Observations on the Breeding Kittiwake

The drought conditions prevailing in the Marismas of the Guadalquivir (SW Spain) during the winter 1982–83 caused a large proportion of the wintering Greylag Goose population to use the same flooded areas throughout the winter at very high densities. Results indicate that Greylags prefer small Scirpus rhizomes to large ones, the geese taking an increasing amount of larger rhizomes as smaller rhizomes are depleted. A combination of factors (abundance, extraction and handling times, nutritive quality) could make smaller rhizomes more profitable food for Greylags than larger rhizomes. The large concentrations of birds in some areas probably determined the depletion of preferred resources and this could have led some geese to steal food from conspecifics; some aspects of this behaviour are discussed.     

Trends in annual and seasonal survival of Pink-footed Geese Anser brachyrhynchus

Adult Pink-footed Geese Anser brachyrhynchus from the Svalbard breeding population were neck-banded and resighted in staging and wintering areas outside the breeding season 1990–1999. We estimated annual and seasonal survival using capture–recapture statistical models. Mean annual survival was 0.829, declining over the study period from 0.90 to 0.79. The annual cycle was partitioned into three periods: summer (1 April–30 September), including both spring and autumn migration, autumn (1 October–31 December), including most shooting, and winter (1 January–31 March). The parsimonious model selected to describe seasonal survival included a declining trend in summer survival, constant autumn and winter survival with lower survival in the severe winter of 1996, and an additive effect of sex on summer and winter survival. Monthly survival was highest during winter. Decreasing summer survival was the main contributor to the overall decline in annual survival, and was attributed mainly to increasing natural mortality on the breeding grounds. Mean annual survival based on the seasonal survival probabilities was 0.835 for females and 0.805 for males. The effect of sex was most pronounced during summer and remains to be explained.     

A demographic comparison of two Nordic populations of Greylag Geese Anser anser

Greylag Geese Anser anser have been neck-banded on an annual basis in Scania, southern Sweden since 1984 and in Norway since 1986 as part of a Nordic Greylag project. This has yielded a large database of resightings, which we used here to estimate and compare survival rates between the two populations by means of mark–recapture models. Estimated adult survival was sex-dependent in the Scanian population, probably a result of differential neck band retention rates in this population. Mean juvenile survival was about 12% higher in the Scanian population (0.603 vs. 0.485). Adult survival in the Norwegian population was 0.728 (males 0.733; females 0.725), and in males and females from the Scanian population was 0.711 (0.752 after accounting for higher neck band loss in males) and 0.771, respectively. Over the course of the study, juvenile survival in the Scanian population increased substantially, and adult survival was constant, whereas both these parameters declined in the Norwegian population. This study demonstrates that the two Nordic populations are demographically distinct and gives further support to the notion that they should be treated as separate management units. The decline of 10% in adult survival in the Norwegian population, the cause of which still remains uncertain, is likely to have had a major impact on the growth of this population.     

Demography and population dynamics of the mouse opossum (Thylamys elegans) in semi-arid Chile: seasonality, feedback structure and climate

Here we present, to the authors' knowledge for the very first time for a small marsupial, a thorough analysis of the demography and population dynamics of the mouse opossum (Thylamys elegans) in western South America. We test the relative importance of feedback structure and climatic factors (rainfall and the Southern Oscillation Index) in explaining the temporal variation in the demography of the mouse opossum. The demographic information was incorporated into a stage-structured population dynamics model and the model's predictions were compared with observed patterns. The mouse opossum's capture rates showed seasonal (within-year) and between-year variability, with individuals having higher capture rates during late summer and autumn and lower capture rates during winter and spring. There was also a strong between-year effect on capture probabilities. The reproductive (the fraction of reproductively active individuals) and recruitment rates showed a clear seasonal and a between-year pattern of variation with the peak of reproductive activity occuring during winter and early spring. In addition, the fraction of reproductive individuals was positively related to annual rainfall, while population density and annual rainfall positively influenced the recruitment rate. The survival rates were negatively related to annual rainfall. The average finite population growth rate during the study period was estimated to be 1.011 +/- 0.0019 from capture-recapture estimates. While the annual growth rate estimated from the seasonal linear matrix models was 1.026, the subadult and adult survival and maturation rates represent between 54% (winter) and 81% (summer) of the impact on the annual growth rate.     

