Richard B. Lanctot

Genetic similarity between mates and extra-pair parentage in three species of shorebirds

Matings between close relatives often reduce the fitness of offspring, probably because homozygosity leads to the expression of recessive deleterious alleles. Studies of several animals have shown that reproductive success is lower when genetic similarity between parents is high, and that survival and other measures of fitness increase with individual levels of genetic diversity. These studies indicate that natural selection may favour the avoidance of matings with genetically similar individuals. But constraints on social mate choice, such as a lack of alternatives, can lead to pairing with genetically similar mates. In such cases, it has been suggested that females may seek extra-pair copulations with less related males, but the evidence is weak or lacking. Here we report a strong positive relationship between the genetic similarity of social pair members and the occurrence of extra-pair paternity and maternity (‘quasi-parasitism’) in three species of shorebirds. We propose that extra-pair parentage may represent adaptive behavioural strategies to avoid the negative effects of pairing with a genetically similar mate.

Prevalence of Influenza A viruses in wild migratory birds in Alaska: Patterns of variation in detection at a crossroads of intercontinental flyways

BackgroundThe global spread of the highly pathogenic avian influenza H5N1 virus has stimulated interest in a better understanding of the mechanisms of H5N1 dispersal, including the potential role of migratory birds as carriers. Although wild birds have been found dead during H5N1 outbreaks, evidence suggests that others have survived natural infections, and recent studies have shown several species of ducks capable of surviving experimental inoculations of H5N1 and shedding virus. To investigate the possibility of migratory birds as a means of H5N1 dispersal into North America, we monitored for the virus in a surveillance program based on the risk that wild birds may carry the virus from Asia.ResultsOf 16,797 birds sampled in Alaska between May 2006 and March 2007, low pathogenic avian influenza viruses were detected in 1.7% by rRT-PCR but no highly pathogenic viruses were found. Our data suggest that prevalence varied among sampling locations, species (highest in waterfowl, lowest in passerines), ages (juveniles higher than adults), sexes (males higher than females), date (highest in autumn), and analytical technique (rRT-PCR prevalence = 1.7%; virus isolation prevalence = 1.5%).ConclusionThe prevalence of low pathogenic avian influenza viruses isolated from wild birds depends on biological, temporal, and geographical factors, as well as testing methods. Future studies should control for, or sample across, these sources of variation to allow direct comparison of prevalence rates.

ASSESSING THE DEVELOPMENT OF SHOREBIRD EGGS USING THE FLOTATION METHOD: SPECIES-SPECIFIC AND GENERALIZED REGRESSION MODELS

Abstract ABSTRACT We modeled the relationship between egg flotation and age of a developing embryo for 24 species of shorebirds. For 21 species, we used regression analyses to estimate hatching date by modeling egg angle and float height, measured as continuous variables, against embryo age. For eggs early in incubation, we used linear regression analyses to predict hatching date from logit-transformed egg angles only. For late incubation, we used multiple regression analyses to predict hatching date from both egg angles and float heights. In 30 of 36 cases, these equations estimated hatching date to within four days of the true hatching date for each species. After controlling for incubation duration and egg size, flotation patterns did not differ between shorebirds grouped by mass (≥100 g or <100 g) or taxonomy (Scolopacidae versus Charadriidae). Flotation progressed more rapidly in species in which both adults incubate the clutch versus species in which only one adult incubates the clutch, although this did not affect prediction accuracy. We also pooled all continuous data and created a generalized regression equation that can be applied to all shorebird species. For the remaining three species, we estimated hatching date using five float categories. Estimates of hatching date using categorical data were, overall, less accurate than those generated using continuous data (by 3%–5% of a given incubation period). Our equations were less accurate than results reported in similar studies; data collected by multiple observers and at multiple sites, as well as low sample sizes for some species, likely increased measurement error. To minimize flotation method prediction error, we recommend sampling in early incubation, collecting both egg angle and float height data in late incubation, and developing site- and species-specific regression models where possible.

