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.     

SUBFOSSIL PETREL BONES FROM THE CHATHAM ISLANDS

The breeding community of petrels on the Chatham Islands has previously seemed curiously poor compared to that of analogous stations in the Indian Ocean or South Atlantic (14 species compared with 18 in the Tristan-Gough group). Examination of subfossil deposits from the Chatham Islands probably dating from the early stages of human colonization which occurred within the last few hundred years reveals remains of at least 20 petrels, all of which except one may well have bred in the past, without including storm petrels, of which two species are still known to breed while others may also have bred in the past. The grand total of at least 21 breeding species, many apparently in vast numbers, would probably make the archipelago the most important breeding station for petrels in the world at the time of the first human colonization. It seems possible that in addition to the present breeding species the following may have bred: at least one endemic species of gadfly petrel, the Magenta Petrel Pterodroma magentae , and local populations of at least two other gadfly petrels, two shearwaters of the genus Procellaria and two of the genus Puffinus and a sooty albatross of the genus Phoebetria . Small populations of these, and perhaps other undetected species as well, may still survive in inaccessible places.     

Origins, phytogeny and taxonomy of the gadfly petrels Pterodroma spp

Taxonomic characters of gadfly petrels (Procellariidae: Pterodroma spp.) are reviewed and the genus is redefined. The structure of the upper intestines, which have become helicoidally twisted to varying degrees in most species, is an important character not hitherto used. Results of a phylogenetic study of the genus based mainly on the development of helicoidal intestines agree substantially with findings from studies of the Mallophaga parasitizing these petrels. One species, the Kerguelen Petrel Lugensa brevirostris, previously classified in Pterodroma, is shown to have closer affinity with some fulmar genera. The Tahiti Petrel Pseudobulweria rostrata and its rare or extinct relatives, formerly placed in a superspecies of Pterodroma, are more closely related to the Procellaria and Bulweria petrels, and may be derived from the link between the subfamilies Procellariinae and Fulmarinae. From consideration of the fossil record, anatomical characters and Mallophaga, a phylogeny of the Procellariidae is proposed which supports its monophyletic origin. Gadfly petrels apparently descended from the fulmars through the ancestral lines of Snow Petrels Pagodroma, the Kerguelen Petrel and finally the Blue Petrel Halobaena, which may have given rise separately both to gadfly petrels and to the prions Pachyptila. A late Pliocene origin of Pterodroma in the vicinity of southern New Zealand is possible. The genus Pterodroma is divided into four subgenera, representing four connected radiations, and 29 species are recognised. In probable chronological order, Proaestrelata (new subgenus) contains five species and is restricted to the Pacific Ocean, Cookilaria comprises six species restricted as breeders to the South Pacific, Hallstroma has seven species mainly in the tropics of the Pacific, Indian and South Atlantic Oceans, whereas Pterodroma includes 11 species which extend as breeders from the southwest Pacific west to the South Atlantic and into the North Atlantic. Ecological, physical and physiological adaptations in the evolution of gadfly petrels are discussed.     

Breeding ecology of the White-headed Petrel Pterodroma lessoni on the Kerguelen Islands

The breeding season of the White-headed Petrel Pterodroma lessoni on Mayes Island off the Kerguelen archipelago was studied between December 1985 and February 1987. The birds returned to land in November, and the chicks hatched at the end of January and fledged in May. These incubation and fledging periods are among the longest known in the family Procellariidae. Successful birds normally bred every two years, which has not previously been reported for any member of this family. The breeding ecology is compared to that of other gadfly petrels from sub-antarctic and tropical waters, and the breeding frequency is discussed.     

The modern avifauna of the Pitcairn Islands

Excluding the gadfly petrels Pterodroma spp. and the resident landbirds, this paper details the present status of bird species seen during an expedition to the Pitcairn Islands (Ducie, Henderson, Oeno and Pitcairn) in 1991 and early 1992. Ten species were recorded in the Islands for the first time. Several Southern Ocean petrels were recorded, most in the midwinter period of June and July. The populations of Christmas Island shearwaters breeding on Oeno and Ducie laid in synchrony every 9–10 months. The species is one of few with such a sub-annual, synchronized regime. The majority of other seabirds had an annual breeding cycle, laying between May and October. Around one percent of the world population of the bristle-thighed curlew passes the non-breeding season in the Pitcairn Islands.     

