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.     

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.     

Widespread gene flow between oceans in a pelagic seabird species complex

Global‐scale gene flow is an important concern in conservation biology as it has the potential to either increase or decrease genetic diversity in species and populations. Although many studies focus on the gene flow between different populations of a single species, the potential for gene flow and introgression between species is understudied, particularly in seabirds. The only well‐studied example of a mixed‐species, hybridizing population of petrels exists on Round Island, in the Indian Ocean. Previous research assumed that Round Island represents a point of secondary contact between Atlantic (Pterodroma arminjoniana) and Pacific species (Pterodroma neglecta and Pterodroma heraldica). This study uses microsatellite genotyping and tracking data to address the possibility of between‐species hybridization occurring outside the Indian Ocean. Dispersal and gene flow spanning three oceans were demonstrated between the species in this complex. Analysis of migration rates estimated using bayesass revealed unidirectional movement of petrels from the Atlantic and Pacific into the Indian Ocean. Conversely, structure analysis revealed gene flow between species of the Atlantic and Pacific oceans, with potential three‐way hybrids occurring outside the Indian Ocean. Additionally, geolocation tracking of Round Island petrels revealed two individuals travelling to the Atlantic and Pacific. These results suggest that interspecific hybrids in Pterodroma petrels are more common than was previously assumed. This study is the first of its kind to investigate gene flow between populations of closely related Procellariiform species on a global scale, demonstrating the need for consideration of widespread migration and hybridization in the conservation of threatened seabirds.     

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.     

Molecular evidence for the identity of the Magenta petrel

A lone petrel was shot from the decks of an Italian warship (the ‘Magenta’) while it was sailing the South Pacific Ocean in 1867, far from land. The species, unknown to science, was named the ‘Magenta petrel’ (Procellariiformes, Procellariidae, Pterodroma magentae). No other specimens of this bird were collected and the species it represented remained a complete enigma for over 100 years. We compared DNA sequence of the mitochondrial cytochrome b gene from the Magenta petrel to that of other petrels using phylogenetic methods and ancient DNA techniques. Our results strongly suggest that the Magenta petrel specimen is a Chatham Island taiko. Furthermore, given the collection location of the Magenta petrel, our finding indicates that the Chatham Island taiko forages far into the Pacific Ocean (near South America). This has implications for the conservation of the taiko, one of the world's rarest seabirds.     

Range expansion and hybridization in Round Island petrels (Pterodroma spp.): evidence from microsatellite genotypes

Historical records suggest that the petrels of Round Island (near Mauritius, Indian Ocean) represent a recent, long‐distance colonization by species originating from the Atlantic and Pacific Oceans. The majority of petrels on Round Island appear most similar to Pterodroma arminjoniana, a species whose only other breeding locality is Trindade Island in the South Atlantic. Using nine microsatellite loci, patterns of genetic differentiation in petrels from Round and Trindade Islands were analysed. The two populations exhibit low but significant levels of differentiation in allele frequencies and estimates of migration rate between islands using genetic data are also low, supporting the hypothesis that these populations have recently separated but are now isolated from one another. A second population of petrels, most similar in appearance to the Pacific species P. neglecta, is also present on Round Island and observations suggest that the two petrel species are hybridizing. Vocalizations recorded on the island also suggest that hybrid birds may be present within the population. Data from microsatellite genotypes support this hypothesis and indicate that there may have been many generations of hybridization and back‐crossing between P. arminjoniana and P. neglecta on Round Island. Our results provide an insight into the processes of dispersal and the consequences of secondary contact in Procellariiformes.     

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.     

