The effect of local change in habitat quality on populations of migratory species

1. Habitat deterioration is a major problem world-wide as a result of processes such as change in land use, introduced species, human disturbance and exploitation of food supplies. Many studies have shown that habitat change can have considerable effect on the numbers of individuals using a site. For migratory species, however, the consequences for the total population cannot be deduced from local studies.
2. For a migratory species, the change in total population size Δ N , as a consequence of habitat change in the wintering area, can be calculated from Δ N  =  LM γ d '/( b ' +  d '), where γ is the expected proportional change in the number of birds using a site as a result of the habitat change, L is the area affected, M is the density of individuals using the site prior to habitat change, b ' is the strength of the per capita density-dependent breeding output, and d ' is the strength of the per capita density-dependent winter mortality. Similarly the consequences of habitat change in the breeding area can be calculated from Δ N  =  LM γ b '/( b ' +  d ').
3. The same approach can be used for predicting the consequences of improvements in habitat quality.
4. A worked example is given to illustrate how this approach could be used to predict the consequences for the total population of changes in the food supply of oystercatchers within one estuary.
5. There is a need for more measures of γ, the expected proportional change in the number of birds using a site as a result of various forms of habitat deterioration, and the strengths of density dependence.     

Seasonal matching of habitat quality and fitness in a migratory bird

When species occupy habitats that vary in quality, choice of habitat can be critical in determining individual fitness. In most migratory species, juveniles migrate independently of their parents and must therefore choose both breeding and winter habitats. Using a unique dataset of marked black-tailed godwits (Limosa limosa islandica) tracked throughout their migratory range, combined with analyses of stable carbon isotope ratios, we show that those individuals that occupy higher quality breeding sites also use higher quality winter sites. This seasonal matching can severely inflate inequalities in individual fitness. This population has expanded over the last century into poorer quality breeding and winter habitats and, across the whole population; individual birds tend to occupy either novel or traditional sites in both seasons. Winter and breeding season habitat selection are thus strongly linked throughout this population; these links have profound implications for a wide range of population and evolutionary processes. As adult godwits are highly philopatric, the initial choice of winter habitat by juveniles will be critical in determining future survival, timing of migration and breeding success.     

Corticosterone levels as indicators of habitat quality: effects of habitat segregation in a migratory bird during the non-breeding season

During the non-breeding season, many species of territorial migratory birds exhibit a non-random pattern of habitat distribution, with males and females occupying different habitats. In this study, we examined possible physiological consequences arising from such habitat segregation in one migrant passerine species, the American redstart (Setophaga ruticilla), on its non-breeding grounds in Jamaica, West Indies. For 2 years, we measured concentrations of corticosterone, at the time of capture (baseline) and 30 min after capture (profile of acute corticosterone secretion), in redstarts in two distinct habitats, one occupied predominately by males and one mostly by females. All redstarts in both habitat types exhibited similar concentrations of baseline corticosterone levels in fall (October), whereas in spring (March–April), redstarts in female-biased habitat exhibited significantly higher baseline levels regardless of age or sex. In fall, all individuals in both habitats exhibited significant increases in corticosterone concentration with capture and handling, but in spring only redstarts (both sexes) in male-biased habitat continued to exhibit acute corticosterone secretion. Redstarts in female-biased habitat had elevated baseline corticosterone levels and reduced acute corticosterone secretion. In spring, baseline corticosterone concentration was negatively correlated with body mass, suggesting muscle catabolism associated with high corticosterone concentrations or possibly that birds are leaner as a result of increased foraging effort. These results indicate that redstarts (primarily females) in female-biased habitats suffered a decline in physiological condition, which could in turn influence their departure schedules, migration patterns and even their condition and arrival schedules on the breeding grounds. Thus, segregation of populations into habitats of different quality during the non-breeding period may have ramifications throughout the annual cycle of such migratory species. Furthermore, these results show the usefulness of plasma corticosterone levels as indicators of physiological condition and thus habitat quality for birds during the non-breeding period. Received: 14 November 1997 / Accepted: 9 March 1998     

