Male care,mate switching,and future reproductive success in a double-brooded passerine

Summary In species with male parental care, polygyny may reduce the parental effort provided by a male, and hence impose a cost on the fitness of his mates, because of decreased growth, survival or health of offspring. I examined a cost of polygyny in the green dart-poison frog, Dendrobates auratus, a species with male parental care in which both male polygyny and mate guarding by females occurs (Summers 1989). All D. auratus males seen carrying tadpoles in a marked area were followed and the pools where they deposited their tadpoles were recorded. Males frequently deposited more than one tadpole in the same pool (in 25% of the observed depositions a male deposited a tadpole in a pool where he had previously deposited at least one other tadpole). Experiments involving manipulation of tadpole densities in pools typically utilized by D. auratus (calabash husks and treeholes) showed that increasing tadpole number had a strong negative effect on both growth rate and survivorship, indicating that polygyny can impose a cost on the fitness of a male's offspring. Hence, females do face a potential cost, in terms of reduced offspring growth and survivorship, when their mates mate polygynously and care for the offspring of other females.     

Fitness consequences of dispersal decisions for male mountain gorillas (Gorilla beringei beringei)

The fitness consequences of dispersal decisions are difficult to quantify, especially for long-lived species with complex social systems. To calculate those consequences for male mountain gorillas, from the perspective of both subordinate and dominant males, we used behavioral and demographic data obtained over 30 years from the Virunga Volcano population to develop an agent-based model that simulates the life history events, social structure, and population dynamics of the species. The model included variables for birth rates, mortality rates, dispersal patterns, and reproductive skew. The model predicted an average lifetime reproductive success (LRS) of 3.2 for philopatric males (followers) and 1.6 for emigrants. The benefits of philopatry were most sensitive to opportunities for social queuing and to female transfer preferences, but philopatry remained the best strategy over a wide range of group conditions and hypothetical simulations. The average LRS for dominant males was 4.5 when a subordinate stayed and 4.6 when the subordinate emigrated. The dispersal decision of the subordinate male had little impact on the fitness of the dominant male because it came relatively late in the dominant males reproductive life span, and it changed his group composition only incrementally. The fitness consequences for the dominant male were most sensitive to the degree of reproductive skew. Since subordinates suffer a fitness loss when they leave a group, they should accept whatever reproductive restraint is needed to avoid eviction, and the dominant male does not need to offer concessions for them to stay. The dominant male may offer reproductive concessions for other reasons, such as peace incentives or to confuse paternity, or he may not have complete control of reproduction within his group.Communicated by D. Watts     

Fitness consequences of choosy oviposition for a time-limited butterfly

For the majority of insects, a female's choice of oviposition site(s) greatly influences both the success of individual offspring and her own total fitness. Theory predicts that females most strongly limited by egg number will employ greater oviposition site discrimination than those predominately subject to time limitation. The reproductive success of the butterfly Pieris virginiensis at our Connecticut, USA, field site is strongly time constrained on two fronts. First, during their three-week flight season, only 60% of days and 28% of daytime hours were suitable for flight. Second, larval survival is impacted by the rapid senescence of their spring ephemeral host plant Dentaria diphylla, with eggs laid during the first half of the flight season having approximately three times the survival chance of those laid later. Yet, on average, females choose to oviposit on only half the plants they closely inspect and fly over most ramets without any inspection. Our experiments demonstrate that the preferred host ramets confer an approximate two-fold survival advantage. Females are not choosing plants that senesce later, despite the advantage that such plants would confer. We use empirical data on female behavior and larval performance to parameterize a simulation model. Model results suggest that, despite the notable time limitation in this system, the observed level of female oviposition site preference not only increases individual larval survival, but also total female fitness. Low egg loads in this species may contribute to selection for strong host plant discrimination.     

