Mate guarding and territorial aggression vary with breeding synchrony in golden whistlers (<Emphasis Type="Italic">Pachycephala pectoralis</Emphasis>)

Male paternity assurance behaviour during the female fertile period has been widely documented amongst birds. In contrast, how sex-specific behavioural strategies vary with local breeding synchrony levels remains largely unknown. This is important because, in many species, intra-population patterns of extra-pair fertilisation rates, and hence cuckoldry risk, are known to vary with the number of simultaneously fertile females. Each sex may therefore differ in how they behave towards male conspecifics during different degrees of breeding synchrony. Here I provide evidence of such sex-specific differences in the golden whistler (Pachycephala pectoralis), a species in which within-pair paternity assurance is negatively associated with breeding synchrony. Via simulated territorial intrusions using decoy males, I show that males, but not females, increase levels of aggression to male intruders during periods of low synchrony, possibly because cuckoldry risk is greatest during this period. In addition, males appear to invest more effort into mate guarding after, but not before, territorial intrusions during this period. These inter-sexual differences may reflect conflicts in interest between the sexes, with females consistently showing interest in males during the fertile period regardless of synchrony levels and males investing more resources into expelling intruders when the risk of paternity loss is greatest. This study thus provides evidence that males may be able to detect variation in breeding synchrony and cuckoldry risk and adjust their paternity assurance behaviour accordingly.     

Thelytokous parthenogenesis by queens in the dacetine ant Pyramica membranifera (Hymenoptera: Formicidae)

Thelytokous parthenogenesis in which diploid females are produced from unfertilized eggs, was recently reported for some ant species. Here, we document thelytokous reproduction by queens in the polygynous species Pyramica membranifera. Queens that emerged in the laboratory were kept with or without workers under laboratory conditions. Independent colony founding was successful for a few queens if prey was provided. All artificial colonies, which started with a newly emerged queen and workers produced new workers and some of the colonies also produced female sexuals. Some of the female sexuals shed their wings in the laboratory and started formation of new polygynous colonies. Workers had no ovaries and thus, were obligatorily sterile.     

The functionality of female reciprocal calls in the Iberian midwife toad (<Emphasis Type="Italic">Alytes cisternasii</Emphasis>): female-female acoustic competition?

Female midwife toads (genus Alytes) emit highly variable reciprocal calls of unclear function prior to and during courtship. In some species, female-female competition, expressed as physical fighting, has been reported. Males of Majorcan midwife toads (Alytes muletensis) show phonotactic response to female calls, and females of Iberian midwife toads (Alytes cisternasii) respond differently according to the male call characteristics. In this study, I test the hypothesis of female-female acoustic competition as an additional function of female reciprocal calls. Playback tests indicate that female calls are not clearly involved in female acoustic competition in the Iberian midwife toad, therefore female calls could be directed at males rather than towards competitive females.     

Blue-green eggshell coloration is not a sexually selected signal of female quality in an open-nesting polygynous passerine

It has been proposed that blue-green egg colours have evolved as a post-mating signal of female quality, selected by males allocating their parental effort in response to the strength of this signal. We tested two main assumptions of the sexually selected egg coloration hypothesis: (1) whether the intensity of eggshell blue-green chroma (BGC) reflects female quality; and (2) whether males make their decisions on the level of parental care that they provide according to the intensity of eggshell BGC. As a model species, we chose the facultatively polygynous great reed warbler (Acrocephalus arundinaceus). In this species, females simultaneously paired with the same male, compete for his nest attendance and could benefit from signalling their quality through egg coloration. However, we found no association between the variation in eggshell BGC and the measures of female quality (physical condition, mean egg volume and age). Moreover, great reed warbler males did not adjust their investment (as measured in terms of nest defence against a brood parasite) in relation to the eggshell BGC. We conclude that blue-green egg coloration in this open-nesting passerine is unlikely to have a signalling function. Rather, the large colour variation among clutches of individual females may depend on yearly fluctuations in environmental conditions.     

