Individual differences in feeding efficiencies and feeding preferences of captive great tits

The preferences of eight, adult, hand-reared great tits for feeding on four different types of feeding sites have been tested. There were significant differences between individuals in these preferences, and the preferences for the different objects were correlated with the birds' individual feeding efficiencies on the objects. This correlation was thought to be a result of the birds' experience and learning during training for the experiment. During the tests the birds tended to make runs of visits to the same object type. This effect was more marked for less-preferred objects, and when the birds had just made a successful visit.     

Rank-dependent compromises between growth and predator protection in lizard dominance hierarchies

Juvenile Anolis aeneus lizards often live in dominance hierarchies, and in both the laboratory and field, rank was primarily determined by a juvenile's size relative to the others when it entered a hierarchy. Because juveniles entered hierarchies soon after hatching and left when they reached subadult sizes, most juveniles slowly rose in status during their sojourn in dominance hierarchies. In laboratory experiments, rank and growth rate were positively related when both food and perches were uniformly distributed. However, when perches were non-uniformly distributed, juveniles competed to live on those perch configurations which offered protection from predators. Under these conditions, growth was more weakly related to rank than in the uniform-habitat trials, and high-ranking juveniles living on the preferred (safe) perches had significantly lower growth rates than those living elsewhere. In contrast, low-ranking juveniles in non-uniform habitats experienced no such compromise between growth and safety, and those on the preferred perches tended to grow faster than those living elsewhere. Since most juveniles in the field seem to gradually rise from low to high rank, the behavioural strategies for optimizing growth and survival may change ontogenetically in this species.     

Rival imprinting: interspecifically cross-fostered tits defend their territories against heterospecific intruders

Failure to recognize conspecifics in social interactions such as mate choice and aggressive encounters will often result in reduced fitness. Studies on mate choice show that the ability to recognize conspecifics as mates is not universally present at birth, but often needs to be learned. In contrast, little is known about the ontogeny of intrasexual species recognition. To test whether learning influences the recognition of sexual rivals, we compared the aggressive response towards intruders of interspecifically cross-fostered individuals and controls reared by conspecific parents. We simulated territorial intrusion by presenting either a caged individual or playback song near the nest of breeding pairs of great tits, Parus major, and blue tits, P. caeruleus. Great tits reared by blue tit parents responded much more to blue tit stimuli than did great tit controls, and furthermore showed stronger responses to blue tit stimuli than to those of their own species. Blue tits reared by great tits responded much more to great tit stimuli than did blue tit controls. In contrast, blue tits cross-fostered to coal tits, P. ater, did not respond more to coal tits than did blue tit controls. There was a species difference in the response to conspecifics: blue tits cross-fostered to great tits responded more to conspecifics than did cross-fostered great tits. The results were similar for males and females. We conclude that learning influences intrasexual species recognition in these tits. Copyright 2003 Published by Elsevier Science Ltd on behalf of The Association for the Study of Animal Behaviour.      

Context-dependent visual preferences in starlings and blue tits: mate choice and light environment

There is considerable interest in the role that ultraviolet (UV) cues play in the foraging and mate choice decisions of birds. However, with the exception of the zebra finch, Taeniopygia guttata, it is not yet clear whether ultraviolet preferences are context specific, or whether birds show a general preference for full-avian-spectrum environments (320-700 nm) irrespective of the activity in which they are engaged. We investigated whether European starlings, Sturnus vulgaris, and blue tits, Parus caeruleus, show general (nonresource based) or context-specific preferences for full-spectrum environments. We found that neither species showed a general preference for UV-present (UV+) over UV-deficient (UV−) environments, when those environments contained no resources (experiment 1). Furthermore, neither species showed a UV+ preference when cages contained food, water and perches (starlings; experiment 2) or food, perches and heterospecifics (blue tits; Hunt et al. 1999. Animal Behaviour, 58, 809-815). However, both species did show highly significant preferences for UV+ conditions when viewing potential mates. Such experiments are necessary before one can conclude that particular wavebands have specific relevance to mate choice. In fact, our results suggest that the importance of particular wavelength compositions do indeed vary with behavioural context.     

