Barry D. Smith

Western sandpipers have altered migration tactics as peregrine falcon populations have recovered

The presence of top predators can affect prey behaviour, morphology and life history, and thereby can produce indirect population consequences greater and further reaching than direct depredation would have alone. Raptor species in the Americas are recovering since restrictions on the use of dichlorodiphenyltrichloroethane (DDT) and the implementation of conservation measures, in effect constituting a hemisphere–wide predator–reintroduction experiment, and profound effects on populations of their prey are to be expected. Here, we document changes in the behaviour of western sandpipers (Calidris mauri) at migratory stopover sites over two decades. Since 1985, migratory body mass and stopover durations of western sandpipers have fallen steadily at some stopovers in the Strait of Georgia, British Columbia. Comparisons between years, sites and seasons strongly implicate increasing danger from the recovery of peregrine falcons (Falco peregrinus) as a causal factor. A decade–long ongoing steep decline in sandpiper numbers censused on our study site is explained entirely by the shortening stopover duration, rather than fewer individuals using the site. Such behavioural changes are probably general among migratory shorebird species, and may be contributing to the widespread census declines reported in North America.

Social and maternal factors affecting duckling survival in eiders Somateria mollissima

1. With the aid of a novel survivorship model, an 8-year field study of social and maternal factors affecting duckling survival in eiders (Somateria mollissima) revealed that duckling survival probability varies in accordance with maternal brood-rearing strategy. This variability in survival provides compelling evidence of different annual fitness consequences between females that share brood-rearing and those that tend their broods alone. Consequently, as prebreeding survival is often a major source of individual variation in lifetime reproductive success, a females annual, state-dependent (e.g. condition) choice of a brood-rearing strategy can be a critical fitness decision. 2. Variance in duckling survival among lone tender broods was best explained by a model with significant interannual variability in survival, and survivorship tending to increase with increasing clutch size at hatch. Clutch size was correlated positively with female condition. Hatch date and female body condition together affected duckling survival, but their contributions are confounded. We were unable to identify a relationship between female age or experience and duckling survival. 3. Variance in duckling survival among multifemale brood-rearing coalitions was best explained by a model that included the number of tenders, the number of ducklings and interannual variation in how their ratio affected survivorship. Hatch date did not significantly influence survival. 4. Expected duckling survival is higher in early life for lone tenders when compared with multifemale brood-rearing coalitions. However, as ducklings approach 2-3 weeks of age, two or three females was the optimal number of tenders to maximize daily duckling survival. The survivorship advantage of multifemale brood-rearing coalitions was most evident in years of average survival. 5. The observed frequency distribution of female group sizes corresponds with the distribution of offspring survival probabilities for these groups. Evidence for optimal group sizes in nature is rare, but the most likely candidates may be groups of unrelated animals where entry is controlled by the group members, such as for female eiders. 6. Our study demonstrates that differences in social factors can lead to different predictions of lifetime reproductive success in species with shared parental care of self-feeding young.

LATITUDINAL CLINES IN FOOD DISTRIBUTION CORRELATE WITH DIFFERENTIAL MIGRATION IN THE WESTERN SANDPIPER

We report that a latitudinal cline in intertidal food distribution is associated with the nonbreeding distribution of the Western Sandpiper (Calidris mauri). This novel result is the first to demonstrate a clear relationship between patterns of differential nonbreeding distribution and food availability for any shorebird species. Within each age class and sex, longer-billed Western Sandpipers winter further south. Moreover, females, the longer-billed sex, tend to winter south of males. Thus, both inter- and intra-sexual clines in bill morphology result in an overall gradient of increasing bill length from north to south. Longer-billed birds are able to extract prey that are buried more deeply in the sediment; therefore, we predicted shifts in the vertical distribution of food resources to coincide with the clines in bill morphology across the nonbreeding range. We tested our prediction by measuring biofilm density and the vertical distribution of macrofaunal invertebrates at six nonbreeding sites. Although no latitudinal trend was observed for biofilm, the vertical distribution of invertebrates was consistent with our prediction and revealed that the greatest relative abundance of surface prey occurred at northern nonbreeding sites and declined with decreasing latitude. We discuss the potential implications of these findings in the context of competing evolutionary hypotheses of differential migration and bill dimorphism in shorebirds.

SYMPATRIC GROWTH MORPHS AND SIZE BIMODALITY IN THE GREEN SEA URCHIN (STRONGYLOCENTROTUS DROEBACHIENSIS)

Green sea urchins (Strongylocentrotus droebachiensis) have been a dominant structuring force in boreal, hard-bottom communities and recently have become an important fishery. Despite these roles, relatively little is known about the age, growth, and demographics of field populations. To begin to address these issues, we systematically sampled urchins during 1997 and 1998 at two sites (Allen Island [AI] and Schoodic Peninsula [SP]) in the Gulf of Maine, USA. Basic demographic information (density and size structure) was obtained and a subset of urchins aged by counting annual bands on the interambulacral plates. Size-at-age analyses showed that the population at AI consisted of two sympatric growth morphs, fast growing (fg) and slow growing (sg), each described by its own von Bertalanffy growth parameters for mean size and size variance. The two morphs also differed in maximum observed ages (sg, 6–11 yr, vs. fg, 16–18 yr) and in maximum test diameter sizes (sg, 30–35 mm, vs. fg, 50–70 mm). At SP, a single ...