Influence of reproductive success on breeding frequency in four southern petrels

Patterns of breeding frequency of White-headed Pterodroma lessoni, Great-winged Pterodroma macroptera, White-chinned Procellaria aequinoctialis and Grey Petrels Proceharia cinerea have been studied at Kerguelen and Crozet Islands. The White-headed Petrel (summer breeder) followed an essentially biennial cycle with only 13% of successful individuals breeding again the next year. On the other hand, the Great-winged Petrel (winter breeder) was mainly an annual breeder with 78% of successful birds breeding again the next year. Both White-chinned (summer breeder) and Grey Petrels (winter breeder) showed a higher breeding frequency than Pterodroma species, with 91% and 93%, respectively, of successful birds returning to breed the following season. These contrasting patterns may be related to the timing of energetic constraints encountered during the interbreeding period, principally the moult and the restoration of breeding condition. It is suggested that these post-breeding constraints are more marked in winter and in oceanic waters, leading to breeding frequencies being lower than in summer and in neritic waters. The intermediate patterns of breeding frequency found in the Great-winged Petrel (mainly annual) and the White-headed Petrel (mainly biennial) show that among Procellariiformes there is a continuum between annual and strictly biennial species.     

Use of burrow entrance flaps to minimise interference to Chatham petrel (

The Chatham petrel (Pterodroma axillaris) is an endangered species, restricted to a single population on South East Island, Chatham Islands. The key threat to breeding success is loss of chicks as a result of interference by broad-billed prions (Pachyptila vittata) prospecting for burrows for their oncoming breeding season. The effectiveness in decreasing interference using an artificial burrow entrance flap was investigated. The flap exploits behavioural differences between the species. Chatham petrels have a high incentive to push through a flap due to their investment in their burrow and chick, while prospecting prions are influenced by ease of access when searching for potential burrows. This trial found 90% of Chatham petrels entered their burrows through the artificial flap. Flaps acted as barriers to broad-billed prions, where 22% entered the burrow through the flap (P <0.01) compared to the control burrows. Artificial burrow flaps have the potential to provide a low cost, low labour strategy for protecting the known breeding population of Chatham petrels.     

The breeding biology of the gadfly petrels Pterodroma spp. of the Pitcairn Islands: characteristics, population sizes and controls

This paper reports the breeding biology and nesting seasons of the gadfly petrels which nest on the four islands of the Pitcairn group, Pitcairn, Henderson, Oeno and Ducie. The species currently breeding are Murphy's petrel Pterodroma ultima , Kermadec petrel P. neglecta , Herald petrel P. heraldica and Henderson petrel P. atrata. Of these, Murphy's petrel is the most numerous; an estimated 250000 pairs bred on Ducie, which is probably the major breeding station of the species. Novel basic breeding data for Murphy's petrel are presented. Incubation spells, averaging 19.3 days, are exceptionally long for a petrel. Phoenix petrel P. alba appears to have ceased to breed on the Pitcairn Islands since the 1922 surveys of the Whitney Expedition. Nesting success was low on Henderson Island during the study. For all four breeding species, less than 20% of eggs laid yielded fledglings. Failure occurred at the early chick stage and observations indicated that it was due to predation by Pacific rats Rattus exulans. Although rats are present on Ducie, predation was apparently less severe there. The situation on Oeno may be intermediate. I consider how the populations of Henderson are maintained in the face of this intense predation. The Murphy's petrel population may be sustained by immigration from Ducie while the Herald and Henderson petrel populations could be undergoing a long-term decline on Henderson. It is not clear how the Kermadec petrel population is maintained. The conservation implications of these findings are discussed.     

Assessing the potential impact of wind turbines on the endangered Galapagos Petrel <Emphasis Type="Italic">Pterodroma phaeopygia</Emphasis> at San Cristóbal Island,Galapagos

We evaluated the collision risk of Galapagos Petrels Pterodroma phaeopygia with a wind energy development recently constructed in the highlands of San Cristóbal Island, Galapagos. Trained observers recorded the movements of petrels at dusk and dawn from the wind project site, and from control sites located along ravines that host nesting colonies. Collision mortality was also assessed by monitoring circular plots and transect lines located under human-made structures. Petrel flight activity showed a bimodal pattern, with the majority of the movements recorded in the hours previous to sunrise. Most petrels (96%) moved along major ravines that descend from the highlands to the south–southeastern coast of the island. Significant differences in passage rates were found between the project and control sites, with only five petrels recorded on the site selected for turbine installation. Although our data suggest that wind farms will not be more detrimental to petrels than other existing man-made structures, a word of caution is made because even very low levels of additional mortality might be significant for a species with such low productivity and slow maturation rates. Moreover, some other possible indirect effects on habitat change and disturbance might occur that were not assessed in our study. A post-construction monitoring program should be implemented to adequately assess long-term effects on petrels and to enable these uncertainties to be satisfactorily addressed.     

