Parasites and health affect multiple sexual signals in male common wall lizards, Podarcis muralis

Multiple advertising sexual traits may either advertise different characteristics of male condition or be redundant to reinforce reliability of signals. Research has focused on multiple visual traits. However, in animals that use different multiple additional sensory systems, such as chemoreception, different types of traits might have evolved to signal similar characteristics of a male quality using different sensory channels. We examined whether ventral coloration and chemicals in femoral gland secretions of male common wall lizards, Podarcis muralis, are affected by their health state (blood-parasite load and cell-mediated immune response). Our results indicated that less parasitized lizards had brighter and more yellowish ventral colorations and also femoral secretions with higher proportions of two esters of octadecenoic acid. In addition, lizards with a greater immune response had more saturated coloration and secretions with higher proportions of octadecenoic acid methyl ester. We suggest that these signals would be reliable because only healthier males seemed able to allocate more carotenoids to coloration and presumably costly chemicals to secretions. The use of multiple sensory channels may provide more opportunities to signal a male quality under different circumstances, but also may reinforce the reliability of the signal when both types of traits may be perceived simultaneously.     

Negative correlation between nuptial throat colour and blood parasite load in male European green lizards supports the Hamilton–Zuk hypothesis

During female mate choice, conspicuous male sexual signals are used to infer male quality and choose the best sire for the offspring. The theory of parasite-mediated sexual selection (Hamilton–Zuk hypothesis) presumes that parasite infection can influence the elaboration of sexual signals: resistant individuals can invest more energy into signal expression and thus advertise their individual quality through signal intensity. By preferring these males, females can provide resistance genes for their offspring. Previous research showed that nuptial throat colour of male European green lizard, Lacerta viridis, plays a role in both inter- and intrasexual selections as a condition-dependent multiple signalling system. The aim of this study was to test the predictions of the Hamilton–Zuk hypothesis on male European green lizards. By blood sampling 30 adult males during the reproductive season, we found members of the Haemogregarinidae family in all but one individual (prevalence?=?96 %). The infection intensity showed strong negative correlation with the throat and belly colour brightness in line with the predictions of the Hamilton–Zuk hypothesis. In addition, we found other correlations between infection intensity and other fitness-related traits, suggesting that parasite load has a remarkable effect on individual fitness. This study shows that throat patch colour of the European green lizards not only is a multiple signalling system but also possibly acts as an honest sexual signal of health state in accordance with the Hamilton–Zuk hypothesis.     

Sexually dichromatic coloration reflects size and immunocompetence in female Spanish terrapins, Mauremys leprosa

Many studies have shown the importance of colorful ornamentation in mate choosiness or intrasexual conflict. However, research on color ornaments has focused mainly on birds, lizards or fish, but remains practically unknown in other animal groups such as turtles. In addition, female ornaments and their relation with sexual selection also remain almost unknown. Here, we measured the coloration of the shell and the limb stripes of male and female Spanish terrapins Mauremys leprosa and explored the existence of sexual dichromatism and the relation of color characteristics with body size and health state estimated from the immune response to the injection of an antigen (phytohaemagglutinin test). Our results showed that shell coloration, which could be constrained by natural selection to be cryptic, changed with body size, but did not differ between sexes. In contrast, females had brighter and less ultraviolet-saturated and more orange-saturated limb stripes than males. In females, interindividual variation in limb stripe coloration was related with body size and immune response suggesting that this coloration may inform honestly about multiple traits that could be important in sexual selection. In contrast, coloration of limb stripes of males was duller than in females, and was not related with any trait suggesting that coloration is not important in sexual selection for males.     

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.     

Tail loss compromises immunity in the many-lined skink, Eutropis multifasciata

Tail autotomy incurs energetic costs, and thus, a trade-off in resource allocation may lead to compromised immunity in lizards. We tested the hypothesis that tailless lizards will favor constitutive innate immunity responses over an energetically costly inflammatory response. The influence of fasting and colorful ornamentation was also investigated. We experimentally induced tail autotomy in the lizard Eutropis multifasciata and found that inflammation was suppressed by tail loss, but not further affected by fasting; the suppressive effect of colorful ornamentation was manifested only in males, but not in females. Constitutive innate immunity was not affected by any of these factors. As expected, only costly inflammation was compromised, and a less expensive constitutive innate immunity might be favored as a competent first-line defense during energetically demanding periods. After considering conventional trade-offs among tail regeneration and reproduction, further extending these studies to incorporate disease risk and how this influences escape responses to predators and future reproduction would make worthwhile studies.     

