Prolonged and flexible primary moult overlaps extensively with breeding in beach-nesting Hooded Plovers Thinornis rubricollis

We present the first report of complete overlap of breeding and moult in a shorebird. In southeastern Australia, Hooded Plovers Thinornis rubricollis spend their entire lives on oceanic beaches, where they exhibit biparental care. Population moult encompassed the 6-month breeding season. Moult timing was estimated using the Underhill-Zucchini method for Type 2 data with a power transformation to accommodate sexual differences in rates of moult progression in the early and late stages of moult. Average moult durations were long in females (170.3 ± 14.2 days), and even longer in males (210.3 ± 13.5 days). Breeding status was known for most birds in our samples, and many active breeders (especially males) were also growing primaries. Females delayed the onset of primary moult but were able to increase the speed of moult and continue breeding, completing moult at about the same time as males. The mechanism by which this was achieved appeared to be flexibility in moult sequence. All moult formulae fell on one of two linked moult sequences, one faster than the other. The slower sequence had fewer feathers growing concurrently and also had formulae indicating suspended moults. Switching between sequences via common formulae is possible at many points during the moult cycle, and three of 12 recaptures were confirmed to have switched sequences in the same moult season. Hooded Plovers thus have a prolonged primary moult with the flexibility to change their rate of moult; this may facilitate high levels of replacement clutches that are associated with passive nest defence and low reproductive success. © 2014 British Ornithologists' Union.

Personality in captivity: more exploratory males reproduce better in an aviary population

The existence of animal personality is well-established across a wide range of species, with the majority of evidence for this being obtained from individuals held in captivity. However, there has been little work assessing the influence of commonly-measured personality traits on fitness, which is pertinent when the genetic basis of personality is considered. We measured whether the reproductive behaviour and success of zebra finches in a captive mixed-sex aviary environment was influenced by an aspect of their personality, their exploratory behaviour in a single-sex social aviary. We found that more exploratory males made a greater number of breeding attempts and raised more nestlings than less exploratory males. These results were not confounded by extra-pair paternity, which was not related to personality, or by the individuals that did not initiate any reproductive attempts at all. Our work provides evidence that attributes of personality may influence the degree to which individuals cope with, and thrive in a captive environment and this should be accounted for in both experimental design and the interpretation of results. Furthermore, this suggests that there may be selection on these traits as part of the domestication process.

Winter foraging site fidelity of king penguins breeding at the Falkland Islands

Foraging site fidelity has profound consequences for individual fitness, population processes and the effectiveness of species conservation measures. Accordingly, quantifying site fidelity has become increasingly important in animal movement and habitat selection studies. To assess foraging site fidelity in king penguins (Aptenodytes patagonicus) breeding at the Falkland Islands (51.48°S, 57.83°W), we measured overlap in time spent in foraging areas (at a 0.1° × 0.1° grid resolution) between successive foraging trips and foraging route consistency during the crèche period. In total, 30 complete foraging trips from seven king penguins were recorded between April and October 2010. King penguins predominantly foraged on the highly productive Patagonian slope, to the north of the Falkland Islands [median foraging trip distance 213 km (SD = 215 km) and duration 12.8 days (SD = 14.7 days)]. Overlap in time spent in an area on consecutive foraging trips ranged between 2 and 73 % (mean 27 %, SD = 22 %). Bearing during the outbound portion of foraging trips was typically highly repeatable for individual birds, but foraging trip duration and distance were not. Travel during the outbound phase of foraging trips was consistent with the direction of the northward-flowing Falkland Current that may act as a directional cue or facilitate rapid transit to foraging areas. Flexibility in foraging trip distances and durations may be a response to changes in resource availability and changes in the energetic requirements of adults and chicks over an extended breeding cycle.

Different male vs. female breeding periodicity helps mitigate offspring sex ratio skews in sea turtles

 The implications of climate change for global biodiversity may be profound with those species with little capacity for adaptation being thought to be particularly vulnerable to warming. A classic case of groups for concern are those animals exhibiting temperature-dependent sex-determination (TSD), such as sea turtles, where climate warming may produce single sex populations and hence extinction. We show that, globally, female biased hatchling sex ratios dominate sea turtle populations (exceeding 3:1 in >50% records), which, at-a-glance, reiterates concerns for extinction. However, we also demonstrate that more frequent breeding by males, empirically shown by satellite tracking 23 individuals and supported by a generalized bio-energetic life history model, generates more balanced operational sex ratios (OSRs). Hence, concerns of increasingly skewed hatchling sex ratios and reduced population viability are less acute than previously thought for sea turtles. In fact, in some scenarios skewed hatchling sex ratios in groups with TSD may be adaptive to ensure optimum OSRs.

