Phaeomelanin- and carotenoid-based pigmentation reflect oxidative status in two populations of the yellow warbler (Setophaga petechia)

Carotenoid- and phaeomelanin-based sexual pigmentation may signal a capacity to maintain oxidative balance and viability. However, diverse empirical results leave the association between pigmentation and oxidative stress (OS) unclear. We assessed the hypothesis that population-specific levels of oxidative challenge, or strategies for managing OS, affect relationships between sexual pigmentation and OS. Specifically, intense oxidative challenge in migratory, temperate breeding birds might enhance correlations between pigmentation and OS relative to allied tropical breeders, since quality-based differences in OS may arise only under intense oxidative challenge. Alternatively, in temperate breeders with intense within-season reproductive effort, high-quality birds may invest in reproduction over oxidative balance, dampening negative correlations between pigmentation and OS. To assess these alternatives, we compared prenesting relationships between pigmentation and OS in a migratory, Californian population of yellow warblers (Setophaga petechia brewsteri) and in a resident, Mexican population (Setophaga petechia bryanti, “mangrove warblers”). Yellow warblers displayed higher OS than mangrove warblers. However, year of capture and sex had bigger influences on correlations between pigmentation and OS than population. Males with more intense melanin pigmentation had lower OS among mangrove warblers and yellow warblers captured in 2011, but not among yellow warblers captured in 2012. In females only, lower OS levels were associated with more colorful carotenoid pigmentation. Results suggest that both phaeomelanin- and carotenoid-based pigmentation have the potential to correlate with OS levels, but that the signaling potential of pigmentation may shift with inter-annual variation in environmental conditions and display sex-specific dynamics.     

Responses of female lizards,<Emphasis Type="Italic"> Lacerta monticola</Emphasis>, to males' chemical cues reflect their mating preference for older males

Female mate choice based on visual traits appears to be rare in lizards. Field observations suggest that females of the lizard Lacerta monticola preferred to mate with larger/older males. Although older males are usually green and larger, and younger males brown and smaller, there is some overlap in size and coloration between age classes. Thus, visual cues may not always be reliable indicators of a male's age. We hypothesized that female mate-choice preferences may be based on males' pheromones, which might transmit information about characteristics such as age. In a laboratory experiment, we analyzed the effect of age of males on attractiveness of their scents to females. When we offered scents of two males of different age, females associated preferentially with scents of older males. This suggested that females were able to assess the age of males by chemical signals alone, and that females preferred to be in areas scent-marked by older males. Thus, females may increase their opportunities to mate with males of high quality, or may avoid harassment by sneaking young males. This result agreed with field observations on females mating with old males, and rejection of advances by young males. Our results also suggested that female preference for older males may depend on their own body size. Large females showed a strong preference for older males, whereas smaller females were not so selective. This, together with males' preference for large females, might lead to size-assortative matings. We suggest that the quality and/or quantity of male pheromones could communicate to the female heritable male genetic quality (i.e. age) and thereby serve as the basis of adaptive female choice in lizards.Communicated by W.E. Cooper     

Offspring sex ratios reflect lack of repayment by auxiliary males in a cooperatively breeding passerine

The repayment hypothesis posits that primary sex ratios in cooperative species should be biased towards the helping sex because these offspring “repay” a portion of their cost through helping behavior and therefore are less expensive to produce. However, many cooperatively breeding birds and mammals do not show the predicted bias in the primary sex ratio. Recent theoretical work has suggested that the repayment hypothesis should only hold when females gain a large fitness advantage from the presence of auxiliary adults in the group. When auxiliaries provide little or no fitness advantage, competition between relatives should lead to sex ratios biased towards the dispersing (non-helping) sex. We examined the benefits auxiliaries provide to females and corresponding offspring sex ratios in the red-backed fairy-wren (Malurus melanocephalus), a cooperatively breeding Australian bird with male auxiliary helpers. We found that auxiliaries provide little or no benefit to female reproductive success or survival. As predicted, the population primary sex ratio was biased towards daughters, the dispersing sex, and females with auxiliaries produced female-biased broods whereas females without auxiliaries produced unbiased broods. Moreover, offspring sex ratios were more strongly biased toward females in years when auxiliaries were more common in the population. These results suggest that offspring sex ratios are associated with competition among the non-dispersing sex in this species, and also that females may use cues to assess local breeding opportunities for their offspring.     