Fitness consequences of body-size-dependent host species selection in a generalist ectoparasitoid

In insect parasitoids, offspring fitness is strongly influenced by the adult females choice of host, particularly in ectoparasitoids that attack non-growing host stages. We quantified the fitness consequences of size-dependent host species selection in Dirhinus giffardii, a solitary ectoparasitoid of tephritid fruit fly pupae. We first showed a positive correlation between the size of emerged D. giffardii wasps and the size of their host fruit fly species (in order of decreasing size): Bactrocera latifrons, B. cucurbitae, B. dorsalis or Ceratitis capitata. We then manipulated individual wasps to show that the parasitoid preferred to attack the largest (B. latifrons) to the smallest (C. capitata) host species when provided with a choice, and laid a greater proportion of female eggs in B. latifrons than in C. capitata. There were no differences in developmental time or offspring survival between individuals reared from these two host species. Finally, we compared the foraging efficiency of large versus small wasps (reared from B. latifrons vs C. capitata) under two different laboratory conditions: high versus low host habitat quality, given that realized fecundity in parasitoids may be influenced by either egg-limited or time-limited factors. Under both conditions, large wasps parasitized more hosts than did small ones as a consequence of high searching efficiency in the host-poor habitat, and high capacity for adjusting egg maturation in response to host availability in the host-rich habitat. Considering the flexibility of body growth, the apparent lack of cost of achieving large body size in either development or survival, and the strong dependence of realized reproductive success on a females size, we argue that body size may be a key to understanding evolution of host species selection in ectoparasitoids. We also discuss constraints upon the evolution of size-dependent host species selection in parasitoids.Communicated by D. Gwynne     

Fitness consequences of prolonged copulation in the bowl and doily spider

Summary Bowl and doily spiders, Frontinella pyramitela (Araneae, Linyphiidae), copulate much longer than the 15 min required for insemination. This apparently maladaptive behavior has been satisfactorily explained in the literature by invoking hypotheses that involve sperm competition, resistance to predators and parasites, or foraging enhancement. In this study, the fertility, live progeny production, progeny size, female receptivity, and latency to oviposition of bowl and doily spiders were measured and related to copulation duration. Using these data, we were able to eliminate six hypotheses and support a seventh: in this species, longer copulations result in larger hatchlings in the first clutch of eggs produced by the female. Although the correlation between copulation duration and progeny size is positive and significant, much of the variation in progeny size remains unexplained, and the reason for the variability in copulation durations remains obscure. Offprint requests to: R.B. Suter     

Fitness consequences of within-brood dominance in the cooperatively breeding acorn woodpecker

Social dominance confers potential advantages in terms of access to superior resources, habitats, and breeding opportunities. In the cooperatively breeding acorn woodpecker (Melanerpes formicivorus), within-brood dominance among juveniles is correlated with relative body size as nestlings. Capitalizing on this relationship, we investigated the fitness consequences of dominance by means of paired comparisons of broodmates. We found that (1) larger fledglings retained at least some of their size advantage as adults; (2) overwinter survival of larger, dominant fledglings was significantly greater than subordinates, but was not relatively greater when resources were poor than when they were good; (3) among birds surviving their first winter, there were no differences vis-à-vis dominance in terms of the proportion of birds acting as helpers or inheriting their natal territory. However, larger, dominant males were present in the study area longer than subordinates, suggesting that they either survived better or were more successful at gaining reproductive opportunities; (4) if only one male broodmate became a helper instead of dispersing, he was significantly more likely to be the smaller subordinate, consistent with the view that helping is a best-of-a-bad-job strategy; and (5) there were no significant differences in reproductive success among pairs of male broodmates that cobred together as adults, consistent with prior work failing to detect a phenotypic correlation of reproductive skew. Our results indicate that within-brood dominance relationships established as juveniles have significant effects on first-year survivorship and at least some aspects of adult fitness.     

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.     

Feeding frequency and parental division of labour in the double-brooded great tit Parus major

Summary We studied the relative contribution of each sex and total effort expended in feeding nestlings in the great tit Parus major in relation to artificially altered brood size. A recent model suggests that feeding frequency should reflect the optimal trade-off between parental and fledgling survival, the former being negatively, the latter positively, influenced by high feeding frequencies. In both sexes weight loss was linearly related to feeding frequency. Since fledgling survival increases with nestling weight, the conditions of this model are fulfilled. However, in contrast to the predictions of the model, the total feeding frequency for both sexes combined did not differ between control and enlarged broods, but was lower for reduced ones. This outcome was not the result of a physiologically related inability of the parents to increase their delivery rate. Instead, we suggest that parents with enlarged broods could not find sufficient amounts of prey large enough to be economically worth transporting to the nest. Differences in brood-provisioning rates between the sexes may arise because costs and benefits of feeding nestlings may differ. Females lost more weight than males during the nesting period, but maintained a relatively higher weight during the incubation period. The relationship between weight loss and feeding frequency was similar for both sexes. Male and female brood-feeding frequency was related to brood size in a similar way. This is discussed in light of the great tit's mating system and the fact that the great tit is facultatively double-brooded.     