Neohyadesia microtricha (Acari: Astigmata: Algophagidae): a new species from the sub-Antarctic

The sub-antarctic mite genus Neohyadesia (Acari: Astigmata) is known from two described taxa: N. signyi from Signy Island (South Orkney Islands, South Atlantic Province), and a subspecies, N. s. punctulata from Ile Kerguelen (South Indian Province). This paper describes a second species distinguishable by, in particular, cuticular microtrichae covering the dorsal surface. The new species, N. microtricha sp. nov., has a similarly disjunct sub-antarctic distribution, occurring on both Marion (SIP) and South Georgia (SAP) islands. A systematic synopsis of the sub-antarctic algophagine mites is given. Accepted: 30 June 2000     

Predation risk and moonlight avoidance in nocturnal seabirds

Unlike most seabird families, the vast majority of small petrel species are nocturnal on their breeding grounds. Further, they reduce markedly their activity when the light level increases. Moonlight avoidance might be a consequence of reduction in foraging profitability, as bioluminescent prey do not come to the sea surface on bright nights. Alternatively, petrels may avoid colonies during moonlit nights because of increased predation risk. We studied predation on petrels by Brown Skuas Catharacta antarctica lönnbergi at Kerguelen, and the influence of moonlight on behaviour of both skuas and petrels, to test the 'predation risk' hypothesis. On the study area, Brown Skuas hunt at night and prey heavily upon the Blue Petrel Halobaena caerulea and the Thin-billed Prion Pachyptila belcheri . Predation risk was higher on moonlit nights, as skuas caught more prey, and particularly more Blue Petrels when the light level increased. Nightly intakes of Blue Petrel and Thin-billed Prion by skuas was related to colony attendance of non-breeders rather than that of breeders. Biometry of prey also suggested that skuas caught a higher proportion of non-breeding birds than was present at the colonies. Predation risk was thus greater in non-breeders and on moonlit nights. Colony attendance by non-breeding Blue Petrels and Thin-billed Prions was also reduced during moonlit nights. Vocal activity, which is mainly by non-breeders, was also drastically reduced when the light level increased in the species suffering the highest predation rate. Our results supported the 'predation risk' hypothesis, although the 'foraging efficiency' and the 'predation risk' hypotheses are not mutually exclusive: the former might explain the moonlight avoidance behaviour of breeding, and the latter that of non-breeding individuals.     

Molecular analysis of Southern Ocean skates (Bathyraja) reveals a new species of Antarctic skate

Two regions of mtDNA, cytochrome b and cytochrome c oxidase subunit 1, were sequenced in nine species of Bathyraja from the Southern Ocean and New Zealand. Based on sequence divergence, the species that has been referred to as Bathyraja eatonii from the Antarctic continental shelf and slope is a species distinct from B. eatonii from the Kerguelen Plateau (the type locality) and is a new and undescribed species Bathyraja sp. (cf. eatonii). There was no sequence divergence among samples of Bathyraja sp. (dwarf) from the Ross Sea and the South Atlantic. However, for both Bathyraja sp. (cf. eatonii) and Bathyraja maccaini in the Ross Sea and the South Atlantic Ocean, the DNA sequence divergences indicate differentiation among ocean basins and within Bathyraja sp. (cf. eatonii) divergences are similar to those among recognized species of Bathyraja in the North Pacific Ocean.     

A phylogeographic investigation of the kelp genus Laminaria (Laminariales,Phaeophyceae), with emphasis on the South Atlantic Ocean

The genus Laminaria has a wide distribution range compared with other kelp genera because it is found in both the North and the South Atlantic, on both sides of the North Pacific, as well as in the Mediterranean. Hypotheses behind this biogeographical pattern have been discussed by several authors but have not yet been fully evaluated with time‐calibrated phylogenies. Based on the analysis of four molecular markers (ITS2, rbcL, atp8 and trnWI), our goal was to reassess the Laminaria species diversity in South Africa, assess its relationship with the other species distributed in the South Atlantic and reconstruct the historical biogeography of the genus. Our results confirm the occurrence of a single species, L. pallida, in southern Africa, and its sister relationship with the North Atlantic L. ochroleuca. Both species belonged to a clade containing the other South Atlantic species: L. abyssalis from Brazil, and the Mediterranean L. rodriguezii. Our time‐calibrated phylogenies suggest that Laminaria originated in the northern Pacific around 25 mya, followed by at least two migration events through the Bering Strait after its opening (~5.32 mya). Today, the first is represented by L. solidungula in the Arctic, while the second gave rise to the rest of the Atlantic species. The colonization of the North Atlantic was followed by a gradual colonization southward along the west coast of Europe, into the Mediterranean (~2.07 mya) and two recent, but disconnected, migrations (~1.34 and 0.87 mya) across the equator, giving rise to L. abyssalis in Brazil and L. pallida in southern Africa, respectively.     