Male satin bowerbirds (<Emphasis Type="Italic">Ptilonorhynchus violaceus</Emphasis>) compensate for sexual signal loss by enhancing multiple display features

Numerous studies have focussed on the relationship between female choice and the multiple exaggerated sexual traits of males. However, little is known about the ability of males to actively enhance specific components of their display in response to the loss of one component. We investigated the capacity of male satin bowerbirds (Ptilonorhynchus violaceus) to respond to the loss of one of their sexual signals by performing an experiment in which we removed decorations at their bowers. We found that males compensated for decoration loss by increasing bower construction behaviour and decreasing their latency to bower painting. These results are novel because they suggest that males can assess the quality of their own display and make decisions about how to augment their displays. We discuss these results in the context of previous studies of mate choice in satin bowerbirds, as both of the supplementary behaviours we observed are known correlates of male mating success.     

Individual consistency in exploratory behaviour and mating tactics in male guppies

While behavioural plasticity is considered an adaptation to fluctuating social and environmental conditions, many animals also display a high level of individual consistency in their behaviour over time or across contexts (generally termed ‘personality’). However, studies of animal personalities that include sexual behaviour, or functionally distinct but correlated traits, are relatively scarce. In this study, we tested for individual behavioural consistency in courtship and exploratory behaviour in male guppies (Poecilia reticulata) in two light environments (high vs. low light intensity). Based on previous work on guppies, we predicted that males would modify their behaviour from sneak mating tactics to courtship displays under low light conditions, but also that the rank orders of courtship effort would remain unchanged (i.e. highly sexually active individuals would display relatively high levels of courtship under both light regimes). We also tested for correlations between courtship and exploratory behaviour, predicting that males that had high display rates would also be more likely to approach a novel object. Although males showed significant consistency in their exploratory and mating behaviour over time (1 week), we found no evidence that these traits constituted a behavioural syndrome. Furthermore, in contrast to previous work, we found no overall effect of the light environment on any of the behaviours measured, although males responded to the treatment on an individual-level basis, as reflected by a significant individual-by-environment interaction. The future challenge is to investigate how individual consistency across different environmental contexts relates to male reproductive success.     

Geographical variation in sexual behavior and body traits in a sex role reversed wolf spider

Mating partners need to recognize, assess each other, and exchange information through behavioral events that occur before, during, and after mating. Sexual signals, as well as life history traits, are influenced by selective pressures and environmental factors that can vary across distant geographical areas. Allocosa senex is a sand-dwelling wolf spider which constructs burrows along the sandy coasts of Argentina, Brazil, and Uruguay. Females are the mobile sex that searches for males and initiates courtship. They prefer males which construct longer burrows, and males prefer virgin females in good body condition. The objective of this study was to compare sexual behavior patterns, as well as body characteristics and burrow dimensions, between two geographically distant locations of A. senex, one in Uruguay (Uruguayan location) and the other from central Argentina (Argentinean location). We found differences in the number of male abdominal vibrations, male and female touches during mating, and number of erections of male leg spines, which all were higher in matings of Argentinean pairs. On the other hand, male body mass and female body condition were higher in Uruguayan individuals. The wide distribution of A. senex could be determining variations in the biotic and abiotic features that affect the species, generating differences in the strength of selective forces acting on individuals from the two studied locations.     

Multimodal sexual signals in male ocellated lizards Lacerta lepida: vitamin E in scent and green coloration may signal male quality in different sensory channels

The mechanisms that confer honesty to multiple sexual signals of vertebrates remain less known. We suggest that vitamin E in the scent (femoral secretions) and carotenoids in coloration of Lacerta lepida lizards might advertise quality because they are relevant in metabolism as major antioxidants and have a dietary origin. Results showed that there may be similar trade-offs between immunity and both chemical and visual signaling, as males with more vitamin E in secretions and those with more greenish, darker, and saturated carotenoid-dependent colorations had a higher immune response. This suggests that only high-quality males can divert from metabolism and allocate enough vitamin E to secretions and enough carotenoids to coloration. Therefore, information provided by visual and chemical signals is congruent. The use of multiple sensory channels may allow signaling a male quality under different circumstances or reinforce the reliability of the signal when both signals are perceived simultaneously. However, we also found that characteristics of carotenoid-dependent coloration mirror the amounts of vitamin E in secretions. This might support that carotenoids, which are not true antioxidants in the organism, would just reflect and “inform” on the actual contents of metabolic antioxidant vitamin E, which is part of the true system that prevent oxidation. We suggest that the metabolic roles of vitamins might be the actual basis of honesty of many sexual displays, and that carotenoid-based signals might be just the visible part of the whole system.     

Immune activation affects chemical sexual ornaments of male Iberian wall lizards

Many animals use chemical signals in sexual selection, but it is not clear how these sexual traits might have evolved to signal honestly male condition. It is possible that there is a trade-off between maintaining the immune system and the elaboration of ornaments. We experimentally challenged the immune system of male Iberian wall lizards, Podarcis hispanica, with a bacterial antigen (lipopolysaccharide), without pathogenic effects, to explore whether the immune activation affected chemical ornaments. Immune activation resulted in decreased proportions of a major chemical in femoral secretions (cholesta-5,7-dien-3-ol = provitamin D(3)) known to be selected in scent of males by females and which active form (vitamin D) has a variety of important effects on immune system function. This result suggests the existence of a potential trade-off between physiological regulation of the immune system and the allocation of essential nutrients (vitamins) to sexual chemical ornaments in male lizards.     

