Previous experiments suggest that males spend more time with the more receptive of two novel females or the one with the higher fitness potential. However, males often court individual females repeatedly over a season; for example, male lizards sequentially visit familiar females as they patrol territorial boundaries. It may benefit males to vary display intensity as they move between multiple females. In this study, we explored the factors influencing amount of male courtship to familiar females in the Sagebrush lizard, Sceloporus graciosus. We tested whether males vary the amount of courtship exhibited due to individual differences among males, female reproductive state, or female fitness potential. Each male was allowed to interact separately, but repeatedly, with two females until both females laid eggs. Male courtship behavior with each of the two females was assayed at an intermediate point, after three weeks of interaction. We found that individual differences among males were considerable. The number of male courtship displays was also positively correlated with female latency to lay eggs, with males displaying more often towards females with eggs that had not yet been fertilized. Courtship behavior was not well predicted by the number of eggs laid or by female width, both measures of female quality. Thus, male S. graciosus appear to alter courtship intensity more in response to signals of female reproductive state than in response to variation in potential female fitness.
Sceloporus graciosus; male choice; mate choice; sexual selection; reproductive state; courtship
The energetic resources in an organism’s environment are essential for executing a wide range of life history functions, including immunity and reproduction. Most energetic budgets, however, are limited, which can lead to trade-offs among competing functions. Increasing reproductive effort tends to decrease immunity in many cases; and increasing total energy via supplemental feedings can eliminate this effect. Testosterone (T), an important regulator of reproduction, and food availability are thus both potential factors regulating life-history processes, yet they are often tested in isolation of each other. In this study, we considered the effect of both food availability and elevated T on immune function and reproductive behavior in sagebrush lizards, Sceloporus graciosus, to assess how T and energy availability affect these trade-offs. We experimentally manipulated diet (via supplemental feedings) and T (via dermal patches) in males from a natural population. We determined innate immune response by calculating the bacterial killing capability of collected plasma exposed to E. coli ex vivo. We measured reproductive behavior by counting the number of courtship displays produced in a 20-min sampling period. We observed an interactive effect of food availability and T-patch on immune function, with food supplementation increasing immunity in T-patch lizards. Additionally, T increased courtship displays in control food lizards. Lizards with supplemental food had higher circulating T than controls. Collectively, this study shows that the energetic state of the animal plays a critical role in modulating the interactions among T, behavior and immunity in sagebrush lizards and likely other species.
Context-dependent; Energy allocation; Innate immunity; Life history; Resources; Sceloporus; Trade-offs
Male sexual displays play an important role in sexual selection by affecting reproductive success. However, for such displays to be useful for female mate choice, courtship should vary more among than within individual males. In this regard, a potentially important source of within male variation is adjustment of male courtship effort in response to female traits. Accordingly, we set out to dissect sources of variation in male courtship effort in a fish, the desert goby (Chlamydogobius eremius). We did so by designing an experiment that allowed simultaneous estimation of within and between male variation in courtship, while also assessing the importance of the males and females as sources of courtship variation.
Although males adjusted their courtship depending on the identity of the female (a potentially important source of within-male variation), among-male differences were considerably greater. In addition, male courtship effort towards a pair of females was highly repeatable over a short time frame.
Despite the plasticity in male courtship effort, courtship displays had the potential to reliably convey information about the male to mate-searching females. Our experiment therefore underscores the importance of addressing the different sources contributing to variation in the expression of sexually-selected traits.