Population variables and causes of mortality in a hedgehog (Erinaceous europaeus) population in southern Sweden

A hedgehog population in southern Sweden was studied by capture-mark-recapture during 1972–1979. Each animal was weighed, sexed and marked with one numbered clip in each ear.
The number of subadult and adult hedgehogs was about 20 in the first two years, 1972–1973. Numbers then increased and approximately doubled and the population peaked in 1978. A decrease came in 1979.
The estimated number of juveniles produced varied between years with the lowest figure (15 juv.) in 1974 when the population was low and the highest in 1977 (39 juv.) when the population was high. As no captured subadult female had suckled young, estimates of recruitment rate assumed that females did not reproduce until they were adult. The average number of juveniles per adult female was 2.8. Recruitment rate did not show any correlation with non-juvenile population densities.
For both juveniles and non-juveniles the annual mortality varied greatly between years. Average yearly mortality was 47% for non-juveniles and 34% for juveniles.
For non-juveniles, most of the total mortality occurred during winter, ranging between 26% and 43% (average 33%), and increased with increasing densities of non juveniles. Juvenile winter mortality varied between 6% and 94%, averaging 33%.
Summer mortality of non-juveniles averaged 15%, ranging between 3% and 22%. Traffic kills were the predominant cause of death. Only a few juveniles were found dead during summer and autumn.
There were no statistical differences in age-specific survival rate between males and females and, generally, no differences in survival rate between the age categories within each sex. It is argued that the population size appears to be more influenced by environmental factors, such as food availability, winter nest sites and winter climate, than by density-dependent factors.     

First-year survival of brown trout in three Norwegian streams

Monthly survival rates during the first year of life were estimated for the 1999 cohort of stream-resident brown trout Salmo trutta in three Norwegian streams, using capture-mark-recapture methods and the Cormack–Jolly–Seber model. It was hypothesized that reduced survival would occur during the winter. For one of the study populations, the data did support seasonal variation in survival, with monthly survival rates being lower during winter than during summer (0·65 v . 0·99). For the remaining two populations, there was no evidence for seasonal variations in monthly survival rates, but monthly survival rates were significantly different (0·87 v . 0·99). No evidence was found for size-dependent winter survival. Some marked individuals emigrated from the study sites, suggesting that survival rates were underestimated and that different survival rates among populations were partly due to different emigration rates. Net immigration of brown trout was evident at all three study sites.     

High within‐winter and annual survival rates in a declining Afro‐Palaearctic migratory bird suggest that wintering conditions do not limit populations

For migratory birds, it is necessary to estimate annual and overwinter survival rates, identify factors that influence survival, and assess whether survival varies with age and sex if we are to understand population dynamics and thus inform conservation. This study is one of the first to document overwinter and annual survival from the wintering grounds of a declining Afro‐Palaearctic migrant bird, the Whinchat Saxicola rubetra. We monitored a population of marked individuals for which dispersal was low and detectability was high, allowing accurate estimates of survival. Annual survival was at least 52% and did not differ significantly across demographic groups or with habitat characteristics or residency time in the previous winter. Overwinter survival was very high and monthly survival at least 98% at some sites. Although winter residency varied spatially and with age, lower residency did not correlate with reduced annual survival, suggesting occupancy of multiple wintering sites rather than higher winter mortality of individuals with shorter residency. Our results suggest that mortality occurs primarily outside the wintering period, probably during migration, and that wintering conditions have minimal influence on survival. The similarity between survival rates for all age and sex classes when measured on the wintering grounds implies that any difference in survival with age or sex occurs only during the first migration or during the post‐fledging stage, and that selection of wintering habitat, or territory quality, makes little difference to survival in Whinchats. Our findings suggest that the wintering grounds do not limit populations as much as the migratory and breeding stages, with implications for the conservation of declining Afro‐Palaearctic migrants more widely.     

