Janos C. Hennicke

Fine‐scale foraging behaviour of a medium‐ranging marine predator

1. Movement patterns of predators should allow them to detect and respond to prey patches at different spatial scales, particularly through the adoption of area-restricted search (ARS) behaviour. Here we use fine-scale movement and activity data combined with first-passage time (FPT) analysis to examine the foraging strategy of northern gannets Morus bassanus in the western North Sea, and to test the following hypotheses: (i) birds adopt a hierarchical foraging strategy characterized by nested ARS behaviour; (ii) the locations and characteristics of ARS zones are strongly influenced by physical oceanography; (iii) the initiation of ARS behaviour is triggered by the detection and pursuit of prey; (iv) ARS behaviour is strongly linked to increased foraging effort, particularly within nested ARS areas. 2. Birds on 13 of 15 foraging trips adopted ARS behaviour at a scale of 9.1 +/- 1.9 km, and birds on 10 of these 13 trips adopted a second, nested ARS scale of 1.5 +/- 0.8 km, supporting hypothesis 1 above. ARS zones were located 117 +/- 55 km from the colony and over half were within 5 km of a tidal mixing front ~50 km offshore, supporting hypothesis 2 above. 3. The initiation of ARS behaviour was usually followed after only a short time interval (typically ~5 min) by the commencement of diving. Gannets do not dive until after they have located prey, and so this pattern strongly suggests that ARS behaviour was triggered by prey detection, supporting hypothesis 3 above. However, ~33% of dives in mixed coastal water and 16% of dives in stratified water were not associated with any detectable ARS behaviour. Hence, while ARS behaviour resulted from the detection and pursuit of prey, encounters with prey species did not inevitably induce ARS behaviour. 4. Following the initiation of ARS behaviour, dive rates were almost four times higher within ARS zones than elsewhere and almost three times higher in zones with nested ARS behaviour than in those without, supporting hypothesis 4 above and suggesting that the foraging success of birds was linked to their ability to match the hierarchical distribution of prey.

Hemosporidian blood parasites in seabirds—a comparative genetic study of species from Antarctic to tropical habitats

Whereas some bird species are heavily affected by blood parasites in the wild, others reportedly are not. Seabirds, in particular, are often free from blood parasites, even in the presence of potential vectors. By means of polymerase chain reaction, we amplified a DNA fragment from the cytochrome b gene to detect parasites of the genera Plasmodium, Leucocytozoon, and Haemoproteus in 14 seabird species, ranging from Antarctica to the tropical Indian Ocean. We did not detect parasites in 11 of these species, including one Antarctic, four subantarctic, two temperate, and four tropical species. On the other hand, two subantarctic species, thin-billed prions Pachyptila belcheri and dolphin gulls Larus scoresbii, were found infected. One of 28 thin-billed prions had a Plasmodium infection whose DNA sequence was identical to lineage P22 of Plasmodium relictum, and one of 20 dolphin gulls was infected with a Haemoproteus lineage which appears phylogenetically clustered with parasites species isolated from passeriform birds such as Haemoproteus lanii, Haemoproteus magnus, Haemoproteus fringillae, Haemoproteus sylvae, Haemoproteus payevskyi, and Haemoproteus belopolskyi. In addition, we found a high parasite prevalence in a single tropical species, the Christmas Island frigatebird Fregata andrewsi, where 56% of sampled adults were infected with Haemoproteus. The latter formed a monophyletic group that includes a Haemoproteus line from Eastern Asian black-tailed gulls Larus crassirostris. Our results are in agreement with those showing that (a) seabirds are poor in hemosporidians and (b) latitude could be a determining factor to predict the presence of hemosporidians in birds. However, further studies should explore the relative importance of extrinsic and intrinsic factors on parasite prevalence, in particular using phylogenetically controlled comparative analyses, systematic sampling and screening of vectors, and within-species comparisons.