Male wing color properties predict the size of nuptial gifts given during mating in the Pipevine Swallowtail butterfly (Battus philenor)

In many animals, males bear bright ornamental color patches that may signal both the direct and indirect benefits that a female might accrue from mating with him. Here we test whether male coloration in the Pipevine Swallowtail butterfly, Battus philenor, predicts two potential direct benefits for females: brief copulation duration and the quantity of materials the male passes to the female during mating. In this species, males have a bright iridescent blue field on the dorsal hindwing surface, while females have little or no dorsal iridescence. Females preferentially mate with males who display a bright and highly chromatic blue field on their dorsal hindwing. In this study, we show that the chroma of the blue field on the male dorsal hindwing and male body size (forewing length) significantly predict the mass of material or spermatophore that a male forms within the female’s copulatory sac during mating. We also found that spermatophore mass correlated negatively with copulation duration, but that color variables did not significantly predict this potential direct benefit. These results suggest that females may enhance the material benefits they receive during mating by mating with males based on the coloration of their dorsal hindwing.     

Males do not see only red: UV wavelengths and male territorial aggression in the three-spined stickleback (Gasterosteus aculeatus)

Animal colour signals serve important functions in intraspecific interactions, including species recognition, mate choice and agonistic behaviour. An increasing interest concerns ultraviolet (UV) wavelengths, for instance studies on the effect of UV in mating decisions. More recently, some studies also established that UV signals affect intrasexual interactions. We studied the role of UV during aggressive encounters between male three-spined sticklebacks (Gasterosteus aculeatus), a species in which UV has an effect on female and male mate choice and shoaling behaviour. To that aim, we compared the aggressive response of a territorial male to male intruders, either seen in UV-including (UV+) or UV-lacking (UV-) conditions. Our prediction was that, if UV wavelengths are used in male-male competition, a territorial male should show less competitive behaviour towards an intruder representing a lower threat, i.e. the one presented without UV light. Male sticklebacks showed significantly lower levels of aggression towards male opponents lacking an UV component to their coloration than male opponents possessing this colour component. Discrimination was not influenced by a difference in brightness between the UV+ and UV- stimuli. Finally, we present some reflectance-spectrophotometrical data of two skin regions (cheek and abdomen) of the experimental males and analysed relationships between colorimetric variables, body variables and behaviour. Our study emphasises that UV visual cues are of importance in different communicational tasks in the three-spined stickleback.     

Sexual selection on receptor organ traits: younger females attract males with longer antennae

Sexual selection theory predicts that female choice may favour the evolution of elaborate male signals. Darwin also suggested that sexual selection can favour elaborate receiver structures in order to better detect sexual signals, an idea that has been largely ignored. We evaluated this unorthodox perspective by documenting the antennal lengths of male Uraba lugens Walker (Lepidoptera: Nolidae) moths that were attracted to experimentally manipulated emissions of female sex pheromone. Either one or two females were placed in field traps for the duration of their adult lives in order to create differences in the quantity of pheromone emissions from the traps. The mean antennal length of males attracted to field traps baited with a single female was longer than that of males attracted to traps baited with two females, a pattern consistent with Darwin’s prediction assuming the latter emits higher pheromone concentrations. Furthermore, younger females attracted males with longer antennae, which may reflect age-specific changes in pheromone emission. These field experiments provide the first direct evidence of an unappreciated role for sexual selection in the evolution of sexual dimorphism in moth antennae and raise the intriguing possibility that females select males with longer antennae through strategic emission of pheromones.     

Sex-biased investment in yolk androgens depends on female quality and laying order in zebra finches (Taeniopygia guttata)

The Trivers–Willard hypothesis predicts sex biases in parental investment according to parental condition. In addition, parents may need to sex bias their investment if there is an asymmetry between the sexes in offspring fitness under different conditions. For studying maternal differential investment, egg resources are ideal subjects because they are self contained and allocated unequivocally by the female. Recent studies show that yolk androgens can be beneficial to offspring, so here we test for sex-biased investment with maternal investment of yolk testosterone (T) in zebra finch (Taeniopygia guttata) eggs. From the Trivers–Willard hypothesis, we predicted females to invest more in male eggs in optimum circumstances (e.g. good-condition mother, early-laid egg), and more in female eggs under suboptimal conditions (e.g. poor-condition mother, late-laid egg). This latter prediction is also because in this species there is a female nestling disadvantage in poor conditions and we expected mothers to help compensate for this in female eggs. Indeed, we found more yolk T in female than male eggs. Moreover, in accordance with our predictions, yolk T in male eggs increased with maternal quality relative to female eggs, and decreased with laying order relative to female eggs. This supports our predictions for the different needs and value of male and female offspring in zebra finches. Our results support the idea that females may use yolk androgens as a tool to adaptively manipulate the inequalities between different nestlings.     