In animal communication, complex displays usually have multiple functions and, male and female receivers often differ in their utilization and response to different aspects of these displays. The perceptual variability hypothesis suggests that different aspects of complex signals differ in their ability to be detected and processed by different receivers. Here, we tested whether receiver male and female Sceloporus graciosus lizards differ in visual motion detection by measuring the latency to the visual grasp response to a motion stimulus. We demonstrate that in lizards that largely exhibit complex motions as courtship signals, female lizards are faster than males at visually detecting motion. These results highlight that differential signal utilization by the sexes may be driven by variability in the capacity to detect different display properties.
sex difference; visual performance; motion detection; complex signals; animal communication; Sceloporus graciosus
Conflict between the sexes has traditionally been studied in terms of costs of mating to females and female resistance. However, courting can also be costly to males, especially when females are larger and aggressively resist copulation attempts. We examined male display intensity towards females in the Cape dwarf chameleon, Bradypodion pumilum, in which females are larger than males and very aggressive. We assessed whether aggressive female rejection imposes potential costs on males and whether males vary their display behaviour with intensity of female rejection, female size or relative size differences. Males persisted in courtship after initial female rejection in 84% of trials, and were bitten in 28% of trials. Attempted mounts were positively associated with males being bitten. Males reduced courtship with increased intensity of female rejection. Male courtship behaviour also varied with female size: males were more likely to court and approach smaller females, consistent with the hypothesis that larger females can inflict more damage. These results suggest that, in addition to assessing female willingness to mate, male dwarf chameleons may use courtship displays to assess potential costs of persistence, including costs associated with aggressive female rejection, weighed against potential reproductive pay-offs associated with forced copulation.
mating costs; sexual conflict; antagonistic coevolution; female rejection
Although it is axiomatic that males and females differ in relation to many aspects of reproduction related to physiology, morphology and behaviour, relatively little is known about possible sex differences in the response to cues from the environment that control the timing of seasonal breeding. This review concerns the environmental regulation of seasonal reproduction in birds and how this process might differ between males and females. From an evolutionary perspective, the sexes can be expected to differ in the cues they use to time reproduction. Female reproductive fitness typically varies more as a function of fecundity selection, while male reproductive fitness varies more as a function sexual selection. Consequently, variation in the precision of the timing of egg laying is likely to have more serious fitness consequences for females than for males, while variation in the timing of recrudescence of the male testes and accompanying territory establishment and courtship are likely to have more serious fitness consequences for males. From the proximate perspective, sex differences in the control of reproduction could be regulated via the response to photoperiod or in the relative importance and action of supplementary factors (such as temperature, food supply, nesting sites and behavioural interactions) that adjust the timing of reproduction so that it is in step with local conditions. For example, there is clear evidence in several temperate zone avian species that females require both supplementary factors and long photoperiods in order for follicles to develop, while males can attain full gonadal size based on photoperiodic stimulation alone. The neuroendocrine basis of these sex differences is not well understood, though there are many candidate mechanisms in the brain as well as throughout the entire hypothalamo–pituitary–gonadal axis that might be important.
photoperiodism; circannual rhythms; sex differences; supplementary cues
In systems where territory quality varies, animals are expected to exhibit plasticity in behaviour in order to maximize fitness relative to their present territory quality. This requires assessment of territory quality followed by decision-making in relation to the priority of activities necessary for survival and reproduction. We examined how differences in territory quality of beaugregory damselfish (Stegastes leucostictus) influence the prioritization of courtship and egg defence by comparing behavioural responses of males defending artificial sites (high quality) with males defending natural sites (low quality) when presented with an egg predator, a conspecific female, and a simultaneous choice between both. A significant three-way interaction of territory quality, presentation type and stimulus was observed for time near stimuli. In paired presentations, males defending low-quality territories spent more time near a female and less near an egg predator; while males on high-quality territories spent more time near a predator than a female. Additionally, comparing single and paired presentations reveals that behaviours towards egg predators remain constant while behaviours towards females decrease with paired stimuli. These data suggest that territory quality and ecological context impact decision-making and the relative values of potential reproduction and/or past reproductive effort.
