Cas Eikenaar

• Institute of Avian Research

Migratory birds may either upregulate their immune system during migration as they might encounter novel pathogens or downregulate their immune system as a consequence of trade‐offs with the resource costs of migration. Support for the latter comes not least from a study that reports a positive correlation in autumn migrating birds between fuel sto...

Migrating birds perform extreme endurance exercise when flying. This shifts the balance between the production of reactive oxygen species and the antioxidant defense system toward the former, potentially generating oxidative damages. In between migratory flights, birds make stopovers, where besides accumulating fuel (mainly fats), they are assumed...

Migratory birds may either upregulate their immune system during migration as they might encounter novel pathogens or downregulate their immune system as a consequence of trade-offs with the resource costs of migration. Support for the latter comes not least from a study that reports a positive correlation in autumn migrating birds between fuel sto...

Strenuous physical activity can negatively affect constitutive innate immune function (CIF), the always present first line of defence against pathogens. CIF is non-specific, and thus vital when encountering novel pathogens. A lowered CIF likely increases the risk of infection and disease. Migratory birds engage in truly extreme physical activity du...

Background Birds have extremely elevated metabolic rates during migratory endurance flight and consequently can become physiologically exhausted. One feature of exhaustion is oxidative damage, which occurs when the antioxidant defense system is overwhelmed by the production of damaging reactive oxygen species (ROS). Migrating birds have been shown...

Many songbirds migrate twice a year to exploit seasonally available resources. These journeys are energetically demanding and energy reserves of these small birds are limited. Accordingly, most of the time migrating is spent during stopovers that serve to rest, replenish resources and recover. While external influences, like prevailing weather cond...

During migratory endurance flights, which are energetically very demanding, migrants have to deal with prolonged elevated generation of reactive oxygen species (ROS). To limit the damaging actions that ROS have on lipids and proteins, migrating birds are known to upregulate their antioxidant defence system. However, there may be additional ways to...

Global movement patterns of migratory birds illustrate their fascinating physical and physiological abilities to cross continents and oceans. During their voyages, most birds land multiple times to make so‐called ‘stopovers’. Our current knowledge on the functions of stopover is mainly based on the proximate study of departure decisions. However, s...

To fuel their migratory endurance flights, most birds accumulate large quantities of fat prior to departure. It therefore seems logical that the decision to depart on a migratory flight depends on fuel stores, at least to some extent; very small fuel stores prohibit migratory endurance flight. However, studies linking migrants' estimated fuel store...

Birds have been observed to have dietary preferences for unsaturated fatty acids during migration. Polyunsaturated fatty acids (PUFAs) may increase the exercise performance of migrant birds; however, PUFAs are also peroxidation prone and might therefore incur increased costs in terms of enhanced oxidative damage in migratory individuals. To shed li...

Animals usually show distinct periods of diel activity and non-activity. Circulating baseline levels of glucocorticoid hormones (corticosterone and cortisol) often peak just before or at the transition from the non-active to the active period of the day. This upregulation of glucocorticoids may function to mobilize stored energy and prepare an anim...

Most migrating birds need to stopover in between flights in order to refuel. Lately, additional purposes of stopover have been suggested, including physiological recovery from metabolically demanding migratory flight. One apparently unavoidable, but harmful physiological effect of migratory flight is increased oxidative damage to lipids and protein...

Background: Songbirds following distinct migration strategies (e.g. long- vs. short- to medium-distance migrants) often differ in their speed of migration during autumn and, thus, are assumed to face different time constraints. During migration, most songbird species alternate migratory flights with stopover periods. Many of them restrict these mi...

Migratory flight is physiologically highly demanding and has been shown to negatively affect multiple parameters of constitutive immune function (CIF), an animal's first line of physiological defence against infections. In between migratory flights, most birds make stopovers, periods during which they accumulate fuel for the next flight(s). Stopove...

[This corrects the article DOI: 10.1093/cz/zoz009.][This corrects the article DOI: 10.1093/cz/zoz009.].

Stopover periods in between migratory flights determine the speed of migration and as such may affect birds’ fitness. The decision to leave a stopover site is affected by, for example, energy stores and weather conditions. We investigated whether immunological state influences autumn stopover departure decisions in Northern Wheatear (Oenanthe oenan...

