Current analyses do not resolve whether extra-pair paternity is male or female driven

Recent studies question the importance of indirect genetic effects in explaining female benefits of extra-pair matings in socially monogamous species. Compiling data on 14 wild bird species, Arnqvist and Kirkpatrick (Am Nat 165:S26–S37, 2005) estimated the average direct cost in terms or reduced parental care to be an order of magnitude larger than the potential effect of genetic benefits. This study has sparked a debate regarding potential confounding factors but no consensus appears to have been reached. Here we focus on the implicit assumption that all individuals face the same selective pressures and argue that this assumption is probably too strong in most cases. Using a theoretical model we show that when the amount of resources that a male provides depends on territory quality, his physical condition or prospects for alternative breeding opportunities, a female may respond to such differences by altering her mating behaviour. Such confounding factors may lead to direct fitness effects that result in negative correlations between paternal care and paternity even if females that produce extra-pair young experience a net benefit. Negative correlations can also result when males forcefully seek copulations and females resist them. We discuss the studies included in the analysis in this light, and conclude that current analyses on the net selective pressures remain uninformative. In addition to considering average effects across individuals and species we suggest giving attention to individual differences and the influence of ecological factors such as territory quality and predation pressures on female mating behaviour.     

Testosterone and the allocation of reproductive effort in male house finches (Carpodacus mexicanus)

Testosterone has been proposed to serve as the mediator that controls the relative effort that an individual male bird will devote to mating effort versus parental effort. Here, we demonstrate a testosterone-influenced trade-off between parental and mating efforts in male house finches. Male house finches with experimentally elevated testosterone fed nestlings at a significantly lower rate, but sang at a higher rate than males without manipulated testosterone levels. Females mated to testosterone-implanted males fed nestlings at a significantly higher rate than females mated to males without testosterone implants, resulting in similar feeding rates for both treated and untreated pairs. The effects of testosterone on male house finches, however, were not as dramatic as the effects of testosterone observed in some other socially monogamous species of birds. Because extra-pair copulations are uncommon in house finches and males provide substantial amounts of parental care, these more modest effects may be due to differences in how the allocation of reproductive effort affects the costs and benefits of different reproductive behaviors. Received: 6 June 2000 / Accepted: 17 July 2000     

Female settlement in marsh wrens: is it affected by other females?

Summary The influence of resident females on the settling patterns of subsequent females was examined for marsh wrens (Cistothorus palustris). Proportionally more females responded to a playback of a female intruder during the pre-laying stage than during the laying and incubation stages. The strength of the response was also greatest during pre-laying and laying stages (Table 1). Female wrens did not settle randomly with respect to the presence of other females, but rather settled with bachelor males more often than predicted by chance (Table 3). Female settlement within territories was more asynchronous than settlement between territories. An attempt to fit these data to two models of female settlement suggested that prospecting females were not displaced by residents, but rather avoided settling with mated males (Table 3). Females within territories were also more dispersed than predicted by chance, a pattern that may be enforced through aggression on the part of the resident female. Staggered settlement reduces the overlap in nestling stages of harem mates and so may be a strategy to reduce competition for resources at this critical stage.     

Developmental stress affects the attractiveness of male song and female choice in the zebra finch (Taeniopygia guttata)

Developmental stress has recently been shown to have adverse effects upon adult male song structure in birds, which may well act as an honest signal of male quality to discriminating females. However, it still remains to be shown if females can discriminate between the songs of stressed and non-stressed males. Here we use a novel experimental design using an active choice paradigm to investigate preferences in captive female zebra finches (Taeniopygia guttata). Nine females were exposed to ten pairs of songs by previously stressed and non-stressed birds that had learned their song from the same tutor. Song pairs differed significantly in terms of song complexity, with songs of stressed males exhibiting lower numbers of syllables and fewer different syllables in a phrase. Song rate and peak frequency did not differ between stressed and non-stressed males. Females showed a significant preference for non-stressed songs in terms of directed perching activity and time spent on perches. Our results therefore indicate that developmental stress affects not only the structure of male song, but that such structural differences are biologically relevant to female mate choice decisions.     

