Ecology

Biology, Ecology and Geography Fieldwork Courses in Barcelona, Spain.
Barcelona Field Studies Centre offers biology, ecology, geography and environmental field studies programmes throughout the year.

University of Tennessee, Ecology and Evolutionary Biology
The University of Tennessee, Knoxville was founded in 1794 and was designated the state land-grant institution in 1879.

Behavioral Ecology - recent issues
Behavioral Ecology - RSS feed of recent issues (covers the latest 3 issues, including the current issue)


Female mating preferences for male morphological traits used in species and m...

We tested whether sexually selected morphological traits in the sailfin mollies, Poecilia velifera and Poecilia petenensis, are also used in species recognition. Our first experiment, using live males as stimuli and providing females with olfactory as well as visual cues, found that females of both sailfin species preferred conspecific males to males of shortfin species. However, neither species preferred conspecific males when compared with heterospecific sailfin males, suggesting that premating reproductive isolation is not well developed between them. Our second experiment, providing females with only visual cues when distinguishing between live males, found that females of P. velifera preferred the larger of 2 stimulus males, regardless of whether the larger male was a conspecific or an heterospecific sailfin male. Such a preference for the larger sized male was not found in P. petenensis. To further investigate the role of the dorsal "sailfin" in species recognition, we used model males that varied only in the species identity of their dorsal fins. Females of both sailfin species preferred conspecific models with conspecific sailfins to those with dorsal fins of the shortfin species. In addition, females of P. velifera preferred the model with the largest sailfin, regardless of species identity. Similarly to the live male experiments, females of P. petenensis did not distinguish between conspecific and heterospecific sailfins. Overall, our study suggests that females of P. velifera have a generalized preference for larger males and that species-specific differences in sailfin shape do not lead to premating reproductive isolation between these 2 sailfin species.

Anthropogenic environmental change is escalating in magnitude, rate, and extent, inducing cascading effects across trophic levels. Assessing the nature of these alterations to trophic interactions requires an understanding of how species' demography and behavior are altered by simultaneous, complex pressures. For predator–prey relationships, "landscapes of fear" have been used to measure the trade-off prey animals make between maximizing energy gain and minimizing risk of predation. However, hierarchical foraging theory predicts that the degree to which aggregations of resources are used will depend upon the context in which they occur, not merely on the predation risk associated with those patches. We develop a conceptual framework that synthesizes theories of foraging hierarchies and landscapes of fear to show how predation risk and resource variation may interact to influence foraging behavior. We show, experimentally, that northern brown bandicoots (Isoodon macrourus), do respond to the likely predation risk when making their foraging decisions; however, the food resources in the habitat surrounding the food patch also play a significant role in the degree to which food patches are used. This result has important implications for the accuracy of assessments of landscapes of fear and habitat use using observations of animal foraging behavior.

Female color polymorphisms are common in the cichlid species radiations of Lake Victoria and Lake Malawi. According to theory, when a population harbors variation in sex-determining factors, polymorphism in female-linked coloration might generate individual variation in male mating preferences for female color morphs. We tested whether individual males exhibit consistent mating preferences for female color morphs in the Lake Malawi cichlid Pseudotropheus (Maylandia) ‘zebra gold’, a species polymorphic for female coloration and sex determination. We also explored whether male mating preferences could be predicted by maternal coloration or were acquired by imprinting on siblings' coloration. We found large individual variation in the strength and direction of male preferences for sex-linked female color patterns. Male mating preferences could be predicted by the mother's color morph and were not affected by visual imprinting. These findings represent the first evidence of male choice on sex-linked female coloration in a Lake Malawi cichlid. Our analysis indicates a strong genetic component to male preference for female coloration and large individual variation in the strength and direction of male mating preferences. Within-population variation in innate mating preferences might have important implications in cichlid fish species radiations.

