In summer 2017 I studied the abundance and distribution of marine associated birds and mammals at the Five Finger Lighthouse in Southeast Alaska. My objectives were (1) to identify the areas of highest habitat use by species of conservation concern, (2) to make recommendations for an ecosystem-based management plan for the island, and (3) to initiate a citizen science project. I found higher relative abundance and greater biodiversity of both birds and marine mammals on the south and west facing sectors of the island compared to the north and east facing sectors. I attribute this to the greater habitat complexity on the south and west facing sectors that comprise a near-shore reef, a mixed kelp forest, and a channel between the reef and rocky cliffs, areas used extensively for foraging, nesting, traveling, socializing, and resting by many of the documented species. These findings provided the basis for recommendations to avoid development and to minimize anthropogenic disturbance on the southern and western portions of the island including the adjacent reef and channel. As both the Five Finger Lighthouse ecosystem and management continue to evolve in response to changing environmental conditions and human interests, this study also established a baseline for future study that will inform future adaptive management, document changes over time, and engage community stakeholders in science and conservation.

In summer 2017 I studied the abundance and distribution of marine associated birds and mammals at the Five Finger Lighthouse in Southeast Alaska. My objectives were (1) to identify the areas of highest habitat use by species of conservation concern, (2) to make recommendations for an ecosystem-based management plan for the island, and (3) to initiate a citizen science project. I found higher relative abundance and greater biodiversity of both birds and marine mammals on the south and west facing sectors of the island compared to the north and east facing sectors. I attribute this to the greater habitat complexity on the south and west facing sectors that comprise a near-shore reef, a mixed kelp forest, and a channel between the reef and rocky cliffs, areas used extensively for foraging, nesting, traveling, socializing, and resting by many of the documented species. These findings provided the basis for recommendations to avoid development and to minimize anthropogenic disturbance on the southern and western portions of the island including the adjacent reef and channel. As both the Five Finger Lighthouse ecosystem and management continue to evolve in response to changing environmental conditions and human interests, this study also established a baseline for future study that will inform future adaptive management, document changes over time, and engage community stakeholders in science and conservation.

Semipalmated Sandpiper (Calidris pusilla) populations have undergone significant declines at core nonbreeding sites in northeastern South America. Breeding populations have also declined in the eastern North American Arctic, but appear to be stable or increasing in the central and western Arctic. To identify vulnerable populations and sites, we documented the migratory connectivity of Semipalmated Sandpipers using light-level geolocators, deploying 250 at 8 Arctic sites across the species' breeding range from 2011 to 2015, plus 87 at a single wintering site in northeastern Brazil in 2013 and 2014. We recovered 59 units and resighted 7 more (26% return rate) on the breeding grounds, but none at the nonbreeding site. We recovered only similar to 3% of units deployed in 2013 at eastern Arctic breeding sites, but recovered 33% of those deployed in 2015. Overall, birds with geolocators were 57% as likely to return as those carrying alphanumeric flags. Stopover durations at prairie sites (mean: 8.7 days southbound, 6.7 days northbound) were comparable with durations estimated by local banding studies, but geolocator-tagged birds had longer stopovers than previously estimated at James and Hudson Bay, the Bay of Fundy, and the Gulf of Mexico. Migration routes confirmed an eastern Arctic connection with northeastern South America. Birds from eastern Alaska, USA, and far western Canada wintered from Venezuela to French Guiana. Central Alaskan breeders wintered across a wider range from Ecuador to French Guiana. Birds that bred in western Alaska wintered mainly on the west coasts of Central America and northwestern South America, outside the nonbreeding region in which population declines have been observed. Birds that bred in the eastern Arctic and used the Atlantic Flyway wintered in the areas in South America where declines have been reported, whereas central Arctic-breeding populations were apparently stable. This suggests that declines may be occurring on the Atlantic Flyway and in the eastern Arctic region.

