Search

Morphological identification of digested prey remains from a generalist predator can be challenging, especially when attempting to match degraded remains to taxonomic keys. DNA techniques, whereby prey is sequenced and matched to large public nucleotide sequence databases, are increasingly being used to augment morphological identification. We used “metaBARFcoding” (DNA metabarcoding) to target a region of the cytochrome c oxidase subunit I (COI) mitochondrial gene to identify prey in highly digested regurgitations from Christmas shearwaters Puffinus nativitatis at Hōlanikū (Kure Atoll). Metabarcoding was used to bulk‐process 92 water samples from regurgitations collected from 2009 to 2017, providing an overview of the seabird's diet. We additionally Sanger‐sequenced 100 prey items from 50 randomly chosen regurgitations to verify that metabarcoding characterized key components of the diet. The metabarcoding technique identified 87 unique taxa from 29 families of fish and squid, spanning diverse taxa, including reef‐associated, pelagic–oceanic, and mesopelagic species. Rare prey (frequency of occurrence ≤5% of samples) constituted 66% of the species richness, demonstrating the highly diverse diet of this generalist predator. Overall, 81% of the families detected in the contemporary diet were previously documented in Christmas shearwater diets from the Northwestern Hawaiian Islands. Our results indicate that metabarcoding the COI region is useful in identifying a wide range of taxa from highly digested regurgitations, thus facilitating this approach to study seabird diets.

SummaryPelagic seabird populations have declined strongly worldwide. In the North Atlantic there was a huge reduction in seabird populations following the European colonization of the Azores, Madeira and Canary archipelagos but information on seabird status and distribution for the subtropical region of Cabo Verde is scarce, unavailable or dispersed in grey literature. We compiled and compared the historical and current distribution of all seabird species breeding in the Cabo Verde archipelago, updated their relative abundance, investigated their inland habitat preferences, and reviewed their threats. Currently, the breeding seabird community in Cabo Verde is composed of Bulwer’s Petrel Bulweria bulwerii, White-faced Storm-petrel Pelagodroma marina aedesorum, Cape Verde Shearwater Calonectris edwardsii, Cape Verde Storm-petrel Hydrobates jabejabe, Cape Verde Petrel Pterodroma feae, Boyd's Shearwater Puffinus lherminieri boydi, Brown Booby Sula leucogaster, and Red-billed Tropicbird Phaethon aethereus. One breeding species is currently extinct, the Magnificent Frigatebird Fregata magnificens. The relative abundance of Cape Verde Shearwater, Boyd’s Shearwater, Cape Verde Petrel, and Cape Verde Storm-petrel was determined from counts of their nocturnal calls in Santo Antão, São Vicente, Santa Luzia, Branco, Raso and São Nicolau. Cape Verde Petrel occurred only on mountainous islands (Santo Antão, São Nicolau, Santiago, and Fogo) from mid-to high elevations. Larger species such as the Cape Verde Shearwater and Boyd’s Shearwater exhibited a wider distribution in the archipelago, occurring close to the coastline but at lower densities on populated islands. Small procellariforms such as the Cape Verde Storm-petrel occurred at high densities only on rat-free islets and in steep areas of main islands where introduced cats and rats are unlikely to occur. The main threats to seabird populations in Cabo Verde range from predation by introduced predators, habitat alteration or destruction, and some residual human persecution.

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., We conducted a multilocus phylogeographic study on a widespread shearwater and sequenced three mitochondrial and six nuclear markers on all extant populations. We found sharp differentiation among populations separated by the African continent with both markers, while only mitochondrial markers allowed characterizing the five nominal lineages. We propose that Atlantic populations originated from the Indian Ocean. Within the Atlantic, a stepping‐stone process accounts for the current distribution.

International audience; The eradication of invasive mammals on islands is important for protecting seabird populations and insular ecosystems. However, the impacts of such eradications are insufficiently known because monitoring of potentially beneficiary species is often sporadic and limited. We performed a survey of all seabird species on Saint-Paul Island, southern Indian Ocean, 20 years after successful eradication of invasive black rat (Rattus rattus) and European rabbit (Oryctolagus cuniculus). Using complementary sampling designs including adaptive cluster sampling, stratified random sampling and entire sampling, we estimated population changes and colonization by new seabird species. A total of 13 seabird species were found breeding on Saint-Paul post-eradication compared to six before the eradication. Among the seven species that colonized the island, five (MacGillivray’s prion Pachyptila macgillivrayi, fairy prion P. turtur, white-bellied storm petrel Fregetta grallaria, Antarctic tern Sterna vittata, sooty tern Onychoprion fuscatus) had relictual populations breeding on a nearby islet, and one (brown skua Catharacta antarctica) was a new breeding species. We also found breeding subantarctic little shearwaters Puffinus elegans. For species that were breeding on the Saint Paul pre-eradication, the mean annual population growth rate was 1.030 ± 0.093 (SE). Species known to be vulnerable to rat predation (prions, great-winged petrel Pterodroma macroptera, flesh-footed shearwater Puffinus carneipes, subantarctic little shearwater, white-bellied storm petrel, Antarctic tern) had the highest population growth rates. Two decades after the eradication of invasive mammals on a remote oceanic island, seabird populations were high beneficiaries. These findings further highlight the importance of invasive mammal eradication on islands as a conservation tool. Results are encouraging for the planned eradication of invasive mammals from nearby Amsterdam Island, and suggest this will mainly benefit terns and small burrowing petrels.