Western sandpiper (Calidris mauri) chicks are precocial and leave the nest shortly after hatch to forage independently. Chicks require thermoregulatory assistance from parents (brooding) for 5–7 days posthatch, and parents facilitate chick survival for 2–3 weeks posthatch by leading and defending chicks. Parental vocal signals are likely involved in protecting chicks from predators, preventing them from wandering away and becoming lost and leading them to good foraging locations. Using observational and experimental methods in the field, we describe and demonstrate the form and function of parent–chick communication in the western sandpiper. We document 4 distinct calls produced by parents that are apparently directed toward their chicks (brood, gather, alarm, and freeze calls). Through experimental playback of parental and non–parental vocalizations to chicks in a small arena, we demonstrated the following: 1) chicks respond to the alarm call by vocalizing relatively less often and moving away from the signal source, 2) chicks respond to the gather call by vocalizing relatively more often and moving toward the signal source, and 3) chicks respond to the freeze call by vocalizing relatively less often and crouching motionless on the substrate for extended periods of time. Chicks exhibited consistent directional movement and space use to parental and non–parental signals. Although fewer vocalizations were given in response to non–parental signals, which may indicate a weaker response to unfamiliar individuals, the relative number of chick calls given to each type of call signal was consistent between parental and non–parental signals. We also discovered 2 distinct chick vocalizations (chick-contact and chick-alarm calls) during arena playback experiments. Results indicate that sandpiper parents are able to elicit antipredatory chick behaviors and direct chick movement and vocalizations through vocal signals. Future study of parent–offspring communication should determine whether shorebird chicks exhibit parental recognition though vocalizations and the role of chick vocalizations in parental behavior.

Much of the research examining the effects of ambient noise on communication has focused on adult birds using acoustic signals in mate attraction and territory defense. Here, we examine the effects of noise exposure on young birds, which use acoustic signals to solicit food from parents. We found that nestling tree swallows (Tachycineta bicolor) exposed to playbacks of white noise, within natural amplitude levels, from days 3 to 15 posthatch had begging calls with higher minimum frequencies and narrower frequency ranges than control nestlings raised in nests without added noise. Differences in begging call structure also persisted in the absence of noise. Two days after the noise was removed, experimental nestlings produced calls that were narrower in frequency range and less complex than control nestlings. We found no difference in growth between experimental and control nestlings. Our results suggest that long-term noise exposure affects the structure of nestling begging calls. These effects persist in the absence of noise, suggesting that noise may affect how calls develop.

The occurrence of multiple phenotypes within a sex of a single species has long puzzled behavioral ecologists. Male red-backed fairy-wrens Malurus melanocephalus exhibit 3 behaviorally distinct types in their first breeding season: breed in bright nuptial plumage, breed in dull plumage, or remain as an unpaired auxiliary (helper) with dull plumage. The retention of dull plumage by auxiliaries and dull breeders is an example of delayed plumage maturation (DPM), a widespread phenomenon in birds whose costs and benefits are not well understood. At a mechanistic level, DPM might allow dull males either to deceptively mimic females (female mimicry hypothesis) or to honestly signal their subordinate status (status-signaling hypothesis). DPM might function via either mechanism to provide ultimate benefits relative to developing nuptial plumage by increasing reproductive success, survival, or both. In this study, we tested the hypothesis that DPM is related to increased male survival in the red-backed fairy-wren via either female mimicry or status signaling. Aviary-based experiments revealed that dull males were perceived as male, which is consistent with the status-signaling hypothesis but contradicts the female mimicry hypothesis. Further aviary and field-based experiments also revealed that dull males were socially subordinate to bright males and received less aggression than bright males, further evidence for status signaling. However, male survival was not related to plumage coloration or breeding status. These findings indicate that male plumage coloration signals social status but that dull plumage does not afford a net survival advantage, perhaps because plumage color is a conditional strategy.

Males of many species can breed in distinct alternative phenotypes; for example, in many birds some males breed in dull plumage while others breed in bright plumage. Because females often appear to prefer brighter males, it is unclear why some males breed in dull plumage. Males in dull plumage might enjoy enhanced within-pair reproductive success if they can gain access to better breeding territories, or they might have relatively high extrapair reproductive success if they are better able to intrude on the territories of other males. To test these possibilities, we examined the reproductive consequences of plumage color in the red-backed fairy-wren (Malurus melanocephalus), a species in which males can breed in either bright plumage or dull plumage or serve as nonbreeding auxiliaries. Male plumage color was distributed bimodally and was loosely associated with age, such that some males molted into bright plumage a year or more earlier than others. Both male phenotypes were cuckolded at similar rates, but bright males sired significantly more extrapair young than did dull males, and this effect was independent of age. Thus, 1-year-old males who bred in dull plumage had low seasonal reproductive success compared with same-aged males who bred in bright plumage. These results suggest that males may not reap any fitness benefits by breeding in dull coloration, compared with breeding in bright plumage, but rather may be constrained to breed in suboptimal plumage by the timing of plumage acquisition.