------------------------------------------------------------------------------------------------------Description of the dataset "Supplementary Data 3 - Study sites.csv"--------------------------------------------------------------------------------------------------------The dataset- is used in the paper "Unexpected diversity in socially synchronized rhythms of shorebirds" Nature 2016 by M. Bulla et al- contains estimates of mean female and male wing length for each population of biparental shorebirds from a specific study site, plus the locations of the study site, whether the locations had tide, and whether the tide was used by the population for foraging, and how the incubation was monitored.--------------------------------------------------------------------------------------------------------Questions can be directed to: Martin Bulla (bulla.mar@gmail.com)--------------------------------------------------------------------------------------------------------Values are separated by comma.--------------------------------------------------------------------------------------------------------1. scinam : scientific name of the species2. sp : four letter abbreviation of the species's English name3. study_site : name of the study site4. site_abbreviation : four letter abbreviation of the study site5. type : was the study site at the breeding ground (breeding) or not (wintering)6. lat : latitude of the study site (decimal)7. lon : longitude of the study site (decimal)8. tidal_habitat : is the study site at primarily tidal habitat (y=yes, n=no)9. tidal_used : if the study site is at primarily tidal habitat, do the birds use it for foraging (y=yes, n=no)10. incubation_monitoring : method used to monitor incubation (for details see the paper's Extended Data Table 4)11. sexing_method : identifies the method used to sex individuals to estimate the mean female and male wing length12. pop_wing_f : mean female wing length for the population13. f_wing_N : sample size used for the female mean estimate14. pop_wing_m : mean male wing length for the population15. m_wing_N : sample size used for the male mean estimate16. data_source : is the mean wing estimate based on the primary data ("our primary data") or literature (citation))--------------------------------------------------------------------------------------------------------WHEN USING THIS DATA, PLEASE CITE:Bulla et al (2016). Supplementary Data 3 - Study sites: location, population wing length, monitoring method, tide.figshare. https://doi.org/10.6084/m9.figshare.1536260. Retrieved ADD DATETIME.--------------------------------------------------------------------------------------------------------

Geolocators are useful for tracking movements of long-distance migrants, but potential negative effects on birds have not been well studied. We tested for effects of geolocators (0.8–2.0 g total, representing 0.1–3.9 % of mean body mass) on 16 species of migratory shorebirds, including five species with 2–4 subspecies each for a total of 23 study taxa. Study species spanned a range of body sizes (26–1091 g) and eight genera, and were tagged at 23 breeding and eight nonbreeding sites. We compared breeding performance and return rates of birds with geolocators to control groups while controlling for potential confounding variables.

Socially synchronized rhythms in shorebirds were assessed during biparental incubation under natural circumstances and were exceptionally diverse, often not following the 24-h day, whereby risk of predation, not starvation, determined some of the variation in incubation rhythms. All organisms have biorhythms, but in social species these have to be synchronized between individuals within a community. Here Martin Bulla et al. address the issue of how parents synchronize their biorhythms when both are caring for their offspring. Using data from 729 nests of 91 populations of 32 species of shorebirds in which parents synchronize their schedules to achieve continuous incubation of the eggs, they show that even under similar environmental conditions and despite day-long environmental cues, social synchronization can generate far more diverse behavioural rhythms than expected from studies of captive birds. The risk of predation, not starvation, might be a key determinant of biorhythmic diversity. The behavioural rhythms of organisms are thought to be under strong selection, influenced by the rhythmicity of the environment1,2,3,4. Such behavioural rhythms are well studied in isolated individuals under laboratory conditions1,5, but free-living individuals have to temporally synchronize their activities with those of others, including potential mates, competitors, prey and predators6,7,8,9,10. 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)6,7,8,9,11. The behavioural rhythms that emerge from such social synchronization and the underlying evolutionary and ecological drivers that shape them remain poorly understood5,6,7,9. Here we investigate these rhythms in the context of biparental care, a particularly sensitive phase of social synchronization12 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 captivity5,6,7,9. The risk of predation, not the risk of starvation, may be a key factor underlying the diversity in these rhythms.