Natural selection for earlier male arrival to breeding grounds through direct and indirect effects in a migratory songbird

For migratory birds, the earlier arrival of males to breeding grounds is often expected to have fitness benefits. However, the selection differential on male arrival time has rarely been decomposed into the direct effect of male arrival and potential indirect effects through female traits. We measured the directional selection differential on male arrival time in the pied flycatcher (Ficedula hypoleuca) using data from 6 years and annual number of fledglings as the fitness proxy. Using structural equation modeling, we were able to take into account the temporal structure of the breeding cycle and the hierarchy between the examined traits. We found directional selection differentials for earlier male arrival date and earlier female laying date, as well as strong selection differential for larger clutch size. These selection differentials were due to direct selection only as indirect selection for these traits was nonsignificant. When decomposing the direct selection for earlier male arrival into direct and indirect effects, we discovered that it was almost exclusively due to the direct effect of male arrival date on fitness and not due to its indirect effects via female traits. In other words, we showed for the first time that there is a direct effect of male arrival date on fitness while accounting for those effects that are mediated by effects of the social partner. Our study thus indicates that natural selection directly favored earlier male arrival in this flycatcher population.

Employing sea-level rise scenarios to strategically select sea turtle nesting habitat important for long-term management at a temperate breeding area

Management strategies to protect endangered species primarily focus on safeguarding habitats currently perceived as important (due to high-density use, rarity or contribution to the biological cycle), rather than sites of future ecological importance. This discrepancy is particularly relevant for species inhabiting beaches and coastal areas that may be lost due to sea-level rise over the next 100 years through climate change. Here, we modelled four sea-level rise (SLR) scenarios (0.2, 0.6, 0.9 and 1.3 m) to determine the future vulnerability and viability of nesting habitat (six distinct nesting beaches totalling about 6 km in length) at a key loggerhead sea turtle (Caretta caretta) rookery (Zakynthos, Greece) in the Mediterranean. For each of the six nesting beaches, we identified (1) the area of beach currently used by turtles, (2) the area of the beach anticipated to become inundated under each SLR, (3) the area of beach anticipated to become unsuitable for nesting under each SLR, (4) the potential for habitat loss under the examined SLR, and (5) the extent to which the beaches may shift in relation to natural (i.e. cliffs) and artificial (i.e. beach front development) physical barriers. Even under the most conservative 0.2 m SLR scenario, about 38% (range: 31–48%) total nesting beach area would be lost, while an average 13% (range: 7–17%) current nesting beach area would be lost. About 4 km length of nesting habitat (representing 85% of nesting activity) would be lost under the 0.9 m scenario, because cliffs prevent landward beach migration. In comparison, while the other 2 km of beach (representing 15% nests) is also at high risk, it has the capacity for landward migration, because of an adjoining sand-dune system. Therefore, managers should strengthen actions on this latter area, as a climatically critical safeguard for future sea turtle nesting activity, in parallel to regularly assessing and revising measures on the current high-use nesting habitats of this important Mediterranean loggerhead population.

Geographical variation and population structure in the White-rumped Sandpiper Calidris fuscicollis as shown by morphology, mitochondrial DNA and carbon isotope ratios

We studied the population structure of a high arctic breeding wader bird species, the White-rumped Sandpiper Calidris fuscicollis. Breeding adults, chicks and juveniles were sampled at seven localities throughout the species' breeding range in arctic Canada in 1999. The mitochondrial control region was analysed by DNA sequencing, feathers were analysed for carbon isotope ratios (C13/C12) by isotope ratio mass spectrometry, and morphological measurements were analysed using principal component analyses, taking the effect of sex into account (identified by molecular genetic methods). In general, our results support the notion that the White-rumped Sandpiper is a monotypic species with no subspecies, and most of the morphological and genetic variation occurs within sites. Nevertheless, some differences between sites were found. Birds from the two northernmost sites (Ellesmere and Devon Islands) had relatively longer bill and wing and shorter tarsus than birds sampled further south, possibly reflecting genetic differences between populations. The carbon isotope ratios were higher at the easternmost site (Baffin Island), revealing differences in the isotope content of the food. The mtDNA sequences showed no significant differentiation between sites and no pattern of isolation-by-distance was found. Based on the mtDNA variation, the species was estimated to have a long-term effective population size of approximately 9,000 females. The species shows no clear evidence of any population expansion or decline. Our results indicate that carbon isotope ratios, and possibly also certain mtDNA haplotypes, may be useful as tools for identifying the breeding origin of White-rumped Sandpipers on migration and wintering sites.

Breeding habitat selection in an obligate beach bird: a test of the food resource hypothesis

The food resource hypothesis of breeding habitat selection in beach-nesting birds suggests that birds breed at sites with more prey to meet the increased energetic requirements associated with breeding. We compare prey resources using pitfall traps and core samples at breeding sites and absence sites of the eastern population of hooded plover, Thinornis rubricollis rubricollis, which, in this part of its range, is a threatened obligate beach bird. Breeding sites had higher abundances, equivalent species richness, and different assemblages of invertebrate prey compared with absence sites. Assemblages at breeding sites were characterised by more isopods, and fewer beetles of the family Phycosecidae. Breeding habitat selection by plovers appears to be associated with selection for sites with more food, and any process that degrades food resources at a site (e.g. kelp harvesting or marine pollution events) may reduce the likelihood of occupancy of that site by breeding birds.

