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Speciation through homoploid hybridization (HHS) is considered extremely rare in animals. This is mainly because the establishment of reproductive isolation as a product of hybridization is uncommon. Additionally, many traits are underpinned by polygeny and/or incomplete dominance, where the hybrid phenotype is an additive blend of parental characteristics. Phenotypically intermediate hybrids are usually at a fitness disadvantage compared with parental species and tend to vanish through backcrossing with parental population(s). It is therefore unknown whether the additive nature of hybrid traits in itself could lead successfully to HHS. Using a multi-marker genetic data set and a meta-analysis of diet and morphology, we investigated a potential case of HHS in the prions (Pachyptila spp.), seabirds distinguished by their bills, prey choice, and timing of breeding. Using approximate Bayesian computation, we show that the medium-billed Salvin’s prion (Pachyptila salvini) could be a hybrid between the narrow-billed Antarctic prion (Pachyptila desolata) and broad-billed prion (Pachyptila vittata). Remarkably, P. salvini’s intermediate bill width has given it a feeding advantage with respect to the other Pachyptila species, allowing it to consume a broader range of prey, potentially increasing its fitness. Available metadata showed that P. salvini is also intermediate in breeding phenology and, with no overlap in breeding times, it is effectively reproductively isolated from either parental species through allochrony. These results provide evidence for a case of HHS in nature, and show for the first time that additivity of divergent parental traits alone can lead directly to increased hybrid fitness and reproductive isolation.

Studies of avian migration increasingly use stable isotope analysis to provide vital trophic and spatial markers. However, when interpreting differences in stable isotope values of feathers, many studies are forced to make assumptions about the timing of moult. A fundamental question remains about the consistency of these values within and between feathers from the same individual. In this study, we examine variation in carbon and nitrogen isotopes by sub-sampling feathers collected from the wings of adults of two small congeneric petrel species, the broad-billed Pachyptila vittata and Antarctic prion P. desolata. Broad-billed prion feather vane material was enriched in super(15)N compared to feather rachis material, but there was no detectable difference in delta super(13)C. Comparison of multiple samples taken from Antarctic prion feathers indicated subtle difference in isotopes; rachis material was enriched in super(13)C compared to vane material, and there were differences along the length of the feather, with samples from the middle and tip of the feather depleted in super(15)N compared to those from the base. While the greatest proportion of model variance was explained by differences between feathers and individuals, the magnitude of these within-feather differences was up to 0.5 ppt in delta super(15)N and 0.8 ppt in delta super(13)C. We discuss the potential drivers of these differences, linking isotopic variation to individual-level dietary differences, movement patterns and temporal dietary shifts. A novel result is that within-feather differences in delta super(13)C may be attributed to differences in keratin structure within feathers, suggesting further work is required to understand the role of different amino acids. Our results highlight the importance of multiple sampling regimes that consider both within- and between-feather variation in studies using stable isotopes.

AIM: Pelagic seabirds exploit large areas of ocean when acting as central‐place foragers during the breeding season, and ranges are even more extensive outside the breeding period. Spatial niche partitioning is known to occur among species that breed sympatrically, but is less apparent during the non‐breeding period when there is increased potential for overlap among closely related species from neighbouring island groups. This applies to several species of prion, Pachyptila spp., in the Southern Ocean; although extremely abundant, their at‐sea distribution was virtually unknown because they are difficult to distinguish while at sea. To understand spatial niche partitioning at large scales, we investigated the year‐round distribution of thin‐billed prions (Pachyptila belcheri) from the Falkland Islands (Islas Malvinas) and Antarctic prions (Pachyptila desolata) from South Georgia. LOCATION: South Atlantic Ocean. METHODS: Recently, geolocation devices have become small enough to be deployed on small seabirds. During 2009–10, we tracked 20 thin‐billed prions and 9 Antarctic prions with miniaturized geolocators. We applied ecological niche models to compare environmental conditions in the habitat utilized year‐round. RESULTS: We show that two prion species from the south‐west Atlantic Ocean have divergent patterns of migration, and that this has resulted in nearly complete spatial segregation (0–5% overlap by month in the 95% kernel density polygons). Nineteen of 20 thin‐billed prions migrated to an area > 3000 km east of their breeding site, whereas all Antarctic prions migrated a much shorter distance, and to the north‐west. The non‐breeding distribution of thin‐billed prions included the waters around South Georgia, but only when the Antarctic prions were absent. The models highlighted large differences in the realized niche between the two species, and between the habitat characteristics of breeding and non‐breeding areas of thin‐billed and Antarctic prions. MAIN CONCLUSIONS: Our results are consistent with the prediction that spatial niche partitioning occurs at large scales, allowing the co‐existence of related species. The methods applied here will enable predictive maps of the distributions of other prion populations to be created, once data become available from other breeding sites in the Southern Ocean.

