Unexpected diversity in socially synchronized rhythms of shorebirds.

The behavioural rhythms of organisms are thought to be under strong selection, influenced by the rhythmicity of the environment. Such behavioural rhythms are well studied in isolated individuals under laboratory conditions, but free-living individuals have to temporally synchronize their activities with those of others, including potential mates, competitors, prey and predators. Individuals can temporally segregate their daily activities (for example, prey avoiding predators, subordinates avoiding dominants) or synchronize their activities (for example, group foraging, communal defence, pairs reproducing or caring for offspring). The behavioural rhythms that emerge from such social synchronization and the underlying evolutionary and ecological drivers that shape them remain poorly understood. Here we investigate these rhythms in the context of biparental care, a particularly sensitive phase of social synchronization where pair members potentially compromise their individual rhythms. Using data from 729 nests of 91 populations of 32 biparentally incubating shorebird species, where parents synchronize to achieve continuous coverage of developing eggs, we report remarkable within- and between-species diversity in incubation rhythms. Between species, the median length of one parent's incubation bout varied from 1-19 h, whereas period length-the time in which a parent's probability to incubate cycles once between its highest and lowest value-varied from 6-43 h. The length of incubation bouts was unrelated to variables reflecting energetic demands, but species relying on crypsis (the ability to avoid detection by other animals) had longer incubation bouts than those that are readily visible or who actively protect their nest against predators. Rhythms entrainable to the 24-h light-dark cycle were less prevalent at high latitudes and absent in 18 species. Our results indicate that even under similar environmental conditions and despite 24-h environmental cues, social synchronization can generate far more diverse behavioural rhythms than expected from studies of individuals in captivity. The risk of predation, not the risk of starvation, may be a key factor underlying the diversity in these rhythms.

Cryptic species and independent origins of allochronic populations within a seabird species complex (Hydrobates spp.).

Humans are inherently biased towards naming species based on morphological differences, which can lead to reproductively isolated species being mistakenly classified as one if they are morphologically similar. Recognising cryptic diversity is needed to understand drivers of speciation fully, and for accurate estimates of global biodiversity and assessments for conservation. We investigated cryptic species across the range of band-rumped storm-petrels (Hydrobates spp.): highly pelagic, nocturnal seabirds that breed on tropical and sub-tropical islands in the Atlantic and Pacific Oceans. In many breeding colonies, band-rumped storm-petrels have sympatric but temporally isolated (allochronic) populations; we sampled all breeding locations and allochronic populations. Using mitochondrial control region sequences from 754 birds, cytochrome b sequences from 69 birds, and reduced representation sequencing of the nuclear genomes of 133 birds, we uncovered high levels of genetic structuring. Population genomic analyses revealed up to seven unique clusters, and phylogenomic reconstruction showed that these represent seven monophyletic groups. We uncovered up to six independent breeding season switches across the phylogeny, spanning the continuum from genetically undifferentiated temporal populations to full allochronic species. Thus, band-rumped storm-petrels encompass multiple cryptic species, with non-geographic barriers potentially comprising strong barriers to gene flow.

Recurrent hybridization and recent origin obscure phylogenetic relationships within the 'white-headed' gull (Larus sp.) complex.