Differential use of conspecific-derived information by sexual and asexual parasitic wasps exploiting partially depleted host patches

When foraging partially depleted patches (i.e., a fraction of hosts are already parasitized), female parasitoids must decide: 1—whether to superparasitize, and 2—whether to stay in their current patch (thus missing the opportunity of finding a better patch elsewhere). To make these decisions, parasitoids may rely on different cues, produced both by the environment and by conspecifics. Animals thriving in different environments may differ in cues they use. In the solitary parasitoid Venturia canescens, thelytokous (asexual) and arrhenotokous (sexual) individuals are found in two contrasting environments. Thelytokous females, from anthropogenic conditions, are known to cope with superparasitism in an adaptive way. On the other hand, little is known about superparasitism by arrhenotokous females. We compared the host exploitation strategies of thelytokous and arrhenotokous females in partially depleted patches. Hosts parasitized by thelytokous females were more frequently avoided than those parasitized by arrhenotokous females, suggesting a stronger chemical marking of the former. Only thelytokous females used information from conspecifics for patch-leaving decisions. The conformity of the differences in the behavior of thelytokous and arrhenotokous females with the environmental conditions they experience in their habitat is discussed.     

Experimental analysis of flow and turbulence in the wake of neighboring emergent vegetation patches with different densities

Environmental Fluid Mechanics - Patches of vegetation in natural water bodies grow close to each other and may affect each other’s wake pattern with significant implications on nutrient...     

Food or sex—males and females in a sex role reversed pipefish have different interests

In a sex role reversed pipefish, Syngnathus typhle, we found that basic life history allocations were directly influenced by sexual selection. We investigated time allocation to foraging and mating, respectively, in a choice experiment, giving males and females, of small or large body size, a choice between food and a potential partner. We found that males were more interested in foraging than mating, i.e., were more frequently observed in front of the food than in front of the partner, whereas females were more interested in the potential partner. This reflects sexual selection operating differently on the two sexes, as males and females are relatively similar in other life history traits, such as growth, mortality, age of maturity, dispersal, and parental expenditure. Moreover, large individuals allocated more time to mating activities, small to feeding. Individuals more interested in mating compared to food were subsequently more critical when given a choice between a large (high-quality) and a small (low-quality) partner, whereas individuals more interested in food were not selective. These findings are consistent with our predictions: sex-role reversed males can be relatively sure of achieving one or more matings, and should allocate more time to feeding and, hence, to parental investment, growth and/or future reproduction. Females, on the other hand, have more uncertain mating prospects and should allocate time to imminent reproductive activities, thereby foregoing other life history traits such as growth and future egg production. By this, they also sacrifice future fecundity and attractiveness.     

Armaments and ornaments in the rock sparrow: a possible dual utility of a carotenoid-based feather signal

Females often base their mating preferences on male sexual secondary traits that are used to settle contests among males. Such traits are likely to be honest indicators of male quality if they are constantly used during costly male–male agonistic interactions. Carotenoid signals have been shown to work as a handicap because they are costly to produce. However, the role of carotenoids as “honest” signals during male contests is less clear, and it is not known whether a carotenoid-based trait can serve in both male–male competition and female choice. In this study, we studied the dual function of a carotenoid feather ornament in the rock sparrow (Petronia petronia), a bird species in which both sexes have a yellow throat patch whose size positively correlates with phenotypic measures. First, we investigated, in a field study, whether the size of a male’s yellow patch correlates with his ability to acquire a territory. Second, we tested the signal function of the yellow patch in two male–male interaction in captivity experiments. Finally, we measured female preference for males differing in throat patch size in a mate choice experiment. Our experiments revealed that the size of a male’s throat patch positively correlated with the number of nest boxes he was able to defend. Moreover, in controlled conditions, males with relatively large yellow patches had earlier access to food than those with small patches. Also, in an experiment in which a dummy rock sparrow with an experimentally manipulated yellow patch was positioned near a feeder, latency to feed by focal birds positively correlated with dummy patch size. Lastly, in a dichotomous mate choice experiment, females showed a proximity preference for males whose patch was experimentally enlarged. Taken together, these results suggest that the same carotenoid feather signal may be used in both male–male competition and female choice in this passerine bird.     

