is more likely to find empty Antarctic prion burrows than empty blue petrel burrows, since the former species only start breeding a month later. However, squatting in a prion’s burrow is risky for blue petrels as the legitimate owner very often returns and claims the burrow back, thus ruining the squatter’s breeding attempt. We present here results of a survey of two sympatric colonies of blue petrels and Antarctic prions on Kerguelen Island. Our data show that blue petrel squatters preferentially occupy blue petrel empty burrows. To investigate potential underlying mechanisms behind this preference, we used a simple Y‐maze design to show that blue petrels can discriminate and prefer their specific odour over the prion odour. Our results confirm the existence of alternative burrowing strategies in blue petrels and suggest that squatters could use olfaction to avoid the less suitable Antarctic prion burrows.     

Greater flamingos Phoenicopterus roseus are partial capital breeders

Capital breeding refers to a strategy in which birds use body stores for egg formation, whereas income breeders obtain all resources for egg formation at breeding sites. Capital breeding should occur more in large‐bodied species because the relative cost of carrying stores for egg formation becomes smaller with increasing body size. Based on a comparison between stable isotopes of carbon and nitrogen in potential prey at wintering sites and eggs, we examined whether greater flamingos use nutrients stored earlier in the year for egg production. Our results suggest that the greater flamingo is a partial income breeder, since prey for egg formation were obtained both in overwintering sites and at the breeding site. This may be because there are selective pressures for nesting females to lay soon after arriving at the breeding site, which may be facilitated by arriving at the breeding site with developed ovarian follicles.     

Attraction of petrels to artificial lights in the Canary Islands: effects of the moon phase and age class

The extent and intensity of artificial night lighting has increased with urban development worldwide. The resulting light pollution is responsible for mortality among many Procellariiformes species which show nocturnal activity on their breeding grounds. Here, we report light‐induced mortality of Procellariiformes during a 9‐year study (1998–2006) on Tenerife, the largest island of the Canary archipelago. A total of 9880 birds from nine species were found grounded, the majority being Cory’s Shearwaters Calonectris diomedea (93.4%). For this species the majority of grounded birds were fledglings (96.4%), which fall apparently while leaving their nesting colony for the first time; for the smaller species (storm‐petrels) adult birds were more often grounded than fledglings. For almost all species, grounding showed a seasonal pattern linked with their breeding cycle. Certain phases of the moon influenced grounding of Cory’s Shearwater, with the extent of grounding being reduced during phases of full moon. The percentage of fledglings attracted to lights in relation to the fledglings produced annually varied between species and years (0–1.3% for the Madeiran Storm‐petrel Oceanodroma castro; 41–71% for Cory’s Shearwater). Mean adult mortality rates also varied between species (from 0.4% for the European Storm‐petrel Hydrobates pelagicus and the Cory’s Shearwater, to 2.3% for the Manx Shearwater Puffinus puffinus). Here we show that light‐induced mortality rates are of concern, at least for petrels and small shearwaters. Thanks to efforts involving civil cooperation, 95% of grounded birds have been returned to the wild. To minimize the impact of artificial lights on petrels we recommend several conservation measures: continuing rescue campaigns, alteration of light signatures and reduction of light emissions during the fledging peaks. Furthermore, we recommend that a monitoring program for petrel populations be implemented, as well as further studies to assess the fate of released fledglings and continued research to address why petrels are attracted to lights.     

Playback experiments indicate absence of vocal recognition among temporally and geographically separated populations of Madeiran Storm-petrels Oceanodroma castro