Species complexes that have undergone recent radiations are often characterized by extensive allele sharing due to recent ancestry and (or) introgressive hybridization. This can result in discordant evolutionary histories of genes and heterogeneous genomes, making delineating species limits difficult. Here we examine the phylogenetic relationships among a complex group of birds, the white-headed gulls (Aves: Laridae), which offer a unique window into the speciation process due to their recent evolutionary history and propensity to hybridize. Relationships were examined among 17 species (61 populations) using a multilocus approach, including mitochondrial and nuclear intron DNA sequences and microsatellite genotype information. Analyses of microsatellite and intron data resulted in some species-based groupings, although most species were not represented by a single cluster. Considerable allele and haplotype sharing among white-headed gull species was observed; no locus contained a species-specific clade. Despite this, our multilocus approach provided better resolution among some species than previous studies. Interestingly, most clades appear to correspond to geographic locality: our BEAST analysis recovered strong support for a northern European/Icelandic clade, a southern European/Russian clade, and a western North American/canus clade, with weak evidence for a high latitude clade spanning North America and northwestern Europe. This geographical structuring is concordant with behavioral observations of pervasive hybridization in areas of secondary contact. The extent of allele and haplotype sharing indicates that ecological and sexual selection are likely not strong enough to complete reproductive isolation within several species in the white-headed gull complex. This suggests that just a few genes are driving the speciation process.

North or south? Phylogenetic and biogeographic origins of a globally distributed avian clade.

Establishing phylogenetic relationships within a clade can help to infer ancestral origins and indicate how widespread species reached their current biogeographic distributions. The small plovers, genus Charadrius, are cosmopolitan shorebirds, distributed across all continents except Antarctica. Here we present a global, species-level molecular phylogeny of this group based on four nuclear (ADH5, FIB7, MYO2 and RAG1) and two mitochondrial (COI and ND3) genes, and use the phylogeny to examine the biogeographic origin of the genus. A Bayesian multispecies coalescent approach identified two major clades (CRD I and CRD II) within the genus. Clade CRD I contains three species (Thinornis novaeseelandiae, Thinornis rubricollis and Eudromias morinellus), and CRD II one species (Anarhynchus frontalis), that were previously placed outside the Charadrius genus. In contrast to earlier work, ancestral area analyses using parsimony and Bayesian methods supported an origin of the Charadrius plovers in the Northern hemisphere. We propose that major radiations in this group were associated with shifts in the range of these ancestral plover species, leading to colonisation of the Southern hemisphere.

Genetic and phenotypic characterization of a hybrid zone between polyandrous Northern and Wattled Jacanas in Western Panama.

BACKGROUND: Hybridization provides a unique perspective into the ecological, genetic and behavioral context of speciation. Hybridization is common in birds, but has not yet been reported among bird species with a simultaneously polyandrous mating system; a mating system where a single female defends a harem of males who provide nearly all parental care. Unlike simple polyandry, polyandrous mating is extremely rare in birds, with only 1% of bird species employing this mating system. Although it is classically held that females are "choosy" in avian hybrid systems, nearly-exclusive male parental care raises the possibility that female selection against heterospecific matings might be reduced compared to birds with other mating systems. RESULTS: We describe a narrow hybrid zone in southwestern Panama between two polyandrous freshwater waders: Northern Jacana, Jacana spinosa and Wattled Jacana, J. jacana. We document coincident cline centers for three phenotypic traits, mtDNA, and one of two autosomal introns. Cline widths for these six markers varied from seven to 142 km, with mtDNA being the narrowest, and five of the six markers having widths less than 100 km. Cline tails were asymmetrical, with greater introgression of J. jacana traits extending westward into the range of J. spinosa. Likewise, within the hybrid zone, the average hybrid index of phenotypic hybrids was significantly biased towards J. spinosa. Species distribution models indicate that the hybrid zone is located at the edge of a roughly 100 km wide overlap where habitat is predicted to be suitable for both species, with more westerly areas suitable only for spinosa and eastward habitats suitable only for J. jacana. CONCLUSION: The two species of New World jacanas maintain a narrow, and persistent hybrid zone in western Panama. The hybrid zone may be maintained by the behavioral dominance of J. spinosa counterbalanced by unsuitable habitat for J. spinosa east of the contact zone. Although the two parental species are relatively young, mitochondrial cline width was extremely narrow. This result suggests strong selection against maternally-inherited markers, which may indicate either mitonuclear incompatibilities and/or female choice against heterospecific matings typical of avian hybrid systems, despite jacana sex role reversal.