Relationships between otolith microstructure, otolith growth, somatic growth and ontogenetic transitions in two cohorts of windowpane

Otolith increments in larval and juvenile windowpane Scophthalmus aquosus can provide an estimate of daily age for spring-spawned individuals held under summer conditions. Otolith increments for spring- and autumn-spawned individuals occurred at intervals >1 day under winter conditions. A significant decrease in the slope of the linear relationship between otolith size and somatic size at 40 mm L_S coincided with significant habitat, morphological and behavioural transitions. In smaller, field-captured windowpane belonging to spring- and autumn-spawned cohorts, the formation of accessory growth centres coincided with a transitional settlement period and the completion of eye migration (c. 8–20 mm L_S). Back-calculated growth rate estimates for spring-spawned windowpane were significantly faster than those for autumn-spawned windowpane and these differences could produce differential rates of survival for the two cohorts during the first year of life.     

Survival of British Sedge Warblers Acrocephalus schoenobaenus in relation to west African rainfall

Fluctuations in the population levels and annual adult survival rates of British Sedge Warblers Acrocephalus schoenobaenus since the late 1960s are strongly correlated with indices of wet season rainfall in the west African winter quarters. Population changes are unrelated to estimates of breeding productivity in the previous year. Habitat availability in the winter quarters has probably been the main factor limiting the size of the Sedge Warbler population in Britain during the period of study.     

Seasonal mortality and sequential density dependence in a migratory bird

Migratory bird populations may be limited during one or more seasons, and thus at one or more places, but there is a dearth of empirical examples of this possibility. We analyse seasonal survival in a migratory shellfish‐eating shorebird (red knot Calidris canutus islandica) during a series of years of intense food limitation on the nonbreeding grounds (due to overfishing of shellfish stocks), followed by a relaxation period when destructive harvesting had stopped and food stocks for red knots recovered. For the estimation of seasonal survival from the 15 yr‐long near‐continuous capture–resight dataset, we introduce a ‘rolling window’ approach for data exploration, followed by selection of the best season definition. The average annual apparent survival over all the years was 0.81 yr^?1. During the limitation period, survival probability of adult red knots was low in winter (0.78 yr^?1), but this was compensated by high survival in summer (0.91 yr^?1). During the relaxation period survival rate levelled out with a winter value of 0.81 yr^?1 and a summer survival of 0.82 yr^?1. The fact that during the cockle‐dredging period the dip in survival in winter was completely compensated by higher survival later in the annual cycle suggests sequential density dependence. We conclude that seasonal compensation in local survival (in concert with movements to areas apparently below carrying capacity) allowed the islandica population as a whole to cope, in 1998–2003, with the loss of half of the suitable feeding habitat in part of the nonbreeding range, the western Dutch Wadden Sea. As a more general point, we see no reason why inter‐seasonal density dependence should not be ubiquitous in wildlife populations, though its limits and magnitude will depend on the specific ecological contexts. We elaborate the possibility that with time, and in stable environments, seasonal mortality evolves so that differences in mortality rates between seasons would become erased.     

Are badgers ‘Under The Weather’? Direct and indirect impacts of climate variation on European badger (Meles meles) population dynamics

Weather conditions, and how they in turn define and characterize regional climatic conditions, are a primary limit on global species diversity and distribution, and increasing variability in global and regional climates have significant implications for species and habitat conservation. A Capture–Mark–Recapture study revealed that badger (Meles meles) life history parameters interact in complicated ways with annual variability in the seasonality of temperature and rainfall, both in absolute and in phenological terms. A strong predictive relationship was observed between survival and both temperature and late‐summer rainfall. This link at the population dynamics level was related to individual body‐weight increases observed between summer and autumn. In addition, fecundity was correlated with spring rainfall and temperature. We investigated and confirmed that relationships were consistent with observed variation in the intensity of a parasitic infection. Finally, fecundity during any given year correlated with conditions in the preceding autumn. Badger survival also correlated with late winter weather conditions. This period is critical for badgers insofar as it coincides with their peak involvement in road traffic accidents (RTAs). RTA rate during this period was linked strongly to temperature, underlining the intricate ways in which a changing climate might interact with anthropogenic agents to influence species' population processes. Equinoctial conditions produced significant population driver effects. That is, while summers will always be relatively warm compared with winters, spring and autumn weather can be more variable and functionally delimit the ‘productive’ vs. nonproductive period of the year in terms of badger behavioural and physiological cycles. This study highlights how appropriately informed conservation strategies, mindful of trends in climatic conditions, will become ever‐more essential to ensure the survival of many species globally.     