Distant populations of animals may share their non-breeding grounds or migrate to distinct areas, and this may have important consequences for population differentiation and dynamics. Small burrow-nesting seabirds provide a suitable case study, as they are often restricted to safe breeding sites on islands, resulting in a patchy breeding distribution. For example, Thin-billed prions Pachyptila belcheri have two major breeding colonies more than 8,000 km apart, on the Falkland Islands in the south-western Atlantic and in the Kerguelen Archipelago in the Indian Ocean. We used geolocators and stable isotopes to compare at-sea movements and trophic levels of these two populations during their non-breeding season, and applied ecological niche models to compare environmental conditions in the habitat. Over three winters, birds breeding in the Atlantic showed a high consistency in their migration routes. Most individuals migrated more than 3000 km eastwards, while very few remained over the Patagonian Shelf. In contrast, all Indian Ocean birds migrated westwards, resulting in an overlapping nonbreeding area in the eastern Atlantic sector of the Southern Ocean. Geolocators and isotopic signature of feathers indicated that prions from the Falklands moulted at slightly higher latitudes than those from Kerguelen Islands. All birds fed on low trophic level prey, most probably crustaceans. The phenology differed notably between the two populations. Falkland birds returned to the Patagonian Shelf after 2-3 months, while Kerguelen birds remained in the nonbreeding area for seven months, before returning to nesting grounds highly synchronously and at high speed. Habitat models identified sea surface temperature and chlorophyll a concentration as important environmental parameters. In summary, we show that even though the two very distant populations migrate to roughly the same area to moult, they have distinct wintering strategies: They had significantly different realized niches and timing which may contribute to spatial niche partitioning.

The hydrothermal vent mussel Bathymodiolus azoricus lives in an intimate symbiosis with two types of chemosynthetic Gammaproteobacteria in its gills: a sulfur oxidizer and a methane oxidizer. Despite numerous investigations over the last decades, the degree of interdependence between the three symbiotic partners, their individual metabolic contributions, as well as the mechanism of carbon transfer from the symbionts to the host are poorly understood. We used a combination of proteomics and genomics to investigate the physiology and metabolism of the individual symbiotic partners. Our study revealed that key metabolic functions are most likely accomplished jointly by B. azoricus and its symbionts: (1) CO 2 is pre-concentrated by the host for carbon fixation by the sulfur-oxidizing symbiont, and (2) the host replenishes essential biosynthetic TCA cycle intermediates for the sulfur-oxidizing symbiont. In return (3), the sulfur oxidizer may compensate for the host’s putative deficiency in amino acid and cofactor biosynthesis. We also identified numerous ‘symbiosis-specific’ host proteins by comparing symbiont-containing and symbiont-free host tissues and symbiont fractions. These proteins included a large complement of host digestive enzymes in the gill that are likely involved in symbiont digestion and carbon transfer from the symbionts to the host.