Egg sampling as a possible alternative to blood sampling when monitoring the exposure of yellow-legged gulls (Larus michahellis) to avian influenza viruses.

We explored whether antibody detection in egg yolks could serve as an alternative to antibody detection in plasma samples when monitoring yellow-legged gulls (Larus michahellis) for exposure to avian influenza viruses (AIVs). We tested female plasma and eggs for anti-AIV antibodies and used the data we obtained to check whether the two sample types yielded the same antibody status (positive or negative) and to compare the antibody prevalence estimated from the blood data with that estimated from the yolk data. Our results showed that sampling one egg per clutch, regardless of that egg's position in the laying sequence, is sufficient to provide an unbiased estimate of antibody prevalence across clutches. The results also showed that almost 25% of the clutches laid by positive females contained only antibody-negative eggs, which suggests that yolk samples might underestimate female antibody prevalence. However, this result may stem from differences in the methods used to assess plasma versus yolk antibody status. Further research is needed to clarify this issue; while the number of false negatives could be reduced by adapting antibody detection techniques, it may be that they are an unavoidable consequence of natural avian maternal transfer dynamics.

Sexual selection on brain size in shorebirds (Charadriiformes).

Natural selection is considered a major force shaping brain size evolution in vertebrates, whereas the influence of sexual selection remains controversial. On one hand, sexual selection could promote brain enlargement by enhancing cognitive skills needed to compete for mates. On the other hand, sexual selection could favour brain size reduction due to trade-offs between investing in brain tissue and in sexually selected traits. These opposed predictions are mirrored in contradictory relationships between sexual selection proxies and brain size relative to body size. Here, we report a phylogenetic comparative analysis that highlights potential flaws in interpreting relative brain size-mating system associations as effects of sexual selection on brain size in shorebirds (Charadriiformes), a taxonomic group with an outstanding diversity in breeding systems. Considering many ecological effects, relative brain size was not significantly correlated with testis size. In polyandrous species, however, relative brain sizes of males and females were smaller than in monogamous species, and females had smaller brain size than males. Although these findings are consistent with sexual selection reducing brain size, they could also be due to females deserting parental care, which is a common feature of polyandrous species. Furthermore, our analyses suggested that body size evolved faster than brain size, and thus the evolution of body size may be confounding the effect of the mating system on relative brain size. The brain size-mating system association in shorebirds is thus not only due to sexual selection on brain size but rather, to body size evolution and other multiple simultaneous effects.

[Food habits and winter diet of Charadrius melodus (Charadriiformes: Charadriidae) in Boca Ciega, Tamaulipas, Mexico]. / Hábitos alimenticios y dieta invernal de Charadrius melodus (Charadriiformes: Charadriidae) en Boca Ciega, Tamaulipas, México.

The Piping Plover (Charadrius melodus) is a migratory endangered species that arrives, along with a great number of other winter migratory birds, to Boca Ciega every year. In spite of the importance of this ecosystem, these species, are threatened by the current habitat change caused by the dredging activities in the area. With the aim to generate new information about the importance of this area during winter, we studied C melodus activities during the winter season in Laguna Madre, from December 2009 to March 2010. Our objectives were: 1) determine the importance of the area during winter, 2) describe C. melodus ethology, feeding substrate preferences and food items, 3) to analyze and describe the sympatric diversity associated with C melodus. A total of ninety nine individuals were observed during the monitoring. The Cochran and Kendall test showed a high significance of the species with the substrate and signs tests using a binomial distribution that indicated a high preference for algal type of substrate. The highest activity recorded for this species during this winter season was feeding. The principal food items found in sediments were larvae of Diptera: Chironomidae and Ephydridae. The sympatric species of C. melodus were two families of Charadriiforms: Scolopacidae (nine species) and Charadriidae (two species). We concluded that this is an important area for feeding, protection and rest sites for this species, and its protection and management is recommended.

