Do great tits rely on inadvertent social information from blue tits? A habitat selection experiment

Sympatric species sharing requirements are competitors, but recent evidence suggests that heterospecifics may also be used as a source of information. The heterospecific habitat copying hypothesis proposes that individuals of one species might use information inadvertently produced by the breeding performance of individuals of other species to assess habitat quality whenever the two species share needs. In this study, we provide the first experimental test of this hypothesis by examining whether the manipulated reproductive success of blue tits (Cyanistes caeruleus) is used as heterospecific inadvertent social information (ISI) in breeding-habitat selection by sympatric great tits (Parus major). The reproductive success of blue tits was manipulated 1year at the scale of patches by transferring nestlings from decreased to increased patches. No evidence was found of great tits using the reproductive success of blue tits as a source of heterospecific ISI. However, dispersal decisions by adult great tits correlated with information on con- and heterospecific densities, which constitute other sources of ISI. As density and breeding performance are tightly intertwined forms of information, the difficulty in distinguishing between them might lead great tits to use heterospecific ISI more in the form of density than breeding performance when making dispersal decisions.     

Spatial scale of local breeding habitat quality and adjustment of breeding decisions

Experimental studies provide evidence that, in spatially and temporally heterogeneous environments, individuals track variation in breeding habitat quality to adjust breeding decisions to local conditions. However, most experiments consider environmental variation at one spatial scale only, while the ability to detect the influence of a factor depends on the scale of analysis. We show that different breeding decisions by adults are based on information about habitat quality at different spatial scales. We manipulated (increased or decreased) local breeding habitat quality through food availability and parasite prevalence at a small (territory) and a large (patch) scale simultaneously in a wild population of Great Tits (Parus major). Females laid earlier in high-quality large-scale patches, but laying date did not depend on small-scale territory quality. Conversely, offspring sex ratio was higher (i.e., biased toward males) in high-quality, small-scale territories but did not depend on large-scale patch quality. Clutch size and territory occupancy probability did not depend on our experimental manipulation of habitat quality, but territories located at the edge of patches were more likely to be occupied than central territories. These results suggest that integrating different decisions taken by breeders according to environmental variation at different spatial scales is required to understand patterns of breeding strategy adjustment.     

Terrestrial habitat selection and strong density-dependent mortality in recently metamorphosed amphibians

To predict the effects of terrestrial habitat change on amphibian populations, we need to know how amphibians respond to habitat heterogeneity, and whether habitat choice remains consistent throughout the life-history cycle. We conducted four experiments to evaluate how the spatial distribution of juvenile wood frogs, Rana sylvatica (including both overall abundance and localized density), was influenced by habitat choice and habitat structure, and how this relationship changed with spatial scale and behavioral phase. The four experiments included (1) habitat manipulation on replicated 10-ha landscapes surrounding breeding pools; (2) short-term experiments with individual frogs emigrating through a manipulated landscape of 1 m wide hexagonal patches; and habitat manipulations in (3) small (4-m2); and (4) large (100-m2) enclosures with multiple individuals to compare behavior both during and following emigration. The spatial distribution of juvenile wood frogs following emigration resulted from differences in the scale at which juvenile amphibians responded to habitat heterogeneity during active vs. settled behavioral phases. During emigration, juvenile wood frogs responded to coarse-scale variation in habitat (selection between 2.2-ha forest treatments) but not to fine-scale variation. After settling, however, animals showed habitat selection at much smaller scales (2-4 m2). This resulted in high densities of animals in small patches of suitable habitat where they experienced rapid mortality. No evidence of density-dependent habitat selection was seen, with juveniles typically choosing to remain at extremely high densities in high-quality habitat, rather than occupying low-quality habitat. These experiments demonstrate how prediction of the terrestrial distribution of juvenile amphibians requires understanding of the complex behavioral responses to habitat heterogeneity. Understanding these patterns is important, given that human alterations to amphibian habitats may generate extremely high densities of animals, resulting in high density-dependent mortality.     