The use of stable isotopes in animal ecology depends on accurate descriptions of isotope dynamics within individuals. The prevailing assumption that laboratory-derived isotopic parameters apply to free-living animals is largely untested. We used stable carbon isotopes (delta C-13) in whole blood from migratory Dunlin (Calidris alpina arcticola) to estimate an in situ turnover rate and individual diet-switch dates. Our in situ results indicated that turnover rates were higher in free-living birds, in comparison to the results of an experimental study on captive Dunlin and estimates derived from a theoretical allometric model. Diet-switch dates from all 3 methods were then used to estimate arrival dates to the Arctic; arrival dates calculated with the in situ turnover rate were later than those with the other turnover-rate estimates, substantially so in some cases. These later arrival dates matched dates when local snow conditions would have allowed Dunlin to settle, and agreed with anticipated arrival dates of Dunlin tracked with light-level geolocators. Our study presents a novel method for accurately estimating arrival dates for individuals of migratory species in which return dates are difficult to document. This may be particularly appropriate for species in which extrinsic tracking devices cannot easily be employed because of cost, body size, or behavioral constraints, and in habitats that do not allow individuals to be detected easily upon first arrival. Thus, this isotopic method offers an exciting alternative approach to better understand how species may be altering their arrival dates in response to changing climatic conditions.

Five subspecies of Dunlins (Calidris alpina) that breed in Beringia are potentially sympatric during the non-breeding season. Studying their ecology during this period requires techniques to distinguish individuals by subspecies. Our objectives were to determine (1) if five morphometric measures (body mass, culmen, head, tarsus, and wing chord) differed between sexes and among subspecies (C. a. actites, arcticola, kistchinski, pacifica, and sakhalina), and (2) if these differences were sufficient to allow for correct classification of individuals using equationsderivedfrom discriminantfunctionanalyses. We conductedanalyses usingmorphometric data from 10DunlinpopulationsbreedinginnorthernRussiaandAlaska,USA.Univariatetestsrevealedsignificantdifferences between sexes in most morphometric traits of all subspecies, and discriminant function equations predicted the sex of individuals with an accuracy of 83-100% for each subspecies. We provide equations to determine sex and subspecies of individuals in mixed subspecies groups, including the (1) Western Alaska group of arcticola and pacifica (known to stage together in western Alaska) and (2) East Asia group of arcticola, actites, kistchinski, and sakhalina (known to winter together in East Asia). Equations that predict the sex of individuals in mixed groups had classification accuracies between 75% and 87%, yielding reliable classification equations. We also provide equations that predict the subspecies of individuals with an accuracy of 22-96% for different mixed subspecies groups. When the sex of individuals can be predetermined, the accuracy of these equations is increased substantially. Investigators are cautioned to consider limitations due to age and feather wear when using these equations during the non- breeding season. These equations will allow determination of sexual and subspecies segregation in non-breeding areas, allowing implementation of taxonomic-specific conservation actions.

Shorebirds are exceptionally diverse in their social mating systems, and genetic analyses of a few species have revealed further variation in genetic mating systems. The ecological and evolutionary causes of this variation are largely unknown. We studied the mating system of American Golden-Plovers (Pluvialis dominica) at Barrow, Alaska, between 2004 and 2009. In this arctic population, birds bred in socially monogamous pairs, as has been seen in other populations, yet rates of philopatry and mate fidelity were low. We used microsatellite DNA markers to analyze parentage and found that most offspring originated from monogamous matings, but 8% of 131 offspring and 16% of 37 nests were the result of nonmonogamous matings. One nest was ambiguous; it was either a full clutch sired through extra-pair copulation or represented an instance of social polygyny. Despite wide variation in synchrony of breeding and density of nesting both within and among breeding seasons, these ecological variables were unrelated to mating patterns. We speculate that the low rate of pair fidelity and the high rate of extra-pair paternity are linked, because the lack of familiarity between partners reduces benefits of monogamy. Further study of breeding behavior and rates of philopatry are needed to understand why, among socially monogamous shorebirds, the American Golden-Plover has high rates of extra-pair paternity.