Skull morphology of bottlenose dolphins from different ocean populations with emphasis on South America

The bottlenose dolphin, genus Tursiops, is cosmopolitan occurring in tropical and temperate regions, with morphological variation between and within different oceans. Since the genus' taxonomy has been under discussion for a long time, this work aimed at analyzing the cranial variability of T. truncatus from different regions of the world. Geometric Morphometrics analyses were performed in 201 skulls of adult specimens, on dorsal, ventral, and lateral views, from the Eastern North Pacific, Eastern North Atlantic, Eastern South Atlantic, and Western South Atlantic oceans. The results indicate differences between individuals that inhabit the Atlantic and Pacific oceans. Within the Atlantic Ocean, there is an evident longitudinal differentiation of specimens from the eastern and western regions. A latitudinal separation was also observed, considering specimens from the North and South Atlantic Ocean. In the Western South Atlantic statistical differences were found between two morphological groups, identified as T. gephyreus (sensu Lahille, 1908) and T. truncatus, and the cross-validation presented 98% as minimum confidence for correct classification of these two groups. The present study provides strong morphological support to consider these two lineages as separate species.     

The geographic scale of speciation in a marine snail with high dispersal potential

Aim We use the Stramonita haemastoma species complex (Muricidae) to investigate the geographic scale of speciation in a marine snail with a long pelagic larval duration (PLD) of 2–3 months and, consequently, high dispersal potential. We aim to: (1) delimit species within Stramonita, (2) discover the phylogenetic relationship among them, (3) map their distributions, and (4) infer the age and likely cause of speciation events. Location Tropical intertidal of the Atlantic and eastern Pacific Oceans. Methods We use one nuclear and two mitochondrial genes to construct a molecular phylogeny of the S. haemastoma species complex. We first test the monophyly of the genus and of the species complex, and then use statistical methods to delimit species within the complex. We incorporate information from museum collections and the literature to map distributions and to look for diagnostic morphological traits. We use fossils to date our phylogeny. Results The genus Stramonita is monophyletic and restricted to the tropical and warm‐temperate Atlantic and eastern Pacific oceans. The genus is composed of Stramonita delessertiana and six members of the S. haemastoma complex: S. haemastoma, Stramonita rustica, Stramonita floridana, Stramonita canaliculata, Stramonita biserialis and Stramonita brasiliensis (new species described herein). These species are supported by reciprocal monophyly in mitochondrial gene trees, together with independent evidence from morphology, distribution and the nuclear gene. The species are almost entirely allopatric, with only three instances of sympatry. Two species have unusually wide distributions, consistent with their long PLD; one of these is amphi‐Atlantic. Main conclusions Despite the long PLD of Stramonita, speciation has occurred within the Atlantic, both in response to barriers operating at the largest geographical scale (the width of Atlantic, but not the Amazon barrier) and at a smaller scale within the western Atlantic.     

Influence of the diet on the bioaccumulation of heavy metals in zooplankton-eating petrels at Kerguelen archipelago,Southern Indian Ocean

Concentrations of cadmium, mercury, copper and zinc were measured in muscle, kidney and liver tissues and in the main prey of five species of zooplankton-eating petrels: blue petrel (Halobaena caerulea), thin-billed (Pachyptila belcheri) and Antarctic (P. desolata) prions, and South Georgian (Pelecanoides georgicus) and common (Pelecanoides urinatrix) diving petrels. Since some of these species are closely related species with respect to body size, timing of moult and life span, their diet has been examined to evaluate its influence on heavy-metal bioaccumulation. Inter-specific differences were significant for Hg concentrations in the liver and for Zn concentrations in both liver and kidney tissues. Blue petrels exhibited the highest Hg concentrations in the liver (3.9±2.02 µg.g^–1 wet weight). No significant differences were found in Cd concentrations between species. Exposure to heavy metals through the most important prey species in the diet during the breeding period was evaluated. The most evident result was the influence of fish prey on Hg levels. Although crustacean species exhibit different cadmium concentrations, the diet composition does not appear to be discriminant for Cd bioaccumulation within the small petrel community at Kerguelen.     