Condition-dependent expression of melanin-based coloration in the Eurasian kestrel

Melanin is the most common pigment in animal integuments and is responsible for some of the most striking ornaments. A central tenet of sexual selection theory states that melanin-based traits can signal absolute individual quality in any environment only if their expression is condition-dependent. Significant costs imposed by an ornament would ensure that only the highest quality individuals display the most exaggerated forms of the signal. Firm evidence that melanin-based traits can be condition-dependent is still rare in birds. In an experimental test of this central assumption, we report condition-dependent expression of a melanin-based trait in the Eurasian kestrel (Falco tinnunculus). We manipulated nestling body condition by reducing or increasing the number of nestlings soon after hatching. A few days before fledging, we measured the width of sub-terminal black bands on the tail feathers. Compared to nestlings from enlarged broods, individuals raised in reduced broods were in better condition and thereby developed larger sub-terminal bands. Furthermore, in 2?years, first-born nestlings also developed larger sub-terminal bands than their younger siblings that are in poorer condition. This demonstrates that expression of melanin-based traits can be condition-dependent.     

Detrimental effects of carotenoid pigments: the dark side of bright coloration

Carotenoid pigments produce yellow, orange, and red integumentary color displays that can serve as reliable signals of health and condition. In many birds and fish, individuals gain competitive or mating advantages by ingesting and utilizing large quantities of carotenoid pigments. Carotenoid pigments serve as antioxidants, performing important functions as free-radical scavengers. The beneficial effects of carotenoid pigments are well documented, but rarely have researchers considered potential detrimental effects of high-level accumulation of carotenoids. We maintained American goldfinches (Carduelis tristis) on high- or low-carotenoid diets through molt and tested for damage to the liver and skeletal muscle. High intake of carotenoids had no measurable effect on liver enzymes but caused an increase in creatine kinase, an indicator of skeletal muscle breakdown, and a reduction in vertical flight performance, a measure of skeletal muscle integrity. The detrimental effects of high-level carotenoid accumulation were approximately equivalent to the negative effects of removing carotenoids from the diet. The adverse effects observed in this study have important implications for theories of the function and evolution of colorful plumage.     

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.     

Now you see me,now you don’t: iridescence increases the efficacy of lizard chromatic signals

The selective forces imposed by primary receivers and unintended eavesdroppers of animal signals often act in opposite directions, constraining the development of conspicuous coloration. Because iridescent colours change their chromatic properties with viewer angle, iridescence offers a potential mechanism to relax this trade-off when the relevant observers involved in the evolution of signal design adopt different viewer geometries. We used reflectance spectrophotometry and visual modelling to test if the striking blue head coloration of males of the lizard Lacerta schreibeiri (1) is iridescent and (2) is more conspicuous when viewed from the perspective of conspecifics than from that of the main predators of adult L. schreibeiri (raptors). We demonstrate that the blue heads of L. schreiberi show angle-dependent changes in their chromatic properties. This variation allows the blue heads to be relatively conspicuous to conspecific viewers located in the same horizontal plane as the sender, while simultaneously being relatively cryptic to birds that see it from above. This study is the first to suggest the use of angle-dependent chromatic signals in lizards, and provides the first evidence of the adaptive function of iridescent coloration based on its detectability to different observers.     

Response of red deer stags (Cervus elaphus) to playback of harsh versus common roars

Red deer stags (Cervus elaphus) give two distinct types of roars during the breeding season, the “common roar” and the “harsh roar.” Harsh roars are more frequent during contexts of intense competition, and characterized by a set of features that increase their perceptual salience, suggesting that they signal heightened arousal. While common roars have been shown to encode size information and mediate both male competition and female choice, to our knowledge, the specific function of harsh roars during male competition has not yet been studied. Here, we investigate the hypothesis that the specific structure of male harsh roars signals high arousal to competitors. We contrast the behavioral responses of free ranging, harem-holding stags to the playback of harsh roars from an unfamiliar competitor with their response to the playback of common roars from the same animal. We show that males react less strongly to sequences of harsh roars than to sequences of common roars, possibly because they are reluctant to escalate conflicts with highly motivated and threatening unfamiliar males in the absence of visual information. While future work should investigate the response of stags to harsh roars from familiar opponents, our observations remain consistent with the hypothesis that harsh roars may signal motivation during male competition, and illustrate how intrasexual selection can contribute to the diversification of male vocal signals.     