Intercontinental tests of the effects of habitat patch type on the distribution of chitons within and among patches in intertidal boulder field landscapes

Patchy distributions within landscapes may be caused by migration in response to different types of habitat patches. Intertidal boulder fields are landscapes in which boulders are discrete habitat patches, often with chitons attached to their under-surfaces. Chiton densities and associations with patch edges differed between boulders overlying coarse- versus fine-sediment types, with greater densities occurring over fine sediments. We tested whether adult migration caused between-boulder distributions by measuring immigration and emigration following experimental replacement of coarse sediments with fine sediment under boulders. We also assessed whether the manipulations altered chiton positions relative to patch edges, and large-scale generality was tested by including data from two continents. The manipulations did not influence the association of chitons with edges or amounts of emigration, but chitons did display positive density-dependent immigration that mirrored their distributional patterns, indicating the importance of immigration. Strikingly, all results were consistent between continents despite involving different species and even genera of chitons. By using boulder fields as a small-scale, easily manipulated landscape, we show that, even in sedentary organisms, patchy distributions within landscapes can be caused by migration alone, without the need to invoke mortality or larval recruitment.     

When to pay the cost of reproduction? A brood size manipulation experiment in great tits (<Emphasis Type="Italic">Parus major</Emphasis>)

Parental investment (PI) theory assumes that optimal parental investment is a function of expected cost/benefit ratio of current versus future reproduction. This study tests the prediction of PI theory that in a species with high adult mortality, paying of reproductive costs can be induced or prevented by manipulation of brood reproductive value (RV). Analysis of recruitment rates in a 10-year study of great tits (Parus major) showed that brood RV was affected by both fledgling number and quality (body mass and tarsus length). A 4-year brood size manipulation experiment (±2 hatchlings) resulted in reduction of fledgling quality in enlarged versus control and reduced broods, while the fledgling number did not differ between manipulation categories due to increase of nestling mortality with brood size manipulation score. Hence, the experiment resulted in reduction of brood RVs in enlarged broods. Females rearing enlarged broods survived better than those with control and reduced broods, while male survival was not affected by the experiment. This indicates that paying the survival costs for reproduction was a part of normal life history for studied female great tits, and that decisions whether to pay this cost were based on the estimation of brood RV. Recruitment rate was lowest (but extremely male-biased) in reduced broods, while control and increased broods did not differ in the number of locally recaptured offspring.     

From eavesdropping on performance to copying the behavior of others: a review of public information use

Public information (PI), a form of indirect social information, is used by individuals to estimate the quality of environmental parameters. It can be acquired in two ways. One way is by noting the performance of others. The other way it can be acquired is by noting the behavioral decisions of other individuals. Performance-based PI has been observed most often in the context of food and breeding patch estimation, as well as by individuals eavesdropping on contests between others in the context of assessing the fighting ability of opponents and the quality of mates. Evidence for the acquisition of PI from behavioral decisions of others comes mostly from studies of copying behavior, although recent work suggests that it also occurs when individuals estimate the tendency of others to be altruistic. PI use appears to be widespread across many taxa, although most work has been conducted on birds and fish. Absent from the literature are clear examples of PI use in mammals. The use of PI appears to often depend on its cost of acquisition and whether it contradicts an individual’s personal prior experience. PI can be an important benefit of associating with others.     

Ecological Trap for Desert Lizards Caused by Anthropogenic Changes in Habitat Structure that Favor Predator Activity

Abstract: Anthropogenic habitat perturbation is a major cause of population decline. A standard practice managers use to protect populations is to leave portions of natural habitat intact. We describe a case study in which, despite the use of this practice, the critically endangered lizard Acanthodactylus beershebensis was locally extirpated from both manipulated and natural patches within a mosaic landscape of an afforestation project. We hypothesized that increased structural complexity in planted patches favors avian predator activity and makes these patches less suitable for lizards due to a heightened risk of predation. Spatial rarity of natural perches (e.g., trees) in arid scrublands may hinder the ability of desert lizards to associate perches with low‐quality habitat, turning planted patches into ecological traps for such species. We erected artificial trees in a structurally simple arid habitat (similar to the way trees were planted in the afforestation project) and compared lizard population dynamics in plots with these structures and without. Survival of lizards in the plots with artificial trees was lower than survival in plots without artificial trees. Hatchlings dispersed into plots with artificial trees in a manner that indicated they perceived the quality of these plots as similar to the surrounding, unmanipulated landscape. Our results showed that local anthropogenic changes in habitat structure that seem relatively harmless may have a considerable negative effect beyond the immediate area of the perturbation because the disturbed habitat may become an ecological trap.     