Carry-over effects,sequential density dependence and the dynamics of populations in a seasonal environment

Most animal populations have distinct breeding and non-breeding periods, yet the implications of seasonality on population dynamics are not well understood. Here, we introduce an experimental model system to study the population dynamics of two important consequences of seasonality: sequential density dependence and carry-over effects (COEs). Using a replicated seasonal population of Drosophila, we placed individuals at four densities in the non-breeding season and then, among those that survived, placed them to breed at three different densities. We show that COEs arising from variation in non-breeding density negatively impacts individual performance by reducing per capita breeding output by 29–77%, implying that non-lethal COEs can have a strong influence on population abundance. We then parametrized a bi-seasonal population model from the experimental results, and show that both sequential density dependence and COEs can stabilize long-term population dynamics and that COEs can reduce population size at low intrinsic rates of growth. Our results have important implications for predicting the successful colonization of new habitats, and for understanding the long-term persistence of seasonal populations in a wide range of taxa, including migratory organisms.     

Predicting the consequences of carry-over effects for migratory populations

Migratory animals present a unique challenge for predicting population size because they are influenced by events in multiple stages of the annual cycle that are separated by large geographic distances. Here, we develop a model that incorporates non-fatal carry-over effects to predict changes in population size and show how this can be integrated with predictive models of habitat loss and deterioration. Examples from Barn swallows, Greater snow geese and American redstarts show how carry-over effects can be estimated and integrated into the model. Incorporation of carry-over effects should increase the predictive power of models. However, the challenge for developing accurate predictions rests both on the ability to estimate parameters from multiple stages of the annual cycle and to understand how events between these periods interact to influence individual success.     

Effects of habitat quality and size on extinction in experimental populations

Stochastic population theory makes clear predictions about the effects of reproductive potential and carrying capacity on characteristic time-scales of extinction. At the same time, the effects of habitat size and quality on reproduction and regulation have been hotly debated. To trace the causal relationships among these factors, we looked at the effects of habitat size and quality on extinction time in experimental populations of Daphnia magna. Replicate model systems representative of a broad-spectrum consumer foraging on a continuously supplied resource were established under crossed treatments of habitat size (two levels) and habitat quality (three levels) and monitored until eventual extinction of all populations. Using statistically derived estimates of key parameters, we related experimental treatments to persistence time through their effect on carrying capacity and the population growth rate. We found that carrying capacity and the intrinsic rate of increase were each influenced similarly by habitat size and quality, and that carrying capacity and the intrinsic rate of increase were in turn both correlated with time to population extinction. We expected habitat quality to have a greater influence on extinction. However, owing to an unexpected effect of habitat size on reproductive potential, habitat size and quality were similarly important for population persistence. These results support the idea that improving the population growth rate or carrying capacity will reduce extinction risk and demonstrate that both are possible by improving habitat quality or increasing habitat size.     

Consequences of habitat loss and change to populations of wintering migratory birds: predicting the local and global effects from studies of individuals

Population models show that the response of migratory bird populations to a reduction in the area, or change in the quality, of their winter feeding areas depends critically on the shape (linear or non-linear) and strength (slope) of the relationship describing how the proportion dying from starvation changes with population density. Yet the parameters that define such density-dependent functions are extremely difficult to estimate from direct measurements of mortality at different population sizes. Even if they could be estimated, there would be uncertainty as to whether or not the relationship would remain the same, and thus provide reliable predictions, in the new circumstances for which forecasts are required. This paper summarizes studies of the Oystercatcher Haematopus ostralegus which aim to derive the shape and strength of the winter density-dependent mortality and emigration function for one estuary, under existing and new circumstances, from the responses of individual birds to each other and to their spatially and seasonally varying food supply. Based on these studies, an individuals-based, physiologically structured game theoretic distribution model has been built which predicts the carrying capacity and numbers of birds dying at different population sizes. The model also can be applied to Oyster-catchers occupying several estuaries distributed throughout their wintering range and can thus be extended to the entire biogeographical or global population. In addition, it can be used to identify some easily measured behavioural and ecological parameters that identify those species, from a wide range of taxa, that are most likely to be affected by habitat loss and change.     

