Michael Schaub

Fuel deposition of three passerine bird species along the migration route

Abstract The rate at which migrant birds replenish their energy stores at intermittent stopovers largely determines overall migration speed, the manner in which migration proceeds and success of migration. In this study, data on the fuel deposition rate (FDR) of three long-distance migrants from 17 ringing sites along their autumn migration route were used to examine: (1) effects of endogenous factors on FDR, and (2) how relationships between exogenous factors and FDR affect the organisation of migration. We developed a model to estimate FDR from retrapped birds which takes into account time of day and various other factors which might influence FDR. The two endogenous factors, moult and current energy stores, generally reduced FDR. This may result in lower departure energy loads and more stopovers than expected from optimal migration theory. Differences between species with respect to seasonal, year-to-year and geographical patterns of FDR could be related to differences in availability and predictability of food resources, and help to explain differences in the organisation of migration. A low FDR in northern and central Europe could be related to low, but predictable, food resources and an early departure during moult of the reed warbler (Acrocephalus scirpaceus); FDRs varying between years were related to large spatial and year-to-year variation in the density of the main prey of the sedge warbler (Acrocephalus schoenobaenus); and a high FDR in the garden warbler (Sylvia borin) was related to abundant food resources, due to a switch from a purely invertebrate diet to a mixed diet including fruits which are abundant over large areas of Europe and north Africa. This study demonstrated that the organisation of migration is the outcome of a complex interplay of the seasonal timing of moult, food availability and predictability and a seasonal switch in diet, and can be modified by individual birds in response to a limited amount of time in which to migrate.

Year-Round Tracking of Small Trans-Saharan Migrants Using Light-Level Geolocators

Since 1899 ringing (or banding) remained the most important source of information about migration routes, stopover sites and wintering grounds for birds that are too small to carry satellite-based tracking systems. Despite the large quantity of migrating birds ringed in their breeding areas in Europe, the number of ring recoveries from sub-Saharan Africa is very low and therefore the whereabouts of most small bird species outside the breeding season remain a mystery. With new miniaturized light-level geolocators it is now possible to look beyond the limits of ring recovery data. Here we show for the first time year round tracks of a near passerine trans-Saharan migrant, the European Hoopoe (Upupa epops epops). Three birds wintered in the Sahel zone of Western Africa where they remained stationary for most of the time. One bird chose a south-easterly route following the Italian peninsula. Birds from the same breeding population used different migration routes and wintering sites, suggesting a low level of migratory connectivity between breeding and wintering areas. Our tracking of a near passerine bird, the European Hoopoe, with light-level geolocators opens a new chapter in the research of Palaearctic-African bird migration as this new tool revolutionizes our ability to discover migration routes, stopover sites and wintering grounds of small birds.

ESTIMATING SURVIVAL AND TEMPORARY EMIGRATION IN THE MULTISTATE CAPTURE–RECAPTURE FRAMEWORK

The open population Cormack-Jolly-Seber (CJS) capture–mark–recapture model for estimating survival allows for random temporary emigration from the sampling area, but Markovian temporary emigration can bias estimates of survival. We explore a multistate capture–recapture model that has been proposed for coping with Markovian temporary emigration. We provide a comprehensive assessment of the performance of this model using computer algebra and simulation. We found that most model parameters were identifiable unless survival, emigration, and immigration were all time dependent. Simulation results showed that intrinsically identifiable parameters were estimated without bias and that precision of survival estimates was always high. When temporary emigration was Markovian, precision of estimates of emigration, immigration, and recapture probabilities was acceptable; otherwise it was not. Test component 2.Ct of the goodness-of-fit test for the CJS model had good power to detect Markovian temporary emigration. We conclude that the multistate model works well when temporary emigration is Markovian (i.e., when the CJS model should not be used) and when survival and recapture probabilities are high.

