Summary 1.We tested ecological hypotheses about timing of breeding and reproductive effort in birds, by analysing > 15-year data sets for individually marked females in three species of Latvian ducks (northern shoveler, tufted duck, common pochard). 2. Duckling survival and recruitment declined with advancing hatch date in pochard and tufted duck, after controlling for effects of female age and other factors with path analysis, a novel finding which indicates that fitness advantages associated with early hatching extended beyond the prefledging period. Logistic regression analysis suggested further that individual duckling prefledging survival was moderate in the earliest phase of the breeding season, greatest in mid-season and lowest later on. 3. However, selection acting against early hatched ducklings was surpassed by strong directional selection favouring recruitment of the earliest hatching females. The absolute and relative numbers of female recruits produced by a breeding female declined sharply with advancing hatch date in all species. 4. Unlike previous studies, an hypothesized intraspecific trade-off between duckling mass and brood size was detected, being very robust in two of three species. 5. Unexpectedly, female age effects on recruitment were manifested only indirectly by several pathways, the most important being the earlier hatching dates of older females. Size-adjusted body mass (i.e. condition index) was positively related to reproductive success, and was 2-8-fold more influential than female size (indexed by wing length). 6. Overall, fecundity-independent variables (e.g. hatching date, weather, indices of duckling production and habitat quality) generally had 2-10 times greater influence on recruitment rates than did fecundity-dependent variables such as female size or condition, duckling mass and brood size, suggesting a critical role for external environmental factors vs. individual female-specific traits in the recruitment process.Key-words : body condition and reproductive success, directional selection on timing of breeding, egg size-clutch size trade-off, pre-fledging survival, female recruitment.

Questions about individual variation in "quality" and fitness are of great interest to evolutionary and population ecologists. Such variation can be investigated using either a random effects approach or an approach that relies on identifying observable traits that are themselves correlated with fitness components. We used the latter approach with data from 1,925 individual females of three species of ducks (tufted duck, Aythya fuligula; common pochard, Aythya ferina; northern shoveler, Anas clypeata) sampled on their breeding grounds at Engure Marsh, Latvia, for over 15 years. Based on associations with reproductive output, we selected two traits, one morphological (relative body condition) and one behavioral (relative time of nesting), that can be used to characterize individual females over their lifetimes. We then asked whether these traits were related to annual survival probabilities of nesting females. We hypothesized quadratic, rather than monotonic, relationships based loosely on ideas about the likely action of stabilizing selection on these two traits. Parameters of these relationships were estimated directly using ultrastructural models embedded within capture-recapture-band-recovery models. Results provided evidence that both traits were related to survival in the hypothesized manner. For all three species, females that tended to nest earlier than the norm exhibited the highest survival rates, but very early nesters experienced reduced survival and late nesters showed even lower survival. For shovelers, females in average body condition showed the highest survival, with lower survival rates exhibited by both heavy and light birds. For common pochard and tufted duck, the highest survival rates were associated with birds of slightly above-average condition, with somewhat lower survival for very heavy birds and much lower survival for birds in relatively poor condition. Based on results from this study and previous work on reproduction, we conclude that nest initiation date and body condition covary with both reproductive and survival components of fitness. These associations lead to a positive covariance of these two fitness components within individuals and to the conclusion that these two traits are indeed correlates of individual quality.

Summary 1.We used long-term capture-recapture-recovery data and a modelling approach developed by Burnham (1993) to test a priori predictions about sources of variation in annual survival rates and fidelity within a population of individually marked females in three species of European ducks from a breeding ground study site in Latvia. 2. True annual survival was higher for diving ducks (tufted duck 0·72, common pochard 0·65) and lower for northern shoveler (0·52). Survival of female diving ducks was positively correlated with mean winter temperatures at Western European wintering areas, the relationship being much stronger for pochard. 3. We present the first unbiased estimates of breeding fidelity and permanent emigration in European ducks. Estimated fidelity rates were high (0·88 -1·0) and emigration rates low (0-0·12) for all three species, and we found strong evidence for age-specific differences in fidelity of pochards. Unusual long-distance (up to 2500 km) breeding dispersal movements that we found in female tufted ducks have not been documented in any other European waterfowl and are most probably a result of saturated nesting habitats. 4. Fidelity was a function of patch reproductive success in the previous year for all three species providing support for the idea that patch success is an important cue influencing fidelity. 5. Fidelity probability increased to 1·0 for shovelers during the last 12 years of study following provision of critical improvements in nesting habitats and suggested that habitat conditions and reproductive success determined site fidelity and settling patterns for shoveler and probably also influenced fidelity of the two other species. In predictable habitats, fidelity is a parameter that reflects the integration of fitness components and is thus a good quantity for assessing the effectiveness of habitat management actions.