Individual diet specialization is known to occur in populations of generalist predators, where specific individuals develop specialist feeding strategies. Diet specialization has been reported in many raptor species, and it may be an important driver of intraspecific population structure. Here, we quantify the diet of five breeding pairs of Peregrine Falcons Falco peregrinus from an offshore island determined from prey remains collected over four breeding seasons. Three prey species accounted for 69.8 % of total prey frequency, with Manx Shearwater Puffinus puffinus the primary prey accounting for 47.3 % by frequency and 40.8 % by biomass. Herring Gull Larus argentatus was the second most important prey species by frequency (13.8 %) and biomass (29.8 %) followed by Domestic Pigeon Columba livia (frequency = 8.7 %, biomass = 7.0 %). Predation frequency on specific prey groups varied substantially between breeding pairs and months. Two pairs specialized on Manx Shearwater, one pair specialized on Herring Gull and Manx Shearwater, with the remaining two pairs having a relatively generalist diet of Manx Shearwaters, Domestic Pigeon and small passerines. Predation on Manx Shearwaters increased throughout the breeding season with a peak in total diet frequency of 63.8 % in July, with a concurrent decrease in Herring Gull predation frequency. Higher percentage of Manx Shearwater in the diet was able to explain 87 % of the variation in a narrower dietary breadth for the Peregrine pairs. Our results suggest individual diet specialization may be important for understanding population density in insular raptor populations.

There is increasing evidence for impacts of light pollution on the physiology and behaviour of wild animals. Nocturnally active Procellariiform seabirds are often found grounded in areas polluted by light and struggle to take to the air again without human intervention. Hence, understanding their responses to different wavelengths and intensities of light is urgently needed to inform mitigation measures. Here, we demonstrate how different light characteristics can affect the nocturnal flight of Manx shearwaters Puffinus puffinus by experimentally introducing lights at a colony subject to low levels of light pollution due to passing ships and coastal developments. The density of birds in flight above the colony was measured using a thermal imaging camera. We compared number of flying shearwaters under dark conditions and in response to an artificially introduced light, and observed fewer birds in flight during ‘light-on’ periods, suggesting that adult shearwaters were repelled by the light. This effect was stronger with higher light intensity, increasing duration of ‘light-on’ periods and with green and blue compared to red light. Thus, we recommend lower light intensity, red colour, and shorter duration of ‘light-on’ periods as mitigation measures to reduce the effects of light at breeding colonies and in their vicinity.

Erysipelothrix rhusiopathiae, the causative agent of the disease erysipelas, is a gram-positive bacillus, and an opportunistic pathogen in diverse species of animals. In New Zealand, E. rhusiopathiae has killed endangered birds on offshore islands, including Kākāpō (Strigops habroptilus), Takahē (Porphyrio hochstetteri), and Kiwi (Apteryx spp.). The source of infection is uncertain, and the prevalence of E. rhusiopathiae among wild birds is currently unknown. During October 2018 to December 2018, we surveyed dead and live seabirds that visit two of New Zealand's offshore islands used for Kākāpō conservation with the goal of determining the prevalence of E. rhusiopathiae. Bone marrow from dead birds was cultivated on selective agar, and organisms cultured were identified using matrix-assisted laser desorption/ionization with time-of-flight mass spectrometry. The prevalence of E. rhusiopathiae was calculated in different species for each island. We surveyed live birds using PCR with Erysipelothrix spp.-specific primers on blood samples. The prevalence of E. rhusiopathiae in dead seabirds on Whenua Hou and Te Hauturu-o-Toi was 3.4% (3/86) and 11.4% (5/44), respectively. On Whenua Hou, E. rhusiopathiae was detected in Sooty Shearwaters (Puffinus griseus; 5.9%, 2/34) and Mottled Petrels (Pterodroma inexpectata; 2.7%, 1/36) while it was detected only in Cook's Petrels (Pterodroma cookie; 13.5%, 5/37) on Te Hauturu-o-Toi. Blood samples were collected from two seabird species; only one of 50 Mottled Petrels (2.0%) was positive for the presence of Erysipelothrix spp. Our findings confirm that burrowing seabirds are possible reservoirs of E. rhusiopathiae on both islands studied and may be the source of spillover to other species on the island. The differences in observed prevalence suggest the species composition of the reservoir of E. rhusiopathiae may vary geographically.