Comparing the seasonal survival of resident and migratory oystercatchers: carry‐over effects of habitat quality and weather conditions

Events happening in one season can affect life‐history traits at (the) subsequent season(s) by carry‐over effects. Wintering conditions are known to affect breeding success, but few studies have investigated carry‐over effects on survival. The Eurasian oystercatcher Haematopus ostralegus is a coastal wader with sedentary populations at temperate sites and migratory populations in northern breeding grounds of Europe. We pooled continental European ringing‐recovery datasets from 1975 to 2000 to estimate winter and summer survival rates of migrant and resident populations and to investigate long‐term effects of winter habitat changes. During mild climatic periods, adults of both migratory and resident populations exhibited survival rates 2% lower in summer than in winter. Severe winters reduced survival rates (down to 25% reduction) and were often followed by a decline in survival during the following summer, via short‐term carry‐over effects. Habitat changes in the Dutch wintering grounds caused a reduction in food stocks, leading to reduced survival rates, particularly in young birds. Therefore, wintering habitat changes resulted in long‐term (>10 years) 8.7 and 9.4% decrease in adult annual survival of migrant and resident populations respectively. Studying the impact of carry‐over effects is crucial for understanding the life history of migratory birds and the development of conservation measures.     

Non-breeding season habitat quality mediates the strength of density-dependence for a migratory bird

Our understanding of when natural populations are regulated during their annual cycle is limited, particularly for migratory species. This information is needed for parametrizing models that can inform management and conservation. Here, we use 14 years of data on colour-marked birds to investigate how conspecific density and habitat quality during the tropical non-breeding period interact to affect body condition and apparent annual survival of a long-distance migratory songbird, the American redstart (Setophaga ruticilla). Body condition and survival of birds in high-quality mangrove habitat declined as density increased. By contrast, body condition improved and survival did not vary as density increased in adjacent, lower quality scrub habitat, although mean condition and survival were almost always lower than in mangrove. High rainfall enhanced body condition in scrub but not in mangrove, suggesting factors such as food availability outweighed consequences of crowding in lower quality habitat. Thus, survival of overwintering redstarts in mangrove habitat, disproportionately males, appears to be regulated by a crowding mechanism based on density-dependent resource competition. Survival of individuals in scrub, mostly females, appears to be limited by density-independent environmental factors but not regulated by crowding. The contrasting effects of density and food limitation on individuals overwintering in adjacent habitats illustrate the complexity of processes operating during the non-breeding period for migratory animals, and emphasize the need for long-term studies of animals in multiple habitats and throughout their annual cycles.     

Macroecology of habitat choice in long-distance migratory birds

Patterns of habitat choice in ecological communities are not only influenced by present-day selective forces but also by historical processes, such as the biogeographical history of the lineages they are composed of. Nevertheless, it has been very difficult to test historical factors. The possible tropical origin of long-distance migratory birds provides an opportunity for such a test. If habitat choice of long-distance migrants is inherited from their tropical ancestors then Nearctic long-distance migrants might have acquired their habitat choice from Neotropical forest species and European long-distance migrants from African savannah species. Here we use a macroecological approach to show that this hypothesis can be confirmed. Long-distance migrants in the Nearctic are found in forested habitat types, while those in Europe are found in open ones. In comparison, the habitat choice of residents and short-distance migrants (in genera without long-distance migration) does not differ between the Nearctic and Europe. These results demonstrate that habitat choice in temperate bird communities can be explained by the tropical history of long-distance migrants. Thus, habitat choice seems to be shaped not only by local mechanisms, but also by processes acting on much larger spatial and temporal scales.     