Assessing the role of body mass and sex on apparent adult survival in polygynous passerines: a case study of cetti's warblers in central Portugal

Adult survival, an important fitness component, is usually 1) lower in lighter individuals due to their reduced ability to survive winter conditions compared to heavier ones, especially in resident species at northern temperate latitudes and 2) lower in females compared with males due to higher reproductive costs incurred by females. In this paper, a capture–mark–recapture dataset of 649 cetti's warblers Cettia cetti ringed seasonally at two wetlands in central Portugal over an 11‐yr period (2000–2010) was modelled in a multi‐state framework to examine the influence of these individual covariates on apparent adult survival, while controlling for the presence of transient individuals in our study area. The probability of change in mass state (ψ_{Light→Heavy}, ψ_{Heavy→Light}) during the annual cycle was also estimated. Overall, birds survived better during spring–summer (breeding/moulting periods) compared with autumn–winter, but there was no effect of body mass on apparent adult survival probability. The modelling detected a significant interaction between sex and season, in which resident females survived better than resident males in spring–summer (?_RF= 0.857 ± 0.117 and ?_RM= 0.698 ± 0.181) while the opposite pattern was found in autumn–winter (?_RM= 0.440 ± 0.086 and ?_RF= 0.339 ± 0.084). In addition, cetti's warblers had a tendency to lose mass in spring–summer (ψ_{Heavy → Light}= 0.560 ± 0.063) and to regain mass in autumn–winter (ψ_{Light→Heavy}= 0.701 ± 0.069). This pattern of body mass change during the annual cycle may reflect energetic costs to reproduction and moulting, and/or a response to increased starvation risk during winter. High body mass, however, did not increase adult survival in this population presumably due to the relatively mild winter weather prevailing in central Portugal. Survival estimates are more likely to be explained by important ecological and behavioural differences between the two sexes in polygynous passerines. Our results highlight that studies aiming to identify the main factors shaping survival and individual fitness in polygynous species should be conducted during different phases of their annual cycle.     

The effect of climate change on partial migration – the blue tit paradox

Climate change has proven to affect various aspects of the migration of birds. In response to milder winters making the habitat more profitable and increasing the survival of residents, the migratory fraction of partially migratory populations has been predicted to decline. We studied the blue tit Parus caeruleus , a common partial migrant in southern Sweden. The numbers migrating at Falsterbo, a migratory passage site in SW Sweden, has increased during the last decades, in parallel with increasing winter and annual temperatures. Migration data from Falsterbo were compared with yearly indices of the size of the breeding population as estimated by the Swedish National Bird Monitoring Programme. Over the study period 1975–2004, also the breeding population has increased in size. The proportion of blue tits migrating each year did not change over the study period, or possibly even increased slightly, which is in contrast to how climate change has been predicted to influence populations containing both migratory and resident individuals. The most important factors determining the intensity of blue tit migration in a given year was the size of an important winter food source, the beech mast crop (more migrants at lower crops) and the size of the breeding population (more migrants at higher densities).     

Body mass and winter mortality in red deer calves: disentangling sex and climate effects

Understanding population dynamics of large mammals requires studies of variation in the age and sex–specific demographic parameters over time and the factors causing this variation. Here, we have focused on the variation in body mass of 8-10–month old red deer calves, in relation to climate and sex over a 20-yr period (1977–1997). We investigated the relationship between body mass and over–winter mortality during 1985 and 1986 and thereby, the phenotypic selection on body mass. We found a high variability from year to year in calf body mass. Males were consistently heavier than females. No interaction between sex and year was detected. The body mass of individuals from the same cohort shot during the annual hunting season and snow depth in January each explained ca 20% of the variability in calf body mass. Body mass loss during winter did not differ between sexes, but increased with body mass and varied from year to year. The probability of surviving was strongly related to body mass in each sex. For a given body mass relative to the sex–specific mean, males had a lower probability of survival than females. Hence, males had to be 1 kg larger than their mean in order to achieve the same survival as average-sized females. Our results suggested a directional phenotypic selection on body mass that led to an increasing body mass dimorphism in calves. The strength of this selection and the sex difference in the shape of the selection curve may depend, however, on the severity of winter and on sexual dimorphism in body mass at the beginning of winter.     