Infection by Haemoproteus Parasites in Four Species of Frigatebirds and the Description of a New Species of Haemoproteus (Haemosporida: Haemoproteidae)

Abstract: Among seabirds, the fregatids stand out with a high prevalence of blood parasites. Four of 5 species in this family have been found to be infected with Haemoproteus; however, complete species descriptions with molecular phylogeny are lacking. Seventy-five samples from 4 species of frigatebirds, i.e., Fregata andrewsi, Fregata minor, Fregata magnificens, and Fregata aquila, were screened for infections caused by species of Haemoproteus. Four different parasite haplotypes were found infecting frigatebirds based on the sequencing of a fragment of the cytochrome b gene. Two haplotypes belong to the subgenus Parahaemoproteus, and the other 2 correspond to haplotypes within the subgenus Haemoproteus. The more prevalent and cosmopolitan Parahaemoproteus haplotype (FregPHae1) was phylogenetically grouped with other Haemoproteus parasites infecting non-passerine birds, but it could not be detected from the single sample from F. aquila. The other Parahaemoproteus haplotype (FregPHae2) was not phylogenetically clustered with parasites infecting non-passerine birds, and it was sequenced from a single (1 each) F. andrewsi and F. minor. Blood smears from F. andrewsi infected only by FregPHae1 haplotype showed sufficient gametocytes to allow description of a new species, Haemoproteus valkiūnasi sp. nov. In contrast to Haemoproteus iwa, the only previously known blood parasite infecting frigatebirds and described from F. minor from Galapagos Islands, parasites from F. andrewsi (1) are shorter with no contact of gametocyte with host cell membrane, (2) have fewer pigment granules, and (3) have wider microgametocytes, with a smaller host nuclear displacement. In contrast, patent single infections corresponding to the cosmopolitan haplotype of the subgenus Haemoproteus (FregHae1) were also found in samples from 1 F. andrewsi, 1 F. minor, and 1 F. aquila. In all these cases, the number of microgametocytes was very low, resembling H. iwa, which lacks microgametocytes in the original description. Macrogametocytes of haplotype FregHae1 in F. andrewsi differ significantly from all the characteristics measured from H. valkiūnasi. In addition, it also differs from all characteristics of H. iwa despite being genetically identical in the analyzed fragment.

Comparison of trip duration, activity pattern and diving behaviour by Red-tailed Tropicbirds (Phaethon rubricauda) during incubation and chick-rearing

Abstract In the tropical marine environment food resources are scarce and often patchily distributed. Consequently, nesting tropical seabirds must balance travelling to productive areas and sharing incubation or providing food to the chick. Changes in foraging behaviour occur when adults switch from feeding only for themselves during incubation to feeding a chick and fulfilling their own energy requirements. Externally attached data-devices were used to compare the duration of foraging trips, flight activity and depth of dives of incubating and chick-rearing Red-tailed Tropicbirds (Phaethon rubricauda) on Christmas Island, Indian Ocean. Adults with chicks had a bimodal pattern of foraging, with alternating short and long foraging trips (mean duration±s.d. of short trips = 3.0 ± 4.1 h, of long trips = 57.4 ± 41.1 h), whereas incubating adults undertook only longer trips (mean duration ± s.d. = 152.9 ± 47.2 h). This strategy is stimulated by partner behaviour, rather than body mass as reported for many other seabirds, and may enable the optimisation of simultaneous food delivery to chicks and self-feeding when local resources are poor. The proportion of time spent in flight (flight activity) by adults with chicks averaged 90.1% during short trips whereas incubating birds spent only 62.4% of their trips flying. Overall mean depth of foraging dives by adults with chicks averaged 0.6 ± 0.3 m during short trips; mean maximum depth of dives by incubating adults were recorded using maximum-depth recorders and were significantly deeper at 10.7 ± 8.7 m, reaching a maximum depth of 25.6 m.