Does colour matter? The importance of colour in avoidance learning, memorability and generalisation

Aposematic species exploit the ability of predators to associate, for example, conspicuous colouration with the unprofitability of prey. We tested the importance of colour for avoidance learning, memory and generalisation in wild-caught great tits (Parus major). First, we determined the birds’ initial colour preferences for red, yellow, orange and grey artificial prey items. The birds showed some preferences, as they were more willing to eat grey prey as their first choice, but these were not strong preferences. We then trained birds to discriminate red, yellow or variable (red and yellow) signals from grey where colours signalled palatable and unpalatable food. In general, the birds learned the discrimination task equally well, irrespective of which colours signalled unpalatability, and subsequently remembered the distinction between previously palatable and previously unpalatable colours in the memorability test. We did not find strong evidence that variability in the signal affected learning or memory. Our results suggest that, in a task where birds must discriminate between palatable and unpalatable prey, it does not matter which specific colour signals unpalatability, although this might be context-dependent. To study whether training also affects responses to unconditioned stimuli, we included orange prey items in the memorability test. Although orange had been palatable in the initial preference test, the birds ate fewer orange prey items after they had been trained to avoid red, yellow or both colours (variable signal) as unpalatable prey, but did not change their preference when trained that these colourful signals were palatable. This indicates that generalisation occurred more readily after a negative experience than a positive experience, a situation that would potentially allow imperfect mimicry to occur.     

Different habitats, different habits? Response to foraging information in the parasitic wasp Venturia canescens

The parasitic wasp, Venturia canescens (Gravenhorst) (Hymenoptera: Ichneumonidae), has two reproductive modes, namely, thelytoky or arrhenotoky, and occurs in habitats with highly variable or relatively stable host abundances, respectively. Since information processing is costly, we expected that information indicating resource availability would be mainly used by the thelytokous wasps and less so by the arrhenotokous type. This idea was explored by two different approaches. In a study on patch-time allocation, we used females from ten populations and measured patch-residence times of individuals that visited multiple patches at different encounter rates. In a more detailed approach, thelytokous and arrhenotokous females from a single location were observed continuously while foraging, and all behaviors were recorded. Wasps of both reproductive modes (i.e., both habitat types) used information for the assessment of habitat quality. However, the way that the information was used differed between them. Whereas thelytokous females used foraging information to maximize their efficiency at high patch-encounter rates, arrhenotokous females merely reduced the number of offspring produced without changing patch times. The behavior of the arrhenotokous females should result in a spreading of offspring across the habitat and, thus, reduced sib-mating. The foraging strategy of these wasps might therefore be an adaptation to reduce costs associated with inbreeding.     

Roles of male residence and relative size in the social behavior of Iberian rock lizards, Lacerta monticola

Many studies on contest competition used residency asymmetry as a discrete variable. However, the probability of winning an interaction may change as a continuous function of the value of the location where the encounter occurs. We performed a field study to examine the importance of location within a home range and relative body size to the outcomes of agonistic interactions between male lizards, Lacerta monticola. The distances to activity centers (the most used locations based on a density function of sightings) and relative size play important roles in agonistic interactions and had interacting effects in natural conditions. On the other hand, previous studies with lizards suggested that inferior competitors are able to avoid agonistic interactions in the field. Thus, we staged encounters in the laboratory to examine the behavioral responses of smaller individuals. The responses of each focal smaller male were measured in its own home cage (resident), in the cage of a larger male (intruder) and in a cage in which no male was previously present (control). The predominant behavioral tactics of smaller males were avoidance when they are the intruders and displaying when they are the residents. Submissive displays by smaller males may help reduce the costs of agonistic encounters.     