[Temporal and spatial distribution of shorebirds (Charadriiformes) at San Ignacio Lagoon, Baja California Sur, Mexico]. / Distribución espacial y temporal de aves playeras (Orden: Charadriiformes) en Laguna San Ignacio, Baja California Sur, Mexico.

Baja California Peninsula has several wetlands that represent important ecosystems for shorebirds. San Ignacio Lagoon is one of these sites, and supports 10% of the total abundance of shorebirds reported in this Peninsula. Since there is few information about this group in this area, we studied spatial and temporal changes in abundance and distribution of shorebirds in San Ignacio Lagoon. For this, we conducted twelve monthly censuses (October 2007-September 2008) on the entire internal perimeter of the lagoon, which we divided into four areas: two at the North and two at the South. We observed a seasonal pattern, with the lowest abundance in May (1 585 birds) and the highest in October (47 410). The most abundant species were Marbled Godwits (Limosa fedoa; 55% of the total records), Western Sandpipers (Calidris mauri; 23%), and Willet (Tringa semipalmata; 10%). All three species were more abundant in autumn; for both, the Marbled Godwit and Willet, we observed their highest numbers in winter and spring, while the Western Sandpiper showed noticeable oscillations, reaching a maximum in early winter (December). In summer, Marbled Godwit and Willet were the only birds present but in lower numbers. Here present the first records of the Pacific Red Knot (Calidris canutus roselaari) in the area. Bird abundance and species richness were influenced seasonally by migration and spatially by sites in the lagoon. The greatest shorebird abundance was in the South area of the lagoon, probably because of better accessibility to food. Our results allowed the inclusion of San Ignacio Lagoon in the Western Hemisphere Shorebirds Reserve Network (WHSRN) as a site of international importance.

Variable and complex food web structures revealed by exploring missing trophic links between birds and biofilm.

Food webs are comprised of a network of trophic interactions and are essential to elucidating ecosystem processes and functions. However, the presence of unknown, but critical networks hampers understanding of complex and dynamic food webs in nature. Here, we empirically demonstrate a missing link, both critical and variable, by revealing that direct predator-prey relationships between shorebirds and biofilm are widespread and mediated by multiple ecological and evolutionary determinants. Food source mixing models and energy budget estimates indicate that the strength of the missing linkage is dependent on predator traits (body mass and foraging action rate) and the environment that determines food density. Morphological analyses, showing that smaller bodied species possess more developed feeding apparatus to consume biofilm, suggest that the linkage is also phylogenetically dependent and affords a compelling re-interpretation of niche differentiation. We contend that exploring missing links is a necessity for revealing true network structure and dynamics.

Eight independent nuclear genes support monophyly of the plovers: the role of mutational variance in gene trees.

Molecular phylogenies of Charadriiformes based on mtDNA genes and one to three nuclear loci do not support the traditional placement of Pluvialis in the plovers (Charadriidae), assigning it instead to oystercatchers, stilts, and avocets (Haematopodidae and Recurvirostridae). To investigate this hypothesis of plover paraphyly, the relationships among Pluvialis and closely related families were revisited by sequencing two individuals of all taxa except Peltohyas for eight independent single copy nuclear protein-coding loci selected for their informativeness at this phylogenetic depth. The species tree estimated jointly with the gene trees in the coalescent programme (*)BEAST strongly supported plover monophyly, as did Bayesian analysis of the concatenated matrix. The data sets that supported plover paraphyly in Baker et al. (2007) and Fain and Houde (2007) reflect two to four independent gene histories, and thus discordance with the plover monophyly species tree might have arisen by chance through stochastic mutational variance. For the plovers we conclude there is no conclusive evidence of coalescent variance from ancient incomplete lineage sorting across the interior branch leading to Pluvialis in the species tree. Rather, earlier studies seem have been misled by faster evolving mtDNA genes with high mutational variance, and a few nuclear genes that had low resolving power at the Pluvialis sister group level. These findings are of general relevance in avian phylogenetics, as they show that careful attention needs to be paid to the number and the phylogenetic informativeness of genes required to obtain accurate estimates of the species tree, especially where there is mutational heterogeneity in gene trees.