A number of lines of evidence suggest that temporally segregated sympatric populations of Madeiran Storm‐petrels Oceanodroma castro breeding in the Azores are reproductively isolated and morphologically and genetically distinct from each other. Within the Galapagos Islands, similar sympatric populations may also be isolated from each other, as individuals are not known to switch breeding seasons. The taxonomic relationships among populations of this species that are seasonally and spatially separated are unclear and in need of revision. In this study, playback experiments were used to determine the level of vocal response among prospecting Madeiran Storm‐petrels at colonies in the Azores, Galapagos and Cape Verde islands to recordings from different populations. Vocalizations of all populations studied here differ in their structural characteristics and in all but one case prospecting Storm‐petrels showed far greater response to playback of burrow calls from their own colony type than to calls recorded at other seasonally or geographically distinct colonies. Additionally, the level of response to foreign colony types was no different to playback of vocalizations of an unrelated control species present at the same location. Although not all combinations of geographical and seasonal populations could be examined, the finding that prospecting hot‐season (breeding April–August) Storm‐petrels in the Azores did not differ in their response level to playback from Azores cool‐season (breeding August–March) storm‐petrels and Cory's Shearwaters Calonectris diomedea is of particular significance and suggests the existence of a pre‐mating isolation mechanism that would prevent interbreeding between these two sympatric populations. Furthermore, Azores hot‐season Storm‐petrels showed a similar absence of response to playback from Galapagos dry‐season (May–July) populations, indicating that they are also taxonomically distant from this group. Madeiran Storm‐petrels in the Cape Verde islands showed a low response rate to Azores hot‐season vocalizations, which did not differ from the response to unrelated controls. These data provide further evidence that the hot‐season Azores population represents a distinct taxon that is reproductively isolated from the sympatrically breeding cool‐season population, as well as from more distant populations in the Cape Verde and Galapagos islands.     

Use of cloacal sexing techniques in mark-recapture estimates of breeding population size in Wilson's stormpetrel Oceanites oceanicus at South Georgia

Summary Many stormpetrel species breed in habitats where their populations cannot be estimated by direct counts of burrows or birds; mark-recapture experiments have been confounded by the presence of many wandering non-breeders. With a population of Wilson's Stormpetrel Oceanites oceanicus at Bird Island, South Georgia, we tried to estimate the proportion of breeding females in samples obtained during a mark-recapture experiment. These were identified by measurements of the cloaca, which greatly enlarges at egg-laying. A concurrent experiment with individually marked birds determined that breeding females could be discriminated from males and non-breeders for c 30 days after laying. The technique is probably applicable to other petrels, though it will work best with those that lay most synchronously. The overall population estimate was 4841–5515 birds (SE 856–1417); estimates of breeding females gave a population of 2300 paris early in the incubation period and 1400 pairs near hatching.     

Pelagic distribution and numbers of the Antarctic petrelThalassoica antarctica in the Weddell Sea during spring

Estimates of the population size of the Antarctic petrelThalassoica antarctica are hampered by its breeding locations in remote nunatak areas of the Antarctic continent. Studies at sea can provide additional information on numbers. Seabird censuses during spring 1992 showed few Antarctic petrels in the western part of the Weddell Sea, but high numbers east of the South Sandwich Islands. The vast majority of the birds occurred in a band of 300 km north to 150 km south of the outer ice edge. Numbers at sea fluctuated in agreement with synchronized colony attendance patterns in the pre-breeding phase and peaked shortly before egglaying when colonies are completely deserted. In this period, densities of Antarctic petrels in the marginal ice zone suggest that at least 2.7±0.5 million individuals occur in the Weddell Sea. It is likely that similar numbers occur immediately east of the study area. The distribution of the petrels matches the location, but not the size, of known breeding colonies in Dronning Maud Land and Coats Land, which suggests that important colonies in this area remain to be discovered. The observations imply that Antarctic petrels in the Weddell Sea commute over ice a surprisingly large distance of at least 2,000 km for a single pre-breeding visit to the colonies.     

Migratory connectivity in a declining bird species: using feather isotopes to inform demographic modelling

Aim Conservation programmes for endangered migratory species or populations require locating and evaluating breeding, stopover and wintering areas. We used multiple stable isotopes in two endangered European populations of wrynecks, Jynx torquilla L., to locate wintering regions and assess the degree of migratory connectivity between breeding and wintering populations. Location Switzerland and Germany. Methods We analysed stable nitrogen (δ¹⁵N), carbon (δ¹³C) and hydrogen (δD) isotopes from wing feathers from two populations of wrynecks to infer their wintering origins and to assess the strength of migratory connectivity. We tested whether variation in feather isotopic values within the Swiss population was affected by bird age and collection year and then considered differences in isotopic values between the two breeding populations. We used isotopic values of summer‐ and winter‐grown feathers to estimate seasonal distributions. Finally, we calculated a species‐specific δD discrimination factor between feathers and mean annual δD values to assign winter‐grown feathers to origin. Results Bird age and collection year caused substantial isotopic variation in winter‐grown feathers, which may be because of annually variable weather conditions, movements of birds among wintering sites and/or reflect asynchronous moulting or selection pressure. The large isotopic variance in winter‐grown feathers nevertheless suggested low migratory connectivity for each breeding population, with partially overlapping wintering regions for the two populations. Main conclusions Isotopic variance in winter‐grown feathers of two breeding populations of wrynecks and their geographical assignment point to defined, albeit overlapping, wintering areas, suggesting both leapfrog migration and low migratory connectivity. On this basis, integrative demographic models can be built looking at seasonal survival patterns with links to local environmental conditions on both breeding and wintering grounds, which may elucidate causes of declines in migratory bird species.     