Individual dispersal status influences space use of conspecific residents in the common lizard, Lacerta vivipara

The effects of immigration on the behaviour of residents may have important implications for the local population characteristics. A manipulative laboratory experiment with yearlings of the common lizard (Lacerta vivipara) was performed to test whether the introduction of dispersing or philopatric individuals influences the short-term spacing behaviour of resident individuals. Staged encounters were carried out to induce interactions within dyads. The home cage of each responding individual was connected by a corridor to an unfamiliar “arrival cage” to measure the latency to leave their own home cage after each encounter. Our results showed that the time that pairs spent in close proximity was longer when a dispersing individual was introduced in the home cage. The latency to leave the home cage was longer after the introduction of a dispersing individual. These response variables were not influenced by the relative body sizes of contestants nor by the level of aggression towards each other. In contrast, the aggressive response was significantly influenced by the residency asymmetry established experimentally (“owner” of the home cage vs introduced individual). Our results suggest that the space use by resident individuals is influenced by the dispersal status of conspecifics. The potential ultimate causes driving this effect are discussed.     

Heritability of the algal-foraging ability: an indirect benefit of female mate preference for males' carotenoid-based coloration in the guppy, Poecilia reticulata

Several sexual selection theories assume certain benefits of female mate preference. The direct benefit, i.e., the direct contribution from males to their offspring and females, has been well tested empirically. However, the indirect benefit, i.e., the male's genetic contribution to their offspring, has been poorly demonstrated. Female preference for males' carotenoid-based coloration is known in some animals. Since animals must acquire carotenoids through foods, it is often hypothesized that the brightness of the carotenoid-based coloration is a reliable indicator of the male's foraging ability. Hence, females' indirect benefits, such as greater foraging ability in their offspring, through mate preference for the carotenoid-based coloration are assumed. However, the heritability of the foraging ability for foods that serve as carotenoid resources has not been tested. In this study, a maze experiment was performed in guppies (Poecilia reticulata) to examine the heritability of the foraging ability for algae, carotenoid resources in nature. The latency for completing algal-foraging tasks in this experiment showed high individual variation. Heritable estimates of the foraging ability were substantial (h ² = 0.57 – 0.66) and significant, suggesting a genetic contribution to the foraging ability from fathers to their offspring. This result may support the hypothesis that indirect benefits influence the evolution of female choice.     

Conspicuous displays in cryptic males of a polytypic poison-dart frog

The evolution of aposematism is linked to increased opportunities for conspicuous sexual displays since detection by potential predators is no longer disadvantageous. Therefore, phenotypic divergence in aposematic species leading to relatively cryptic forms is expected to constrain such opportunities, by restoring the trade-off between natural and sexual selection on the boldness of sexual displays. We asked if and how a derived phenotype of the poison-dart frog Oophaga granulifera that appears relatively cryptic to potential predators exhibits conspicuous sexual displays for potential mates. We used visual modeling of frog contrasts against their natural backgrounds to test if for conspecifics green frogs appear less conspicuous than red frogs as they do for birds. We conducted behavioral observations of focal red and green males to determine if green frogs adjust their display behavior to the availability of potential mates. Dorsal brightness is known to influence female preferences in at least one poison frog species. We found that, despite being less visible under some measures, green frogs may appear as bright as red frogs for conspecifics but not birds, when viewed on dark backgrounds. Additionally, green males called more intermittently than red males when advertising to distant females, but they exhibited a dramatic increase in calling activity in proximity of a female and were as active as red males in this context. Together, our results suggest that green frogs retain context-dependent conspicuousness to conspecifics despite the evolution of relative crypsis to potential predators.     

Reproductive success of parasitized males in a marine reef fish

Parasitism is hypothesized to reduce reproductive success in heavily parasitized males because females may preferentially mate with less parasitized males (parasite-mediated sexual selection) or parasites may compromise male competitiveness. In marine systems, this hypothesis is largely unexplored. This paper provides the first confirmed record of a copepod ectoparasite (Caligus buechlerae Hewitt 1964) on the common triplefin (Forsterygion lapillum) and evaluates the hypothesis that males parasitized with C. buechlerae experience lower reproductive success than unparasitized males (as determined by the presence and area of eggs within male nests). We found that 38 % of males we surveyed were infected with at least one C. buechlerae, with a median of two individuals per infected male. About 32 % of males were defending eggs, with 62.5 % of those males infected with at least one parasite. Males of greater total length (TL) were both more likely to be infected and more likely to be defending eggs. However, when statistically accounting for the effects of TL, parasite infection had no effect on the probability of defending eggs, or the average surface area of eggs when present. Positive covariation in fish length, the presence of eggs and parasite infection observed here potentially suggest that the importance of parasitic infection on reproductive success may depend upon the strength of selection for larger male body size. Our study is one of the few studies to investigate the effects of ectoparasites on reproductive success in reef fish and also provides a quantitative measure of infection for a widespread species within New Zealand.     