Sperm allocation in relation to male traits, female size, and copulation behaviour in freshwater crayfish species

Sperm competition is a well-recognised agent in the evolution of sperm and ejaculate structure, as well as variation in female quality. Models of the evolution of ejaculate expenditure predict that male body condition, female fecundity and the risk and intensity of sperm competition may be the ultimate factors shaping optimal ejaculate size. We investigated sperm allocation in Austropotamobius italicus, a freshwater crayfish exhibiting a coercive mating system and external fertilisation, in relation to male and female traits and copulation behaviour under laboratory conditions. We found that mating males were sensitive to female size and produced larger ejaculates when mating with larger females, which were more fecund in terms of number of eggs produced. We found no evidence for female egg production being sperm-limited, as the number of eggs was not dependent on male sperm expenditure. Copulation duration and number of ejaculations reliably predicted the amount of sperm transferred, and both these behavioural measures positively covaried with female body size. These results indicate that male freshwater crayfish can modulate their sperm expenditure in accordance with cues that indicate female fecundity. In addition, a novel finding that emerged from this study is the decrease in sperm expenditure with male body size, which may either suggest that large, old male crayfish are better able than small males to economise sperm at a given mating to perform multiple matings during a reproductive season, or that they experience senescence of their reproductive performance.     

Spacing of leks in relation to female home ranges,habitat requirements and male attractiveness in the great snipe (Gallinago media)

Summary In an attempt to examine three main hypotheses on the evolution of leks, data on female home ranges, distance between leks and male site fidelity in the great snipe (Gallinago media) were obtained. In a 30 × 40 km area in central Sweden, the locations of 12 leks were identified. Six were within the study area (7 × 11 km) in which probably all leks were known. In the study area, nearest neighbor distances between leks were longer than the predicted drawing area (diameter of female home ranges plus the detection range of leks). Furthermore, in only 1 of 13 radio-tracked females did the estimated home range enclose many leks. These data do not fit the prediction from the hotspot hypothesis on the dispersion of leks, namely, that the distance between leks should be less than the drawing area and hence an average female home range should enclose more than one lek. The observed distance between leks was about the same as predicted by the female preference hypothesis. This hypothesis also predicts females visit mainly one lek; however, we found they sometimes visit two and nest close to a third. We propose that males may first settle according to hotspot rules, but females will resettle according to preferences for certain males and/or larger leks. In this way some hotspot leks are abandoned, and the distance between leks is increased as males become more tightly clumped. However, when all males and females have settled, the leks still existing would be on hotspots. In the female preference hypothesis, males are assumed to aggregate on leks because females prefer clustered males as they can be more easily compared than if they are dispersed. Alternatively, in the attractiveness hypothesis, it is suggested that females prefer certain males, and hence unattractive males surround the attractive are forced to join leks in order to come close to females. Our data on male site fidelity support the attractiveness hypothesis because successful and dominant males return to the same lek and territory, both within and between years, whereas unsuccessful males move to other leks. Attractiveness of certain males may explain why according to hotspot rules in this species males are more clumped than expected. Offprint requests to: J. Höglund     

Female-female competition is influenced by forehead patch expression in pied flycatcher females