In many animals, catabolic and anabolic periods are temporally separated. Migratory birds alternate energy expenditure during flight with energy accumulation during stopover. The size of the energy stores at stopover affects the decision to resume migration and thus the temporal organization of migration. We now provide data suggesting that it is n...

Wild animals typically suffer from stress when brought into captivity. This stress is characterized by elevated circulating glucocorticoid levels and weight loss. We here describe for the first time a case where a wild animal, the long-distance migrating northern wheatear, does not show signs of stress when caged. We captured these birds on a stopo...

The spectacular natural phenomenon of avian migration is evidently shaped by physical factors, but we know little about the underlying physiological regulation. This contrast is especially apparent for the process of departure on a migratory flight. The decision to resume migration is shaped by a suite of departure cues from innate rhythms, and int...

In migratory songbirds, older individuals are thought to be more efficient migrants than younger individuals. Age‐specific differences in migratory efficiency have been reported mainly in respect of arrival timing, energy stores, rate of energy accumulation, departure behaviour, and departure direction. Yet, these traits were rarely assessed simult...

Most migratory songbirds travel between their breeding areas and wintering grounds through a series of nocturnal flights. The timing of their departures defines the potential flight duration and thus the distance covered during a migratory night. Yet, migratory songbirds show substantial variation in their nocturnal departure timing. With this stud...

Migration is energetically demanding and physiologically challenging. Migrating birds, for example, need to boost their antioxidant defenses to defeat the pro-oxidants produced during high energetic activity. The enhanced antioxidant defense possibly withdraws limited resources (e.g., energy or micronutrients) from other physiological functions, su...

Most migrating birds make stopovers to replenish fuel stores. The decision to resume migration from stopover to a large extent shapes the temporal organization of migration. This decision is known to be shaped by a suite of intrinsic and extrinsic factors such as the bird's fuel stores and current weather conditions. However, how departures from st...

Most migrating birds make multiple stopovers to fuel and/or rest. The decision to resume migration from stopover is based on various cues, such as time within the season and wind conditions. There are hints that the strength of these departure cues shapes corticosterone level, which in its turn appears to regulate the timing of departure. We here p...

To support migratory endurance flight, birds accumulate large amounts of fat by hyperphagia (fueling). Whereas the factors influencing migrants’ motivation to fuel are well described, the physiological mechanism regulating fueling is largely unknown. Hormones are likely involved and arguably the best studied with respect to food intake and fueling...

In birds, accumulating energy is far slower than spending energy during flight. During migration, birds spend, therefore, most of the time at stopover refueling energy used during the previous flight. This elucidates why current energy stores and actual rate of accumulating energy are likely crucial factors influencing bird’s decision when to resum...

Cooperative breeding – in which some sexually mature individuals forgo independent breeding, join a group as subordinate and help to raise the offspring of others – occurs in at least 3% (mammals) and 9% (birds) of vertebrates. Because helping others is costly, this behaviour contradicts the concept of ‘selfish’ natural selection. The intriguing ev...

Bird migration entails replenishing fuel stores at stopover sites. There, individuals make daily decisions whether to resume migration, and must also decide their time of departure. Variation in departure timing affects the total time required to complete a migratory journey, which in turn affects fitness through arrival time at the breeding and wi...

Most avian migrants alternate flight bouts, characterized by high metabolic rates, with stopovers, periods of fuel replenishment through hyperphagia. High-energy metabolism and excessive calorie intake shift the balance between damaging prooxidants and antioxidants toward the former. Hence, migration likely affects the oxidative balance of birds. M...

Migrating animals typically make stopovers to rest and replenish the fuel used during previous travel bouts. In birds, fat or fast refuelling individuals are generally more likely to depart from stopover than lean or slow refuelling birds. The departure decision, however, involves more than the day-to-day departure likelihood. Also, the time of nig...

Animals need a well-functioning immune system to protect themselves against pathogens. The immune system, however, is costly and resource trade-offs with other demands exist. For migratory animals several (not mutually exclusive) hypotheses exist. First, migrants reduce immune function to be able to allocate resources to migration. Second, migrants...

For migrating birds, late arrival at the breeding grounds reduces reproductive success. Hence, while en route, late migrants are expected to accelerate migration. Most time can be saved by shortening stopovers, periods of rest and fuel replenishment (refuelling) in between flight bouts. In several field studies late migrants have indeed been observ...