The value of small size: loss of forest patches and ecological thresholds in southern Madagascar.

Many services generated by forest ecosystems provide essential support for human well-being. However, the vulnerability of these services to environmental change such as forest fragmentation are still poorly understood. We present spatial modeling of the generation of ecosystem services in a human-dominated landscape where forest habitat patches, protected by local taboos, are located in a matrix of cultivated land in southern Madagascar. Two ecosystem services dependent on the forest habitats were addressed: (1) crop pollination services by wild and semidomesticated bees (Apoidea), essential for local crop production of, for example, beans, and (2) seed dispersal services based on the presence of ring-tailed lemurs (Lemur catta). We studied the vulnerability of these ecosystem services to a plausible scenario of successive destruction of the smallest habitat patches. Our results indicate that, in spite of the fragmented nature of the landscape, the fraction of the landscape presently covered by both crop pollination and seed dispersal services is surprisingly high. It seems that the taboo system, though indirectly and unintentionally, contributes to upholding the generation of these services by protecting the forest patches. Both services are, however, predicted to be very vulnerable to the successive removal of small patches. For crop pollination, the rate of decrease in cover was significant even when only the smallest habitat patches were removed. The capacity for seed dispersal across the landscape displayed several thresholds with habitat patch removal. Our results suggest that, in order to maintain capacity for seed dispersal across the landscape and crop pollination cover in southern Androy, the geographical location of the remaining forest patches is more crucial than their size. We argue that in heavily fragmented production landscapes, small forest patches should increasingly be viewed as essential for maintaining ecosystem services, such as agricultural production, and also should be considered in the ongoing process of tripling the area of protected habitats in Madagascar.     

Allometric scaling predicts preferences for burned patches in a guild of East African grazers

The high herbivore diversity in savanna systems has been attributed to the inherent spatial and temporal heterogeneity related to the quantity and quality of food resources. Allometric scaling predicts that smaller-bodied grazers rely on higher quality forage than larger-bodied grazers. We replicated burns at varying scales in an East African savanna and measured visitation by an entire guild of larger grazers ranging in size from hare to elephant. We found a strong negative relationship between burn preference and body mass with foregut fermenters preferring burns to a greater degree than hindgut fermenters. Burns with higher quality forage were preferred more than burns with lower quality forage by small-bodied grazers, while the opposite was true for large-bodied grazers. Our results represent some of the first experimental evidence demonstrating the importance of body size in predicting how large herbivores respond to fire-induced changes in plant quality and quantity.     

Site fidelity and territorial movements of males in a rapidly declining population of yellow-headed blackbirds

We tested several hypotheses to explain low between-year territory fidelity in a breeding population of yellow-headed blackbirds (Xanthocephalus xanthocephalus). During a 5-year study the population of territorial males declined by two-thirds and some of the marshes that supported territories significantly deteriorated. Individual males held territories and bred for an average of 1.9 years. Of males that bred for at least 2 years, 30% skipped owning a territory in the study area during at least 1 year of their breeding lifetimes. Our information suggests that they may have bred outside of the area in those years. Of males with territories in two or more breeding seasons, 60% changed breeding marshes at least once. Males changed territories during 42.9% of between-year opportunities to do so. We found no support for the hypotheses that male yellow-headed blackbirds: (1) are more likely to move when territory density is low; (2) are likely to abandon territories that are deteriorating; or (3) change territories to improve their reproductive success. We suggest three non-mutually exclusive explanations for the yellow-headed blackbird's weak site fidelity: (1) it is a response to habitat deterioration and to other factors that may be causing the population's decline; (2) the males, being migratory, make fresh settlement decisions each year after they arrive on the breeding grounds in the general vicinity of their previous year's breeding; (3) yellow-headed blackbirds may have evolved in, and be adapted to, highly unstable habitats, moving frequently in response to changes in local breeding site conditions. Correspondence to: L.D. Beletsky