Ecological genetics of an island population of the House mouse (Mus musculus)

The House mouse ( Mus musculus L.) population on the small Welsh island of Skokholm was founded in the 1890s when a few animals were inadvertently introduced. It has thrived ever since. The animals breed from late March to early October, during which time the population increases 8–10 fold. Winter survival is related to temperature: the spring population can be as low as 100 individuals after a cold winter; if the winter is warm, so many animals survive that high numbers may occur in the following autumn. There are no resident predators of the mice, and no other small mammals on the island.
The population was sampled on 20 occasions between 1957 and 1972, and genetical changes measured by the incidences of non-metrical skeletal variants (which describe a substantial proportion of the genome) and (from 1968–72) by allele and genotype frequencies of electrophoretically detected genes. Two sorts of changes occurred: an irregular but increasing deviation from the genetical composition of the population at the beginning of the study period, and an annual cycle with frequencies changing in one direction during the breeding season and the opposite direction in the winter. Samples collected in the autumn were more similar to each other than spring caught samples, which conforms with independent evidence about the differing environmental contributors to death in different winters, as compared with general homogeneity of mortality factors during the summers. It is concluded that the genetical composition of the population is largely dependent on natural selection, although the agents and intensity of selection change with both seasons and years.     

Non‐additive effects of density‐dependent and density‐independent factors on brown trout vital rates

Interactions between density‐dependent and density‐independent processes can lead to variation in both growth and survival rates. Detecting such effects, however, will often require sampling on an individual level and at the appropriate spatial and temporal scale. This study documents substantial variation in survival and growth of stream‐dwelling brown trout Salmo trutta from a small Norwegian stream. The data is based on seasonal capture–recaptures of individually marked trout on fixed stations during eight years. The fish were small‐sized, rarely reaching sizes larger than 20 cm and ages older than seven years. Density varied between 0.2–0.8 fish m^?2. Variation in survival and recapture probabilities was analysed using program MARK. Apparent survival (the probability of being alive and present within the study area) generally decreased with increasing trout density and increasing drought level (measured as lowest observed water flow) during both winter and summer. Further, there was a significant interaction effect between density and water flow, indicating that density‐dependent effects on survival predominated when environmental conditions were benign (no drought), while density‐independent processes were most important under harsh environmental conditions (drought). Observed length‐at‐age during autumn indicated a more or less linear growth trajectory throughout life, and no effect of density, water flow or temperature was found. However, using the individual‐based capture–recapture data to estimated specific growth rate, significant positive effects of water flow and temperature and a negative effect of density were identified. Thus, the capture–recapture data suggest a strong potential for population regulation at the rather low densities found in this stream, and regulation may occur both through effects on survival and growth.     

Estimates of annual survival,growth, and recruitment of a white-tailed ptarmigan population in Colorado over 43 years

Long-term datasets for high-elevation species are rare, and considerable uncertainty exists in understanding how high-elevation populations have responded to recent climate warming. We present estimates of demographic vital rates from a 43-year population study of white-tailed ptarmigan (Lagopus leucura), a species endemic to alpine habitats in western North America. We used capture-recapture models to estimate annual rates of apparent survival, population growth, and recruitment for breeding-age ptarmigan, and we fit winter weather covariates to models in an attempt to explain annual variation. There were no trends in survival over the study period but there was strong support for age and sex effects. The average rate of annual growth suggests a relatively stable breeding-age population ( \( \bar{\lambda } \)  = 1.036), but there was considerable variation between years for both population growth and recruitment rates. Winter weather covariates only explained a small amount of variation in female survival and were not an important predictor of male survival. Cumulative winter precipitation was found to have a quadratic effect on female survival, with survival being highest during years of average precipitation. Cumulative winter precipitation was positively correlated with population growth and recruitment rates, although this covariate only explained a small amount of annual variation in these rates and there was considerable uncertainty among the models tested. Our results provide evidence for an alpine-endemic population that has not experienced extirpation or drastic declines. However, more information is needed to understand risks and vulnerabilities of warming effects on juveniles as our analysis was confined to determination of vital rates for breeding-age birds.     