The principle of competitive exclusion postulates that ecologically-similar species are expected to partition their use of resources, leading to niche divergence. The most likely mechanisms allowing such coexistence are considered to be segregation in a horizontal, vertical or temporal dimension, or, where these overlap, a difference in trophic niche. Here, by combining information obtained from tracking devices (geolocator-immersion and time depth recorders), stable isotope analyses of blood, and conventional morphometry, we provide a detailed investigation of the ecological mechanisms that explain the coexistence of four species of abundant, zooplanktivorous seabirds in Southern Ocean ecosystems (blue petrel Halobaena caerulea, Antarctic prion Pachyptila desolata, common diving petrel Pelecanoides urinatrix and South Georgian diving petrel P. georgicus). The results revealed a combination of horizontal, vertical and temporal foraging segregation during the breeding season. The stable isotope and morphological analyses reinforced this conclusion, indicating that each species occupied a distinct trophic space, and that this appears to reflect adaptations in terms of flight performance. In conclusion, the present study indicated that although there was a degree of overlap in some measures of foraging behaviour, overall the four taxa operated in very different ecological space despite breeding in close proximity. We therefore provide important insight into the mechanisms allowing these very large populations of ecologically-similar predators to coexist.

All animals are associated with microorganisms; hence, host-microbe interactions are of fundamental importance for life on earth. However, we know little about the molecular basis of these interactions. Therefore, we studied the deep-sea Riftia pachyptila symbiosis, a model association in which the tubeworm host is associated with only one phylotype of endosymbiotic bacteria and completely depends on this sulfur-oxidizing symbiont for nutrition. Using a metaproteomics approach, we identified both metabolic interaction processes, such as substrate transfer between the two partners, and interactions that serve to maintain the symbiotic balance, e.g., host efforts to control the symbiont population or symbiont strategies to modulate these host efforts. We suggest that these interactions are essential principles of mutualistic animal-microbe associations. The deep-sea tubeworm Riftia pachyptila lacks a digestive system but completely relies on bacterial endosymbionts for nutrition. Although the symbiont has been studied in detail on the molecular level, such analyses were unavailable for the animal host, because sequence information was lacking. To identify host-symbiont interaction mechanisms, we therefore sequenced the Riftia transcriptome, which served as a basis for comparative metaproteomic analyses of symbiont-containing versus symbiont-free tissues, both under energy-rich and energy-limited conditions. Our results suggest that metabolic interactions include nutrient allocation from symbiont to host by symbiont digestion and substrate transfer to the symbiont by abundant host proteins. We furthermore propose that Riftia maintains its symbiont by protecting the bacteria from oxidative damage while also exerting symbiont population control. Eukaryote-like symbiont proteins might facilitate intracellular symbiont persistence. Energy limitation apparently leads to reduced symbiont biomass and increased symbiont digestion. Our study provides unprecedented insights into host-microbe interactions that shape this highly efficient symbiosis. IMPORTANCE All animals are associated with microorganisms; hence, host-microbe interactions are of fundamental importance for life on earth. However, we know little about the molecular basis of these interactions. Therefore, we studied the deep-sea Riftia pachyptila symbiosis, a model association in which the tubeworm host is associated with only one phylotype of endosymbiotic bacteria and completely depends on this sulfur-oxidizing symbiont for nutrition. Using a metaproteomics approach, we identified both metabolic interaction processes, such as substrate transfer between the two partners, and interactions that serve to maintain the symbiotic balance, e.g., host efforts to control the symbiont population or symbiont strategies to modulate these host efforts. We suggest that these interactions are essential principles of mutualistic animal-microbe associations.