Stable social aggregations in an Australian lizard,Egernia stokesii

Social groups in many animal species contain family members, and are maintained by parental care, and then tolerance of related individuals in the group. Lizards rarely show prolonged parental care, and rarely form stable social aggregations, although cohesive groups have been reported in the Australian skink genus, Egernia. A population of Egernia stokesii was surveyed for six seasons on Camel Hill, South Australia. At this site individuals occupied rock crevices for refuges on an isolated rocky outcrop of about 1.5 ha. The population was divided into 17 stable social groups, each containing 2-17 individual lizards. Surveyed juveniles took more than 5 years to reach mature size, and most juveniles and subadults remained in the social group of their parents for that period and longer. There were 2-8 permanent adult members of each group. Group members shared common crevice refuges, basked close together (and sometimes on top of each other) and defecated in common scat piles. There was low mortality after the first 2 years of life, and low dispersal either into or out of the population. Some individuals were "floaters" that did not belong to a social group. Over the study a number of these became established in groups. The social structure of these lizards resembles the family groups reported in many species of birds and mammals where group members help to raise the offspring of relatives. The low level of parental care in lizards suggests that the evolution of this form of social organisation in lizards has resulted from different processes than in other vertebrate taxa.     

Silk wrapping of nuptial gifts as visual signal for female attraction in a crepuscular spider

An extensive diversity of nuptial gifts is known in invertebrates, but prey wrapped in silk is a unique type of gift present in few insects and spiders. Females from spider species prefer males offering a gift accepting more and longer matings than when males offered no gift. Silk wrapping of the gift is not essential to obtain a mating, but appears to increase the chance of a mating evidencing a particularly intriguing function of this trait. Consequently, as other secondary sexual traits, silk wrapping may be an important trait under sexual selection, if it is used by females as a signal providing information on male quality. We aimed to understand whether the white color of wrapped gifts is used as visual signal during courtship in the spider Paratrechalea ornata. We studied if a patch of white paint on the males’ chelicerae is attractive to females by exposing females to males: with their chelicerae painted white; without paint; and with the sternum painted white (paint control). Females contacted males with white chelicerae more often and those males obtained higher mating success than other males. Thereafter, we explored whether silk wrapping is a condition-dependent trait and drives female visual attraction. We exposed good and poor condition males, carrying a prey, to the female silk. Males in poor condition added less silk to the prey than males in good condition, indicating that gift wrapping is an indicator of male quality and may be used by females to acquire information of the potential mate.     

Effects of height and habitat type on egg parasitism by Trichogramma minutum and T. pretiosum (Hymenoptera: Trichogrammatidae)

Heliothis virescens (Fabricius) eggs were sampled to determine the effects of height and habitat type (soybean field vs. uncultivated vegetation) on rates of p arasitism by Trichogramma minutum Riley and T. pretiosum Riley. The potentially confounding effects of plant associations and host density and distribution were eliminated by sampling with laboratory-produced eggs deposited on cloths, rather than with eggs naturally deposited by endemic hosts.A 3-way contingency table analysis showed that parasitism (whether the egg cloths were parasitized or not) was significantly associated with height and habitat type in 1981, with higher rates of parasitism in the uncultivated habitat and at the greater height. Parasitism was independent of height and habitat in 1982. Rates of parasitism by the two Trichogramma species were not clearly associated with habitat type, but were associated with height. T. minutum consistently parasitized egg cloths positioned at greater heights than those parasitized by T. pretiosum. While the behavioral mechanisms responsible are unknown, this inter-specific difference could have important implications with regard to selecting the most suitable of these two Trichogramma species for a particular inundative release program.     

The effects of different predator species on antipredator behavior in the Trinidadian guppy, Poecilia reticulata

Different types of predators often elicit different antipredator responses in a common type of prey. Alternatively, some prey species may adopt a general response, which provides limited protection from many different types of predator. The Trinidadian guppy, Poecilia reticulata, is faced with a wide range of different predators throughout its range and is known to display varying levels of antipredator behavior depending on the predator assemblage. Pike cichlids, Crenicichla frenata, are regarded as the primary aquatic guppy predator in streams in the northern mountain range in Trinidad. As such, they are seen to be responsible for many of the differences in morphology, life history traits, and behavior between guppy populations from areas with few predators and those from areas with many pike cichlids. In this study we investigated how guppies responded when faced with different predator species using three common aquatic predators. We exposed shoals of ten guppies to one out of four treatments: no predator (control), pike cichlid, acara cichlid (Aequidens pulcher), and wolf fish (Hoplias malabaricus); and we made behavioral observations on both focal individuals and the shoal as a whole. Guppies showed significantly greater levels of predator inspection and shoaling behavior, foraged less, spent more time in the surface water, and stayed in significantly larger shoals when faced with pike cichlids than in other treatments. We discuss these results in the context of multiple predator effects.