Many animals bear colors and patterns to reduce the risk of predation from visually hunting predators, including warning colors, camouflage, and mimicry. In addition, various species possess paired circular features often called "eyespots," which may intimidate or startle predators preventing or postponing an attack. Most explanations for how eyespots work assert that they mimic the eyes of the predators own enemies. However, recent work has indicated that spots may reduce the risk of predation based purely on how conspicuous they are to a predator's visual system. Here, we use a field technique involving artificial prey marked with stimuli of various shapes, numbers, and sizes, presented to avian predators in the field, to distinguish between the eye mimicry and conspicuousness theories. In 3 experiments, we find that the features which make effective antipredator wing markings are large size and higher numbers of spots. Stimuli with circles survived no better than those marked with other conspicuous shapes such as bars, and changing the spatial construction of the spots to increase the level of eye mimicry had no effect on the protective value of the spots. These experiments support other recent work indicating that conspicuousness, and not eye mimicry, is important in promoting avoidance behavior in predators and that eyespots on real animals need not necessarily, as most accounts claim, mimic the eyes of other animals.

The benefits of prey grouping may be offset by increased detectability. With a focus on visual detectability, I investigated the potential costs of 2 traits, mine size and group size (number of mines per leaf), of a leaf-mining species, Antispila nysaefoliella (Lepidoptera: Heliozelidae), on the risk of visual detection by parasitoids and the ability to evade attack and capture after detection. Through field experimentation using artificial leaves and mines coated with a nontoxic adhesive spray for trapping insects, I found that the visual cues from groups of mines caused a significant increase in the number of parasitoids captured on experimental leaves (with artificial mines) relative to control leaves (without artificial mines). However, mine size did not. The observational evidence not only supports these patterns but also shows that the per capita risk of parasitism declines with large groups. These results provide evidence of a trade-off between avoiding visual detection and escaping attack after detection.

The theory of species coexistence predicts avoidance between species that compete for similar resources. Recent studies, however, have suggested that facilitation is also possible if competitor density provides information about resources. Optimal solution to trade-off between competition and facilitation is predicted to occur at intermediate competitor densities. We tested this hypothesis by experimentally creating a density range of resident tit species (Parus spp.), and measured the response of a competitively subordinate migratory bird, the collared flycatcher (Ficedula albicollis) in terms of habitat preference (settlement order and density), offspring investment (clutch size and primary sex ratio of offspring), and reproductive success (number and condition of nestlings). We show that most habitat choice and investment decisions of flycatchers were unimodally related to tit density, whereas reproductive success decreased linearly with increasing density. Flycatchers thus made mismatched investment decisions at the artificial tit densities because manipulation disassociated the natural correlation between habitat quality and population density. Apparently low and high tit densities were perceived as indication of poor quality habitat in terms of low amount or quality of resources/high mortality risk and high costs of competition, respectively. This demonstrates that competitor density can be used in assessing overall habitat quality in habitat selection and offspring investment decisions, integrating information on resources and competition.

Males and females often experience different optima in mating rate, which may cause evolution of female resistance to matings and male counter adaptations to increase mating rate. Males of the spider Pisaura mirabilis display a spectacular mating behavior involving a nuptial gift and thanatosis (death feigning). Thanatosis in a sexual context is exceptional and was suggested to function as an antipredation strategy toward potentially cannibalistic females. If thanatosis serves as a protection strategy, males should death feign in response to female aggression or when they are more vulnerable to attack. We tested these predictions in a factorial design: males that were handicapped (1 leg removed) and hence vulnerable and control males were paired with females that were more or less aggressive intrinsically (measured toward prey). In mating trials, we recorded the tendency of males to death feign, copulation success, and copulation duration. In addition, we investigated the effect of female mating status (virgin or mated) on these male mating components. Intrinsically aggressive females showed increased mating aggression toward males. Neither female aggressiveness, mating status, nor male vulnerability increased the propensity of males to perform thanatosis. Instead, death-feigning males were more successful in obtaining copulations and gained longer copulations. Hence, our results suggest that thanatosis functions as an adaptive male mating strategy to overcome female resistance. All males were capable of performing thanatosis although some males use it more frequently than others, suggesting a cost of death feigning which maintains the variation in thanatosis during courtship.