There is increasing evidence that sexual selection operates in females and not only in males. However, the function of female signals in intrasexual competition has been little studied in species with conventional sex roles. In the Iberian populations of the pied flycatcher (Ficedula hypoleuca), some females express a white forehead patch, a trait that in other European populations, only males exhibit and has become a classical example in studies of sexual selection. Here, we investigated whether the expression of this trait plays a role in female-female competition during early breeding stages. To test this hypothesis, we simulated territorial intrusions by challenging resident females with stuffed female decoys expressing or not a forehead patch. We found that resident females directed more attacks per trial and maintained closer distances to non-patched decoys than to patched ones. Also, patched females were more likely to attack the decoy than non-patched females. Interestingly, females were more aggressive against the decoys when their mate was absent. This may indicate that females relax territory vigilance in the presence of their mate or that males interfere in the interaction between competing females. The behavior of resident males was also observed, although it was not affected by decoy’s patch expression. Our findings suggest that the forehead patch plays a role in female intrasexual competition. If the forehead patch signals fighting ability, as it does in males, we may interpret that non-patched females probably avoided repeating costly agonistic encounters with the most dominant rivals.     

Differential deposition of antimicrobial proteins in blue tit (Cyanistes caeruleus) clutches by laying order and male attractiveness

Female birds can influence offspring fitness by varying the relative quantities of egg components they deposit within and between clutches. Antimicrobial proteins (lysozyme, ovotransferrin, and avidin) are significant components of the avian albumen and likely aid in defense of embryos from microbial infection. Within clutches, females may enhance antimicrobial defense of early-laid eggs to protect them from the high risk of infection incurred before the onset of incubation. Among entire clutches, females may invest more resources in young sired by more attractive males because they have higher reproductive value. We tested these hypotheses by quantifying antimicrobial protein distribution within and among clutches in blue tit eggs. Contrary to our hypothesis, clutches showed no differential deposition of lysozyme or avidin within clutches, but eggs laid in the middle of the sequence had higher concentrations of ovotransferrin than eggs in the beginning and end. Consistent with our second hypothesis, we found that females produced eggs with higher concentrations of lysozyme (although not ovotransferrin or avidin) when mated to more attractive (more UV-reflective) males. Furthermore, females mated to polygynous males deposited less lysozyme than those mated to monogamous males. These data suggest that allocation of lysozyme at the clutch level may be a maternal effect mediated by male qualities.     

Paternity alone does not predict long-term investment in juveniles by male baboons

Adult male chacma baboons (Papio hamadryas ursinus) form preferential associations, or friendships, with particular lactating females. Males exhibit high levels of affiliative contact with their friends’ infants and defend them from potentially infanticidal attacks (Palombit et al. 1997). Little is known about males’ associations with juveniles once they have passed the period of infanticidal risk. We conducted an observational, experimental, and genetic study of adult male and juvenile chacma baboons in the Moremi Reserve, Botswana. We identified preferential associations between males and juveniles and used behavioral data and a playback experiment to explore whether those associations have potential fitness benefits for juveniles. We determined whether males preferentially invest in care of their own offspring. We also determined how often males invest in care of their former friends’ offspring. The majority of juveniles exhibited preferential associations with one or two males, who had almost always been their mother’s friend during infancy. However, in only a subset of these relationships was the male the actual father, in part because many fathers died or disappeared before their offspring were weaned. Male caretakers intervened on behalf of their juvenile associates in social conflicts more often than they intervened on behalf of unconnected juveniles, and they did not appear to differentiate between genetic offspring and unrelated associates. Playbacks of juveniles’ distress calls elicited a stronger response from their caretakers than from control males. Chacma males may provide care to unrelated offspring of former friends because the costs associated with such care are low compared with the potentially high fitness costs of refusing aid to a juvenile who is a possible offspring.     

Boldness is influenced by sublethal interactions with predators and is associated with successful harem infiltration in Madagascar hissing cockroaches