Multiple gene sequences resolve phylogenetic relationships in the shorebird suborder Scolopaci (Aves: Charadriiformes).

Shorebirds (Charadriiformes) are a diverse assemblage of species renowned for their variation in behavior, morphology, and life-history traits, but comparative studies of trait variation remain limited by the lack of a well-supported phylogeny based on DNA sequences. In this study we build upon previous shorebird phylogenies to construct the first sequence-based species-level phylogeny for the Scolopaci, one of three shorebird suborders. We sampled 84 species in the Scolopaci, and collected data for five genes (one nuclear and four mitochondrial) via PCR and sequencing or from GenBank. The phylogeny was estimated using Bayesian inference on a partitioned dataset of 6365 aligned base pairs, and was well-supported except for the radiations within Tringa and Calidris. The shanks and phalaropes are sister to the snipes, woodcocks and dowitchers, which in turn are sister to the sandpipers. The godwits and curlews are successive sister-groups to these clades, and the morphologically disparate taxa (jacanas, painted snipes, seedsnipes, and the Plains-wanderer) are the basal sister-group in the Scolopaci. We show that Tringa, Gallinago, and Calidris are paraphyletic assemblages, and thus are in need of taxonomic revision. The clade of Calidridine sandpipers has very short internal branches indicative of a relatively recent rapid radiation, and will require a gene tree/species tree approach to resolve relationships among species.

Reproductive success of South American terns (Sterna hirundinacea) from Cardos Islands, Florianópolis, SC, Brazil.

Sterna hirundinacea (Lesson, 1831) is a migratory seabird that breeds in the Pacific Coast (from Peru to Chile) and along the Atlantic coast of South America from Espírito Santo (Brazil) to Terra del Fuego (Argentina). This paper describes the reproductive success of South American Terns on Cardos Island, Florianopolis, Brazil in the breeding seasons of 2003, 2005 and 2006. The colony was formed in mid-May in 2003 and early April in other years, with the total number of nests ranging from 1,852 in 2006 to 2,486 in 2005. Hatching success was estimated at 76.39% in 2006, 62.73% in 2003 and 41.1% in 2005, the lowest value that could be attributed to predation by hawks Caracara plancus, lizards Tupinambis merianae and black vulture Coragyps atratus. The chicks hatched in July in 2003, and in June 2005 and 2006, and fledging success was 50.94%, 35.96 and 53.47% respectively. Cardos Island has been constantly used as a breeding site by South American Terns, and therefore represents an important area for conservation of this species. This success could be attributed to low pressure of Kelp gulls (Larus dominicanus), the main predator of seabirds along the Brazilian coast.

[Analysis of the complete mitochondrial genome sequence of Larus brunnicephalus (Aves, Laridae)].

The complete sequence of mitochondrial genome of Larus brunnicephalus was determined using long PCR and conserved primers walking approaches. The results showed that the entire mitochondrial genome of L. brunnicephalus is 16,769 bp in length, which has been deposited in GenBank with the accession number JX155863. The mitochondrial genomic organization and gene order of L. brunnicephalus were consistent with that of Gallus gallus, which contains 13 protein coding genes (PCGs), 22 tRNA, 2 rRNA, and a control region. Except for COI gene using GTG and ND3 gene with ATT as the initiation codon, all other 11 PCGs of the mtDNA in L. brunnicephalus started with the typical ATG codon. AGG, TAG, TAA, or AGA were used in 11 PCGs as usual termination codons, except for COIII and ND4 genes with incomplete termination codon (T). The secondary structures of 22 tRNAs were predicted and it is found that the tRNASer (AGN) lacks DHU arm and tRNAPhe contains the fourth types of permutation in the TψC arm. It is predicted that the secondary structures of 12S rRNA and 16S rRNA include 4 structural domains with 47 helics and 6 domains with 60 helics, respectively. F-box, E-box, D-box, C-box, B-box, Bird similarity-box, and CSB-boxes (1-3), which were found in the control regions of other bird species were also present in L. brunnicephalus. The sequence in the starting regions of H-strand replication (OH) and the bidirectional light and heavy-strand transcription promoters (LSP/HSP) in the control region were also predicted. Result of phylogeny analysis supports that L. brunnicephalus should be categorized into the Masked gulls species.