Searching behavior and use of sampling information by free-ranging bison (Bos bison)

I examined the searching behavior of free-ranging plains bison (Bos bison bison) in their natural habitat, and determined whether their assessment of food patch quality was influenced by the short-term sampling information acquired during search. Bison used area-concentrated search during their winter foraging activity. Their movements between areas of suitable food patches were influenced by local environmental conditions, being sometimes less sinuous, and at other times more sinuous, than expected from a correlated random walk model. Bison also systematically avoided digging in areas where plants of low profitability lay under the snow. Where they dug, there was evidence that a bison's perception of food quality varied during a foraging bout, and was therefore influenced by short-term sampling information. After controlling for forage quality, I found that small feeding craters were more likely to be preceded by samples of high quality food patches. My observations suggest that bison take advantage of the structural characteristics of their environment during searching activity, and base foraging decisions on local rather than global availability.     

Great spotted cuckoo fledglings are disadvantaged by magpie host parents when reared together with magpie nestlings

The post-fledging period is a critical phase for juvenile survival, and parental care provided during this period is a key component of avian reproductive performance. Very little is known about the relationships between foster parents and fledglings of brood parasites. Here, we present the results of a 5-year study about the relationships between fledglings of the non-evictor brood parasitic great spotted cuckoo (Clamator glandarius) and its magpie (Pica pica) foster parents. Sometimes, great spotted cuckoo and magpie nestlings from the same nest can fledge successfully, but most often parasitic nestlings outcompete host nestlings and only cuckoos leave the nest. We have studied several aspects of cuckoo post-fledging performance (i.e. feeding behaviour, parental defence and fledgling survival) in experimental nests in which only cuckoos or both magpie and cuckoo nestlings survived until leaving the nest. The results indicate that great spotted cuckoo fledglings reared in mixed broods together with magpie nestlings were disadvantaged by magpie adults with respect to feeding patterns. Fledgling cuckoos reared in mixed broods were fed less frequently than those reared in only cuckoo broods, and magpie adults approached less frequently to feed cuckoos from mixed broods than cuckoos from only cuckoo broods. These results imply that the presence of host's own nestlings for comparison may be a crucial clue favouring the evolution of fledgling discrimination; and furthermore, that the risk of discrimination at the fledgling stage probably is an important selection pressure driving the evolution of the arms race between brood parasites and their hosts.     

From population sources to sieves: the matrix alters host-parasitoid source-sink structure

Field experiments that examine the impact of immigration, emigration, or landscape structure (e.g., the composition of the matrix) on the source sink dynamics of fragmented populations are scarce. Here, planthoppers (Prokelisia crocea) and egg parasitoids (Anagrus columbi) were released among host-plant patches that varied in structural (caged, isolated, or in a network of other patches) and functional (mudflat matrix that impedes dispersal vs. brome-grass matrix that facilitates dispersal) connectivity. Planthoppers and parasitoids on caged patches exhibited density-dependent growth rates, achieved high equilibrium densities, and rarely went extinct. Therefore, experimental cordgrass patches were classified as population sources. Because access to immigrants did not result in elevated population densities, source populations were not also pseudosinks, i.e., patches whose densities occur above carrying capacity due to high immigration. Planthoppers and parasitoids in open patches in mudflat had dynamics similar to those in caged patches, but went extinct in 4-5 generations in open patches in brome. Brome-embedded patches leaked emigrants at a rate that exceeded the gains from reproduction and immigration; populations of this sort are known as population sieves. For species whose suitable patches are becoming smaller and more isolated as a result of increased habitat fragmentation, emigration losses are likely to become paramount, a condition favoring the formation of population sieves. An increase in the proportion of patches that are sieves is predicted to destabilize regional population dynamics.     