Detection distance influences escape behaviour in two parids, Parus major and P. caeruleus

When birds are attacked by aerial predators they should benefit by adjusting their escape to the prevailing attack situation. One important factor likely to affect escape decisions of prey, to our knowledge not previously studied, is the distance at which the attacking predator is detected. We investigated if great tits Parus major and blue tits P. caeruleus alter their escape behaviour to two different detection distances (2.3 m and 1m) by simulating surprise attacks using a predator model. Both species used the information about detection distance when escaping by increasing the escape angle at the shorter detection distance. In addition, blue tits adjusted to the shorter detection distance by dodging sideways more frequently. Great tits escaped initially steeper and faster than blue tits, whereas blue tits increased escape angle and speed more than great tits along the measured distance after taking wing.     

Great tits (Parus major) flexibly learn that herbivore‐induced plant volatiles indicate prey location: An experimental evidence with two tree species

1. When searching for food, great tits (Parus major) can use herbivore‐induced plant volatiles (HIPVs) as an indicator of arthropod presence. Their ability to detect HIPVs was shown to be learned, and not innate, yet the flexibility and generalization of learning remain unclear.
2. We studied if, and if so how, naïve and trained great tits (Parus major) discriminate between herbivore‐induced and noninduced saplings of Scotch elm (Ulmus glabra) and cattley guava (Psidium cattleyanum). We chemically analyzed the used plants and showed that their HIPVs differed significantly and overlapped only in a few compounds.
3. Birds trained to discriminate between herbivore‐induced and noninduced saplings preferred the herbivore‐induced saplings of the plant species they were trained to. Naïve birds did not show any preferences. Our results indicate that the attraction of great tits to herbivore‐induced plants is not innate, rather it is a skill that can be acquired through learning, one tree species at a time.
4. We demonstrate that the ability to learn to associate HIPVs with food reward is flexible, expressed to both tested plant species, even if the plant species has not coevolved with the bird species (i.e., guava). Our results imply that the birds are not capable of generalizing HIPVs among tree species but suggest that they either learn to detect individual compounds or associate whole bouquets with food rewards.

     

Field and laboratory observations on the foraging and feeding techniques of blue tits (Parus caeruleus) and coal tits (P. ater) in relation to their habitats

I have tried to show (a) that results in the literature suggest that blue tits and coal tits survive better in their usual habitats of broad-leaved and conifer woods respectively; (b) that the higher mortality in the ‘wrong’ habitat may be attributed at least in part to an inability to acquire an adequate supply of food; (c) from my own and other people's field observations, that wild coal tits and blue tits use different feeding techniques; (d) from my own laboratory experiments, that each species is ‘best’ at the feeding techniques which it in general uses in the wild, and also that each species prefers to use the feeding techniques at which it is ‘best’; (e) that the feeding techniques of coal tits may be adapted to exploit the type of food supply found in conifers, while those of blue tits may be adapted for feeding in broad-leaved trees.     

Changes in the flocking behaviour of wintering English titmice with time,weather and supplementary food

Over the course of one winter, the food supply of birds living in a deciduous woodland in southern England was supplemented and unsupplemented during alternating periods. In the presence of substantial predation pressure from hawks, the sociality of blue tits, Parus caeruleus, and great tits, P. major, showed significant partial correlations with several weather and temporal factors when the woodland was unsupplemented. Such correlations between social behaviour and abiotic factors diminished significantly when the birds had access to extra food. Blue tits and great tits without access to supplemental food flocked significantly more often with other species while foraging than when they were food-supplemented. Long-tailed tits, Aegithalos caudatus, ignored the artificial food and foraged in mixed-species flocks to the same extent in both unsupplemented and food-supplemented periods. Results disprove the hypothesis that mixed-species foraging groups are caused by increased predation protection alone, and they support the hypotheses that mixed-species foraging groups are caused by increased foraging efficiency alone or by a combination of increased foraging efficiency and increased protection from predators.     