Phylogenetic relationships between tuna species of the genus Thunnus (Scombridae: Teleostei): Inconsistent implications from morphology,nuclear and mitochondrial genomes

In order to infer phylogenetic relationships between tuna species of the genus Thunnus, partial sequences of the mitochondrial cytochrome b and ATPase genes were determined in all eight species. Supplemental restriction analysis on the nuclear rRNA gene was also carried out. Pacific northern bluefin tuna (Thunnus thynnus orientalis) was found to have mtDNA distinct from that of the Atlantic subspecies (T. t. thynnus) but very similar to that from the species albacore (T. alaluga). In contrast, no differentiation in nuclear genome was observed between the Atlantic and Pacific northern bluefin tunas. The Atlantic northern bluefin and southern bluefin tunas possessed mtDNA sequences very similar to species of yellowfin tuna group and not so similar to albacore and bigeye tunas which were morphologically assigned to the bluefin tuna group. The molecular data indicate that (1) mtDNA from albacore has been incorporated into the Pacific population of northern bluefin tuna and has extensively displaced the original mtDNA, and (2) albacore is the earliest offshoot, followed by bigeye tuna in this genus, which is inconsistent with the phylogenetic relationships between these tuna species inferred from morphology. Correspondence to: S. Chow     

SUMMER DISTRIBUTION OF PELAGIC BIRDS OFF THE COAST OF ARGENTINA

Eighteen species of sea–birds were observed from CSS 'Hudson' in the South Atlantic between the Rio de la Plata and Tierra del Fuego in late January and late February 1970. Major feeding areas were noted off Mar del Plata, the Valdes Peninsula and Tierra del Fuego, Argentina, the first two in rich modified or unmodified water of the subantarctic Falkland Current, associated with upwelling. The southernmost area had most birds, the northernmost fewest. A temperate–latitude assemblage including Yellow–nosed Albatross and Cory's Shearwater occurred off Mar del Plata in warmed water of subantarctic (Falkland Current) origin. South of 39°S these two species were replaced by Black–browed Albatross and Great Shearwater in cold offshore water of the Falkland Current. Off the Valdes Peninsula Black–browed Albatross, Great and Sooty Shearwaters, White–chinned Petrels and family groups of Magellanic Penguins occurred near the edge of highly–productive water of the Falkland Current. The largest feeding area, east of Tierra del Fuego, had Magellanic and Rockhopper Penguins, Great Shearwaters, King Cormorants, South American Terns and jaegers, but no oceanographic information is available. At 50°S prions and diving petrels appeared for the first time, marking a possible oceanographic discontinuity or proximity to colonies on the Falkland Islands. Several common species of the South Atlantic were not seen on the transect including Cape Pigeon, Southern Fulmar, Soft–plumaged Petrel, Schlegel's Petrel and Pediunker. Kelp Gull, South American Tern, Manx Shearwater and jaegers were rare or absent far from land. New southern limits for the western South Atlantic Ocean were established for Cory's and Manx Shearwaters, and the Great Shearwater was observed for the first time in Chilean waters.     

A cladistic phylogeny of the genus Littorina (Gastropoda): implications for evolution of reproductive strategies and for classification

Gross anatomical characters of all 18 species of Littorina are used to construct a phylogenetic hypothesis for the genus, by the method of cladistic analysis. The resulting cladogram suggests that of the four subgenera of Littorina, one (Littorina) is paraphyletic. It is uncertain whether the genus Mainwaringia should be included in Littorina. It is shown that the non-planktotrophic Littorina species in the northern Atlantic comprise a monophyletic group, with the sister-species L. kurila and/or L. subrotundata in the northern Pacific. Invasion of the Atlantic by a minimum of two Pacific species, across the Arctic migration route established during the late Cenozoic, is sufficient to account for the modern distribution of the subgenera Littorina and Neritrema. The importance of the cladogram as a basis for hypotheses of adaptation is illustrated by a discussion of spawn and development in Littorina.     