Male antenna morphology and its effect on scramble competition in false garden mantids

Well-developed antennae are crucial for many insects, but especially for scramble competitors, who race to find their mates using female sex cues. In these systems, the ability of males to locate females quickly is thought to be under strong selection. A rarely tested assumption is that males with more sensory structures are able to locate females faster. In the present study, we used the false garden mantid Pseudomantis albofimbriata to investigate male antennal morphology and its effect on male efficiency in finding a mate. We used scanning electron microscopy to describe the major sensilla types and their arrangement along the length of male antennae. We also conducted field enclosure trials relating male antennal morphology to scramble competition in this system. We identified six different types of antennal sensilla (cheatic, trichoid, basiconic, grooved peg, ceolocapitular and campaniform) on male P. albofimbriata antennae. As expected, males who had more trichoid sensilla located females quicker than did males with fewer sensilla. Results of the current study suggest that antenna morphology plays a significant role in mate location and hence scramble competition in the P. albofimbriata mating system.     

Effects of life-history traits on parasitism in a monogamous mammal, the eastern rock sengi (Elephantulus myurus)

The distribution of parasites is often characterised by substantial aggregation with a small proportion of hosts harbouring the majority of parasites. This pattern can be generated by abiotic and biotic factors that affect hosts and determine host exposure and susceptibility to parasites. Climate factors can change a host’s investment in life-history traits (e.g. growth, reproduction) generating temporal patterns of parasite aggregation. Similarly, host age may affect such investment. Furthermore, sex-biased parasitism is common among vertebrates and has been linked to sexual dimorphism in morphology, behaviour and physiology. Studies exploring sex-biased parasitism have been almost exclusively conducted on polygynous species where dimorphic traits are often correlated. We investigated the effects of season and life-history traits on tick loads of the monogamous eastern rock sengi (Elephantulus myurus). We found larger tick burdens during the non-breeding season possibly as a result of energetic constraints and/or climate effects on the tick. Reproductive investment resulted in increased larval abundance for females but not males and may be linked to sex-specific life-history strategies. The costs of reproduction could also explain the observed age effect with yearling individuals harbouring lower larval burdens than adults. Although adult males had the greatest larval tick loads, host sex appears to play a minor role in generating the observed parasite heterogeneities. Our study suggests that reproductive investment plays a major role for parasite patterns in the study species.     

Carotenoid-based plumage coloration predicts resistance to a novel parasite in the house finch

The Hamilton-Zuk hypothesis proposes that the bright colours displayed by many species of birds serve as signals of individual resistance to parasites. Despite the popularity of this hypothesis, only one previous study has tested whether plumage coloration predicts how individuals respond to a disease challenge. We inoculated 24 male house finches (Carpodacus mexicanus) of variable plumage hue with a novel bacterial pathogen, Mycoplasma gallicepticum (MG). We found no relationship between plumage hue and time to first symptoms following inoculation, but we found a significant negative relationship between plumage hue and clearance of disease: males with redder plumage cleared MG infection significantly better than did males with yellower plumage. The hue of carotenoid-based plumage coloration has been shown to be a primary criterion in female mate choice in the house finch. These observations suggest that one benefit to females for choosing redder males is obtaining mates with better resistance to parasites.     

Eunuchs as better fighters?

Male–male competition for females can significantly affect a male’s reproductive success and hence his fitness. Game theory predicts that an individual should avoid fighting when its future reproductive potential is high, but should fight forcefully when its future reproductive potential is insignificant. When mates are scarce, extreme competition and fatal fighting is expected. We recently showed that Nephilengys malabarensis eunuchs, i.e. sterile spider males that lost their genitals during copulation, become more aggressive during male–male contests. Here, we add crucial comparative data by exploring eunuch fighting behaviour in Nephilengys livida from Madagascar, specifically by testing the ‘better fighter hypotheses’ in a laboratory setting. Similar to N. malabarensis, N. livida copulations resulted in total male castration with the severed palp plugging the female genitals in 70.83% cases, which mostly (63.63%) prevented subsequent copulations. Unexpectedly, however, N. livida eunuchs exhibited lower aggressiveness than virgin males. We interpret these results in the light of different mating biology between the so far studied species known for the eunuch phenomenon, which might reflect differing plug effectiveness due to variation in genital anatomy in N. livida, N. malabarensis and Herennia multipuncta. However, detected differences in aggressive behaviour of N. livida versus N. malabarensis eunuchs might also be explained by the species’ ecology, with lower population densities resulting in a relaxed male–male competition making excessive aggression and mate guarding redundant. This study thus questions the generality of overt aggressiveness in mated males with no reproductive value, and highlights the importance of understanding the natural history of species in the question.     