Significance Procellariiform seabirds homing from distant foraging locations present a natural situation in which the homing route can become obstructed by islands or peninsulas because birds will not travel long distances over land. By measuring initial orientation from Global Positioning System (GPS) tracks during homing, we found that the Manx shearwater fails to encode such obstacles while homing, implying a navigation system that encodes the direction of home rather than a learned route. Nonetheless, shearwaters timed their journeys home, implying that their navigational system provides them with information about both direction and distance home, providing evidence that for routine, yet long-distance navigation, seabirds probably ascertain homeward direction by comparing their current position and the location of home with 2 or more intersecting field gradients., While displacement experiments have been powerful for determining the sensory basis of homing navigation in birds, they have left unresolved important cognitive aspects of navigation such as what birds know about their location relative to home and the anticipated route. Here, we analyze the free-ranging Global Positioning System (GPS) tracks of a large sample (n = 707) of Manx shearwater, Puffinus puffinus, foraging trips to investigate, from a cognitive perspective, what a wild, pelagic seabird knows as it begins to home naturally. By exploiting a kind of natural experimental contrast (journeys with or without intervening obstacles) we first show that, at the start of homing, sometimes hundreds of kilometers from the colony, shearwaters are well oriented in the homeward direction, but often fail to encode intervening barriers over which they will not fly (islands or peninsulas), constrained to flying farther as a result. Second, shearwaters time their homing journeys, leaving earlier in the day when they have farther to go, and this ability to judge distance home also apparently ignores intervening obstacles. Thus, at the start of homing, shearwaters appear to be making navigational decisions using both geographic direction and distance to the goal. Since we find no decrease in orientation accuracy with trip length, duration, or tortuosity, path integration mechanisms cannot account for these findings. Instead, our results imply that a navigational mechanism used to direct natural large-scale movements in wild pelagic seabirds has map-like properties and is probably based on large-scale gradients.

Passive acoustic monitoring is increasingly being used as a cost‐effective way to study wildlife populations, especially those that are difficult to census using conventional methods. Burrow‐nesting seabirds are among the most threatened birds globally, but they are also one of the most challenging taxa to census, making them prime candidates for research into such automated monitoring platforms. Passive acoustic monitoring has the potential to determine presence/absence or quantify burrow‐nesting populations, but its effectiveness remains unclear. We compared passive acoustic monitoring, tape‐playbacks and GPS tracking data to investigate the ability of passive acoustic monitoring to capture unbiased estimates of within‐colony variation in nest density for the Manx Shearwater Puffinus puffinus. Variation in acoustic activity across 12 study plots on an island colony was examined in relation to burrow density and environmental factors across 2 years. As predicted fewer calls were recorded when wind speed was high, and on moon‐lit nights, but there was no correlation between acoustic activity and the density of breeding birds within the plots as determined by tape‐playback surveys. Instead, acoustic indices correlated positively with spatial variation in the in‐colony flight activity of breeding individuals detected by GPS. Although passive acoustic monitoring has enormous potential in avian conservation, our results highlight the importance of understanding behaviour when using passive acoustic monitoring to estimate density and distribution.

We investigated how foraging habits vary among three ecologically distinct wide-ranging seabirds. Using amino acid δ15N proxies for nutrient regime (δ15NPhe) and trophic position (Δδ15NGlu-Phe), we compared Newell’s shearwater (Puffinus newelli) and Laysan albatross (Phoebastria immutabilis) foraging habits over the past 50–100 years, respectively, to published records for the Hawaiian petrel (Pterodroma sandwichensis). Standard ellipses constructed from the isotope proxies show that inter-population and interspecific foraging segregation have persisted for several decades. We found no evidence of a shift in nutrient regime at the base of the food web for the three species. However, our data identify a trophic decline during the past century for Newell’s shearwater and Laysan albatross (probability ≥ 0.97), echoing a similar decline observed in the Hawaiian petrel. During this time, Newell’s shearwaters and Hawaiian petrels have experienced population declines and Laysan albatross has experienced range extension and apparent population stability. Counting other recent studies, a pattern of trophic decline over the past century has now been identified in eight species of pelagic seabirds that breed in the Hawaiian Islands. Because our study species forage broadly across the North Pacific Ocean and differ in morphological and behavioral traits and feeding methods, the identified trophic declines suggest a pervasive shift in food web architecture within the past century.

Seabirds care for their offspring in remote breeding colonies where foraging sites are distant and may be unpredictable, and where chicks are left unaccompanied for extended periods during their parents’ foraging trips, leaving them vulnerable to predation or starvation. One way to mitigate this risk is for individuals to coordinate parenting duties with their partner. Many biparental and cooperatively breeding species are now known to coordinate their care, though the mechanisms underlying this are not well understood. In Manx shearwaters (Puffinus puffinus), both parents alternate shifts of incubation in a coordinated manner. To resolve which processes may underlie this routine, we imposed a wing-loading handicap on parents to reduce their foraging efficiency, forcing them to choose between an extended foraging trip or to return to the nest before their condition has recovered to optimal levels. We found that handicapped parents took significantly longer trips than normal, to which their partner responded by lengthening their incubation shift, suggesting shift durations are not pre-determined. However, the duration of foraging trips and the mass at which foraging birds returned to the nest appeared to be mediated by the condition of the partner. These results suggest that while foraging trip duration is largely driven by the need for the foraging bird to recoup its own condition losses, information-transfer between the parents may facilitate a more cooperative mechanism whereby the decisions made by foraging birds still account for the condition of their partner.