We report the trophic structure of a myctophid assemblage by measuring the isotopic niches of 14 species living in Kerguelen waters, southern Indian Ocean. Most of the species show distinct isotopic niches that differ by at least one of the two niche axes (δ¹³C habitat and δ¹⁵N trophic position), indicating trophic partitioning within the assemblage. Strong niche segregation occurs within each of the three most common genera of myctophids (Electrona, Gymnoscopelus, and Protomyctophum), illustrating the different mechanisms (habitat and dietary segregation) that allow coexistence of closely related species. Calculated trophic levels (TLs) of myctophids ranged from 3.3 to 4.2, showing that they are secondary and tertiary consumers in the pelagic ecosystem. The positive relationship between TL and standard length of fish points out a structuring effect of size, with larger species (Gymnoscopelus spp.) occupying a higher trophic position than smaller species (Krefftichthys anderssoni and Protomyctophum spp.). Myctophids occupy an intermediate trophic position between macrozooplanktonic crustaceans and seabirds and marine mammals within the pelagic ecosystem. However, the TLs of large myctophids overlap those of crustacean-eating seabirds [e.g., Eudyptes spp. (crested penguins) and Pachyptila belcheri]. The isotopic niche of myctophids indicates that Aptenodytes patagonicus (king penguin) adults prey upon K. anderssoni when they feed for themselves, thus exemplifying the usefulness of isotopic datasets on potential prey of predators to depict trophic relationships.

Mollusks, and particularly gastropods, are one of the major taxonomic groups at vents. In these ecosystems, devoid of light, chemoautotrophic bacteria are at the base of the food web and symbiotic association between metazoa and these bacteria is numerous. Nevertheless, apart few “large-size” well-known species, the “small-size” gastropods (shell <5 mm), although very abundant, remain poorly studied regarding symbioses. We investigated here Cyathermia naticoides (Warén and Bouchet in Zool Scr 18(1), 1989), a small coiled gastropod found in abundance on the East Pacific Rise among Riftia pachyptila tubes, and usually inferred to graze on tubeworm bacterial cover, and/or filter feeding. Among mollusks, symbioses are well known in large species and almost exclusively rely on sulfide or methane-oxidizing proteobacterial endosymbionts, occurring within the host tissues in gill epithelial bacteriocytes. Combining several approaches (molecular biology, microscopy, stable isotopes analyses), we described here an unusual symbiosis, where autotrophic filamentous Epsilonproteobacteria are located extracellularly, at the base of host gill filaments. Numerous endocytotic lysosome-like structures were observed in the gill epithelium of the animal suggesting bacteria may contribute to its nutrition through intracellular digestion by gill cells. Additional food source by non-symbiotic proteobacteria grazed on R. pachyptila tubes could complete the diet. The possible role of temperature in the selection of Epsilon- vs Gammaproteobacterial partners is discussed.

Toxoplasma gondii is an intracellular protozoan with worldwide distribution and is of significant importance for One Health due to its zoonotic potential. Although its presence in terrestrial environments is well documented, little is known about its circulation and genetic diversity in the marine ecosystem. This study investigated the occurrence of T. gondii in seabirds rescued from the southern coast of Brazil, focusing on bioassay and molecular characterization. Tissue samples (the heart, brain, liver, lung, and pectoral muscle) from 76 seabirds were received. Of these, 26 were from Larus dominicanus , one Macronectes giganteus , one Pachyptila vittata , five Phalacrocorax brasilianus , two Procellaria aequinoctialis , two Puffinus puffinus , 35 Spheniscus magellanicus , two Sterna hirundo , and two Sula leucogaster . The samples of each bird were processed in pool for peptic digestion. The product of each digestion was inoculated (1 mL/SC) into three mice per bioassay/bird. DNA was extracted for molecular characterization using the PCR–RFLP technique with markers SAG1 , 5′-SAG2 , 3′-SAG2 , Alt. SAG2 , SAG3 , BTUB , GRA6 , c22-8 , c29-2 , L358 , PK1 , and Apico . Brain cysts were detected in 16 bioassays. The ToxoDB #170 genotype was identified as an isolate of kelp gull ( Larus dominicanus ) while another bioassay from the same species did not match the database, suggesting a previously undescribed genotype. Infection occurred through ingestion of sporulated oocysts present in the environment or in fish, bivalves, and crustaceans, which have demonstrated parasitic bioaccumulation. This work highlights the dispersal and infection capacity of different classes of T. gondii hosts and the role of birds as environmental sentinels for marine contamination.