No abstract

Senescent decline in survival probabilities of animals is a topic about which much has been written but little is known. Here, we present formal tests of senescence hypotheses, using 1373 recaptures from 8877 duckling (age 0) and 504 yearling Common Pochards (Aythya ferina) banded at a Latvian study site, 1975-1992. The tests are based on capture-recapture models that explicitly incorporate sampling probabilities that, themselves, may exhibit time-and age-specific variation. The tests provided no evidence of senescent decline in survival probabilities for this species. Power of the most useful test was low for gradual declines in annual survival probability with age, but good for steeper declines. We recommend use of this type of capture-recapture modeling and analysis for other investigations of senescence in animal survival rates.

Number of breeding attempts is a strong correlate of lifetime reproductive success (LRS) in birds, but the relative importance of potentially interacting factors affecting LRS has rarely been fully evaluated. We considered simultaneously five main factors hypothesized to influence LRS (age at first breeding, nesting date, number of breeding attempts, female traits, brood parasitism) by analyzing with path analysis 22-year data sets for 1,279 individually marked females and their offspring in tufted duck ( Aythya fuligula), common pochard ( A. ferina) and northern shoveler (Anas clypeata). We recaptured marked offspring as breeding adults (n=496 females) and obtained more complete estimates of LRS by incorporating information about banded ducklings of both sexes shot by hunters > or =12 months after banding (n=138). In tufted ducks and especially pochard (both diving duck species), late-hatched females tended to delay nesting until 2-years old. Most females (tufted duck, 74%; pochard, 71%; shoveler, 59%) apparently produced no breeding-age offspring. Number of breeding attempts (i.e., longevity) was the strongest correlate of LRS in all species, after controlling effects of age at first breeding, relative nest initiation date, wing length and body mass. Percentage of females producing recruits increased gradually with number of breeding attempts for all three species. Also, as expected, females nesting early in the breeding season had higher LRS than late-nesting individuals. In shoveler, female-specific characteristics of relatively longer wings and heavier late incubation body mass had positive effects on LRS, the latter feature being more common in 2-year-old nesters. In diving ducks, no relationships were detected between LRS and female-specific traits like wing length or body mass, and nor did acceptance of parasitic eggs have any deleterious impact on fitness estimates. Overall, number of fledged ducklings and LRS were related in tufted duck, weakly associated in pochard and unrelated in shoveler, implying that fledging success is not always a reliable measure of LRS.

Summary 1.We used up to 35 years of capture-recapture data from nearly 3300 individual female ducks nesting on Engure Marsh, Latvia, and multistate modelling to test predictions about the influence of environmental, habitat and management factors on breeding dispersal probability within the marsh. 2. Analyses based on observed dispersal distances of common pochards and tufted ducks provided no evidence that breeding success in year t influenced dispersal distance between t and t + 1. 3. Breeding dispersal distances (year t to t + 1) of pochards and tufted ducks were associated with a delay in relative nest initiation dates in year t + 1. The delay was greater for pochards ( c . 4 days) than for tufted ducks ( c . 2 days) when females dispersed > 0·8 km. 4. Northern shovelers and tufted ducks moved from a large island to small islands at low water levels and from small islands to the large island at high water levels before the construction of elevated small islands . Following this habitat management (1983-94), breeding fidelity was extremely high and not influenced by water level in the marsh for either species. 5. Because pochard nesting habitats in black-headed gull colonies were saturated during the entire study period, nesting females moved into and out of colonies with similar probabilities. Local survival probabilities and incubation body masses were higher for both yearlings ( SY ) and adults ( ASY ) nesting within gull colonies, suggesting that these females were of better quality than females nesting outside of the colonies. 6. Tufted ducks showed higher probabilities of moving from islands to emergent marshes when water levels were higher both before and after habitat management. However, rates of movement for a given water level were higher during the period before management than after. 7. Both pochards and tufted ducks exhibited asymmetric movement with respect to proximity to water, with higher movement probabilities to near-water nesting locations than away from these locations. 8. Multistate capture-recapture models provided analyses that were useful in investigating sources of variation in breeding dispersal probabilities.

No abstract