Shearwaters are among the most diverse groups of Procellariiformes, and the systematics of genera Calonectris, Ardenna and Puffinus (sensu stricto) is particularly complex. Here we present the first morphology-based phylogenetic hypothesis for the group, using 104 osteological characters (23 continuous and 81 discrete) scored for 20 extant species of shearwaters, representing 62.5% of species currently recognized. Both equal and implied weighting were used for phylogenetic analyses, and the final hypothesis presented herein is the strict consensus of the most stable trees produced by the latter. Calonectris was recovered as monophyletic (Relative Bremer Support = 75), while all other members of shearwaters were included in a single, monophyletic grouping: Puffinus (sensu lato) (RBS = 76). The monophyly of Ardenna, as suggested by all molecular-based hypotheses, was not recovered, whereas a monophyletic Puffinus (sensu stricto) is present within the larger clade. The final hypothesis, wherein shearwaters fall into the sister taxa Calonectris and Puffinus (sensu lato), is consistent with earlier classifications based on osteological evidence.

Puffinus (sensu lato) The final hypothesis found all other members of shearwaters to form a single, monophyletic grouping: Puffinus (sensu lato). We did not recover the split of this clade into two subgroups (i.e. genera Ardenna and Puffinus) as suggested by molecular-based hypotheses (e.g. Austin 1996; Heidrich et al. 1998; Austin et al. 2004; Penhallurick & Wink 2004; Pyle et al. 2011). The Puffinus (sensu lato) clade, as recovered herein, is relatively well-supported (RBS = 76) and established by eight unambiguous synapomorphies, plus six optimized by either ACCTRAN or DELTRAN. Some of these characters include the distinctly curved pars intermedia of the mandible, the dorsoventrally enlarged spina externa of the sternum and the absence of fossa renalis (characters 50, 56, and 90, respectively). Despite minor conflicts, the present hypothesis recalls early morphological studies (Kuroda 1954; Wragg 1985), although the results cannot be directly compared as they did not use the same phylogenetic approach as we have; however, we can nevertheless discuss the congruence of these different studies. The need for recognition of two genera among Puffinus (sensu lato) has been suggested in many phylogenetic hypotheses based on molecular data, all using the same marker (e.g. Austin 1996; Heidrich et al. 1998; Austin et al. 2004; Penhallurick & Wink 2004). However, monophyly of the currently accepted Ardenna is not supported in the present analysis. Instead, its members form successive sister taxa to Puffinus (sensu stricto). The topology of the final hypothesis differs significantly from the molecular hypotheses, wherein Ardenna and Puffinus (sensu stricto) are shown to be two different clades, sometimes not even sister genera (e.g. Heidrich et al. 1998; Nunn & Stanley 1998; Ramirez et al. 2010; Pyle et al. 2011). Alternatively, the morphological, pre-cladistic hypotheses of Kuroda (1954) and Wragg (1985) suggested that a single ���ancestral��� taxon has sequentially given rise to each phylogenetic subgroup within the Puffinus (sensu lato) clade. The final hypothesis, (Ardenna pacifica (A. bulleri (A. creatopus (A. carneipes (A. gravis ((A. grisea + A. tenuirostris) (Puffinus nativitatis (P. puffinus (P. mauretanicus (P. opisthomelas (P. gavia (P. huttoni (P. yelkouan (P. lherminieri + P. assimilis)))))))))))))), appears to match this perception. Kuroda (1954), who studied shearwaters based on different types of data, including external morphology, osteology, and behaviour, proposed a classification with five monophyletic subgroups within Puffinus (sensu lato). According to him, A. pacifica and A. bulleri are closely related, classified as the ��� Thyellodroma subgroup���, the most ���ancestral��� group among all Puffinus (sensu lato). Wragg (1985), using only morphological characters, found the same result. Although A. pacifica and A. bulleri formed a clade in the equally weighted tree, they did not group together in the implied weighting tree, as A. pacifica appears to be the sister taxon to all other species, followed by A. bulleri. A similartopology, regarding both species, was presented in the molecular-based hypothesis by Heidrich et al. (1998). All other molecular studies support the grouping of (A. pacifica + A. bulleri), which is typically regarded as sister to a clade comprising all other species of Ardenna (e.g. Penhallurick & Wink 2004; Pyle et al. 2011; Welch et al. 2014). The next species, sister taxon to the remaining species of Puffinus (sensu lato), is A. creatopus, followed by A. carneipes. Kuroda (1954) proposed that these species be combined in the ��� Hemipuffinus subgroup���, although, in the present hypothesis, they are sequentially arranged. Wragg (1985) did not discuss this subgroup, as his proposal showed a polytomy regarding the position of A. creatopus, A. carneipes, and the clade formed by the remaining species of Puffinus (sensu lato). Molecular-based hypotheses typically consider A. creatopus and A. carneipes as sister taxa; however, the subgroup relationship among the other Ardenna species is sometimes conflicting. Most studies indicate A. gravis as sister taxon to the ��� Hemipuffinus subgroup��� (e.g. Austin et al. 2004; Penhallurick & Wink 2004; Kawakami et al. 2018), but the grouping (A. grisea (A. creatopus + A. carneipes)) was suggested by Tennyson & Shepherd (2017), and Ramirez et al. (2010) was unable to define the position of ��� Hemipuffinus ��� within the genus due to a polytomy. Although Heidrich et al. (1998) did not include A. carneipes in their molecular analyses, which could have changed their resulting trees, the topology they presented for the relationship (A. pacifica (A. bulleri + A. creatopus)) is similar to the present hypothesis. The last three Ardenna species, following Kuroda (1954), were divided into two other subgroups: ��� Ardenna ��� for A. gravis; and ��� Neonectris ��� for A. grisea and A. tenuirostris (Puffinus nativitatis was also placed in the latter). According to Kuroda (1954), ��� Ardenna ��� is closely related to the group formed by ��� Neonectris ��� and all other Puffinus (sensu stricto). As recovered herein, A. gravis is sister to ((A. grisea + A. tenuirostris) + (Puffinus (sensu stricto))). Kuroda (1954) placed P. nativitatis in his ��� Neonectris ��� subgroup based on external morphology and behaviour. Afterwards, osteological analysis by Wragg (1985) led to the hypothesis that P. nativitatis was sister to all other Puffinus (sensu stricto) species, making ��� Neonectris ��� a polyphyletic group. Wragg���s (1985) hypothesis is now supported by molecular-based studies (e.g. Austin 1996; Heidrich et al. 1998; Tennyson & Shepherd 2017), in which Puffinus (sensu lato) is split into two genera, and P. nativitatis is part of Puffinus (sensu stricto). Nevertheless, the present analysis found (A. grisea + A. tenuirostris) to be monophyletic, supported by two unambiguous synapomorphies, plus four according to either ACCTRAN or DELTRAN. Furthermore, this group appears to be the sister clade to Puffinus (sensu stricto). This relationship was previously suggested by Wragg (1985); however, it is not present in most molecular hypotheses. Austin (1996) and Heidrich et al. (1998) placed