Parent-offspring resemblance in degree of sociality in a passerine bird

The tendency to aggregate with conspecifics is a common type of social behavior, and interspecific differences in degree of sociality may indicate that evolutionary changes in sociality have been shaped by past selection pressures. A pre-requisite for such evolutionary change is that the behavior in question has an additive genetic basis. I investigated parent-offspring resemblance in two characteristics of sociality (colony size and nearest neighbor distance) in the semi-colonial barn swallow Hirundo rustica. Heritability estimates of these phenotypic characters were statistically significant. Offspring that were transferred to foster nests as part of brood size manipulation or cross-fostering experiments resembled their original parents more than their foster parents with respect to sociality. There was little evidence of phenotypic characteristics of mothers being significantly related to colony size or nearest neighbor distance of their offspring. Maintenance of genetic variation in sociality in this species may be related to the facts that the relationship between reproductive success and colony size differs among years, and that individuals differing in phenotypic quality have different optimal colony sizes.     

Evidence of obligate female promiscuity in a socially monogamous passerine

Extrapair paternity in birds has been the focus of an increasing number of studies over the last decades. Nevertheless, there is little knowledge about extrapair copulation (EPC) behavior. In this study, we investigate whether female bluethroats (Luscinia s. svecica) with no extrapair offspring (EPO) in their broods really have been sexually monogamous or if they have copulated with extrapair males. We used an experimental approach to prevent transfer of sperm from the social male during copulation by fitting a rubber tube around the cloaca of randomly chosen males. If the females mated to these males do not participate in EPCs, they will produce infertile eggs, whereas females that copulate with extrapair males will only produce EPO. We found that 87% (n=15) of the experimental pairs compared to only 36% (n=51) of unmanipulated pairs produced EPO. Our result therefore suggests that females having no EPO may still have copulated with extrapair males and that promiscuity may be an obligate sexual strategy among bluethroat females.Communicated by S. Pruett-Jones     

Testing a post-copulatory pre-zygotic reproductive barrier in a passerine species pair

Sexual selection may drive speciation, but most research focuses on pre-copulatory sexual selection, overlooking post-copulatory processes. Post-copulatory sexual selection in allopatric populations could drive divergence in post-copulatory pre-zygotic (PCPZ) phenotypes, limiting gene flow upon secondary contact. Here, we performed in vitro experiments examining one potential PCPZ barrier between two closely related passerine species, house sparrows (Passer domesticus) and Spanish sparrows (Passer hispaniolensis). In birds, crossing in the vagina may be particularly challenging for sperm, so we tested the effect of female reproductive tract fluids on sperm swimming speed and motility. If a PCPZ barrier exists at this stage of the fertilization process, heterospecific female fluids are predicted to reduce sperm swimming speed or motility relative to conspecific female fluid. We found that house sparrow female fluids affected the two species’ sperm asymmetrically, depending on the control sperm velocity and male species. Overall, however, sperm performed equally in conspecific and heterospecific female fluids, and the species had similar sperm morphology and sperm swimming performance. Low divergence in PCPZ phenotypes between species, perhaps because post-copulatory sexual selection is stabilizing or only moderately strong in these taxa, may be insufficient to cause an overall PCPZ barrier. Reinforcement may be unlikely to drive PCPZ barriers for this species pair, because relatively effective pre-copulatory barriers exist between the species, and because hybrids can be quite successful. Testing the role of PCPZ barriers in birds with more divergent PCPZ phenotypes will improve our understanding of speciation in passerines.     