Communal nursing in wild house mice is not a by-product of group living: Females choose

Communal nursing, the provision of milk to non-offspring, has been argued to be a non-adaptive by-product of group living. We used 2 years of field data from a wild house mouse population to investigate this question. Communal nursing never occurred among females that previously lacked overlap in nest box use. Females nursed communally in only 33 % of cases in which there was a communal nursing partner available from the same social group. Solitarily nursing females were not socially isolated in their group; nevertheless, high spatial associations prior to reproduction predict which potential female partner was chosen for communal nursing. An increase in partner availability increased the probability of communal nursing, but population density itself had a negative effect, which may reflect increased female reproductive competition during summer. These results argue that females are selective in their choice of nursing partners and provide further support that communal nursing with the right partner is adaptive.     

Skin lipids of the striped plateau lizard (<Emphasis Type="Italic">Sceloporus virgatus</Emphasis>) correlate with female receptivity and reproductive quality alongside visual ornaments

Sex pheromones can perform a variety of functions ranging from revealing the location of suitable mates to being honest signals of mate quality, and they are used in the mate selection process by many species of reptile. In this study, we determined whether the skin lipids of female striped plateau lizards (Sceloporus virgatus) can predict the reproductive quality of females, thereby having the potential to serve as pheromones. Using gas chromatography/mass spectrometry, we identified 17 compounds present in skin lipids of female lizards. Using principal component analysis to compare the skin lipid profile of receptive and non-receptive females, we determined that an uncharacterized compound may allow for chemical identification of receptive mates. We also compared extracted principal components to measures of female fitness and reproductive qualities and found that the level of two 18 carbon fatty acids present in a female’s skin lipids may indicate her clutch size. Finally, we compared the information content of the skin lipids to that of female-specific color ornaments to assess whether chemical and visual cues transmit different information or not. We found that the chroma of a female’s orange throat patch is also related to her clutch size, suggesting that chemical signals may reinforce the information communicated by visual ornamentation in this species which would support the “backup signals” hypothesis for multiple signals.     

Sexual behavior,cannibalism, and mating plugs as sticky traps in the orb weaver spider <Emphasis Type="BoldItalic">Leucauge argyra</Emphasis> (Tetragnathidae)

Unpublished field observations in Leucauge argyra, a tropical orb weaver spider, suggest the occurrence of conspicuous mating plugs that could reduce or prevent remating attempts. Otherwise, the sexual behavior of this species remains unknown. The aims of this study were to describe the courtship behavior and copulation in L. argyra and investigate mating plug formation in this species. Fourteen virgin females and 12 plugged females were exposed to up to three males and checked for mating plug formation. Of the 12 virgins that copulated, nine produced plugs (five immediately after copulation), and the five plugged females that copulated produced another mating plug immediately after copulation. We did not detect the transfer of any male substance during copulation but observed a whitish liquid emerging from female genital ducts. Plug formation was positively associated with male twanging during courtship. One virgin and four plugged females cannibalized males. In seven trials with virgins and in three trials with plugged females, the male’s palp adhered to a substance that emerged from female genital ducts and spread on her genital plate. The male had to struggle energetically to free his glued palp; two of these males were cannibalized while trying to release their palps. Females seem to determine copulation duration by altering the timing of mating plug formation and through sexual cannibalism. This is the first case reported of a mating plug as a sticky trap for males.     