Behavioural and Physiological Reactions to Environmental Variation in Bird Migration: a Review

Many features of the migration of passerine birds are endogenously programmed and have a strong genetic basis. At least in inexperienced migrants, the general seasonal course of migration is endogenously controlled, such as the onset, the temporal pattern, the direction of migration and the seasonal pattern of energy stores. This leads to the conclusion that an endogenous spatiotemporal migration programme guides inexperienced migrants from their place of birth to their first winter quarters (summarized in Berthold 1996).

Spreading free-riding snow sports represent a novel serious threat for wildlife

Stress generated by humans on wildlife by continuous development of outdoor recreational activities is of increasing concern for biodiversity conservation. Human disturbance often adds to other negative impact factors affecting the dynamics of vulnerable populations. It is not known to which extent the rapidly spreading free-riding snow sports actually elicit detrimental stress (allostatic overload) upon wildlife, nor what the potential associated fitness and survival costs are. Using a non-invasive technique, we evaluated the physiological stress response induced by free-riding snow sports on a declining bird species of Alpine ecosystems. The results of a field experiment in which radiomonitored black grouse (Tetrao tetrix) were actively flushed from their snow burrows once a day during four consecutive days showed an increase in the concentration of faecal stress hormone (corticosterone) metabolites after disturbance. A large-scale comparative analysis across the southwestern Swiss Alps indicated that birds had higher levels of these metabolites in human-disturbed versus undisturbed habitats. Disturbance by snow sport free-riders appears to elevate stress, which potentially represents a new serious threat for wildlife. The fitness and survival costs of allostatic adjustments have yet to be estimated.

Sex‐dependent selection on an autosomal melanic female ornament promotes the evolution of sex ratio bias

Sex-dependent selection often leads to spectacularly different phenotypes in males and females. In species in which sexual dimorphism is not complete, it is unclear which benefits females and males derive from displaying a trait that is typical of the other sex. In barn owls (Tyto alba), females exhibit on average larger black eumelanic spots than males but members of the two sexes display this trait in the same range of possible values. In a 12-year study, we show that selection exerted on spot size directly or on genetically correlated traits strongly favoured females with large spots and weakly favoured males with small spots. Intense directional selection on females caused an increase in spot diameter in the population over the study period. This increase is due to a change in the autosomal genes underlying the expression of eumelanic spots but not of sex-linked genes. Female-like males produced more daughters than sons, while male-like females produced more sons than daughters when mated to a small-spotted male. These sex ratio biases appear adaptive because sons of male-like females and daughters of female-like males had above-average survival. This demonstrates that selection exerted against individuals displaying a trait that is typical of the other sex promoted the evolution of specific life history strategies that enhance their fitness. This may explain why in many organisms sexual dimorphism is often not complete.

An assessment of integrated population models: bias, accuracy, and violation of the assumption of independence

Understanding population dynamics requires accurate estimates of demographic rates. Integrated population models combine demographic and survey data into a single, comprehensive analysis and provide more coherent estimates of vital rates. Integrated population models rely on the assumption that different data sets are independent, which is frequently violated in practice. Moreover, the precision that can be gained using integrated modeling compared to conventional modeling is only known from empirical studies. The present study used simulation methods to assess how the violation of the assumption of independence affects the statistical properties of the parameter estimators. Further, the gains in precision and accuracy from the model were explored under varying sample sizes. For capture-recapture, population survey, and reproductive success, we generated independent and dependent data that were analyzed with integrated and conventional models. We found only a minimal impact of the violation of the assumption of independence on the parameter estimates. Furthermore, we observed an overall gain in precision and accuracy when all three data sets were analyzed simultaneously. This was particularly pronounced when the sample size was small. These findings contribute to clearing the way for the application of integrated population models in practice.