A. tenuirostris as sister taxon to (A. gravis + A. grisea), whereas other hypotheses propose (A. tenuirostris (A. grisea (A. gravis))) (e.g. Penhallurick & Wink 2004; Pyle et al. 2011; Welch et al. 2014). The conflicting placement of these species has also produced some unresolved relationships within the genus (i.e. Austin et al. 2004; Ramirez et al. 2010; Tennyson & Shepherd 2017; Kawakami et al. 2018). Molecular-based studies have consistently presented a monophyletic Puffinus (sensu stricto), regardless of its position in relation to Ardenna and Calonectris (e.g. Nunn & Stanley 1998; Austin et al. 2004; Pyle et al. 2011; Tennyson & Shepherd 2017). The monophyly of Puffinus (sensu stricto) in the present analysis is congruent with these hypotheses (RBS = 82), although its members are placed in the same larger Puffinus (sensu lato) clade as Ardenna species in both equal and implied weight analyses. In the present hypothesis, the monophyly of Puffinus (sensu stricto) was supported by 12 synapomorphies from discrete characters (i.e., nine unambiguous, plus three according to eitherACCTRAN or DELTRAN), as well as 20 from continuous characters. Among other characters discussed by Kuroda (1954), these synapomorphies include the cranially positioned apex carinae of the sternum and the slender, pointed crista cnemialis cranialis of tibiotarsus (characters 62 and 98, respectively). Puffinus nativitatis, according to the present analysis, is sister to all other Puffinus (sensu stricto) species, congruent with both morphological (e.g. Wragg 1985) and molecular-based hypotheses (e.g. Austin 1996; Nunn & Stanley 1998; Penhallurick & Wink 2004). It should be noted, however, that P. subalaris, which has been found to group with P. nativitatis as sister taxon to the remaining Puffinus (sensu stricto) (Austin et al. 2004; Pyle et al. 2011), was not included in these analyses. In the present analyses, six of nine included species of Puffinus (sensu stricto) are from the group identified as ���Manx-type shearwaters���, and two others are part of the ���Little/Audubon���s complex��� (Onley & Scofield 2007). The hypothesis for the relationship of these eight species presented herein, (Puffinus puffinus (P. mauretanicus (P. opisthomelas (P. gavia (P. huttoni (P. yelkouan (P. lherminieri + P. assimilis))))))), is highly congruent with earlier morphological studies (Kuroda 1954; Wragg 1985). Hypotheses based on molecular data have consistently shown some phylogenetic associations among these species; however, their placement within the genus is unclear. In most studies, (P. gavia + P. huttoni) is the sister group to the remaining species (e.g. Austin et al. 2004; Pyle et al. 2011; Tennyson & Shepherd 2017), yet some conflicts can be found (e.g. Heidrich et al. 1998). In the implied-weight hypothesis herein, P. puffinus occupies this position, sequentially followed by P. mauretanicus. In the molecular-based analyses, P. mauretanicus is sister to P. yelkouan (e.g. Austin 1996; Pyle et al. 2011; Kawakami et al. 2018), but in the present hypothesis, P. yelkouan is sister to (P. lherminieri + P. assimilis), whereas P. mauretanicus appears to fall in between P. puffinus and P. opisthomelas. Heidrich et al. (1998) and Austin et al. (2004) have, alternatively, suggested a monophyletic (P. puffinus (P. mauretanicus + P. yelkouan)). Regarding P. opisthomelas, the species is, herein, placed between P. mauretanicus and P. gavia, but in some molecular hypotheses it is found among species from the ���Little-Audubon���s complex��� (e.g. Austin et al. 2004; Pyle et al. 2011; Tennyson & Shepherd 2017). The only two members of this group included in the present analyses, P. lherminieri and P. assimilis, are sister taxa, grouped in the least inclusive clade of the final hypothesis., Published as part of Salvagni, Thamara & Carlos, Caio J., 2021, Phylogenetic relationshipsof the shearwater taxaPuffinus (sensu lato) and Calonectris (Aves: Procellariiformes: Procellariidae) based on osteological characters, pp. 243-292 in Zootaxa 4963 (2) on pages 272-273, DOI: 10.11646/zootaxa.4963.2.2, http://zenodo.org/record/4700791, {"references":["Austin, J. J. (1996) Molecular phylogenetics of Puffinus shearwaters: preliminary evidence from mitochondrial cytochrome b gene sequences. Molecular Phylogenetics and Evolution, 6 (1), 77 - 88. https: // doi. org / 10.1006 / mpev. 1996.0060","Heidrich, P., Amengual, J. & Wink, M. (1998) Phylogenetic relationships in Mediterranean and North Atlantic Puffinus shearwaters (Aves: Procellariidae) based on nucleotide sequences of mtDNA. Biochemical Systematics and Ecology, 26 (2), 145 - 170. https: // doi. org / 10.1016 / S 0305 - 1978 (97) 00085 - 9","Austin, J. J., Bretagnolle, V. & Pasquet, E. (2004) A global molecular phylogeny of the small Puffinus shearwaters and implications for systematics of the Little-Audubon's shearwater complex. The Auk, 121 (3), 847 - 864. https: // doi. org / 10.1093 / auk / 121.3.847","Penhallurick, J. & Wink, W. (2004) Analysis of the taxonomy and nomenclature of the Procellariiformes based on complete nucleotide sequences of the mitochondrial cytochrome b gene. Emu, 104 (2), 125 - 147. https: // doi. org / 10.1071 / MU 01060","Pyle, P., Welch, A. J. & Fleischer, R. C. (2011) A new species of shearwater (Puffinus) recorded from Midway Atoll, northwestern Hawaiian Islands. The Condor, 113 (3), 518 - 527. https: // doi. org / 10.1525 / cond. 2011.100117","Kuroda, N. H. (1954) On the Classification and Phylogeny of the Order Tubinares, Particularly the Shearwaters (Puffinus), with Special Considerations on their Osteology and Habit Differentiation. N. H. Kuroda, Tokyo, 179 pp.","Wragg, G. M. (1985) The comparative biology of Fluttering Shearwater and Hutton's Shearwater and their relationship to other shearwater species. Unpublished M. Appl. Sc. Thesis, University of Canterbury, Christchurch, 163 pp.","Nunn, G. B. & Stanley, S. E. (1998) Body size effects and rates of cytochrome b evolution in tube-nosed seabirds. Molecular Biology and Evolution, 15 (10), 1360 - 1371. https: // doi. org / 10.1093 / oxfordjournals. molbev. a 025864","Ramirez, O., Illera, J. C., Rando, J. C., Gonzalez-Solis, J., Alcover, J. A. & Lalueza-Fox, C. (2010) Ancient DNA of the extinct Lava Shearwater (Puffinus olsoni) from the Canary Islands reveals incipient differentiation within the P. puffinus complex. Plos One, 5 (12), e 16072. https: // doi. org / 10.1371 / journal. pone. 0016072","Welch, A. J., Olson, S. L. & Fleischer, R. C. (2014) Phylogenetic relationships of the extinct St Helena petrel, Pterodroma rupinarum Olson, 1975 (Procellariiformes: Procellariidae), based on ancient DNA. Zoological Journal of the Linnean Society, 170 (3), 494 - 505. https: // doi. org / 10.1111 / zoj. 12078","Kawakami, K., Eda, M., Izumi, H., Horikoshi, K. & Suzuki, H. (2018) Phylogenetic position of endangered Puffinus lherminieri bannermani. Ornithological Science, 17 (1), 11 - 18. https: // doi. org / 10.2326 / osj. 17.11","Tennyson, A. J. D. & Shepherd, L. D. (2017) DNA reveals the relationships of the extinct Scarlett's Shearwater Puffinus spelaeus (Procellariiformes: Procellariidae). Journal of Ornithology, 158, 379 - 384. https: // doi. org / 10.1007 / s 10336 - 016 - 1416 - 5","Onley, D. & Scofield, P. (2007) Albatrosses, Petrels and Shearwaters of the World. Princeton University, Princeton, 240 pp."]}