Carry‐over effects of winter habitat quality on en route timing and condition of a migratory passerine during spring migration

We examined how conditions prior to migration influenced migration performance of two breeding populations of black‐and‐white warblers Mniotilta varia by linking information on the migrant's winter habitat quality, measured via stable carbon isotopes, with information on their breeding destination, measured via stable hydrogen isotopes. The quality of winter habitat strongly influenced the timing of migration when we accounted for differential timing of migration between breeding populations. Among birds migrating to the same breeding destination, males and females arriving early to the stopover site originated from more mesic habitat than later arriving birds, suggesting that the benefits of occupying high‐quality mesic habitat during the winter positively influence the timing of migration. However, male warblers arriving early to the stopover site were not in better migratory condition than later arriving conspecifics that originated from poor‐quality xeric winter habitat, regardless of breeding destination. The two breeding populations stopover at the study site during different time periods, suggesting that the lower migratory condition of early birds is not a function of the time of season, but potentially a migrant's migration strategy. Strong selection pressures to arrive early on the breeding grounds to secure high‐quality breeding territories may drive males from high‐quality winter habitat to minimize time at the expense of energy. This migration strategy would result in a smaller margin of safety to buffer the effects of adverse weather or scarcity of food, increasing the risk of mortality. The migratory condition of females was the same regardless of the timing of migration or breeding destination, suggesting that females adopt a strategy that conserves energy during migration. This study fills an important gap in our understanding of the linkages between winter habitat quality and factors that influence the performance of migration, the phase of the annual cycle thought to be limiting most migratory bird populations.     

Influence of water quality, habitat and species richness on brown trout populations

The influence of water quality, physical habitat and species richness on the occurrence, density and size of brown trout at 216 stream sites in southern Sweden was studied. Discriminant analysis showed that the occurrence of trout at a locality was largely determined by oxygen conditions and medium-sized substrata. At localities where trout occurred, the density of + trout was highest in narrow streams with high oxygen concentrations. For older trout, >0+ in age, stream size and temperature were negatively related to density. Biotic factors also appeared to affect trout density, as trout density was inversely related to abundance of predators and coexisting species. Even intraspecific competition appeared to be important as length of 0+ trout was inversely related to trout density. It is suggested that improvements of water quality may be an effective way to restore sea trout populations in southern Sweden, especially in narrow streams where smolt production has the highest potential.     

Migration phenology and winter habitat quality are related to circulating androgen in a long‐distance migratory bird

In migratory birds, the timing of departure from wintering grounds is often dependant on the quality of habitat on an individual's territory and may influence individual fitness, resulting in an interaction of life history stages across large geographical distances. American redstart Setophaga ruticilla males who overwinter in high quality habitats arrive early to breed and subsequently produce more offspring than late arrivers. Since many migratory species overlap vernal migration with the physiological transition to breeding, we examined if breeding preparation plays a role in this seasonal interaction. We tested the hypothesis that early arriving male redstarts from high quality winter habitats are in superior breeding condition by simultaneously measuring winter habitat quality (stable‐carbon isotopes) and breeding preparation (circulating androgen, cloacal protuberance (CP) diameter) upon arrival at breeding grounds. Compared with late arrivers, early arriving males were from higher quality winter habitats and had higher androgen, but smaller CPs. Males arriving with higher androgen were in more advanced physiological migratory condition, as measured by haematocrit. Early arrivers were more likely to successfully breed, but there was no significant relationship between androgen upon arrival and breeding success. One possible explanation for these relationships is that androgen measured during arrival is most relevant in a migratory context, such that birds with high androgen may benefit from effects on migratory condition, positively influencing fitness through earlier arrival.     

Homing fidelity and reproductive rate for migratory populations

Theoretical Ecology - Short-term and long-term population growth rates can differ considerably. While changes in growth rates can be driven by external factors, we consider another source for...     