Discord reigns among nuclear, mitochondrial and phenotypic estimates of divergence in nine lineages of trans-Beringian birds.

Proposals for genetic thresholds for species delimitation assume that simple genetic data sets (e.g. mitochondrial sequence data) are correlated with speciation; i.e. such data sets accurately reflect organismal lineage divergence. We used taxonomically stratified phenotypic levels of differentiation (populations, subspecies and species) among nine avian lineages using paired, trans-Beringian samples from three lineages each in three orders (Anseriformes, Charadriiformes, and Passeriformes) to test this assumption. Using mitochondrial DNA sequence data and nuclear genomic data (amplified fragment length polymorphisms), we found a lack of concordance between these two genomes in their respective estimates of divergence and little or no relationship between phenotype (taxonomic relatedness) and genetic differentiation between taxon pairs. There are several possible reasons for the discord observed (e.g. selection on one of the genomes or perhaps lineage sorting), but the implications are that genetic estimates of lineage divergence may not be correlated with estimates from other parts of the genome, are not well correlated with the speciation process and are thus not reliable indicators of species limits.

Use of fatty acid analysis to determine dispersal of caspian terns in the Columbia River Basin, USA.

Lethal control, which has been used to reduce local abundances of animals in conflict with humans or with endangered species, may not achieve management goals if animal movement is not considered. In populations with emigration and immigration, lethal control may induce compensatory immigration, if the source of attraction remains unchanged. Within the Columbia River Basin (Washington, U.S.A.), avian predators forage at dams because dams tend to reduce rates of emigration of juvenile salmonids (Oncorhynchus spp.), artificially concentrating these prey. We used differences in fatty acid profiles between Caspian Terns (Hydroprogne caspia) at coastal and inland breeding colonies and terns culled by a lethal control program at a mid-Columbia River dam to infer dispersal patterns. We modeled the rate of loss of fatty acid biomarkers, which are fatty acids that can be traced to a single prey species or groups of species, to infer whether and when terns foraging at dams had emigrated from the coast. Nonmetric multidimensional scaling showed that coastal terns had high levels of C(20) and C(22) monounsaturated fatty acids, whereas fatty acids of inland breeders were high in C18:3n3, C20:4n6, and C22:5n3. Models of the rate of loss of fatty acid showed that approximately 60% of the terns collected at Rock Island Dam were unlikely to have bred successfully at local (inland) sites, suggesting that terns foraging at dams come from an extensive area. Fatty acid biomarkers may provide accurate information about patterns of dispersal in animal populations and may be extremely valuable in cases where populations differ demonstrably in prey base.

Closely related gull species show contrasting foraging strategies in an urban environment.

The expansion of urban landscapes has both negative and positive effects on wildlife. Understanding how different species respond to urbanization is key to assessing how urban landscapes influence regional wildlife behavior and ecosystem structure. Gulls are often described as strong urban adapters, but few studies have explored species-specific differences in habitat use. Here, we use GPS tracking in conjunction with stable isotope analysis (SIA) to quantify the habitat use and trophic ecology of great black-backed gulls (Larus marinus) and herring gulls (L. argentatus) in an urbanized area. Non-Metric Multidimensional Scaling (NMDS) of foraging locations revealed significant differences in the habitat use between species. Great black-backed gulls foraged primarily in marine habitats and herring gulls foraged primarily in specific urban habitats (e.g., landfills, dumpsters) and showed higher site fidelity in terms of the proportion of foraging sites revisited. Further, great black-backed gulls had significantly higher δ15N and δ13C than herring gulls, reflecting the use of marine, rather than urban, food sources. This study highlights the variability in urban habitat utilization among closely related species, assesses stable isotope signatures of urban diets in wild birds, and discusses ecological implications of the relative contribution of urban and marine foraging.