One of the contemporary challenges of the behavioral syndromes literature is to identify how individual variation in behavior is determined, and whether this variation impacts ecological success. Although variation in experience is an obvious potential driver of intraspecific behavioral variation, predicting the impact of experience on suites of correlated behavioral traits is less intuitive. Specifically, if experience impacts traits that shape individuals' success in multiple contexts (e.g., foraging and anti-predator behavior), then experience could generate cross-contextual performance trade-offs associated with behavioral spillover. In the present study, we explore how sublethal experience with predators impacts various aspects of male behavioral tendencies in the Madagascar hissing cockroach, Gromphadorhina portentosa. First, we found that males' activity level and boldness were correlated together in the form of a behavioral syndrome. Second, we found that repeated sublethal interactions with predators shifted male boldness but not activity level, thus suggesting that the syndrome's constituent traits can respond to experience at least semi-independently. Third, we discovered that although predator exposure only influenced boldness, we found that boldness was highly correlated with males' ability to obtain rewarding positions in the harems of rival males. Taken together, our data suggest (but do not yet confirm) that although sublethal exposure to predators influences only a narrow subset of male's behavioral tendencies, these effects could still have nonintuitive consequences for males' success in functionally dissimilar ecological contexts (i.e., social and sexual encounters).     

Intraspecific variation in the red alga Portieria hornemannii : monoterpene concentrations are not influenced by nitrogen or phosphorus enrichment

The red alga Portieria hornemannii (Lyngbye) Silva was selected to test the effects of enhanced nutrient availability on the production of carbon-based secondary metabolites, because of its notable site-to-site variation in monoterpene production. On Guam, the major secondary metabolite produced by this alga is ochtodene, a cyclic monoterpene. Quantitative high-performance liquid-chromatography analysis of the extracts of P. hornemannii collected from six sites on Guam showed that both ochtodene and triglyceride concentrations differed significantly among sites. Internal nitrogen and phosphorus content of the algae did not correlate with the observed variation in chemistry. Experimental enhancement of N-alone, P-alone or N + P in the field for 5 wk failed to induce a significant change in ochtodene concentrations in the alga, while triglyceride concentrations increased significantly in the N + P treatment. Ochtodene and triglyceride concentrations did not change among similar treatments in shaded (18 d) and unshaded (11 d) fertilization experiments conducted in the laboratory. Variation in ochtodene concentrations in P. hornemannii cannot be attributed to N and P availability; however, the decrease in ochtodene and triglyceride concentrations during the shaded laboratory experiment suggests that light may be a factor influencing monoterpene biosynthesis. The difference in ochtodene concentration between the initial and final sets of field controls collected for the unshaded laboratory experiment suggests that temporal variation might also contribute to differences observed among the algae at the different sites. Received 11 June 1996 / Accepted 20 July 1996     

Correlated evolutionary changes in Drosophila female genitalia reduce the possible infection risk caused by male copulatory wounding

Male and female genitalia often show corresponding morphologies indicating coevolutionary processes, but ongoing debate exists as to the driving mechanisms of such coevolution between the sexes. Using interspecific matings between Drosophila santomea and Drosophila yakuba, this study provides evidence that micron-scale differences in male and female genital morphology and resultant mismatches between them during copulation cause wounding and infection risk to females. Male D. yakuba possesses a pair of sclerotized spikes (ventral branches of the basal processes) on the aedeagus, while these projections are much more rounded in D. santomea. D. yakuba females possess a pair of cavities with sclerotized platelets that receive the male genital spikes during copulation, whereas D. santomea females lack these cavities. The pointed genital spikes cause wounding in almost all females of D. santomea mating with male D. yakuba. Furthermore, when fluorescent-labeled microbeads were applied to male genitalia, they invaded the female hemocoel through the copulatory wounds significantly more frequently in heterospecific matings. Although the sclerotized platelets in the genital cavities prevented precise quantification of the extent of wounding in D. yakuba females, a similarly enhanced risk of microbial invasion was detected for the reciprocal heterospecific cross. All beads detected in the female hemocoel were immobilized by hemolymph clotting, indicating an effective immune response. The observed enhanced mating costs due to genital mismatch support the view that corresponding morphologies in female genitalia represent counteradaptations to harmful male behaviors, instead of being evolutionary adaptations to discriminate the species or genetic quality of male mates.     