Evaluating the effects of foraging habitat restoration on shorebird reproduction: the importance of performance criteria and comparative design

Coastal development and engineering projects preclude ecosystem processes that provide habitat for beach nesting birds. Management for coastal species may depend on actions that attempt to restore important habitat features and mitigate disturbance effects. However, species response to restoration or other management actions may be difficult to predict or measure. At Jones Beach State Park, on Long Island, New York, a 0.49 ha restoration project provided moist substrate foraging habitat for breeding Piping Plovers (Charadrius melodus) from 2002 to 2005. We examined whether foraging habitat restoration affected Piping Plover breeding population size, productivity, and fledgling production within 300 m of the restoration site. We found a positive relationship between habitat restoration and the number of fledglings produced per year. However, foraging habitat restoration did not significantly increase the number of Plover pairs breeding at Jones Beach. Our ability to evaluate restoration effects on Plovers depended on: 1) use of multiple performance criteria; 2) a design that allowed comparison of pre- and post-restoration data; and 3) a spatial control that allowed comparison of similar areas that were near and far from the restoration site. Despite the small size of the restoration project, there were measurable benefits to Plovers, indicating that foraging habitat restoration may be an effective tool for species recovery.     

Effects of Habitat Size and Patch Isolation on Reproductive Success of the Serpentine Morning Glory

Abstract: We examined the effects of habitat area and patch isolation on reproductive success in serpentine morning glory ( Calystegia collina [Convolvulaceae]), a primarily self-incompatible clonal plant endemic to serpentine outcrops in northern California's coast ranges. Within a 4000-km ² region, we compared the reproductive success of C. collina on 16 small (<5 ha) and 7 large ( >300 ha) outcrops. Flower and fruit production were significantly higher on large serpentine outcrops than on small outcrops. Fruit production also was positively correlated with the soil's ratio of calcium to magnesium. Successful pollination was positively affected by flower density and the number of other flowering patches within 100 m of a C. collina patch. The number of nearby flowering patches was considerably higher on large than on small outcrops. Flowers on large outcrops did not receive significantly more bee visitors than flowers on small outcrops, suggesting that pollination success is related to the quality rather than the quantity of pollen deposited. Fruit production by plants on both small and large outcrops was enhanced by the experimental addition of pollen from other patches, but not by the addition of pollen from the same patch. These findings demonstrate that the size of habitat may have strong effects on the reproductive success of locally endemic plants by enhancing opportunities for successful sexual reproduction. They also warn against the presumption that naturally patchy plant species are invulnerable to the effects of habitat fragmentation.     

Conservation of Fragmented Populations

In this paper we argue that landscape spatial structure is of central importance in understanding the effects of fragmentation on population survival. Landscape spatial structure is the spatial relationships among habitat patches and the matrix in which they are embedded. Many general models of subdivided populations make the assumptions that (1) all habitat patches are equivalent in size and quality and (2) all local populations (in the patches) are equally accessible by dispersers. Models that gloss over spatial details of landscape structure can be useful for theoretical developments but will almost always be misleading when applied to real-world conservation problems. We show that local extinctions of fragmented populations are common. From this it follows that recolonization of local extinctions is critical for regional survival of fragmented populations. The probability of recolonization depends on (1) spatial relationships among landscape elements used by the population, including habitat patches for breeding and elements of the inter-patch matrix through which dispersers move, (2) dispersal characteristics of the organism of interest, and (3) temporal changes in the landscape structure. For endangered species, which are typically restricted in their dispersal range and in the kinds of habitat through which they can disperse, these factors are of primary importance and must be explicitly considered in management decisions.     

Possible causes and consequences of philopatry and breeding dispersal in red-backed shrikes <Emphasis Type="BoldItalic">Lanius collurio</Emphasis>

Studies of animal breeding dispersal have often focused on possible causes, whereas its adaptive significance has received less attention. Using an information-theoretic approach, we assessed predictions of four hypotheses relating to causes and consequences of breeding dispersal in a migratory passerine, the red-backed shrike Lanius collurio. As predicted by the reproductive performance hypothesis, probability of breeding dispersal in females (though not in males) decreased with increasing annual average number of fledglings produced in the past year, but there was no association with conspecific reproductive performance in either sex. The site choice hypothesis, stating that individuals disperse to improve breeding site quality, received support in males only, as dispersal probability was positively associated to a measure indicating low territory quality. The social constraints hypothesis, referring to dispersal in relation to intraspecific interactions, received little support in either sex. The predation risk hypothesis was hardly supported either. Consequences of dispersal were marginal in both sexes because neither fledgling production in females, nor territory quality in males improved after dispersal. In addition, males settled on territories closer to the forest edge than those occupied predispersal, which is opposite to the prediction of the predation risk hypothesis. We conclude that own reproductive success was the major factor determining dispersal behavior in females, whereas territory quality and possibly predation risk were most important in males. Overall, breeding dispersal appeared not to be adaptive in this dense population inhabiting an optimal habitat.     