Working through polytomies: auklets revisited.

Polytomies, or phylogenetic "bushes", are the result of a series of internodes occurring in a short period of evolutionary time (which can result in data that do not contain enough information), or data that have too much homoplasy to resolve a bifurcating branching pattern. In this study we used the Aethia auklet polytomy to explore the effectiveness of different methods for resolving polytomies: mitochondrial DNA gene choice, number of individuals per species sampled, model of molecular evolution, and AFLP loci. We recovered a fully-resolved phylogeny using NADH dehydrogenase subunit 2 (ND2) sequence data under two different Bayesian models. We were able to corroborate this tree under one model with an expanded mtDNA dataset. Effectiveness of additional intraspecific sampling varied with node, and fully 20% of the subsampled datasets failed to return a congruent phylogeny when we sampled only one or two individuals per species. We did not recover a resolved phylogeny using AFLP data. Conflict in the AFLP dataset showed that nearly all possible relationships were supported at low levels of confidence, suggesting that either AFLPs are not useful at the genetic depth of the Aethia auklet radiation (7-9% divergent in the mtDNA ND2 gene), perhaps resulting in too much homoplasy, or that the Aethia auklets have experienced incomplete lineage sorting at many nuclear loci.

How did the Great Auk raise its young?

The extant auks show three strategies of chick rearing--precocial (chicks leave the nest site when a few days old), intermediate (young raised to a mass of around 20% of adult mass) and semi-precocial (young raised to a mass of around 65% of adult mass). It is not known which strategy the extinct Great Auk used. In this paper, we investigate this issue by a novel combination of a time and energy budget model and phylogenetic comparison. The first approach indicates that for reasonable estimates of the equation parameters, the Great Auk could have followed an intermediate strategy. For a limited range of parameters, the Great Auk could have followed the semi-precocial strategy. Phylogenetic comparison shows that it is unlikely that the Great Auk followed a precocial strategy. The results suggest that the Great Auk followed an intermediate strategy as does its presumed closest extant relative the Razorbill.

Intraseasonal patterns in shorebird nest survival are related to nest age and defence behaviour.

Nest survival may vary throughout the breeding season for many bird species, and the nature of this temporal variation can reveal the links between birds, their predators, and other components of the ecosystem. We used program Mark to model patterns in nest survival within the breeding season for shorebirds nesting on arctic tundra. From 2000 to 2007, we monitored 521 nests of five shorebird species and found strong evidence for variation in nest survival within a nesting season. Daily nest survival was lowest in the mid-season in 5 of 8 years, but the timing and magnitude of the lows varied. We found no evidence that this quadratic time effect was driven by seasonal changes in weather or the abundance of predators. Contrary to our prediction, the risk of predation was not greatest when the number of active shorebird nests was highest. Although nest abundance reached a maximum near the middle of the breeding season, a daily index of shorebird nest activity was not supported as a predictor of nest survival in the models. Predators' access to other diet items, in addition to shorebird nests, may instead determine the temporal patterns of nest predation. Nest survival also displayed a positive, linear relationship with nest age; however, this effect was most pronounced among species with biparental incubation. Among biparental species, parents defended older nests with greater intensity. We did not detect a similar relationship among uniparental species, and conclude that the stronger relationship between nest age and both nest defence and nest survival for biparental species reflects that their nest defence is more effective.