Nest desertion is not predicted by cuckoldry in the Eurasian penduline tit

Engagement in extra-pair copulations is an example of the abundant conflicting interests between males and females over reproduction. Potential benefits for females and the risk of cuckoldry for males are expected to have important implications on the evolution of parental care. However, whether parents adjust parental care in response to parentage remains unclear. In Eurasian penduline tits Remiz pendulinus, which are small polygamous songbirds, parental care is carried out either by the male or by the female. In addition, one third of clutches is deserted by both male and female. Desertion takes place during the egg-laying phase. Using genotypes of nine microsatellite loci of 443 offspring and 211 adults, we test whether extra-pair paternity predicts parental care. We expect males to be more likely to desert cuckolded broods, whereas we expect females, if they obtain benefits from having multiple sires, to be more likely to care for broods with multiple paternity. Our results suggest that parental care is not adjusted to parentage on an ecological timescale. Furthermore, we found that male attractiveness does not predict cuckoldry, and we found no evidence for indirect benefits for females (i.e., increased growth rates or heterozygosity of extra-pair offspring). We argue that male Eurasian penduline tits may not be able to assess the risk of cuckoldry; thus, a direct association with parental care is unlikely to evolve. However, timing of desertion (i.e., when to desert during the egg-laying phase) may be influenced by the risk of cuckoldry. Future work applying extensive gene sequencing and quantitative genetics is likely to further our understanding of how selection may influence the association between parentage and parental care.     

The role of thymoquinone as antioxidant protection on oxidative stress induced by imidacloprid in male and female Swiss albino mice

The aim of the present study was to evaluate the possible protective effects of thymoquinone (TQ), an antioxidant agent, against imidacloprid (IMI)-induced oxidative stress in male and female mice. In total, 48 Swiss Albino male and female mice were fed a standard rodent diet and divided into 3 equal groups: the animals in the control group (vehicle treated) were given corn oil, the second group were orally administered 15 mg/kg/day IMI alone, and the third group were orally administered 15 mg/kg/day IMI and with TQ at 10 mg/kg/day for 21 days. During the experimental period, there were no significant changes between initial body weights and final body weights of IMI treated male and female mice. IMI produced significant increase in blood, liver, kidney, and heart malondialdehyde (MDA) levels and decrease in blood and liver glutathione (GSH) levels. In addition, IMI treatment decreased erythrocyte, liver, and kidney superoxide dismutase (SOD) activity in male mice and decreased erythrocyte and liver SOD activity in female mice. Erythrocyte catalase (CAT) activities were found to be low in male and female mice. However, treatment with TQ reversed IMI-induced oxidative stress, lipid peroxidation, and activities of antioxidant enzymes. Moreover, TQ exhibited protective action against the IMI-induced histopathological changes in tissues of male and female mice. In conclusion, TQ was found to be effective in protecting mice against IMI-induced oxidative stress by enhancing antioxidant defense mechanisms.     

Female gregariousness in Western Chimpanzees (Pan troglodytes verus) is influenced by resource aggregation and the number of females in estrus

Among social animals, group size is constrained by competition over resources. Because female reproductive success is limited by access to food resources, and that of males by access to fertile females, chimpanzee females are proposed to be less social than males and to maintain weak intrasexual relations. Findings from Taï National Park, Côte d’Ivoire, challenged this view, as chimpanzee females were described as generally gregarious, and close intrasexual bonds were common. Here, in a new analysis that focuses on the South Group of chimpanzees in Taï forest, we reevaluate the proposed differences in female association patterns between the Taï and East African populations. We find that mean party size and dyadic association index between females has decreased in Taï, although the level of dyadic associations remains high compared with East African chimpanzees. We attribute the decrease in female gregariousness to the decline in community size over the last 10 years. In addition, we use a multivariate approach to analyze social and ecological factors influencing party size in females. We show that female gregariousness increased when the fruit resources were more clumped and with increased number of females in estrus present. Party size of mothers with sons, however, was smaller with increasing number of sexually receptive females. The results of our model and the reviewed findings of other studies support the socioecological model because food distribution affects female gregariousness, but social and demographic aspects are equally influencing female grouping tendencies.     