10 pages, 4 figures.-- Data availability: All relevant data are publicly available in the Seabird Tracking Database: http://seabirdtracking.org/mapper/?dataset_id=879 and http://seabirdtracking.org/mapper/?dataset_id=1409, Foraging distribution of flying seabirds is constrained by environmental factors influencing individual decision-making. This must be particularly true during the breeding period, when individuals face additional limitations imposed by their central-place foraging behaviour. We used GPS data loggers and Argos PTTs to track the foraging flights of Balearic shearwaters (Puffinus mauretanicus) during the chick-rearing period of 2011–2014 in the Balearic Islands (Western Mediterranean). We identified main areas used by tracked birds and characterised their productivity patterns. Based on a spatial seascape approach of flight costs varying with time, shaped by environmental processes as winds, we also estimated flight costs to reach foraging grounds in outward and return trips from the colony. Individuals repeatedly used the closest areas on the Iberian continental shelf. However, sporadic and favourable wind conditions facilitated low-cost flight to more distant and equally productive areas of the western North African shelf, Funding support was provided to LPO and SEO/BirdLife through Interreg project FAME, LIFE + project INDEMARES, and the Ibiza Preservation Fund, and to the IEO through a Marie Curie Career Integration Grant (CONPELHAB; PCIG09-GA-2011-293774). ML was funded by a Juan de la Cierva (JCI-2010-07639) and a Ramón y Cajal (RYC-2012-09897) postdoctoral contract. FR was funded by a Juan de la Cierva (JCI-2015-24531) postdoctoral contract, With the funding support of the ‘Severo Ochoa Centre of Excellence’ accreditation (CEX2019-000928-S), of the Spanish Research Agency (AEI)