Patterns of brood division and an absence of behavioral plasticity in a neotropical passerine

Studies of parental behavior in various habitats provide an opportunity to gain insight into how different environments may mold strategies of parental care. Brood division by parents has been hypothesized to occur facultatively within and among species. Brood division occurs when each parent cares for specific offspring within a brood. We studied brood division in a neotropical passerine, the western slaty antshrike (Thamnophilus atrinucha). Our results present a unique picture of a highly specialized example of avian brood division. Division was a fixed behavioral pattern in the population studied: all broods divided by fledging and remained divided during the entire post-fledging period. Brood division before fledging, a previously unreported phenomenon, occurred in 40% of nests observed. Parents that preferentially fed a certain offspring (defined as their focal offspring) in the nest fed the same individual after fledging. Each parent fed only its focal offspring in broods of one and two. The male parent cared for the heavier offspring and the first offspring to leave the nest. Siblings were segregated spatially during the time of highest predation risk. These observations suggest that a consistently high risk of predation on offspring has favored initial spatial segregation and inflexibility of brood division behavior in this species. Factors other than predation risk alone may explain the observed patterns of long-term, perfect brood division. Because high predation is common and relatively predictable in the tropics, selection for fixed brood division may be stronger in tropical birds than in the temperate zone.     

Context-dependent genetic benefits of extra-pair mate choice in a socially monogamous passerine

Extra-pair paternity is common in socially monogamous passerines; however, despite considerable research attention, consistent differences in fitness between within-pair offspring (WPO) and extra-pair offspring (EPO) have not been demonstrated. Recent evidence indicates that differences between maternal half-siblings may depend on environmental conditions, but it is unclear whether the influence of paternal genetic contribution should be most apparent under comparatively poor or favourable conditions. We compared phenotypic characteristics of WPO and EPO in 30 mixed-paternity broods of the tree swallow (Tachycineta bicolor) in relation to experimentally increased nest temperature (n = 13 heated nests; 17 control nests) and natural abundance of haematophagous parasites (Protocalliphora spp.). This allowed us to test the hypothesis that genetic benefits of extra-pair mating are environment dependent. EPO grew their ninth primary feathers faster than WPO regardless of nest temperature or parasite load and had significantly longer ninth primary feathers at fledging when parasite abundance was low, and when they were positioned early in the hatching sequence relative to WPO. In contrast, WPO under similar conditions did not differ from EPO in any phenotypic trait measured. These results indicate that the fitness benefits of extra-pair mating are likely to be context dependent, and that genetic effects on some phenotypic traits may be more apparent when conditions are relatively favourable.     

Sex-specific effects of the local social environment on juvenile post-fledging dispersal in great tits

An individual’s decision to disperse from the natal habitat can affect its future fitness prospects. Especially in species with sex-biased dispersal, we expect the cost–benefit balance for dispersal to vary according to the social environment (e.g., local sex ratio and density). However, little is known about the social factors affecting dispersal decisions and about the temporal and spatial patterns of the dispersal process. In our study, we investigated experimentally the effects of the social environment on post-fledging dispersal of juvenile great tits by simultaneously manipulating the density and sex ratio of fledglings within forest plots. We expected young females in the post-fledging period mainly to compete for resources related to food and, as they are subordinate to males, we predicted higher female dispersal from male-biased plots. Juvenile males compete for vacant territories already in late summer and autumn; thus, we predicted increased male dispersal from high density and male-biased plots. We found that juvenile females had a higher probability to leave male-biased plots and had dispersed further from male-biased plots in the later post-fledging phase when juvenile males start to become territorial and more aggressive. Juvenile males were least likely to leave male-biased plots and had smallest dispersal distances from female-biased plots early after fledging. The results suggest that the social environment differentially affected the costs and benefits of philopatry for male and female juveniles. The local sex ratio of individuals is thus an important social trait to be considered for understanding sex-specific dispersal processes.     

Long-term effects of yolk androgens on phenotype and parental feeding behavior in a wild passerine

Early growth conditions, such as exposure to maternally derived androgens in bird eggs, have been shown to shape offspring in ways that may have important long-term consequences for phenotype and behavior. Using an experimental approach, we studied the long-term effects of yolk androgens on several phenotypic traits and parental behavior in adult and female collared flycatchers (Ficedula albicollis). We elevated yolk androgen levels and monitored the experimental recruits the following breeding seasons. Androgen treatment had a sex-dependent effect on adult body condition, yolk androgen-treated males being heavier than control males when controlling for size, a result which may be caused potentially by selective mortality, physiological differences, or different life-history strategies. Androgen treatment did not however affect the expression of sexually selected plumage ornaments (forehead and wing patch size), UV coloration, or parental feeding rate in either sex. Our results suggest that yolk androgens are unlikely to affect sexual selection via plumage characteristics or contribute to breeding success via altered parental care. Yolk androgens do not seem to act as a means for female collared flycatchers to enhance the attractiveness of their sons. The lower return rate previously observed for androgen-treated male offspring compared to controls may therefore not be due to lower mating or breeding success, but may rather reflect lower survival or higher dispersal propensity of yolk androgen-treated males.     