In urban environments, anthropogenic noise may mask bird song, especially the notes occurring at lower frequencies (1-2 kHz). Birds living in urban environments may modify their songs, particularly the low-frequency portions, to minimize masking by anthropogenic noise. Such modifications have been observed in Great Tits (Parus major) in The Netherlands, as well as in some mammals. We studied Song Sparrows (Melospiza melodia), which are common in both urban and rural environments in much of North America, and recorded the songs of 28 free-living males in Portland, Oregon. We also measured the amplitude and spectrum of ambient noise at singing locations. Song Sparrows singing at noisier locations exhibited higher-frequency low notes and had relatively less energy (amplitude) in the low-frequency range of their songs (1-4 kHz), where most anthropogenic noise also occurred. Although the mechanism(s) producing the correlation are as yet undetermined, the observed match between song and noise may result from behavioral plasticity. We discuss explanations for these patterns and how to test them.

We investigated whether female zebra finches (Taeniopygia guttata) would alter their mate preferences after observing the choices of other females. Experimental trials consisted of four 30-min stages: (A) acclimation, (B) observer female chooses between two males, (C) observer female watches a model female interact with her nonpreferred male from stage B, and (D) observer female again chooses between the two males. Control trials were identical except that there was no model female in stage C. Females in both experimental and control trials spent significantly more time with the nonpreferred male in stage D than they had in stage B; thus, our experiment appeared to reveal no evidence of mate choice copying. There was, however, a significant positive relationship between the increase in the time that an observer female in experimental trials spent with her nonpreferred male in stage D and the number of interactions that she had previously observed between the model female and that male in stage C. A second experiment of similar design, where observer females were prevented from observing the model female and nonpreferred male interact in stage B, demonstrated that the behaviour of the observer female in the first experiment was not simply a response to changes in male behaviour in stage D. While our experiment does not support the hypothesis that female zebra finches copy each other's mate preferences, our findings suggest that public information may influence the behaviour of female zebra finches toward potential mates.

How and why do the mating opportunities of males and females differ in natural population of animals? Previously we showed that females have higher mating opportunities than males in the Kentish plover Charadrius alexandrinus. Both parents incubate the eggs, and males provide more brood care than females; thus it is not obvious why the females find new mates sooner than the males. In this study we investigated whether the sex-biased mating opportunities stem from biased offspring sex ratios. We determined the sex of newly hatched, precocial chicks using CHD gene markers. Among fully sexed broods, 0.461 +/- 0.024 (SE) of chicks (454 chicks in 158 broods) were male, and this sex ratio was not significantly different from unity. The proportion of males at hatching decreased significantly over the breeding season, which occurred consistently in all 3 years of the study. Large chicks were more likely to be males than females. Neither parental age nor body size of male and female parents was related to brood sex ratio. We also sexed a number of chicks that were caught after they left their nest (range of estimated ages 0-17 days) and found that the proportion of males increased with brood age. This relationship remained highly significant when controlling statistically for hatching date. As brood size decreased due to mortality after the chicks left their nest, these results suggest that the mortality of daughters was higher than that of the sons shortly after hatching. Taken together, our results show that the female-biased mating opportunities in the Kentish plover are not due to biased brood sex ratio at hatching but, at least in part, are due to female-biased chick mortality soon after hatching.