SEX RATIOS OF DUNLIN WINTERING AT TWO LATITUDES ON THE PACIFIC COAST

Abstract Latitudinal clines in sex ratio during the nonbreeding season occur in some shorebirds of the Scolopacidae. We compared populations of nonbreeding Dunlin (Calidris alpina pacifica) from two latitudes along the Pacific flyway: the Fraser River Delta, British Columbia, and Bolinas Lagoon, California, to determine whether, and to what degree, they exhibited sex ratios consistent with a latitudinal cline. Dunlin are plumage monomorphic, so we used a maximum likelihood model to estimate overall and monthly sex ratios for each population based on culmen length distributions. Sex ratios in the Fraser River Delta were corrected for sex differences in habitat use. Monthly sex ratios were similar at the two sites but varied throughout the winter, likely reflecting differences in seasonal movement patterns between the sexes. Both populations showed an overall bias toward males (Fraser = 61% males, Bolinas = 65% males). Since there is no evidence to support the possibility of a skew toward males in C. a. pacifica as a whole, our data are consistent with some form of latitudinal cline in the sex ratio of C. a. pacifica. However, additional data from the Oregon coast, southern California, and Mexico are required to resolve this question. We also tested the hypothesis that mean body size within each sex is larger at the higher-latitude site (Fraser River Delta), but this did not appear to be the case.

MALLARD DUCKLING SURVIVAL IN THE GREAT LAKES REGION

Abstract Survival of young in waterfowl is poorly understood, particularly in regions outside of the traditional prairie breeding areas. Further, traditional methods of survival estimation lack the ability to statistically characterize between the extremes of random and catastrophic mortality events. We estimated Mallard (Anas platyrhynchos) duckling survival rates for 121 broods at nine study sites across the Great Lakes region from 2001–2003, using a novel statistical method that allows for the partitioning of random and correlated mortality processes. Results indicated that survival increased rapidly with age, did not change with hatching dates, did not differ among years, but varied across site-by-year replicates. Rates of random mortality were found to vary among site-years, while rates of correlated mortality varied little across site-years. In contrast to most studies of Mallard duckling survival, seasonal increases in duckling survival were not detected. We speculate that the observed patterns in survival rates with hatching date are related to productivity in Great Lakes brood-rearing wetlands and temperate regional climate.

Bimodality in Size Distributions: The Red Sea Urchin Strongylocentrotus Franciscanus as an Example

We use a model based on the size-structured von Foerster equation to de- scribe how size-dependent growth and mortality rates, pulsed recruitment, and variability in growth affect the shape of a size distribution. The deterministic, equilibrium size dis- tribution with constant recruitment increases with size when the difference between mor- tality rate and the rate at which growth rate decreases with size is positive (growth dom- inated), and decreases when it is negative (mortality dominated). Pulsed recruitment causes modes whose relative amplitudes are indicated by the corresponding constant recruitment case. For typical animal growth patterns, the distance between pulses decreases with age. Pulses merge and can be selectively obscured by variability in growth so that their relative amplitudes no longer correspond to the constant recruitment case. We use this information to evaluate why bimodality occurs in size distributions of the red sea urchin, Strongylo- centrotus franciscanus, in some habitats, but not others. The mode at larger sizes, which occurs in all habitats, arises because the distribution is mortality dominated and the final sizes of individuals vary. The upper half of a second mode at smaller sizes is caused by higher mortality rates at sizes greater than the peak of that mode. The lower half may be due either to a refuge from predation under the spine canopy of adults or to sampling selectivity.

A CLUTCH AND BROOD SURVIVAL MODEL THAT DISCRIMINATES RANDOM AND CORRELATED MORTALITY

Quantitative conservation methodologies such as Population Viability Anal- ysis (PVA) require reliable estimates of life history parameters such as breeding success. The utility of such metrics for egg-laying species is complicated by the fact that the mortality of eggs and juveniles can occur both randomly and independently over time, or catastroph- ically, as in the sudden loss of a clutch or brood. Not knowing the nature of mortality caused by either or both of abiotic (e.g., weather) and biotic (e.g., predation) events limits our ability to confidently assess a populations demography and sustainability, or rank competing hypotheses. To address this deficiency, we describe a statistical model that estimates egg and juvenile survival rates continuously from laying to fledging based on periodic observations of individual clutches and broods. Adjunct data on environmental or predation threats can be included in the model as covariate series potentially affecting juvenile survival. Our model can statistically characterize mortality between the extremes of random and catastrophic mortality and can determine if unwitnessed mortalities occurred independently or were correlated (i.e., overdispersed, where catastrophe is extreme over- dispersion). Overdispersion is estimated as a parameter of the beta-binomial probability distribution of survival outcomes, which differs from its treatment in Program MARK where overdispersion is an a posteriori diagnostic referred to as c. We used data for the sea duck Barrows Goldeneye to illustrate our model. Specifically, we contribute to the argument that a larger brood confers a fitness advantage to a tending hen by concluding that brood size on hatch day is positively correlated with a juveniles probability of surviving to fledge.