To court or not to court: reproductive decisions by male fiddler crabs in response to fluctuating food availability

For males, courting and foraging are often behavioral alternatives, which take time and consume energy. When males have a possibility of mating with receptive females, there may be a behavioral trade-off between courtship and feeding; the outcome of which may be affected by male physiological condition and food availability. Although many mathematical models and empirical studies suggest that the expression of male courtship signals are condition-dependent, decisions about courtship and mating strategies in relation to food availability have not attracted much attention. In this study, we tested whether daily changes in food availability affect males’ decisions about whether to court. We conducted experiments with the fiddler crab Uca lactea by providing males with additional food every other day. In food-supplemented enclosures, males did not increase courtship activity on the days when food was supplemented. However, they built more courtship structures (semidomes) and waved more on the days when they were not given additional food. Male size had a strong influence on the number of days the males courted. We also tested whether the frequency of surface mating, as an alternative reproductive tactic, decreased when food was supplemented. Contrary to our expectation, the number of males that exhibited the surface-mating tactic increased when food was supplemented whereas the number of mate-searching females did not change. Our findings in this field study suggest that reproductive decisions by male fiddler crabs are affected by fluctuating food availability and present body condition, and the alternative mating tactic of this species may be more frequently used by males under good condition.     

Eggshell spotting in brood parasitic shiny cowbirds (<Emphasis Type="Italic">Molothrus bonariensis</Emphasis>) is not linked to the female sex chromosome

For avian brood parasites in which individual females are host-specialists, the arms race between hosts and parasites has favored egg color polymorphism in the parasite, with female lineages laying mimetic eggs that resemble those of the host species they parasitize. Female sex-linked inheritance of egg color fosters evolutionary stability of egg polymorphism if female lineages show both consistent eggshell color and host use. This co-evolutionary relationship is unlikely to occur if individual brood parasites use different hosts or if egg color is not maternally inherited. The shiny cowbird (Molothrus bonariensis) is an extreme generalist brood parasite that shows a very high degree of egg polymorphism. We tested whether egg spotting in this species has female sex-linked inheritance. If genetic factors controlling the expression of egg spotting were present on the female-specific W chromosome, we expected co-segregation between spotting patterns and mtDNA haplotypes, as both W and mtDNA are maternally inherited. In contrast to the known maternal inheritance of spotting patterns in great tits, we found no associations between eggshell spotting and mtDNA haplotypes, which suggests that eggshell spotting is not maternally inherited in this cowbird species.     

Evidence that lipid can be digested by the dumpling squid Euprymna tasmanica, but is not stored in the digestive gland

The aims of this study were to assess quantitatively the enzymatic ability of squid to digest lipids and the ability of the digestive gland to accumulate lipid classes associated with storage. This was achieved through two manipulative experiments using the dumpling squid, Euprymna tasmanica. Firstly, we measured lipase activity and determined the presence and location of lipid vacuoles within the digestive gland; secondly we identified and quantified lipid classes in the digestive gland. Given the levels of lipase activity, we provided evidence for the first time that a squid species is capable of digesting lipid at levels comparable to invertebrates known to use dietary lipid. A poor relationship between feeding activity and lipase secretion suggests that enzyme production is continuous. The second experiment found no evidence that lipid was stored in the digestive gland; most of the lipid present in the gland was either structural or a dietary by-product. The implication of these findings is that for this species lipid is most probably being immediately digested and used for growth and reproduction rather than being stored in the digestive gland. We consider that the role and storage of lipid is likely to vary among different cephalopod species, but not predictably as function of their lifestyle. Therefore, potential locations for lipid storage, other than the digestive gland, need to be considered and using changes in the relative size of the digestive gland as a measure of condition needs to be interpreted with care.