The relationship between social status and resting metabolic rate in great tits (Parus major) and pied flycatchers (Ficedula hypoleuca)

The oxygen consumption of great tits (Parus major) in winter flocks and of male pied flycatchers (Ficedula hypoleuca) in breeding condition, was measured. Resting metabolic rate was significantly correlated with dominance rank, as expressed by the width of the breast-stripe in the great tits and the darkness of the plumage on the head and back, or date of pair-formation, in males of the pied flycatcher. Individuals with the highest metabolic rates were the most dominant ones. Similarly, heart weight, relative to body weight, in dominant great tits was greater than that in lower-ranking birds.     

Geographic variation in bird songs: examination of the effects of sympatric related species on the acoustic structure of songs

Birdsong evolution has influenced by various ecological and social factors. When related species that sing similar songs coexist, the acoustic properties of the songs of one or both species may shift, and the songs may diverge. We investigated geographic variation in the songs of the Japanese tit (Parus minor) and the varied tit (Poecile varius) in the Ryukyu Archipelago, Japan, whose islands harbor either one or both species. The songs of the two species exhibited similar structure, but acoustic measurements differed between them. For example, varied tits sang songs at higher frequency than Japanese tits did. The songs of both species varied geographically. At sites with higher relative densities of varied tits, Japanese tits sang lower frequency songs, indicating that in areas of coexistence, Japanese tits sang songs that had acoustically diverged from those of varied tits. Song variation in varied tits was not related to sympatry with Japanese tits. These asymmetric results suggest that the subordinate Japanese tit modified the acoustic characteristics of its song to avoid harassment by the dominant varied tit. We observed no effects of genetic divergence or local intraspecific density on Japanese tit or varied tit songs. This study used geographic variation to examine hypotheses of song evolution, and the results highlight the importance of character displacement.     

Predicting demographically sustainable rates of adaptation: can great tit breeding time keep pace with climate change?

Populations need to adapt to sustained climate change, which requires micro-evolutionary change in the long term. A key question is how the rate of this micro-evolutionary change compares with the rate of environmental change, given that theoretically there is a ‘critical rate of environmental change’ beyond which increased maladaptation leads to population extinction. Here, we parametrize two closely related models to predict this critical rate using data from a long-term study of great tits (Parus major). We used stochastic dynamic programming to predict changes in optimal breeding time under three different climate scenarios. Using these results we parametrized two theoretical models to predict critical rates. Results from both models agreed qualitatively in that even ‘mild’ rates of climate change would be close to these critical rates with respect to great tit breeding time, while for scenarios close to the upper limit of IPCC climate projections the calculated critical rates would be clearly exceeded with possible consequences for population persistence. We therefore tentatively conclude that micro-evolution, together with plasticity, would rescue only the population from mild rates of climate change, although the models make many simplifying assumptions that remain to be tested.     

Does the temporal mismatch hypothesis match in boreal populations?

The temporal mismatch hypothesis suggests that fitness is related to the degree of temporal synchrony between the energetic needs of the offspring and their food supply. The hypothesis has been a basis in studying the influence of climate warming on nature. This study enhances the knowledge on prevalence of temporal mismatches and their consequences in boreal populations, and questions the role of the temporal mismatch hypothesis as the principal explanation for the evolution of timing of breeding. To test this, we examined if synchrony with caterpillar prey or timing of breeding per se better explains reproductive output in North European parid populations. We compared responses of temperate-origin species, the great tit (Parus major) and the blue tit (Cyanistes caeruleus), and a boreal species, the willow tit (Poecile montanus). We found that phenologies of caterpillars and great tits, but not of blue tits, have advanced during the past decades. Phenologies correlated with spring temperatures that may function as cues about the timing of the food peak for great and blue tits. The breeding of great and blue tits and their caterpillar food remained synchronous. Synchrony explained breeding success better than timing of breeding alone. However, the synchrony effect arose only in certain conditions, such as with high caterpillar abundances or high breeding densities. Breeding before good synchrony seems advantageous at high latitudes, especially in the willow tit. Thus, the temporal mismatch hypothesis appears insufficient in explaining the evolution of timing of breeding.     