Carry-over effects of day length during spring migration

The day lengths to which migratory birds are exposed depend on the timing and course of their journey. While winter day length is known to influence vernal events, it is not clear if birds also use day length during the spring migration as a temporal cue. We addressed this question by exposing captive stonechats (Saxicola torquata) to two different photoperiodic simulations of spring migration routes, following common winter conditions. One group experienced day lengths of the regular (“fast”) migration, and the other group, a “slow”, or more southerly originating, route. The resulting small, temporary differences in day length had lasting effects on the birds. The groups differed in migratory restlessness during and following exposure to different day lengths. “Slow” migrants continued nocturnal activity longer than “fast” migrants. Furthermore, all activities of the ensuing breeding season were delayed in the “slow” migrants, indicating a phase shift in their underlying annual rhythm. “Slow” migrants delayed terminating their reproductive stage by regressing testes and the cloacal protuberance later than the “fast” migrants. Molt started and ended later in “slow” migrants, but the duration of the molt was unaffected by spring day length. Finally, “fast” migrants resumed nightly restlessness earlier than “slow” migrants in late summer. These results demonstrate that Zugunruhe (migratory restlessness) and reproductive windows are not set exclusively during winter but can be modified by day length cues during the spring migration. Because migration modifies the day length exposure of birds, migration routes can have carry-over effects on the timing of breeding season events, including the completion of molt and initiation of autumnal nocturnal activity.

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Einfluss der Tagesl?nge w?hrend des Frühjahrszugs auf Brutzeitraum, Mauser, und Beginn der Herbstzugunruhe

Zusammenfassung Zugv?gel beeinflussen durch ihre Zugroute und Zugzeit die Tagesl?nge, in der sie sich befinden. W?hrend experimentell belegt ist, dass die Wintertagesl?nge das saisonale Verhalten im Frühling beeinflusst, ist bisher unbekannt, ob V?gel auch w?hrend der Frühjahrszugzeit photoperiodische Zeitinformationen nutzen. Um diese Frage zu beantworten, haben wir Schwarzkehlchen (Saxicola torquata) unter den Lichtbedingungen von zwei verschiedenen Zugrouten untersucht. Die V?gel hatten unter einheitlichen Bedingungen überwintert. Anschlie?end erlebte eine Gruppe die Lichtbedingungen ihres regul?ren (“schnellen“) Zugweges, die andere die eines “langsameren“, oder weiter südlich beginnenden Zugweges. Die damit verbundenen geringen Unterschiede in der Tagesl?nge zeigten nachhaltige Wirkung. V?gel der beiden Gruppen unterschieden sich in Zugunruhe w?hrend und nachdem sie unterschiedliche Tagesl?ngen erlebten. “Langsame” Zieher setzten ihre n?chtliche Aktivit?t l?nger fort als “schnelle" Zieher. Darüber hinaus verz?gerten sich bei den “langsamen" Ziehern alle Aktivit?ten der anschlie?enden Brutsaison. Dies deutet auf eine Phasenverschiebung der zugrunde liegenden Jahresrhythmen hin. “Langsame" Zieher verz?gerten den Abschluss ihrer Brutphase, indem sie Hoden und Kloakenprotuberanz sp?ter zurückentwickelten als “schnelle" Zieher. Die Mauser begann und endete sp?ter als bei “schnellen" Ziehern, dauerte aber gleich lang an. Schlie?lich nahmen “schnelle" Zieher die n?chtliche Unruhe früher wieder auf als “langsame" Zieher. Diese Ergebnisse zeigen, dass der Brutzeitraum von Zugv?geln nicht allein im Winter, sondern auch w?hrend des Frühjahrszugs von der Tagesl?nge beeinflusst wird. Somit k?nnen sich Zugzeit und Zugroute auf das anschlie?ende zeitliche Verhalten bis zu Mauserende und Beginn der Herbstzugunruhe auswirken.