Arctic waders and the capital‐income continuum: Further tests using isotopic contrasts of egg components

Birds migrating annually to high‐latitude breeding grounds may benefit from the transport of endogenous nutrient reserves that ultimately contribute to reproduction. Shorebirds represent a diverse group of Arctic breeders that typically arrive on the breeding grounds with body reserves enriched in ¹³C and ¹⁵N due to wintering and staging in marine or estuarine habitats. Such isotopic differences between endogenous macronutrient reserves and local foodwebs allow the use of stable isotopes to test for the source of nutrient allocations to eggs. We examined δ¹³C and δ¹⁵N values in lipid‐free yolk and albumen and δ¹³C values in yolk lipid of first clutches of ten species of sandpiper and plover breeding near Churchill, Manitoba, Canada in 2003. Most birds had egg isotope values indicating a C3 terrestrial biome, which fits primarily an income (exogenous) breeding strategy. Two exceptions were single sandpiper and plover with strong marine isotope values. Among species, strong positive relationships for each isotope between egg tissue components suggest that egg proteins and lipids tended to be derived from the same isotopic source. Correlations of egg δ¹³C values for lipids and proteins approached theoretical relationships expected for exogenous breeding strategies, based on captive studies. Significant positive correlations between clutch initiation date and δ¹³C values of egg lipids and albumen suggest some endogenous nutrient contributions to later laid eggs but the circumstances under which this may occur are unstudied. Where possible, we recommend that researchers use blood and fat biopsies from laying females as a means of anchoring endogenous and exogenous endpoints for modeling of each reproductive event. We encourage the isotopic analysis of egg albumen, yolk and yolk lipids among individuals and species and tests of correlations among these components as a means of inferring origins of nutrients to eggs.     

Males’ calls carry information about individual identity and morphological characteristics of the caller in burrowing petrels

Acoustic communication in burrowing petrels has been poorly studied. However, as for many other bird species, acoustic communication seems to play an essential role in social interactions during the breeding season of these seabirds. Bachelor males call from their burrow, likely to attract females, but also when vocally challenged by other males. Calling in the breeding colony exposes petrels to high predation risks and thus it should provide an important benefit. The present study focuses on the informative content of males’ calls in the blue petrel Halobaena caerulea and the Antarctic prion Pachyptila desolata, two monogamous petrel species producing a single egg per year. We tested the hypotheses that acoustic parameters of a male's calls 1) reflect phenotypic characteristics, and 2) bear an individual vocal signature. To do so, we first tested on both species the relationships between seven morphometric measurements and 11 acoustic parameters using multivariate analyses. Second, we performed a between‐class analysis and calculated the potential of individuality coding (i.e. the ratio between intra‐ and inter‐individual variabilities) for acoustic parameters in both spectral and temporal domains. Results show acoustic parameters (especially energy quartiles, call duration, and syllable or phrase rate) reflect the caller's body size, bill morphology and wing morphology in both species. Considering the seeming pertinence of wing morphology, we suggest wing area may be a more relevant trait to consider than wing length when studying soaring birds. The results support the idea that energy quartiles, phrase rate and call duration also code for individual identity. Information carried by males’ calls might play a role in social interactions, such as burrow defence (e.g. male‐male competition, neighbour‐stranger discrimination) and/or female mate choice.     