Sex-Specific Habitat Utilization and Differential Breeding Investments in Christmas Island Frigatebirds throughout the Breeding Cycle

In seabirds, equal bi-parental care is the rule, as it is considered crucial for raising chicks successfully because seabirds forage in an environment with unpredictable and highly variable food supply. Frigatebirds forage in poor tropical waters, yet males reduce and even stop parental care soon after chick brooding, leaving the female to provision the chick alone for an extended fledging period. Using bird-borne tracking devices, male and female Christmas Island Frigatebirds (Fregata andrewsi) were investigated during the brooding, late chick rearing and post-fledging period to examine whether sexes exhibit foraging strategies that may be linked to differential breeding investments. During brooding, males and females showed similar foraging behaviour under average marine productivity of oceanic waters close to the colony, but males shifted to more distant and more productive habitats when conditions deteriorated to continue with reduced chick provisioning. During the late chick rearing period, females progressively increased their foraging range to the more distant but productive marine areas that only males had visited during brooding. Birds spent the non-breeding period roosting in highly productive waters of the Sunda Shelf. The sex-specific utilisation of three different foraging habitats with different primary productivity (oceanic, coastal, and shelf areas) allowed for temporal and spatial segregation in the exploitation of favourable habitats which seems to enable each sex to optimise its foraging profitability. In addition, post-fledging foraging movements of females suggest a biennial breeding cycle, while limited information on males suggests the possibility of an annual breeding cycle.

Effects of long-term isolation on genetic variation and within-island population genetic structure in Christmas Island (Indian Ocean) seabirds

Seabirds endemic to remote oceanic islands face several unique threats to their population viability. Christmas Island is a small oceanic island with remarkable seabird biodiversity, including two endangered endemics: Christmas Island frigatebirds (Fregata andrewsi) and Abbott’s boobies (Papasula abbotti). Christmas Island seabirds are currently threatened by habitat destruction at breeding sites due to phosphate mining and by the adverse effects of the invasive yellow crazy ant (Anoplolepis gracilipes); however, conservation initiatives are limited by a paucity of genetic data on the breeding seabirds. To address this shortcoming, we collected genetic samples from five breeding seabird species, including the two endemics. We surveyed mitochondrial and nuclear genetic diversity, estimated effective population sizes, and tested whether species had undergone recent or long-term effective population size decline. Effective population sizes for Christmas Island frigatebirds and Abbott’s boobies were both low (approximately 5,000 and 2,100 individuals, respectively). Moreover, four out of the five species appeared to have undergone long-term population decline, suggesting that mitigation of the threats to population viability of Christmas Island seabirds is critical. We also detected cryptic population genetic structure within Abbott’s booby, perhaps due to in situ divergence on Christmas Island or due to the influx of individuals from an extirpated colony. Within-island differentiation may be common among endemic seabird species due to high levels of natal philopatry, and we suggest that careful consideration of population genetic structure on small geographic scales is crucial when designing conservation initiatives.

Resource partitioning between incubating and chick-rearing brown boobies and red-tailed tropicbirds on Christmas Island

BackgroundIn oligotrophic tropical marine environments, the main mechanism explaining the coexistence of sympatric seabirds is segregation by habitat or segregation by prey within the same habitat. Both types of segregation can play a role during the breeding season due to different constraints associated with different phases of the breeding cycle. By using stable isotope analyses, we investigated intra- and interspecific foraging segregation in two tropical seabird species, the red-tailed tropicbird Phaeton rubricauda and the brown booby Sula leucogaster, breeding sympatrically on Christmas Island, Indian Ocean. We compared isotopic values of δ13C and δ15N in blood from incubating and chick-rearing adults of both species.ResultsThe results showed small but significantly interspecific and intraspecific differences in δ13C and δ15N values. Differences in δ13C values suggest spatial segregation in the main foraging grounds between the two species during the breeding season as well as between incubating and chick-rearing brown boobies. In contrast, red-tailed tropicbirds probably exploited similar foraging habitats during both breeding stages. δ15N values did not indicate diet-related differences, neither within nor between species, suggesting a highly opportunistic feeding behavior to cope with the limited prey available in the oligotrophic marine environment.ConclusionsCompetition for prey in breeding red-tailed tropicbirds and brown boobies seems to be reduced by spatial segregation enabling both species to successfully reproduce in sympatry in an oligotrophic tropical marine environment.