Patches of Bare Ground as a Staple Commodity for Declining Ground-Foraging Insectivorous Farmland Birds

Conceived to combat widescale biodiversity erosion in farmland, agri-environment schemes have largely failed to deliver their promises despite massive financial support. While several common species have shown to react positively to existing measures, rare species have continued to decline in most European countries. Of particular concern is the status of insectivorous farmland birds that forage on the ground. We modelled the foraging habitat preferences of four declining insectivorous bird species (hoopoe, wryneck, woodlark, common redstart) inhabiting fruit tree plantations, orchards and vineyards. All species preferred foraging in habitat mosaics consisting of patches of grass and bare ground, with an optimal, species-specific bare ground coverage of 30–70% at the foraging patch scale. In the study areas, birds thrived in intensively cultivated farmland where such ground vegetation mosaics existed. Not promoted by conventional agri-environment schemes until now, patches of bare ground should be implemented throughout grassland in order to prevent further decline of insectivorous farmland birds.

The demographic drivers of local population dynamics in two rare migratory birds

The exchange of individuals among populations can have strong effects on the dynamics and persistence of a given population. Yet, estimation of immigration rates remains one of the greatest challenges for animal demographers. Little empirical knowledge exists about the effects of immigration on population dynamics. New integrated population models fitted using Bayesian methods enable simultaneous estimation of fecundity, survival and immigration, as well as the growth rate of a population of interest. We applied this novel analytical framework to the demography of two populations of long-distance migratory birds, hoopoe Upupa epops and wryneck Jynx torquilla, in a study area in south-western Switzerland. During 2002–2010, the hoopoe population increased annually by 11%, while the wryneck population remained fairly stable. Apparent juvenile and adult survival probability was nearly identical in both species, but fecundity and immigration were slightly higher in the hoopoe. Hoopoe population growth rate was strongly correlated with juvenile survival, fecundity and immigration, while that of wrynecks strongly correlated only with immigration. This indicates that demographic components impacting the arrival of new individuals into the populations were more important for their dynamics than demographic components affecting the loss of individuals. The finding that immigration plays a crucial role in the population growth rates of these two rare species emphasizes the need for a broad rather than local perspective for population studies, and the development of wide-scale conservation actions.

Effect of current reproduction on apparent survival, breeding dispersal, and future reproduction in barn swallows assessed by multistate capture–recapture models

1. Theoretical models predict a negative effect of current reproduction on breeding dispersal, survival and future reproduction, and many studies confirm these predictions. Yet, results of most previous studies may be difficult to interpret because the fate of the affected individuals cannot always be observed. Detection is almost always imperfect and some individuals emigrate from the study area, resulting in biased estimates of both survival and dispersal. 2. Most studies bypass these problems with strong assumptions. We use a multistate capture-recapture model that does not require these assumptions. States are defined based on classes of reproductive success and on observed dispersal events within the study area. By accounting for imperfect detection within the study area, the model allows estimation of the effect of reproductive success on apparent survival, dispersal probabilities within the study area and the annual transition probabilities among classes of reproductive success. Based on an assumption about the estimate of real survival, the model allows the estimation of total dispersal that is not specific to a fixed study area. 3. We applied this model to capture-recapture data of 2262 adult barn swallows (Hirundo rustica) sampled from 1997-2004 in eight local populations in Switzerland. 4. We found that dispersal within the study area decreased with increasing reproductive success in both sexes, that reproductive success was not affected by preceding dispersal and that apparent survival of females but not of males increased with increasing reproductive success. Apparent survival of females with high reproductive success was identical to apparent survival of males suggesting that this estimate of apparent survival (0.48) was close to true survival. Total breeding dispersal was generally higher in females and it increased with decreasing reproductive success in both sexes. Current reproductive success depended on reproductive success in the preceding year suggesting that individual differences were of importance. 5. Our study highlights that reproductive success was an important factor affecting breeding dispersal and population turnover. While unsuccessful males mainly remained in the local populations, many unsuccessful females left them. Population turnover of adult swallows was mainly due to unsuccessful females.