Dynamics of sperm transfer in the ant Leptothorax gredleri

Mating tactics differ remarkably between and within species of social Hymenoptera (bees, wasps, ants) concerning, e.g., mating frequencies, sperm competition, and the degree of male sperm limitation. Although social Hymenoptera might, therefore, potentially be ideal model systems for testing sexual selection theory, the dynamics of mating and sperm transfer have rarely been studied in species other than social bees, and basic information needed to draw conclusions about possible sperm competition and female choice is lacking. We investigated sperm transfer in the ant Leptothorax gredleri, a species in which female sexuals attract males by “female calling.” The analysis of 38 female sexuals fixed immediately or up to 7 days after copulation with a single male each revealed that the sperm is transferred into the female bursa copulatrix embedded in a gelatinous mass, presumably a spermatophore. Sperm cells rapidly start to migrate from the tip of the spermatophore towards the spermatheca, but transfer is drastically slowed down by an extreme constriction of the spermathecal duct, through which sperm cells have to pass virtually one by one. This results in the spermatheca being filled only between one and several hours after mating. During this time, the posterior part of the spermatophore seals the junction between bursa copulatrix and spermathecal duct and prevents sperm loss. The prolonged duration of sperm transfer might allow female sexuals to chose between ejaculates and explain previously reported patterns of single paternity of the offspring of multiply mated queens. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Songs of two starling species: common traits versus adaptations to the social environment

We analysed, for the first time, songs of the African Red-winged Starling Onychognathus morio and compared their general characteristics with those of the European Starling Sturnus vulgaris. Both species are gregarious during the non-breeding season, but European Starlings tend to nest in colonies, form unstable pair-bonds and are occasionally polygynous, whereas Red-winged Starlings form long-term pair-bonds and occupy exclusive nesting territories. Red-winged Starlings produced the same basic song categories as European Starlings: warbles and whistles. These two categories appeared to be involved in similar social interactions in the two species. However, several aspects of song behaviour differed between the two species: Red-winged Starlings, breeding in isolated nests, preferentially used whistles for long-distance communication and showed a simpler organization of warbling song. Whistles in the Red-winged Starling were mostly shared between birds and, in contrast to the European Starling, were not indicators of individual identity. Also in contrast to the European Starling, female song in Red-winged Starlings appeared very important throughout the breeding period. Our results suggest that some song characteristics in the two species are phylogenetically conserved whereas others are affected by the distinct social systems of the two species.     

Olfaction in the female sheep botfly

The nasal botfly Oestrus ovis (Diptera, Cyclorrhapha: Oestridae) is a myiasis-causing insect species, which affects the health of sheep, goats and humans. Gravid females are viviparous and larviposit into the animal’s nostrils. Host-searching and larvipositing flies are visually guided and influenced by climatic conditions, whereas olfaction seemed to play no role in this process. However, here, we show that the antennae of adult O. ovis female flies are relatively small but well developed and inhabited by several types of olfactory sensilla. Further, we show that the antennal lobes of this species receive input from antennal afferents and consist of a clearly defined glomerular organisation. We also give the first evidence of the fly’s ability to detect several synthetic odour compounds. Our findings provide a morpho-functional basis for future investigations on olfactory-mediated behaviour of this insect pest.     

Changes in reproductive life-history strategies in response to nest density in a shell-brooding cichlid, Telmatochromis vittatus

To determine whether the appearance of a reproductively parasitic tactic varies, and how this variation affects territorial males of the Lake Tanganyika cichlid fish Telmatochromis vittatus, we examined the reproductive ecology of territorial males in Mtondwe and compared it with that of a neighboring Wonzye population, where nest density differs from that at Mtondwe. In Wonzye, with high nest density, male tactics change with their body size from a territorial to a non-territorial parasitic tactic called piracy in which they conquer several nests defended by territorial males and take over the nests while females are spawning. These “pirate” males could decrease the costs incurred by travelling among nests by exclusively targeting aggregations of nests in close proximity while avoiding separate nests. Territorial males in Wonzye sacrifice the potential higher attractiveness offered by large nests and instead compete for nests farther from neighbors on which pirates less frequently intrude. In contrast, the Mtondwe population had lower nest density and piracy was absent. Given that the success of piracy depends on the close proximity of nests, nest density is likely responsible for the observed variation in the occurrence of piracy between the two populations. Furthermore, in Mtondwe, territorial males competed for larger nests and were smaller than the territorial males in Wonzye. Thus, this lower nest density may free territorial males from the selection pressures for increased size caused by both defense against nest piracy and the need to develop into pirates as they grow.