Islands worldwide have sufered seabird extinctions after the arrival of humans and the alien species they introduced. On Santa Luzia (Cabo Verde), an uninhabited island of 35 km2 ,the presence of an impressive quantity of petrel bones in coastal dunes suggested the previous existence of an important seabird colony. Yet, these remains had not been identifed and no seabird extinctions have been reported for the island. This paper characterizes the extinct seabird colony of Santa Luzia and discusses the chronology and possible causes of its demise. A total of 130 grid points in a 5 km2 area and 38 supplementary points within and outside the main study area were excavated to collect bone remains. A total of 1318 anatomical elements, identifed as White-faced Storm Petrel Pelagodroma marina eadesorum (85.3% of the minimum number of individuals, MNI), Boyd’s Shearwater Pufnus lherminieri boydi (11.8% MNI) and Cape Verde Storm Petrel Hydrobates jabejabe (2.9% MNI), were found in 18% of the sampling points, within 1.25 km2 . Neither of the two former species currently breeds on Santa Luzia. In addition, two bones of the Cape Verde Shearwater Calonectris edwardsii and Pterodroma sp. were detected in the supplementary points. Radiocarbon dating of White-faced Storm Petrel (n=10) and Boyd’s Shearwater bones (n=13) suggests that the colony probably went extinct during the frst half of the twentieth century. The recent extinction of these species on Santa Luzia might be consequent on the arrival of humans and their domestic animals on the island. We estimate that the extinct seabird populations must have been at least hundreds of thousand strong, far larger than current populations of the same species in Cabo Verde. We suggest that, following feral cat eradication, Santa Luzia has a signifcant potential for seabird restoration Fundação para a Ciência e Tecnologia - FCT info:eu-repo/semantics/publishedVersion