Wintering areas predict age‐related breeding phenology in a migratory passerine bird

Understanding connections between breeding, stopover and wintering grounds for long‐distance migratory birds can provide important insight into factors influencing demography and the strength of carry‐over effects among various periods of the annual cycle. Using previously described, multi‐isotope (δ¹³C, δ¹⁵N, δ²H) feather isoscapes for Africa, we identified the most probable wintering areas for house martins Delichon urbica breeding at Badajoz in southwestern Spain. We identified two most‐probable wintering areas differing in isotopic signature in west Africa. We found that the probability to winter in the isotopic cluster two was related to age and sex of individuals. Specifically, experienced males (i.e. two years or older) winter in the isotopic cluster two with a greater probability than experienced females, whereas first‐year females winter in the isotopic cluster two with a greater probability than first‐year males. In addition, wintering area was correlated with breeding phenology, with individuals wintering in the isotopic cluster two initiating their clutches earlier than those wintering in the isotopic cluster one. For birds wintering in the isotopic cluster two, there was no relationship between age and clutch initiation date. In contrast, young birds wintering in the isotopic cluster one initiated their clutches earlier than experienced birds wintering in this area. There was no significant correlation between wintering area and clutch size or the number of fledglings produced. We hypothesize that the relationship among social status, population density and winter habitat quality should be the most important driver of the carry‐over effect we found for this population.     

Systematics of the Soft-plumaged Petrel Pterodroma mollis (Procellariidae): new insight from the study of vocalizations

The taxonomic status of the populations of the polytypic Soft-plumaged Petrel Pterodroma mollis is controversial. Previous taxonomic studies used morphological characters (measurements and colouration) and univariate statistics. In this study, I add the use of multivariate statistics on morphological (biometry and colouration) and behavioural characters (vocalizations) of the petrels. Vocalizations are especially relevant in nocturnal burrowing petrels (such as the Soft-plumaged Petrel) and act as behavioural isolating mechanisms. The analysis, based on 535 calls from seven of the 11 known breeding localities of the species, showed that physical parameters of the call did not overlap at all between breeding localities in the northern and southern hemispheres but did so within each hemisphere. Conversely, the morphological analysis, based on morphometries from ten localities, displayed wide and significant intra-hemisphere variation, whereas there was no consistent difference between northern and southern parts of the breeding range. Thus analysis of the two sets of characters gave contradictory results. Calls, owing to their role in species-specific recognition in these birds, should be given priority over other characters for species separation. I suggest that only two species should be recognized, Pterodroma mollis and Pterodroma feae, with three subspecies each. I also examined behavioural affinities between P. mollislfeae groups and other gadfly petrels and found that P. mollis/feae were closest to Pterodroma cahow, whilst Pterodroma hasitata on the one hand and Pterodroma phaeopygia and Pterodroma externa (which may comprise two species instead of one) on the other belong to other groups of gadfly petrels.     

Year‐round spatial movements and trophic ecology of Trindade Petrels (Pterodroma arminjoniana)

Trindade Petrels (Pterodroma arminjoniana) are vulnerable gadfly petrels that breed on the remote Trindade Island, located ~1100 km off the Brazilian coast. Little is known about their spatial ecology, and their trophic ecology has only been described for the breeding season. We tagged four Trindade Petrels with global location sensing loggers (GLS) from October 2013 to November 2014 and sampled the blood and feathers (innermost primary and the eighth secondary) of 14 individuals to evaluate their year‐round spatial and isotopic ecology. We examined individual distributions, habitat use and suitability, activity, and isotopic values during the breeding, migration, and non‐breeding periods. Trindade Petrels used areas in the southwest Atlantic Ocean (between 10°N and 50°S in latitude) during the breeding season. They migrated through pelagic waters of the tropical Atlantic to the northwest Atlantic, where they spent the non‐breeding season. Trindade Petrels used mostly tropical to subtropical waters in areas of intermediate to high wind speeds and low marine productivity. Individuals spent more time foraging at night than during the day. During the breeding season, birds in northerly areas had higher carbon‐13 values, and birds that used more pelagic areas foraged on prey at a higher trophic level (higher nitrogen‐15 values) than those in more southern and coastal areas. Isotopic values during the breeding, migration, and non‐breeding periods differed, possibly due to differences among individuals in their at‐sea distribution throughout the year. We confirmed the non‐breeding distribution of Trindade Petrels, which was previously known only from vessel sightings and stranded birds. Our results also suggest a strong temporal segregation in the at‐sea distribution and trophic ecology between two groups of individuals, which might indicate the existence of two separate breeding populations.