Are corticosterone levels a good indicator of food availability and reproductive performance in a kittiwake colony

We evaluated the use of corticosterone to gauge forage availability and predict reproductive performance in black-legged kittiwakes (Rissa tridactyla) breeding in Alaska during 1999 and 2000. We modeled the relationship between baseline levels of corticosterone and a suite of individual and temporal characteristics of the sampled birds. We also provided supplemental food to a sample of pairs and compared their corticosterone levels with that of pairs that were not fed. Corticosterone levels were a good predictor of forage availability in some situations, although inconsistencies between corticosterone levels and reproductive performance of fed and unfed kittiwakes suggested that this was not always the case. In general, higher corticosterone levels were found in birds that lacked breeding experience and in birds sampled shortly after arriving from their wintering grounds. All parameters investigated, however, explained only a small proportion of the variance in corticosterone levels. We also investigated whether corticosterone, supplemental feeding, year of the study, breeding experience, body weight, and sex of a bird were able to predict laying, hatching, and fledging success in kittiwakes. Here, breeding experience, year of the study, and body weight were the best predictors of a birds performance. Corticosterone level and supplemental feeding were good predictors of kittiwake reproductive performance in some cases. For example, corticosterone levels of birds sampled during the arrival stage reliably predicted laying success, but were less reliable at predicting hatching and fledging success. Counts of active nests with eggs or chicks may be more reliable estimates of the actual productivity of the colony. Supplemental feeding had strong effects on kittiwake productivity when natural forage was poor, but had little effect when natural forage was plentiful.

Lekking Without a Paradox in the Buff‐Breasted Sandpiper

Females in lek‐breeding species appear to copulate with a small subset of the available males. Such strong directional selection is predicted to decrease additive genetic variance in the preferred male traits, yet females continue to mate selectively, thus generating the lek paradox. In a study of buff‐breasted sandpipers (Tryngites subruficollis), we combine detailed behavioral observations with paternity analyses using single‐locus minisatellite DNA probes to provide the first evidence from a lek‐breeding species that the variance in male reproductive success is much lower than expected. In 17 and 30 broods sampled in two consecutive years, a minimum of 20 and 39 males, respectively, sired offspring. This low variance in male reproductive success resulted from effective use of alternative reproductive tactics by males, females mating with solitary males off leks, and multiple mating by females. Thus, the results of this study suggests that sexual selection through female choice is weak in buff‐breasted sandpipers. The behavior of other lek‐breeding birds is sufficiently similar to that of buff‐breasted sandpipers that paternity studies of those species should be conducted to determine whether leks generally are less paradoxical than they appear.

Sexing adult black-legged kittiwakes by DNA, behavior, and morphology

-We sexed adult Black-legged Kittiwakes (Rissa tridactyla) using DNA-based genetic techniques, behavior and morphology and compared results from these techniques. Genetic and morphology data were collected on 605 breeding kittiwakes and sex-specific behaviors were recorded for a sub-sample of 285 of these individuals. We compared sex classification based on both genetic and behavioral techniques for this sub-sample to assess the accuracy of the genetic technique. DNA-based techniques correctly sexed 97.2% and sex-specific behaviors, 96.5% of this sub-sample. We used the corrected genetic classifications from this sub-sample and the genetic classifications for the remaining birds, under the assumption they were correct, to develop predictive morphometric discriminant function models for all 605 birds. These models accurately predicted the sex of 73-96% of individuals examined, depending on the sample of birds used and the characters included. The most accurate single measurement for determining sex was length of head plus bill, which correctly classified 88% of individuals tested. When both members of a pair were measured, classification levels improved and approached the accuracy of both behavioral observations and genetic analyses. Morphometric techniques were only slightly less accurate than genetic techniques but were easier to implement in the field and less costly. Behavioral observations, while highly accurate, required that birds be easily observable during the breeding season and that birds be identifiable. As such, sex-specific behaviors may best be applied as a confirmation of sex for previously marked birds. All three techniques thus have the potential to be highly accurate, and the selection of one or more will depend on the circumstances of any particular field study. Received 2 February 2000, accepted 1 April 2000.