Increasing human activities have detrimental consequences on marine ecosystems and their impact can have cumulative effects. Within marine ecosystems, seabirds respond to ecosystem variability and face multiple human pressures, especially threatened species. In long-lived species, juveniles and immatures could represent up to 50% of the total population, but their migratory movements remain largely unknown. Here, we depict the migratory patterns of juvenile Balearic shearwaters Puffinus mauretanicus, the most threatened European seabird, using miniaturised satellite transmitters. At the end of the 2012 breeding season, five tagged juveniles left the breeding colonies of Eivissa Island (western Mediterranean) the first week of July. They moved westwards to reach the Atlantic Ocean between 3 and 13 days afterwards. Juveniles showed a twophase migratory pattern: they first travelled slower close to the breeding colonies, and then moved towards their wintering areas in the Atlantic Ocean by rapid directional movements. Environmental cues (e.g.,marine productivity, water mass distribution, frontal systems) might have a prominent role in driving the migratory patterns of juvenile Balearic shearwaters, moving from warm and poor marine areas in the Mediterranean Sea to cooler and rich non-breeding grounds in the Atlantic Ocean. Based on observational findings, we observed certain spatial overlap of juvenile Balearic shearwaters with areas of high human impact, but the relationship between flying travel speed and both fishing effort and cumulative human impacts were not statistically significant. These results suggest that more research is needed to assess whether the movement patterns of migrating juveniles are affected by human activities. Therefore, understanding the at-sea spatial ecology of juveniles should be a priority for research and conservation due to the importance of this population component in long-lived species, as well as assessing their vulnerability to multiple anthropogenic pressures

Este artículo contiene 21 páginas, 6 figuras, 5 tablas., Seabirds inhabiting vast water masses provide numerous examples where opposing phenomena, such as natal and breeding philopatry vs. vagility have dug cryptic taxonomic boundaries among closely related taxa. The taxonomy of little shearwaters of the North Atlantic Ocean (Little–Audubon’s shearwater complex, Puffinus assimilis–lherminieri) still remains unclear, and complementary information on non-breeding distributions and at-sea behaviour becomes essential to unravel divergent local adaptations to specific habitats. Using miniaturized light-level geolocators from seven study areas in the North Atlantic, we evaluate the spatial and habitat segregation, estimate the timing of their key life-cycle events and describe the at-sea behaviour of three taxa of these little shearwaters year-round to distinguish ecological patterns and specializations that could ultimately unravel potential lineage divergences. We also assess morphometric data from birds that were breeding at each study area to further discuss potential adaptations to specific habitats. Our results show that, while birds from different taxa segregated in space and habitats, they share ecological plasticity, similar annual phenology and diel foraging behaviour. These ecological inconsistencies, while defining the evolutionary stressors faced by these taxa, do not suggest the existence of three Evolutionary Significant Units. However, they confirm the recent evolutionary divergence among the three little shearwaters of the North Atlantic., We acknowledge the Ministerio de Economía y Competitividad (MINECO), the Fondos FEDER (CGL2009-11278/BOS, CGL2013-42585-P), the EU LIFE Programme (LIFE09 NAT/PT/000041) and the British Ornithologists’ Union for funding.RR was supported by postdoctoral contracts of the Juan de la Cierva (JCI-2012–11848) and PLEAMAR (2017/2349) programs from the Spanish MINECO and the European Maritime and Fisheries Fund (EMFF), respectively, VHP by a postdoctoral fellowship of Fundação para a Ciência e a Tecnologia (FCT; SFRH/BPD/85024/2012) and ZZ by a PhD grant (APIF-2012) from Universitat de Barcelona (Spain).

Aim Europe's only globally critically endangered seabird, the Balearic shearwater (Puffinus mauretanicus), is thought to have expanded its postbreeding range northwards into UK waters, though its at sea distribution there is not yet well understood. This study aims to identify environmental factors associated with the species’ presence, map the probability of presence of the species across the western English Channel and southern Celtic Sea, and estimate the number of individuals in this area. Location The western English Channel and southern Celtic Sea. Methods This study analyses strip transect data collected between 2013 and 2017 from vessel‐based surveys in the western English Channel and southern Celtic Sea during the Balearic shearwater's postbreeding period. Using environmental data collected directly and from remote sensors both Generalized Additive Models and the Random Forest machine learning model were used to determine shearwater presence at different locations. Abundance was estimated separately using a density multiplication approach. Results Both models indicated that oceanographic features were better predictors of shearwater presence than fish abundance. Seafloor aspect, sea surface temperature, depth, salinity, and maximum current speed were the most important predictors. The estimated number of Balearic shearwaters in the prediction area ranged from 652 birds in 2017 to 6,904 birds in 2014. Main conclusions Areas with consistently high probabilities of shearwater presence were identified at the Celtic Sea front. Our estimates suggest that the study area in southwest Britain supports between 2% and 23% of the global population of Balearic shearwaters. Based on the timing of the surveys (mainly in October), it is probable that most of the sighted shearwaters were immatures. This study provides the most complete understanding of Balearic shearwater distribution in UK waters available to date, information that will help inform any future conservation actions concerning this endangered species., Areas with consistently high probabilities of shearwater presence were identified at the Celtic Sea front. Our estimates suggest that the study area in southwest Britain supports between 2% and 23% of the global population of Balearic shearwaters. This study provides the most complete understanding of Balearic shearwater distribution in UK waters available to date, information that will help inform any future UK conservation actions concerning this endangered species.