Moose recruitment in relation to bilberry production and bank vole numbers along a summer temperature gradient in Norway

The plant stress hypothesis states that plant stress factors other than herbivory improve herbivore performance due to changes in the content of nutritive or defensive compounds in the plants. In Norway, the bilberry (Vaccinium myrtillus) is important forage for the bank vole (Myodes glareolus) in winter and for the moose (Alces alces) in summer and autumn. The observed peaks in bank vole numbers after years with high production of bilberries are suggested to be caused by increased winter survival of bank voles due to improved forage quality. High production of bilberries should also lead to higher recruitment rates in moose in the following year. We predict, however, that there is an increasing tendency for a 1-year delay of moose indices relative to vole indices with decreasing summer temperatures, because low temperatures prolong the period needed by plants to recover in the vole peak year, and thus positively affect moose reproduction also in the succeeding year. In eight out of nine counties in south-eastern Norway, there was a positive relationship between the number of calves observed per female moose during hunting and a bilberry seed production index or an autumn bank vole population index. When dividing the study area into regions, there was a negative relationship between a moose-vole time-lag index and the mean summer temperature of the region. These patterns suggest that annual fluctuations in the production of bilberries affect forage quality, but that the effect on moose reproduction also depends on summer temperatures.     

Effects of snow cover on the benthic fauna in a glacier-fed stream

1. Alpine streams above the tree line are covered by snow for 6–9 months a year. However, winter dynamics in these streams are poorly known. The annual patterns of macroinvertebrate assemblages were studied in a glacial stream in the Austrian Alps, providing information on conditions under the snow.
2. Snow cover influenced water temperature, the content of benthic organic matter and insect development. Taxa richness and abundance of macroinvertebrates did not show a pronounced seasonal pattern. The duration of the autumn period with stable stream beds was important in determining the abundance and composition of the winter fauna.
3. There were significant differences in species composition between summer and winter. Two potential strategies in larval survival were evident: adaptation to the extreme abiotic conditions in summer (e.g. Diamesa spp.) or avoidance of these conditions and development during winter (e.g. Ephemeroptera and Plecoptera).
4. A comparison of a stream reach with continuous snow cover and a stream reach that remained open throughout winter showed that conditions under snow are suboptimal. At the open stream site, with higher water temperatures and greater food supply (benthic organic matter content), abundance and taxa richness was higher and larval growth was faster. Several taxa were found exclusively at this site.
5. Winter conditions did not provide an entirely homogeneous environment, abiotic conditions changed rapidly, especially at the onset of snowfall and at snowmelt. Continuous monitoring is necessary to recognize spatial and temporal heterogeneity in winter environments and the fauna of alpine streams.     

The brown trout (<Emphasis Type="Italic">Salmo trutta</Emphasis>) in the lake, Øvre Heimdalsvatn: long-term changes in population dynamics due to exploitation and the invasive species,European minnow (<Emphasis Type="Italic">Phoxinus phoxinus</Emphasis>)

Studies of the brown trout (Salmo trutta) population in the Norwegian subalpine lake, Øvre Heimdalsvatn, over a 50-year period have revealed major changes in population dynamics. In 1958, the population density was high, with individuals stagnating in growth at lengths below 30 cm. After heavy exploitation during the years 1958–1969, the number of older fish declined substantially, and growth rates increased significantly. Since 1969, the European minnow (Phoxinus phoxinus) have been observed in the lake, with increasing densities from 1977–1978 to 1999–2000. The age structure of the brown trout population has changed markedly from the 1960s to the period 1993–2006. Annual recruitment significantly declined, from an average number of 3746 individuals in age-class 4 during the period 1958–1966 to an average of 1668 during the period 1993–2006. However, due to lower exploitation rates, the number of old fish was significantly higher in the latter period. The summer diet of brown trout has changed substantially from a dominance of the large crustaceans Lepidurus arcticus and Gammarus lacustris to a high occurrence of European minnows, while L. arcticus has become practically absent from the diet. There was a negative relationship between brown trout biomass and annual length increment. However, despite a brown trout biomass at the same level during the years 1993–2006 as in the 1960s, annual individual growth rates have significantly declined. The reduced recruitment and reduced annual growth rates of the brown trout, as well as changes in the diet, are most likely related to the introduction and establishment of the invasive species, the European minnow.