Who’s the boss? Giant petrel arrival times and interspecific interactions at a seal carcass at sub-Antarctic Marion Island

Most northern giant petrels Macronectes halli arrived before southern giant petrels Macronectes giganteus at a seal carcass at sub-Antarctic Marion Island and left earlier. In interspecific interactions, southern giant petrels initiated and dominated encounters. Level of aggression may be related to the degree of satiation. Dominance by southern giant petrels at seal carcasses may help explain why northern giant petrels commence breeding earlier than southern giant petrels in sympatric situations.     

Seabird associations in Pacific equatorial waters

Seabirds were counted along the same cruise tracks in the eastern tropical Pacific (10°øN-10°øS and 140°ø-90°øW) each spring and autumn for 8 years, 1984–1991. During spring, three species groups were apparent among eight species seen in five or more years: (1) Leach's Storm-petrel Oceanodroma leucorhoa and Wedge-rumped Storm-petrel Oceanodroma tethys , which feed on neuston by surface seizing; (2) Black-winged Petrel Pterodroma nigripennis and White-winged Petrel Pterodroma leucoptera , which feed on neuston and micronekton by surface seizing; (3) Juan Fernandez Petrel Pterodroma externa , Wedge-tailed Shearwater Puffinus pacificus and Sooty Tern Sterna fuscata , which feed on micronekton driven to the surface by foraging tuna. During autumn, only one group was identified from the seven species seen in five or more years: Leach's Storm-petrel associated with either Wedge-rumped Storm-petrel or Juan Fernandez Petrel and/or White-winged Petrel. Annual variability in assemblage composition was more extreme in the autumn and thus was similar to variation in physical and biological factors linked to the El Niño-Southern Oscillation cycle.     

A New Fulmarine Petrel (Aves: Procellariiformes) from the Upper Miocene of the Western Mediterranean

A new sea bird, Pterodromoides minoricensis gen. et sp. nov., similar in size and proportions to species of the modern genus Pterodroma , is described from Late Miocene breccias situated on the north-west coast of Menorca. The cranial osteology is similar to that of fulmarine petrels, but the large orbitonasal opening and characters of the postcranial skeleton justify its classification as a separate genus. At least one other species of procellariid occurs in the accompanying fossil asemblage, suggesting that Late Tertiary ecological conditions favoured sea bird populations in this area of the Mediterranean.     

Regulation of incubation shifts near hatching by giant petrels: a timed mechanism, embryonic signalling or food availability?

In most seabird species incubation shifts shorten when hatching approaches, a behavioural response allowing the chick to be fed soon after hatching. Three mechanisms have been proposed to explain these shortened absences: endogenous timing, a response to embryo signals or a seasonal increase in food availability. I tested these hypotheses by cross-fostering eggs between two sibling species: the northern Macronectes halli and the southern M. giganteus giant petrels; the former normally breeds 5-6 weeks earlier than the latter but both have an incubation period of 60 days. The length of the last trip before hatching was unaffected by the timing of hatching, suggesting that trip length was not related to the increase in food availability with the progress of the season. Furthermore, northern giant petrels reduced their last trip lengths regardless of whether they were incubating their own developed or the southern undeveloped egg (with embryos not developed enough to signal). Parents presumably, therefore, possess endogenous control allowing them to predict hatching date and to reduce trip duration accordingly, regardless of the communication with the embryo. Nevertheless, shorter last trips before hatching in southern giant petrels incubating northern giant petrel eggs suggest that parents use embryo signals for a fine-tuning synchronization of the internal timer to the natural variability in the length of the incubation period.     