An understanding of geographic and phylogenetic variation in passerine life histories is hampered by the scarcity of studies from the Southern Hemisphere. We documented the breeding biology of the White-browed Scrubwren (Sericornis frontalis), an Australia endemic in the Pardalotidae (parvorder Corvida). Like other members of the Pardalotidae, scrubwrens had a long laying interval (two days), a long incubation period (declining from 21 to 17 days through the season), and a long period of postfledging parental care (6 to 7 weeks). Scrubwrens appeared to be typical of the Australian Corvida in having a small clutch size (three eggs) and a long breeding season (5.4 months), and they also had a long interval between breeding attempts (10 days after a failed attempt, 21 days after a successful attempt). Scrubwrens were multibrooded, often raising two broods successfully and occasionally raising three broods. The breeding biology of scrubwrens adds further support to claims of a distinct life-history strategy for members of the Corvida but also reinforces evidence that some "Corvida" life-history traits more specifically are those of the Pardalotidae.

The proportion of extrapair paternity is known to vary greatly among species, but differences between populations of the same species have rarely been considered. We used microsatellite DNA markers to assess parentage of offspring in a subarctic population of the socially monogamous Yellow Warbler Dendroica petechia nesting near Churchill, Manitoba. We found a significantly lower proportion of extrapair young in the Churchill population than in a temperate population nesting approximately 1500 km to the south near the Queen's University Biological Station (QUBS) in Ontario. We show that the Churchill population also had significantly lower nesting density and significantly higher breeding synchrony; both are factors that have been hypothesized to affect extrapair paternity negatively. We suggest that inter-population comparisons can be used to test proximate mechanisms affecting extrapair paternity and in some cases may be better than interspecific or inter-individual comparisons. Towards this end we list eight other species that have had extrapair paternity measured in different populations. Studies that use genetic markers to assess the parentage of offspring have shown that extrapair paternity occurs in a wide range of bird species and that there is considerable interspecific variation in the proportion of offspring fathered by extrapair males (see Westneat and Sherman 1997). Understanding the variation in extrapair paternity has become a focus of mating system research. Nearly all species studied to date, however, have had extrapair paternity measured in only one population. Thus, it is not known how much of the apparent interspecific variation in extrapair paternity might be due to variation between populations within a species. For example, in the Willow Warbler Phylloscopus trochilus, the proportion of extrapair paternity differs considerably between a population in Sweden (0%; Gyllensten et al. 1990) and one in Norway (33%; Bjornstad and Lifjeld 1997). In this study we use microsatellite DNA markers to assess the parentage of offspring in a far northern population of the socially monogamous Yellow Warbler Dendroica petechia. Our first aim was to test whether the proportion of extrapair paternity in this population at the northern edge of the species' range differs from the 33% extrapair paternity in a population 1500 km to the south (Yezerinac et al. 1995). Nesting density and breeding synchrony are the most likely proximate causes of variation in the proportion of extrapair paternity (Westneat and Sherman 1997). However, evidence is mixed regarding the effect of either factor. Between-species comparisons provide no evidence that density affects extrapair paternity, whereas comparison of individuals nesting at different densities in the same population does suggest that density sometimes positively affects extrapair paternity (see Westneat and Sherman 1997). The apparent effect of synchrony on extrapair paternity is also debatable based upon results of interspecific and intraspecific studies (see Westneat and Sherman 1997, Stutchbury 1998, Weatherhead and Yezerinac 1998). In this study we measured nesting density and breeding synchrony within each of the two populations for which extrapair paternity was measured. Our second aim was to determine if there were synchrony and density differences between the populations that might explain any differences in extrapair paternity.