A Framework for Evaluating the Effects of Human Factors on Wildlife Habitat: the Case of Giant Pandas

Abstract: To address the complex interactions between humans and wildlife habitat, we developed a conceptual framework that links human factors with forested landscapes and wildlife habitat. All the components in the framework are integrated into systems models that analyze the effects of human factors and project how wildlife habitat would change under different policy scenarios. As a case study, we applied this framework to the Wolong Nature Reserve in Sichuan Province (southwestern China), the largest home of the giant panda ( Ailuropoda melanoleuca ). We collected ecological and socioeconomic data with a combination of various methods ( field observations, aerial photographs, government documents and statistics, interviews, and household surveys) and employed geographic information systems and systems modeling to analyze and integrate the data sources. Human population size has increased by 66% and the number of households in the reserve has increased by 115% since 1975, when the reserve was established. During the same period, the quality and quantity of the giant panda habitat dramatically decreased because of increasing human activities such as fuelwood collection. Systems modeling predicted that under the status quo, human population in the reserve would continue to grow and cause more destruction of the remaining panda habitat, whereas reducing human birth rates and increasing human emigration rates would lower human population size and alleviate human impacts on the panda habitat. Furthermore, our simulations and surveys suggested that policies encouraging the emigration of young people would be more effective and feasible than relocating older people in reducing human population size and conserving giant panda habitat in the reserve.     

Do fledglings choose wisely? An experimental investigation into social foraging behaviour

Many cooperative bird species have an extended period of post-fledging care. Despite the fact that this period of care can last up to several months, it remains a relatively understudied stage of chick development. This period, when young are actively begging but highly mobile, provides an opportunity for young to maximise the amount of care they receive by selectively choosing particular adults to beg from. In pied babblers Turdoides bicolor (a cooperatively breeding passerine), fledglings closely follow foraging adults and beg for food regularly (a behavioural interaction termed social foraging). Using a combination of natural observations and experimental manipulations, we found that fledgling pied babblers preferentially socially forage with adult care-givers who have high foraging success, since this results in young receiving more food. By supplementally feeding adults to artificially increase their foraging success, we increased the proportion of time that fledglings chose to socially forage with them, confirming that fledglings are selectively choosing dyadic interactions with the best adult foragers. These results indicate that pied babbler fledglings are sensitive to and can respond to short-term changes in adult foraging success, enabling them to maximize their nutritional intake, a behavioural adjustment that has long-term benefits in this system.     

Modeling post-fledging survival of Lark Buntings in response to ecological and biological factors

We evaluated the influences of several ecological, biological, and methodological factors on post-fledging survival of a shortgrass prairie bird, the Lark Bunting (Calamospiza melanocorys). We estimated daily post-fledging survival (n = 206, 82 broods) using radiotelemetry and color bands to track fledglings. Daily survival probabilities were best explained by drought intensity, time in season (quadratic trend), ages < or = 3 d post-fledging, and rank given drought intensity. Drought intensity had a strong negative effect on survival. Rank was an important predictor of fledgling survival only during the severe drought of 2002 when the smallest fledglings had lower survival. Recently fledged young (ages < or = 3 d post-fledging) undergoing the transition from nest to surrounding habitat experienced markedly lower survival, demonstrating the vulnerable nature of this time period. Survival was greater in mid and late season than early season, corresponding to our assumptions of food availability. Neither mark type nor sex of attending parent influenced survival. The model-averaged product of the 22-d survival calculated using mean rank and median value of time in season was 0.360 +/- 0.08 in 2001 and 0.276 +/- 0.08 in 2002. Survival estimates that account for age, condition of young, ecological conditions, and other factors are important for parameterization of realistic population models. Biologists using population growth models to elucidate mechanisms of population declines should attempt to estimate species-specific of post-fledging survival rather than use generalized estimates.     