That individuals attempt to minimize the ratio of mortality risk/growth rate (µ/g) when foraging within individual habitat patches is well established. Do species partition among spatially discrete communities embedded in complex landscapes in a similar manner? We investigated how 3 ovipositing species (2 Hyla treefrogs and a hydrophilid beetle, Tropisternus lateralis) responded to simultaneous gradients of nutrients and predation risk. Species partitioned our experimental metacommunity primarily by reducing oviposition with fish. Tropisternus positively responded to increased nutrients, but the effect decreased with increasing risk, as predicted by µ/g theory. Use of fish habitats by Tropisternus was unrelated to breeding intensity. In contrast, Hyla showed no nutrient response but oviposited with fish only on nights with high breeding activity. Behavioral responses to the spatial distribution of resources and risk among discrete patches generated substantial variation in habitat-specific colonization rates, which has been identified as a primary mechanism generating both community and metacommunity structure.

Cooperatively breeding bell miners (Manorina melanophrys) have numerous male helpers assisting at multiple nests. Helpers are often related to the brood they aid, consistent with kin selection. However, there are also unrelated helpers for which other direct fitness benefits are likely to accrue. Bell miner nestlings can become infested by the larvae of a parasitic fly (Passeromyia indecora), which reduce growth and can be fatal. We investigated the amount of time that breeding pairs and helpers closely inspected nests and preened nestlings, behaviors apparently directed at detecting and removing parasites, a form of helping previously unstudied in a cooperative bird. Female breeders provided the greatest antiparasite effort, with breeding males and helpers not differing in effort regardless of their relatedness to the breeding female or brood. We also experimentally infested nests with nonparasitic flies and larvae. All individuals removed the introduced "parasites" if and when they encountered them. Compared with control sessions, inspection effort increased for all birds immediately after the experimental infestations, but only for a short, 5-min period. Further, we detected no changes in helper antiparasite behaviors after the temporary experimental removal of either breeding females or males. Such consistent helping behavior, independent of relatedness and potential audience effects, suggests that antiparasite behavior in bell miners is not particularly kin directed or operating as a signal of helper quality. Our results instead suggest that helper antiparasite effort appears to represent adaptive investment in the welfare of the brood, consistent with direct fitness benefits from group augmentation.

Previous studies indicate that division of labor can arise spontaneously in social groups. The comparison between normally social populations and forced associations of solitary individuals allows us to dissect the mechanisms by which tasks are distributed within a group and to ask how selection acts on division of labor during the incipient stages of sociality. In some ant species, newly mated queens form cooperative associations during nest initiation, in which individuals specialize on different tasks. The harvester ant Pogonomyrmex californicus shows geographical variation across populations in colony-founding strategies: solitary founding (haplometrosis) and group founding (pleometrosis). This system provides a unique opportunity to investigate how social context affects division of labor during social evolution. We created groups containing normally solitary, normally group founding, or mixed groups of solitary and social queens to examine how social phenotype affects division of labor. We also examined how group size affects task specialization by comparing pairs of queens with groups of 6 queens. Division of labor arose consistently across all associations. Groups of haplometrotic or pleometrotic queens differentiated into an excavation and a brood care specialist. In mixed groups, the haplometrotic queens took the role of excavator whereas the pleometrotic queens mainly tended brood. Our data also show that the intensity of specialization was greater in larger associations, consistent with current models of group size and division of labor. We discuss these data in the context of how emergence and selection act on the evolution of division of labor within incipient social groups.

Leaf-cutting ants culture a mutualistic fungus for which they collect and process a great diversity of fresh plant material as substrate. It has previously been observed that workers show "delayed rejection" behavior toward substrate that is harmful for the fungus but not for the ants: workers initially accept such materials but thereafter avoid its collection. In this study, we investigated delayed rejection behavior toward natural leaves in several 2-choice experiments in laboratory subcolonies of Acromyrmex lundi. We experimentally manipulated leaf suitability for the fungus by infiltrating them with a fungicide (cycloheximide) not detectable to the ants. The ants' delayed rejection behavior was specific toward the respective fungicide-treated plant species. Delayed rejection was also observed in naive ants after contact with the fungus garden containing treated leaves, confirming previous results with artificial bait. The onset of delayed rejection occurred 10 h after incorporation of treated leaves into the fungus garden. Rejection behavior was maintained for at least 9 weeks when incorporation of the previously unsuitable plant species was precluded. However, acceptance resumed after 3 weeks when ants were "forced" to feed on untreated leaves of the previously treated plant species. The observed species-specific, rapidly expressed, and flexible rejection of unsuitable substrate may be a mechanism to successfully avoid the provisioning of the fungus garden with plants containing harmful compounds as they occur in the highly diverse natural habitat of the colonies.