AGE AND SEX DISTRIBUTIONS OF WINTERING SURF SCOTERS: IMPLICATIONS FOR THE USE OF AGE RATIOS AS AN INDEX OF RECRUITMENT

Abstract We assessed age- and sex-specific distribution patterns of Surf Scoters (Melanitta perspicillata) wintering in southern coastal British Columbia, Canada, and evaluated potential biases associated with the use of male age ratios as an index of their recruitment. For surveys conducted during 2000 through 2002, annual variations in male age ratios were evident, with estimates ranging from 0.07 ± 0.02 to 0.13 ± 0.03 (SE; first-year males:total males). Flock composition patterns indicated first-year males did not distribute independently, but tended to associate with other first-year males. With respect to habitat, male age- class proportions did not vary among shoreline substrate types, but higher proportions of first-year males were found in sites with low exposure to wind and waves (<50 km fetch). To determine the efficacy of male age ratios for indexing recruitment, we used a power analysis, which incorporated overdispersion in age-class segregation and determined the sample sizes required for precise estimates of the proportion of first-year male Surf Scoters. Samples of approximately 600–1000 total males were required to obtain 95% confidence limits within 5% of the estimated mean, with sampling accuracy leveling off at about 2% when 6000 or more males were aged. Recruitment among waterfowl species is typically modeled using the ratio of female recruits to breeding-age females. Based on the sex and male age-ratio estimates obtained in this study, we calculated a female age ratio of 0.23 (first-year females:adult females). Distribuciones Invernales de Edad y Sexo en Melanitta perspicillata: Implicancias del Uso de Cocientes de Edad como un Índice de Reclutamiento Resumen. Estimamos los patrones de distribución específicos de edad y sexo de Melanitta perspicillata invernando en la zona costera sur de la Columbia Británica, Canadá, y evaluamos los sesgos potenciales asociados con el uso de cocientes de edad de machos como un índice de su reclutamiento. Durante los muestreos realizados entre el 2000 y el 2002 las variaciones anuales en el cociente de edad de los machos fueron evidentes, con estimaciones que variaron entre 0.07 ± 0.02 (EE) y 0.13 ± 0.03 (machos del primer año: total de machos). Los patrones de composición de las bandadas indicaron que los machos del primer año no se distribuyeron independientemente, sino que tendieron a asociarse con otros machos del primer año. Con relación al hábitat, las proporciones de clases de edad de los machos no variaron entre los tipos de substrato de la línea de costa, pero se encontraron mayores proporciones de machos del primer año en sitios con baja exposición al viento y a las olas (sitios de mar abierto <50 km de ancho). Para determinar la eficacia de los cocientes de edad de los machos como índice de reclutamiento, usamos un análisis de poder, el cual incorporó sobre-dispersión en la segregación de las clases de edad y determinó los tamaños de muestreo necesarios para estimaciones precisas de la proporción de machos del primer año de M. perspicillata. Muestras totales de aproximadamente 600–1000 machos fueron necesarias para obtener límites de confianza del 95% dentro del 5% de la media estimada, con la exactitud de muestreo nivelándose cerca del 2% luego de estimarse la edad de 6000 o más machos. El reclutamiento entre las especies de Anseriformes es modelado típicamente usando el cociente entre nuevas hembras y hembras en edad reproductiva. Basados en las estimaciones de cocientes de sexo y edad de los machos obtenidas en este estudio, calculamos un cociente de edad de las hembras de 0.23 (hembras del primer año: hembras adultas).

Effects of Habitat on Mallard Duckling Survival in the Great Lakes Region

Abstract Habitat provides food and shelter resources for prefledgling waterfowl and thus plays a critical role in their growth, development, and survival. However, few studies have examined whether and how particular elements of habitat affect duckling survival. We investigated relationships of duckling survival rates with distance of overland travel, wetland vegetation composition, water permanency, and surrounding upland vegetation for 116 mallard (Anas platyrhynchos) broods in the Great Lakes region from 2001 to 2003. We found that the probability, on hatch day, that a mallard duckling will survive to 55 days was positively related to the proportion of wetland area that was vegetated and negatively related to the proportion of forest cover within 500 m of duckling locations. We found little support for relationships between duckling survival rates and the proportions of grasslands or seasonal wetlands or to distances traveled overland by broods. Our results suggest that conservation groups and wildlife managers in the Great Lakes region can improve mallard duckling survival rates by managing for, creating, and protecting vegetated wetlands and focusing efforts within lightly-forested areas.