1, The most common form of cooperative breeding in birds involves a pair and their adult offspring ('helpers') provisioning young, Many studies show that pairs with helpers have higher reproductive success than pairs alone, but the differences could be due to confounding variables, like parental or territory quality, rather than to helping behaviour,2, One method of testing whether helping increases reproductive success is to compare the success of intact groups with those from which helpers have been removed. The disadvantage is that variables other than provisioning by, helpers (e,g. group size) are affected, which themselves could affect reproductive success. Alternatively, one can attempt to control statistically for confounding variables, but this is difficult in territorial species because it may be impossible to ensure that all confounding variables are measured.3. We took advantage of natural variability in provisioning behaviour among subordinate white-browed scrubwrens. Sericornis frontalis, to assess the effect of helping. Scrubwrens are a small passerine in the family Pardalotidae in which male offspring may remain on the natal territory; as adults, We compared the reproductive performance of pairs, groups with nonhelping subordinates and groups with helping subordinates, using data from 4-years' study of an individually colour-handed population resident in the Australian National Botanic Gardens, Canberra,4, The total rate of feeds to nestlings vc as greater when helpers were present, but we could detect no increase in nestling weight. Furthermore. we found no effect of helping behaviour on reproductive performance or reproductive success. Our measures included: (i) duration of the nesting cycle and interval between nesting attempts: (ii) success of individual nesting attempts, measured in over 20 ways: and (iii) reproductive success over the whole breeding season, measured as the total number of fledglings and independent young.5, There was also no effect of helping on survival of the female or dominant mole,6, We conclude that helping behaviour does not have a substantial, consistent effect on reproductive success. but cannot eliminate the possibility that it might increase reproductive success by a small amount or in some territories or years. Alternatively, helping behaviour might be the nonadaptive consequence of gene flow from populations in which it is adaptive, Nonetheless, we reject Jamieson's hypothesis that helping behaviour is a nonselected consequence of strong selection on parental behaviour, because, unlike many other species, helping is not an invariant consequence of natal philopatry.

We describe the isolation and genetic characterization of five microsatellite loci in a passerine bird, the yellow warbler Dendroica petechia, and assess their use for various types of population-level analysis using data from two breeding populations. All five loci show levels of variability comparable to those observed in other vertebrates (Hexp = 0.388 – 0.989). One locus, Dpµ05, is highly variable with 46 alleles detected in 41 individuals. All loci appeared to segregate in a Mendelian fashion as judged by patterns of inheritance in known families. However, one locus showed a significant heterozygote deficiency in one population suggesting the possible presence of null alleles at this locus. These markers provide a highly accurate system for determination of parentage in this species: the probability of detecting extrapair fertilization by males given known maternity was 0.999 in each of two separate populations. Comparison of allele frequencies and genetic distances between the two populations showed no evidence for significant differences in allele frequencies at individual loci, whereas the overall genetic distance and FST-value are significantly different from zero suggesting weak differentiation. Finally, cross-species amplification experiments showed that at least one locus appears to amplify products in a wide range of birds including nonpasserine species. Thus, our results demonstrate that these loci will provide a useful set of genetic information for addressing a wide range of population-level analyses in this and other bird species.

We used multi-locus DNA fingerprinting to characterise the genetic mating system of the socially monogamous yellow warbler (Dendroica petechia). Over 2 years there were no instances of brood parasitism, but 59% of families (n = 90) contained extrapair sired young and 37% of offspring (n = 355) were of extra-pair paternity. Most hypotheses for extra-pair mating in monogamous species assume a paternity benefit to extra-pair sires, and focus on the benefit(s) to females. However, the assumption of male benefit has been little tested. Among yellow warblers, known extra-pair sires were just as likely to be cuckolded as any male in the population, and there was at least one reciprocal exchange of extra-pair paternity. Nevertheless, among known extra-pair sires, the paternity gains from extra-pair paternity were, on average, greater than the losses in their own families. These results show there is a paternity benefit to certain males. However, the benefit is not absolute but relative and therefore more difficult to measure. The results also suggest that patterns of extra-pair fertilisation are not determined by female choice alone. Most confirmed extra-pair mates were territorial neighbours, but some resided as far as three territories apart, and greater spatial separation was implied in other cases. Thus, the opportunity for extra-pair mating is great. We estimate that as a result of extra-pair fertilisations, variance in male mating success is increased somewhere between 3-fold and 15-fold over that which would result from within-pair reproduction alone. These findings affirm the potential importance of extra-pair reproduction for sexual selection in monogamous species and they support earlier suggestions that extra-territorial forays by male yellow warblers are for the purpose of extra-pair mating.