CONSERVATION STATUS OF THE BUFF-BREASTED SANDPIPER: HISTORIC AND CONTEMPORARY DISTRIBUTION AND ABUNDANCE IN SOUTH AMERICA

Abstract We present historic and contemporary information on the distribution and abundance of Buff-breasted Sandpipers (Tryngites subruficollis) in South America. Historic information was collated from the literature, area ornithologists, and museums, whereas contemporary data were derived from surveys conducted throughout the main wintering range in Argentina, Uruguay, and Brazil during the austral summers of 1999 and 2001. Variable circular plot sampling was used to estimate population densities. During 1999, the highest concentration of Buff-breasted Sandpipers in Argentina was in southern Bahía Samborombón (General Lavalle District) and areas north of Mar Chiquita coastal lagoon. During 2001, the highest concentrations in Brazil were at Ilha da Torotama and Lagoa do Peixe National Park. During 1999 and 2001, the highest concentrations of Buff-breasted Sandpipers in Uruguay were found along three lagoons (Laguna de Rocha, Laguna de Castillos, and Laguna Garzón) bordering the Atlantic Ocean. Population densities (birds/ha) of Buff-breasted Sandpipers were 0.11 (95% C.I. = 0.04–0.31) in Argentina, 1.62 (0.67–3.93) in Brazil, and 1.08 (0.37–3.18) in Uruguay. High turnover rates at survey sites, due to the formation of large, mobile flocks, contributed to moderately large confidence intervals around our population density estimates. Nevertheless, compared with historic accounts of Buff-breasted Sandpipers, our survey data indicate the population size of this species has declined substantially since the late 1800s and contemporary information suggests the species has continued to decline during the past three decades. Buff-breasted Sandpipers were found almost exclusively in pasturelands and appear to depend heavily upon intensive grazing by livestock, which maintain suitable short grass conditions. We discuss the need for protection of critical areas and proper range management to ensure appropriate habitat remains available for the species, and provide suggestions for future research needs.

HARLEQUIN DUCK RECOVERY FROM THE EXXON VALDEZ OIL SPILL: A POPULATION GENETICS PERSPECTIVE

Concerns about Harlequin Duck (Histrionicus histrionicus) population recov- ery following the Exxon Valdez oil spill led biologists to ask whether birds located in different molting and wintering areas belong to genetically distinct and, thus, demographically in- dependent populations. Owing to the lack of direct observations of movements among ma- rine areas, three classes of genetic markers that differed in mode of inheritance were used to evaluate the degree of genetic differentiation among wintering areas within Prince Wil- liam Sound (PWS) and the Alaska Peninsula and Kodiak Archipelago (APKA). We could not reject the null hypothesis that the wintering aggregations within each region are composed of a single genetically panmictic population. Differences in genotype frequencies among wintering locations within PWS and APKA were low and nonsignificant for all three classes of markers. Furthermore, we saw no evidence for deviations in Hardy-Weinberg equilibrium or gametic disequilibrium between loci within a winter collection site as would be expected if these locales were composed of individuals from reproductively isolated (and genetically distinct) breeding locales. Finally, no evidence for significant structuring was noted between PWS and APKA. Lack of spatial genetic structuring could be due to the cumulative effects of low levels of gene flow over long time periods, low levels of gene flow by immature birds moving between marine habitats, or to episodic dispersal caused by habitat alteration (e.g.

GENETIC PARENTAGE AND MATE GUARDING IN THE ARCTIC-BREEDING WESTERN SANDPIPER

Abstract Extrapair copulations and fertilizations are common among birds, especially in passerines. So far, however, few studies have examined genetic mating systems in socially monogamous shorebirds. Here, we examine parentage in the Western Sandpiper (Calidris mauri). Given that Western Sandpipers nest at high densities on the Arctic tundra, have separate nesting and feeding areas, and show high divorce rates between years, we expected extrapair paternity to be more common in this species compared to other monogamous shorebirds. However, DNA fingerprinting of 98 chicks from 40 families revealed that only 8% of broods contained young sired by extrapair males, and that 5% of all chicks were extrapair. All chicks were the genetic offspring of their social mothers. We found that males followed females more often than the reverse. Also, cuckolded males were separated from their mates for longer than those that did not lose paternity. Although these results suggest a role for male mate guarding, we propose that high potential costs in terms of reduced paternal care likely constrain female Western Sandpipers from seeking extrapair copulations.

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.