Taking the Operant Paradigm into the Field: Associative Learning in Wild Great Tits

Associative learning is essential for resource acquisition, predator avoidance and reproduction in a wide diversity of species, and is therefore a key target for evolutionary and comparative cognition research. Automated operant devices can greatly enhance the study of associative learning and yet their use has been mainly restricted to laboratory conditions. We developed a portable, weatherproof, battery-operated operant device and conducted the first fully automated colour-associative learning experiment using free-ranging individuals in the wild. We used the device to run a colour discrimination task in a monitored population of tits (Paridae). Over two winter months, 80 individuals from four species recorded a total of 5,128 trials. Great tits (Parus major) were more likely than other species to visit the devices and engage in trials, but there were no sex or personality biases in the sample of great tits landing at the devices and registering key pecks. Juveniles were more likely than adults to visit the devices and to register trials. Individuals that were successful at solving a novel technical problem in captivity (lever-pulling) learned faster than non-solvers when at the operant devices in the wild, suggesting cross-contextual consistency in learning performance in very different tasks. There was no significant effect of personality or sex on learning rate, but juveniles’ choice accuracy tended to improve at a faster rate than adults. We discuss how customisable automated operant devices, such as the one described here, could prove to be a powerful tool in evolutionary ecology studies of cognitive traits, especially among inquisitive species such as great tits.     

Optimal trade-offs between competing behavioural demands in the great tit

We present a dynamic optimal control model to show how territorial male great tits (Parus major L.) should combine territorial vigilance with foraging. In the model, territory owners allocate time either to feeding in patches or travelling between them. Travelling is compatible with territorial vigilance, while feeding is not. Owners are assumed to adjust their patch residence and travel times in order to minimize the costs accruing from intruders on their territories, while at the same time reducing their hunger. The relation between patch residence and travel time matches that of Charnov's (1976) marginal value theorem while hunger is high, but as hunger is reduced the dynamic model predicts shorter patch residence times for any observed travel time. We present supporting evidence from an experimental test with captive male great tits.     

Prey selection and the search strategy of the spotted flycatcher (Muscicapa striata): A field study on optimal foraging

Flycatchers switched between different foraging strategies and selected prey so as to maximize energy intake. Changes in diet were influenced by the absolute abundance of the preferred prey and not by that of the alternative prey. Adults could recognize wasps (Vespula) and bees (Bombus) and removed their stings before swallowing them. The differences between adult and nestling diet and factors influencing capture success of prey are described. The flycatchers' behaviour in leaving perches and their choice of a ‘giving-up time’ was consistent with the view that they were maximizing the number of prey they caught in a given time. I conclude that flycatchers often search for and select prey on the basis of maximizing energy intake but I give reasons for not expecting them always to do so.     

Ultraviolet Radiation Influences Perch Selection by a Neotropical Poison-Dart Frog

Ambient ultraviolet-B radiation can harm amphibian eggs, larvae and adults. However, some amphibians avoid UV-B radiation when given the opportunity. The strawberry poison dart frog, Oophaga pumilio, is diurnal and males vocalize throughout the day in light gaps under forest canopies that expose them to solar radiation. Previous studies have demonstrated that males calling from high perches are more successful at mating than those at lower perches. We investigated whether frogs at higher perches receive more ultraviolet-B than those calling from lower perches. We also investigated whether frogs on perches receiving relatively low ultraviolet-B levels maintained their positions for longer compared to individuals calling from perches receiving higher levels of ultraviolet-B. Finally, since it has been hypothesized that some animals utilize levels of UV-A as a visual cue to avoid UV-B damage, we artificially elevated ultraviolet-A levels to examine whether males exposed to artificially elevated ultraviolet-A abandoned their perches sooner compared to males exposed to visible light. We found that frogs called from perches receiving low ultraviolet-B regardless of perch height, and that frogs maintain their positions longer on perches receiving low ultraviolet-B compared to perches receiving even slightly higher ultraviolet-B levels. Exposing the frogs to artificially elevated levels of ultraviolet-A radiation caused males to move off of their perches faster than when they were exposed to a control light source. These experiments suggest that ultraviolet radiation plays an important role in frog behavior related to perch selection, even in rainforests where much of the solar radiation is shielded by the forest canopy.     