Panbiogeographical analysis of distribution patterns in hagfishes (Craniata: Myxinidae)

Aim To analyse the worldwide distribution patterns of hagfishes using panbiogeographical track analysis, and to attempt to correlate these patterns with the tectonic history of the ocean basins. Location Atlantic and Pacific oceans. Method The distributions of 47 out of 70 species of hagfish (in the genera Eptatretus, Myxine, Nemamyxine, Neomyxine, and Paramyxine) were studied by the panbiogeographical method of track analysis. The analysis was performed using distributional data obtained from the collections included in the Ocean Biogeographic Information System (OBIS, http://www.iobis.org ) and FishBase ( http://www.fishbase.org ), with additional records from the literature. Individual tracks were obtained for each species by plotting localities and connecting them by minimum‐spanning trees. Generalized tracks were determined from the spatial overlap between individual tracks. Results Six generalized tracks were found: in the Gulf of Mexico, Caribbean Sea, South‐eastern Atlantic, Western Pacific, North‐eastern Pacific and South‐eastern Pacific. Main conclusions The distribution patterns of myxinids are marked by a high degree of endemism and vicariance, and are correlated with the tectonic features involved in many of the events that led to the development of oceanic basins. The main massing of the group is around the Pacific Basin. In the Atlantic Ocean, the distribution of Myxine glutinosa seems to correspond to a classic trans‐oceanic track and vicariance resulting from the opening of the Atlantic Ocean during the Cretaceous. In the Pacific Ocean, the distribution of the Eptatretus and Paramyxine species is clearly associated with the margins of the Pacific tectonic plate. The generalized tracks of hagfishes are shared by several other groups of marine organisms, including many from shallow tropical waters, implying a common history for this marine biota. Overall, vicariance is a major feature of hagfish distribution, suggesting vicariant differentiation of widespread ancestors as a result of sea‐floor spreading between continents in connection with ocean formation.     

No longer a circumtropical species: revision of the lizardfishes in the Trachinocephalus myops species complex,with description of a new species from the Marquesas Islands

Trachinocephalus, a formerly monotypic and nearly circumtropical genus of lizardfishes, is split into three valid species. Trachinocephalus gauguini n. sp. is described from the Marquesas Islands and is distinguished from the two other species in the genus by having a shorter snout, a narrower interorbital space, larger eye and modally fewer anal‐fin and pectoral‐fin rays. The distribution of Trachinocephalus myops (type species) is restricted to the Atlantic Ocean and the name Trachinocephalus trachinus is resurrected for populations from the Indo‐West Pacific Ocean. Principal component analyses and bivariate plots based on the morphometric data differentiated T. gauguini from the other two species, but a substantial overlap between T. myops and T. trachinus exists. Phylogenetic evidence based on mtDNA COI sequences unambiguously supports the recognition of at least three species in Trachinocephalus, revealing deep divergences between the Atlantic Ocean, Indo‐West Pacific Ocean and Marquesas entities. Additional analyses of species delimitations using the generalized mixed Yule coalescent model and the Poisson tree processes model provide a more liberal assessment of species in Trachinocephalus, indicating that many more cryptic species may exist. Finally, a taxonomic key to identify the three species recognized here is provided.     

Molecular Phylogeny of the Parasitic Dinoflagellate Chytriodinium within the Gymnodinium Clade (Gymnodiniales,Dinophyceae)

The dinoflagellate genus Chytriodinium, an ectoparasite of copepod eggs, is reported for the first time in the North and South Atlantic Oceans. We provide the first large subunit rDNA (LSU rDNA) and Internal Transcribed Spacer 1 (ITS1) sequences, which were identical in both hemispheres for the Atlantic Chytriodinium sp. The first complete small subunit ribosomal DNA (SSU rDNA) of the Atlantic Chytriodinium sp. suggests that the specimens belong to an undescribed species. This is the first evidence of the split of the Gymnodinium clade: one for the parasitic forms of Chytriodiniaceae (Chytriodinium, Dissodinium), and other clade for the free‐living species.