Seeding phenology influences wood mouse seed choices: the overlooked role of timing in the foraging decisions by seed-dispersing rodents

Scatter-hoarding rodents influence the population dynamics of plants by acting as seed predators and dispersers. Therefore, rodent foraging preferences for certain seed traits (species, size, condition) have been extensively studied. However, to what extent these preferences are fixed or they track the temporal changes on seed characteristics due to phenological differences has been seldom explored. We studied the temporal variability in seed preferences by wood mouse (Apodemus sylvaticus), according to phenological changes in seed characteristics of two co-occurring oaks (Quercus ilex and Quercus pubescens). The phenology of acorn abundance and the acorn predation/dispersal patterns by rodents were monitored over an entire seeding season. Results revealed temporal changes in rodent preferences for acorns of the two oaks, matching their different seeding phenology (earlier in Q. pubescens and later in Q. ilex). On the other hand, whatever the species considered, rodents preferred larger and sound acorns along the entire season, although the dispersal of infested ones increased slightly during the peaks of acorn drop. The observed influence of seeding phenology on seed choices by rodents warns about inferring definite conclusions regarding their foraging behavior when arising from short-term experiments. Indeed, this study reveals that foraging preferences may be highly dynamic and context-dependent for some seed traits (e.g., species and condition), rather than fixed behavioral patterns. Plasticity in rodent foraging choices may allow them to successfully exploit different oaks with uncoupled seeding phenologies, while potentially favoring their coexistence.     

Fitness consequences of sexual cannibalism in female<Emphasis Type="Italic"> Argiope bruennichi</Emphasis>

The evolution of sexual cannibalism as the most extreme form of nuptial feeding is still poorly understood. Although increasing evidence suggests that female aggressiveness is related to other aspects of foraging behaviour, it is not clear whether the nutritional value of a male is sufficient to provide an adaptive significance for sexual cannibalism. A widely cited though rarely tested explanation is based on a paternal investment model, and predicts that consumption of a male results in increased female fecundity. The available evidence is either correlational or restricted to species with relatively large and potentially nutritious males, and different studies have come to different conclusions. Here we present a test of the paternal investment hypothesis using the very cannibalistic and highly size-dimorphic spider Argiope bruennichi. After a preset schedule, we had females consume none, one or two males independent of the female's cannibalistic behaviour. Consumption of male bodies did not result in any detectable fitness benefit for the female: neither the number of clutches, nor clutch size or hatching success were affected by consumption of males. The frequency of cannibalism was around 80%, independent of the female mating status. We did not observe male complicity, but cannibalism was associated with prolonged copulation. This suggests a sexually selected benefit of cannibalism for males. We conclude that the paternal investment hypothesis does not explain the existence of sexual cannibalism in A. bruennichi and probably not in other spider species with a pronounced sexual size dimorphism.Communicated by L. Simmons     

Duplex auditory distance assessment in a small passerine bird (Pipilo erythrophthalmus)

Eastern towhees, Pipilo erythrophthalmus, (Emberizidae, Passeriformes) in Florida estimate source sound-pressure level (SPL; i.e., sound amplitude) and often misjudge speaker distance in the field when vocalization source SPL is varied experimentally. Sound frequencies below ~3.5 kHz attenuate reliably with distance in comparison with higher sound frequencies in Florida scrub habitat. As a result, I predicted that towhees should use SPL as an auditory distance cue when they hear stimuli produced with sound frequencies below ~3.5 kHz but use another cue when they hear stimuli produced with sound frequencies above ~3.5 kHz. Subjects often misjudged speaker distance when approaching playbacks of SPL-altered stimuli produced with sound frequencies below ~3.5 kHz but rarely misjudged speaker distance when played SPL-altered stimuli produced with sound frequencies above ~3.5 kHz. I discuss the possibility that towhees employ a duplex sound localization strategy.