A natural experiment identifies an impending ecological trap for a neotropical amphibian in response to extreme weather events

Extreme weather events are predicted to increase as a result of climate change, yet amphibian responses to extreme disturbance events remain understudied, especially in the Neotropics. Recently, an unprecedented windstorm within a protected Costa Rican rainforest opened large light gaps in sites where we have studied behavioral responses of diurnal strawberry poison frogs (Oophaga pumilio) to ultraviolet radiation for nearly two decades. Previous studies demonstrate that O. pumilio selects and defends perches where ultraviolet radiation (UV‐B) is relatively low, likely because of the lethal and sublethal effects of UV‐B. In this natural experiment, we quantified disturbance to O. pumilio habitat, surveyed for the presence of O. pumilio in both high‐disturbance and low‐disturbance areas of the forest, and assessed UV‐B levels and perch selection behavior in both disturbance levels. Fewer frogs were detected in high‐disturbance habitat than in low‐disturbance habitat. In general, frogs were found vocalizing at perches in both disturbance levels, and in both cases, in significantly lower UV‐B levels relative to ambient adjacent surroundings. However, frogs at perches in high‐disturbance areas were exposed to UV‐B levels nearly 10 times greater than males at perches in low‐disturbance areas. Thus, behavioral avoidance of UV‐B may not reduce the risks associated with elevated exposure under these novel conditions, and similarly, if future climate and human‐driven land‐use change lead to sustained analogous environments.     

Testing for effects of climate change on competitive relationships and coexistence between two bird species

Climate change is expected to have profound ecological effects, yet shifts in competitive abilities among species are rarely studied in this context. Blue tits (Cyanistes caeruleus) and great tits (Parus major) compete for food and roosting sites, yet coexist across much of their range. Climate change might thus change the competitive relationships and coexistence between these two species. Analysing four of the highest-quality, long-term datasets available on these species across Europe, we extend the textbook example of coexistence between competing species to include the dynamic effects of long-term climate variation. Using threshold time-series statistical modelling, we demonstrate that long-term climate variation affects species demography through different influences on density-dependent and density-independent processes. The competitive interaction between blue tits and great tits has shifted in one of the studied sites, creating conditions that alter the relative equilibrium densities between the two species, potentially disrupting long-term coexistence. Our analyses show that long-term climate change can, but does not always, generate local differences in the equilibrium conditions of spatially structured species assemblages. We demonstrate how long-term data can be used to better understand whether (and how), for instance, climate change might change the relationships between coexisting species. However, the studied populations are rather robust against competitive exclusion.     

Opposite differential allocation by males and females of the same species

Differential allocation (DA)—the adjustment of an individual''s parental investment in relation to its mate''s attractiveness—is increasingly recognized as an important component of sexual selection. However, although DA is expected by both sexes of parents in species with biparental care, DA by males has rarely been investigated. We have previously demonstrated a decrease in the feeding rates of female blue tits Cyanistes caeruleus when their mate''s UV coloration was experimentally reduced (i.e. positive DA). In this study, we used the same experimental protocol in the same population to investigate DA by male blue tits in relation to their female''s UV coloration. Males mated to UV-reduced females had higher feeding rates than those mated to control females (i.e. negative DA). Thus, male and female blue tits display opposite DA for the same component of parental effort (chick provisioning), the first time that this has been reported for any species.