Validity of Ips chinensis Kurentzov & Kononov confirmed with DNA data

Ips acuminatus has a Eurasian distribution and overlaps with Ips chinensis in southern China.These species are morphologically similar and taxonomic confusion is possible because of the lack of a recent systematic review of these species.Assembly of specimens throughout the ranges of these species allowed for the examination of species boundaries based on monophyly.A phylogeny was reconstructed with cytochrome oxidase Ⅰ DNA sequences for 24 individuals plus three outgroups and morphological diagnostic characters typical for Ips species were reviewed.The strict consensus of 16 most parsimonious trees recovered I.acuminatus and I.chinensis reciprocally monophyletic with an above average pairwise sequence difference (9.6％ compared to 6.1％ for Ips sister species).These clades associated with diagnostic characters of elytral declivity,that is,the position of spine 2.Monophyly,nucleotide differences,and diagnostic characters support the validity of I.chinensis.In addition,high sequence divergence was observed between two clades of I.chinensis which corresponded to northern and southern populations however consistent morphological differences were not found for those clades.     

Phylogeny of "Philoceanus complex" seabird lice (Phthiraptera: Ischnocera) inferred from mitochondrial DNA sequences

The Philoceanus complex is a large assemblage of lice that parasitise procellariiform seabirds (petrels, albatrosses, and their relatives). We obtained mitochondrial 12S rRNA and cytochrome oxidase I DNA sequences from 39 species from diverse hosts and localities. Resolution of deeper relationships between genera was limited, however there is evidence for two major clades, one hosted by albatrosses, the other by petrels. Based on our results, the genera hosted by albatrosses are excellent candidates for detailed analysis of cospeciation. Our results also suggest that a previous estimate of a 5-fold difference in the relative rate of sequence evolution in lice and their avian hosts is an artefact of limited taxonomic sampling.     

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.     

The evolution of north‐east Atlantic gadfly petrels using statistical phylogeography

Macaronesia (north‐east Atlantic archipelagos) has been host to complex patterns of colonization and differentiation in many groups of organisms including seabirds such as gadfly petrels (genus Pterodroma). Considering the subspecies of widely distributed soft‐plumaged petrel for many years, the taxonomic status of the three gadfly petrel taxa breeding in Macaronesia is not yet settled, some authors advocating the presence of three, two or one species. These birds have already been the subject of genetic studies with only one mtDNA gene and relatively modest sample sizes. In this study, using a total of five genes (two mitochondrial genes and three nuclear introns), we investigated the population and phylogeographical histories of petrel populations breeding on Madeira and Cape Verde archipelagos. Despite confirming complete lineage sorting with mtDNA, analyses with nucDNA failed to reveal any population structuring and Isolation with Migration analysis revealed the absence of gene flow during the differentiation process of these populations. It appears that the three populations diverged in the late Pleistocene in the last 150 000 years, that is 10 times more recently than previous estimates based solely on one mtDNA gene. Finally, our results suggest that the Madeira petrel population is ancestral rather than that from Cape Verde. This study strongly advocates the use of nuclear loci in addition to mtDNA in demographical and phylogeographical history studies.     

Breeding of the Trindade Petrel Pterodroma arminjoniana on Round Island, Mauritius

Round Island (19°51'S, 57°47'E), an uninhabited volcanic islet lying 14 miles NNE of the Mauritian mainland, provides locally important breeding grounds for four species of seabirds; the Trindade Petrel Pterodroma arminjoniana , the Wedge-tailed Shearwater Puffinus pacificus , the Red-tailed Tropic Bird Phaethon rubricauda and the White-tailed Tropic Bird Phaethon lepturus. The landing conditions on the island are notoriously difficult and prior to 1975, visits by naturalists had been limited to short stays, a few days at the most. Accounts of the natural history of the island have been made by Vinson (1965), Bullock (1977) and Bullock & North (1984). This paper gives the results of studies on the breeding biology of the Trindade Petrel during two visits to Round Island from 27 July to 2 September 1978, and from 30 July to 18 August 1982.     

Morphometrics and the breeding biology of the Tahiti Petrel Pseudobulweria rostrata

The genus Pseudobulweria includes four extant species, three being classified as critically endangered, while the fourth (Tahiti Petrel P .  rostrata ) is near threatened. Information on the breeding biology of any species of the genus is scarce, even for the Tahiti Petrel. We monitored two Tahiti Petrel nests in New Caledonia and provide the first data on the breeding biology for the genus. Incubation lasts about 55 days, and young fledge at 110 days. Breeding in the southern lagoon of New Caledonia extends from September to April. Very strong interspecific competition for nest cavities occurs with Wedge-tailed Shearwaters Puffinus pacificus . We also document geographical variation in both biometry and breeding phenology over the Pacific breeding range.