In many migratory birds, males precede females during migration and arrival at the breeding sites. Three proximate mechanisms are proposed to explain this phenomenon of protandry: males 1) winter closer to breeding sites, 2) start spring migration earlier, and/or 3) migrate faster than females. So far, the relative contribution of these mechanisms...

In nocturnal songbird migrants, total speed of migration is determined by the time birds stay at stopovers, where they replenish fuel reserves used during previous flights, and by their travel speed. In contrast to factors influencing stopover duration, little is known about individual variation in travel speed which is a combination of birds’ grou...

A widespread phenomenon in migrant birds is that they travel faster in spring than in autumn. During migration birds spend most time at stopover sites and, correspondingly, the faster spring migration is mainly explained by shorter stopovers in spring than autumn. Because a main purpose of stopovers is to replenish the fuel used in flight, a higher...

Numerous studies in both the field and laboratory illustrate factors involved in the endogenous, intrinsic and environmental control of avian migration, but we are lacking an integrated individual-based approach, connecting field and laboratory studies in a single species. The Northern Wheatear Oenanthe oenanthe appeared very feasible for an integr...

A variety of methods have been used to study the relationship between wind conditions and departure decisions of migrant birds at stopover sites. These methods are either costly or suffer from inaccuracy in determining whether or not an individual has resumed migration. Here we present a novel and simple approach to studying the relationship betwee...

Most migrating birds alternate flight bouts with stopovers, during which they rest and replenish the fuel used during flight (refueling). The rate of refueling (fuel deposition rate, FDR) affects stopover duration, and hence is an important determinant of the overall time required for migration. Although environmental and endogenous factors affect...

The decision made by migrating birds to stop refueling and to depart from stopover depends on cues from innate rhythms, intrinsic factors such as fuel reserves, and extrinsic factors such as weather conditions. The physiological mechanism behind this decision, however, is largely unexplored. The transition from refueling to flight involves an incre...

Abstract Field endocrine studies providing new comparisons for inference into the evolutionary and ecological factors shaping organismal physiology are important, often yielding novel physiological insights. Here, we explored factors associated with the sex steroid hormone concentrations and adrenocortical response to capture stress in Hog Island b...

In captivity, migratory birds show increased activity during the time that they would normally migrate. The phenology and intensity of such 'migratory restlessness' has been shown to mirror species- and population-specific migration patterns observed in the wild and has consequently been used as a proxy for the motivation to migrate. Many studies d...

Elevated baseline corticosterone levels function to mobilize energy in predictable life-history stages, such as bird migration. At the same time, baseline corticosterone has a permissive effect on the accumulation of fat stores (fueling) needed for migratory flight. Most migrants alternate flight bouts with stopovers, during which they replenish fu...

Abstract Mating systems can vary among species and populations and thus influence evolutionary trajectories, ecological traits, and population demography. The siring of offspring by an extrapair male, or extrapair paternity (EPP), is a widespread and varied phenomenon in all vertebrate classes. However, we do not understand all of the factors assoc...

Early arrival at the breeding site positively affects the breeding success of migratory birds. During migration, birds spend most of their time at stopovers. Therefore, determining which factors shape stopover duration is essential to our understanding of avian migration. Because the main purpose of stopover is to accumulate fat as fuel for the nex...

The latitudinal increase in extra-pair paternity (EPP) rates in birds suggests broad selective benefits to low EPP rates in the tropics. However, we have few EPP data from tropical birds, particularly from species with close relatives at high latitudes. Here, we report EPP rates in two resident equatorial populations of rufous-collared sparrow Zono...

Migratory birds rely on fueling prior to migratory flights. Fueling in migrants is controlled by intrinsic as well as extrinsic factors. From captive studies we have started understanding the internal mechanisms controlling bird migration. Field studies have demonstrated the effects of external factors, such as food availability, weather, competito...

Migrating birds spend most of their time at stopover sites where they replenish the fuel used during flight, termed refueling. The overall time of migration thus largely depends on the duration of stopovers, and factors shaping stopover duration therefore are of interest. A handful of field studies have shown that the likelihood of departure from s...

Abstract Latitudinal variation in life-history traits has been the focus of numerous investigations, but underlying hormonal mechanisms have received much less attention. Steroid hormones play a central role in vertebrate reproduction and may be associated with life-history trade-offs. Consequently, circulating concentrations of these hormones vary...