Loss of periodicity in breeding success of waders links to changes in lemming cycles in Arctic ecosystems

Lemming population cycles in the Arctic have an important impact on the Arctic food web, indirectly also affecting breeding success in Arctic-nesting birds through shared predators. Over the last two decades lemming cycles have changed in amplitude and even disappeared in parts of the Arctic. To examine the large scale effect of these recent changes we re-analysed published data from the East Atlantic Flyway (EAF), where a relationship between lemming cycles and wader breeding success was earlier found, and new data on breeding success of waders in the East Asian-Australasian Flyway (EAAF). We found that 1) any long-term periodicities in wader breeding success existed only until the year 2000 in the EAAF and until the 1980s in the EAF; 2) studying these patterns at a smaller spatial scale, where the Siberian-Alaskan breeding grounds were divided into five geographical units largely based on landscape features, breeding success of waders from the EAAF was not correlated to an index of predation pressure, but positively correlated to Arctic summer temperatures in some species. We argue that fading out of lemming cycles in some parts of the Arctic is responsible for faltering periodicity in wader breeding success along both flyways. These changed conditions have not yet resulted in any marked changing trends in breeding success across years, and declining numbers of waders along the EAAF are therefore more likely a result of changing conditions at stop-over and wintering sites.

Sequential polyandry through divorce and re-pairing in a cooperatively breeding bird reduces helper-offspring relatedness

Polyandry is an important component of both sexual selection and kin structuring within cooperatively breeding species. A female may have multiple partners within a single reproductive attempt (simultaneous polyandry) or across multiple broods within and/or across years (sequential polyandry). Both types of polyandry confer a range of costs and benefits to individuals and alter the genetic structure of social groups over time. To date, many molecular studies of cooperative breeders have examined the evolution of cooperative breeding in relation to simultaneous polyandry. However, cooperatively breeding vertebrates are iteroparous, and thus sequential polyandry is also likely, but more rarely considered in this context. We examined sequential polyandry in a cooperatively breeding bird that has a low level of within-brood polyandry. Over a 5-year period (2006–2010), we monitored individual mating relationships using molecular markers in a population of individually marked apostlebirds (Struthidea cinerea). Divorce occurred between reproductive seasons in 17 % (8/48) of pairs and appeared to be female-driven. The level of sequential polyandry was also driven by the disappearance of males after breeding, and over 90 % of females, for whom we had suitable data, bred with multiple males over the period of study. This sequential polyandry significantly altered the relatedness of group members to the offspring in the nest. However, in about half of the cases, the second male was related (first- or second-order relative) to the first male of a sequentially polyandrous female and this alleviated the reduction in relatedness caused by polyandry. Our findings suggest that even in species with high within-brood parentage certainty, helper-offspring relatedness values can quickly erode through sequential polyandry.

Effects of sea temperature and stratification changes on seabird breeding success

As apex predators in marine ecosystems, seabirds may primarily experience climate change impacts indirectly, via changes to their food webs. Observed seabird population declines have been linked to climate-driven oceanographic and food web changes. However, relationships have often been derived from relatively few colonies and consider only sea surface temperature (SST), so important drivers, and spatial variation in drivers, could remain undetected. Further, ex - plicit climate change projections have rarely been made, so longer-term risks remain unclear. Here, we use tracking data to estimate foraging areas for 11 black-legged kittiwake Rissa tridac - ty la colonies in the UK and Ireland, thus reducing reliance on single colonies and allowing calculation of colony-specific oceanographic conditions. We use mixed models to consider how SST, the potential energy anomaly (indicating density stratification strength) and the timing of seasonal stratification influence kittiwake productivity. Across all colonies, higher breeding success was associated with weaker stratification before breeding and lower SSTs during the breeding season. Eight colonies with sufficient data were modelled individually: higher productivity was associated with later stratification at 3 colonies, weaker stratification at 2, and lower SSTs at one, whilst 2 colonies showed no significant relationships. Hence, key drivers of productivity varied among colonies. Climate change projections, made using fitted models, indicated that breeding success could decline by 21 to 43% between 1961-90 and 2070-99. Climate change therefore poses a longer-term threat to kittiwakes, but as this will be mediated via availability of key prey species, other marine apex predators could also face similar threats.

Sex-specific arrival times on the breeding grounds: hybridizing migratory skuas provide empirical support for the role of sex ratios

In migratory animals, protandry (earlier arrival of males on the breeding grounds) prevails over protogyny (females preceding males). In theory, sex differences in timing of arrival should be driven by the operational sex ratio, shifting toward protogyny in female-biased populations. However, empirical support for this hypothesis is, to date, lacking. To test this hypothesis, we analyzed arrival data from three populations of the long-distance migratory south polar skua (Catharacta maccormicki). These populations differed in their operational sex ratio caused by the unidirectional hybridization of male south polar skuas with female brown skuas (Catharacta antarctica lonnbergi). We found that arrival times were protandrous in allopatry, shifting toward protogyny in female-biased populations when breeding in sympatry. This unique observation is consistent with theoretical predictions that sex-specific arrival times should be influenced by sex ratio and that protogyny should be observed in populations with female-biased operational sex ratio.