Adaptive superparasitism and patch time allocation in solitary parasitoids : the influence of pre-patch experience

Summary Dynamic optimal diet models predict that host selection decisions and patch time allocation are influenced by the resource value of the habitat. We tested these predictions using the solitary' parasitoid Leptopilina heterotoma. Assuming that travel times between patches, the quality of previously visited patches and parasitoid density affect the parasitoids' estimation of the resource value of the habitat, different treatments were given before introducing parasitoids singly to a patch containing 5 unparasitized and 15 parasitized hosts. The decision to superparasitize is only slightly influenced by the rate of patch encounter. The quality of the previously visited patch has a strong influence. When a poor patch has been visited on the previous day, more superparasitism is observed in the partly depleted patch than when a rich patch has been searched. More superparasitism is also observed when the parasitoids are kept with conspecifics before the experiment than when they are kept alone. Increasing patch residence times are observed as the quality of the previously presented patch decreases. Host selection decisions and patch time allocation are thus clearly influenced by the pre-patch experience of the parasitoid, as predicted by dynamic optimal diet models. This can also explain why females that have never oviposited in unparasitized hosts will superparasitize readily. Correspondence to: M.E. Visser     

Pigment allocation to eggs decreases plasma antioxidants in a songbird

Although evidence is accumulating on the adaptive function of female ornamentation, very little is known about maternal allocation decisions involving sexual signaling and other reproductive functions. Blue egg coloration has been suggested as a sexually selected signal of female quality to males, and some recent studies are in accordance with this hypothesis. Blue eggshell coloration results from the deposition of biliverdin pigment by laying females, which is a potent antioxidant. Thus, egg pigmentation should be costly in terms of antioxidants, an assumption of the signaling hypothesis that has not been tested yet. We induced increased reproductive effort in a set of female pied flycatchers Ficedula hypoleuca through nest removal and measured egg pigmentation and plasma antioxidant levels in relation with a control group. Experimental females showed a negative association between egg color and plasma antioxidant levels, while there was no relationship for control birds. This supports that egg pigmentation is costly in terms of general antioxidant defenses and suggests a tradeoff between the allocations to both traits. Furthermore, experimental females with more colorful eggs raised more fledglings, especially when breeding early. Controls did not show a relationship between egg color and reproductive success. Females laying more colorful eggs could have shifted their allocation decisions towards current reproduction, at the expense of their own antioxidant defenses. Our results highlight that blue egg coloration is a life-history trait, subject to tradeoffs with other attributes, and seems to be especially informative under harsh breeding conditions.     

Short- and long-term consequences of prenatal testosterone for immune function: an experimental study in the zebra finch

Hormone-mediated maternal effects play an important role in the formation of a differentiated phenotype. They have been shown to influence a wide array of offspring traits, both early in life and in adulthood. One important offspring trait that is under the influence of maternal androgens is the immune system. In birds, a growing number of studies show that yolk androgens modulate immune function during the chick stage. However, there is a lack of knowledge regarding long-term effects of prenatal androgens on offspring immunity. In this study, we therefore investigated the influence of prenatal testosterone (T) on several measures of immunity in fledgling and adult zebra finches (Taeniopygia guttata). Cell-mediated immune response (towards phytohaemagglutinin, PHA) of fledglings hatching from control eggs was negatively related to brood size, whereas there was no such association for fledglings hatching from eggs with experimentally elevated T levels (T fledglings). Male control fledglings showed reduced mass gain compared to female control fledglings within 24 h after the PHA injection. This pattern was reversed in T fledglings. Total antibody levels in fledglings were not affected by egg treatment. Neither cell-mediated immunity nor total antibody levels in sexually mature zebra finches were influenced by egg treatment. However, there was an immuno-enhancing effect of elevated egg T on both primary and secondary humoral immune responses toward diphtheria and tetanus antigens in ca 5 and 7 month old zebra finches. In addition, the covariation between different immune components differed between T and control offspring, suggesting that egg treatment may have altered the potential trade-offs between different parts of the immune system. Our results suggest that prenatal androgens could be an important factor contributing to individual variation in immune function even in adulthood.