Relationships between phenotype (testosterone concentration and length of the longest tail feather) and fertilization success of male barn swallows calculated from first nests only (most pairs were double brooded). Fertilization success was split into i) cuckoldry rate, i.e. the proportion of extra-pair young in a male's own first brood, ii) extra-...

In most monogamous bird species, circulating testosterone concentration in males is elevated around the social female's fertile period. Variation in elevated testosterone concentrations among males may have a considerable impact on fitness. For example, testosterone implants enhance behaviours important for social and extra-pair mate choice. Howeve...

Males may face a trade-off between caring for offspring and pursuing additional matings. In birds, the androgen testosterone has been suggested to be a key proximate mediator in this trade-off for several reasons. At the population level, high testosterone is typically associated with the period of intense male--male competition over females, where...

We observed a change in the sex-specific rate of delayed natal dispersal in a stable population of Seychelles warblers over a period of 20 years. At first, females were more likely to delay dispersal in their first year of life than were males, whereas later there was no sex bias in the rate of delayed natal dispersal. Similarly, the female-bias in...

We describe the reproductive cycle of Northern Pacific rattlesnakes (Crotalus oreganus) by quantifying steroid hormone concentrations and observing reproductive behaviors in free-ranging individuals. Additionally, we examined reproductive tissues from museum specimens. Plasma steroid hormone concentrations were quantified for both male and female s...

Knowing why certain individuals refrain from reproduction is fundamental to our understanding of the evolution of cooperative breeding. Although many studies have focused on the role of life history and ecological factors, the proximate mechanisms underlying this behaviour have received considerably less attention. We determined the natural variati...

In many species, territory ownership is a prerequisite for reproduction; consequently, factors that affect success in territory acquisition can have a large impact on fitness. When competing for territories, some individuals may have an advantage if, for example, they are phenotypically superior or more familiar with the site than others. The relat...

Cooperative breeding – in which some adults forgo independent breeding and remain as subordinates within a group helping to raise the offspring of others – occurs in between 3% and 10% of vertebrates. The structure of such systems varies greatly, from pairs with helpers-at-the-nest to communal breeders, and may include young helpers or post-reprodu...

In this study, we test whether patterns of territory inheritance, social mate choice and female-biased natal dispersal act as inbreeding avoidance mechanisms in the cooperatively breeding Seychelles warbler. Our results show that Seychelles warblers do not reduce the likelihood of inbreeding by avoiding related individuals as mates. The occurrence...

The distances that individuals disperse, from their natal site to the site of first breeding and between breeding sites, have important consequences for the dynamics and genetic structure of a population. Nearly all previous studies on dispersal have the problem that, because the study area encompassed only a part of the population, emigration may...

Voor dit proefschrift hebben we de patronen van dispersie van jonge Seychellen zangers onderzocht. Deze kleine zangvogels (15g) leven in groepen bestaande uit een dominant broedpaar met eventueel daarnaast jongen die nog door hun ouders gevoerd worden en/of onafhankelijke jongen die hun dispersie uitgesteld hebben. Een deel van de onafhankelijke jo...

The presence of parents in the natal territory may play an important, but often overlooked, role in natal dispersal and the consequent acquisition of a territory. Living with parents in a territory may confer a fitness advantage to subordinates through, for example, the nepotistic behavior of the parents or indirect benefits gained by helping to ra...

Variation in survival, a major determinant of fitness, may be caused by individual or environmental characteristics. Furthermore, interactions between individuals may influence survival through the negative feedback effects of density dependence. Compared to species in temperate regions, we have little knowledge about population processes and varia...

The amount of time a bird allocates to incubation is likely to be limited by energetic constraints. If food is abundant, energetic constraints may be reduced and the time spent incubating (incubation attendance) may increase. Moreover, the onset of incubation in relation to clutch completion may be advanced, resulting in a higher degree of hatching...

We tested the hypothesis that mother birds counterbalance the negative effects of hatching asynchrony for later-hatched chicks by increasing the yolk androgen concentrations in consecutive eggs of their clutch. In doing so, they may adaptively tune each offspring’s competitive ability and, thus, growth and survival. However, evidence in support of...

Cooperative breeding - in which some adults forgo independent breeding and remain as subordinates within a group helping to raise the offspring of others - occurs in between 3% and 10% of vertebrates. The structure of such systems varies greatly, from pairswithhelpers-at-the-nesttocommunalbreeders,andmayincludeyounghelpersor post-reproductive 'gran...