beaugregory damselfish; courtship; decision-making; Stegastes leucostictus; territory quality
Sexually selected male courtship displays often involve multiple behavioural and physical traits, but little is known about the function of different traits in mate choice. Here, we examine female courtship behaviours to learn how male traits interact to influence female mating decisions. In satin bowerbirds (Ptilonorhynchus violaceus), successful males give highly aggressive, intense behavioural displays without startling females. Males do this by modulating their displays in response to female crouching, which signals the display intensity that females will tolerate without being startled. Females typically visit multiple males for multiple courtships before choosing a mate, and females show differing tolerance for intense displays during their first courtship with each male. We test three hypotheses that may explain this: (i) familiarity with the courting male; (ii) the order of the courtship in mate-searching; and (iii) the attractiveness of the courting male. We found that females are more tolerant of intense displays during first courtships with attractive males; this increased female tolerance may allow attractive males to give higher intensity courtship displays that further enhance their attractiveness. We then examined why this is so, finding evidence that females are less likely to be startled by males with better physical displays (bower decorations), and this reduced startling then contributes to male courtship success. This role of physical displays in facilitating behavioural displays suggests a novel mechanism by which multiple physical and behavioural traits may influence female choice.
Sexual selection theory predicts that females, being the limiting sex, invest less in courtship signals than males. However, when chemical signals are involved it is often the female that initiates mating by producing stimuli that inform about sex and/or receptivity. This apparent contradiction has been discussed in the literature as 'the female pheromone fallacy'. Because the release of chemical stimuli may not have evolved to elicit the male's courtship response, whether these female stimuli represent signals remains an open question. Using techniques to visualise and block release of urine, we studied the role of urine signals during fighting and mating interactions of crayfish (Pacifastacus leniusculus). Test individuals were blindfolded to exclude visual disturbance from dye release and artificial urine introduction.
Staged female-male pairings during the reproductive season often resulted in male mating attempts. Blocking female urine release in such pairings prevented any male courtship behaviour. Artificial introduction of female urine re-established male mating attempts. Urine visualisation showed that female urine release coincides with aggressive behaviours but not with female submissive behaviour in reproductive interactions as well as in intersexual and intrasexual fights. In reproductive interactions, females predominately released urine during precopulatory aggression; males subsequently released significantly less urine during mating than in fights.
Urine-blocking experiments demonstrate that female urine contains sex-specific components that elicit male mating behaviour. The coincidence of chemical signalling and aggressive behaviour in both females and males suggests that urine release has evolved as an aggressive signal in both sexes of crayfish. By limiting urine release to aggressive behaviours in reproductive interactions females challenge their potential mating partners at the same time as they trigger a sexual response. These double messages should favour stronger males that are able to overcome the resistance of the female. We conclude that the difference between the sexes in disclosing urine-borne information reflects their conflicting interests in reproduction. Males discontinue aggressive urine signalling in order to increase their chances of mating. Females resume urine signalling in connection with aggressive behaviour, potentially repelling low quality or sexually inactive males while favouring reproduction with high quality males.
Investigating individual differences in sexual performance in unmanipulated males is important for understanding natural relationships between behavior and morphology, and the mechanisms regulating them. Among male green anole lizards, some court and copulate frequently (studs) and others do not (duds). To evaluate potential factors underlying differences in the level of these behaviors, morphology and androgen receptor expression in neuromuscular courtship and copulatory structures, as well as in the preoptic area and amygdala, were compared in males displaying varying degrees of sexual function. This study revealed that individual differences in behavior among unmanipulated males, in particular the extension of a throat fan (dewlap) used during courtship, were positively correlated with the size of fibers in the associated muscle and with soma size in the amygdala. The physiological response to testosterone, as indicated by the height of cells in an androgen-sensitive portion of the kidney, was also correlated with male sexual behavior, and predicted it better than plasma androgen levels. Androgen receptor expression was not related to the display of courtship or copulation in any of the tissues examined. The present data indicate that higher levels of male courtship behavior result in (or are the result of) enhanced courtship muscle and amygdala morphology, and that androgen-sensitive tissue in studs may be more responsive to testosterone than duds. However, some mechanism(s) other than androgen receptor expression likely confer this difference in responsiveness.
Testosterone; Courtship; Copulation; Reptile; Preoptic area; Amygdala
The laboratory mouse is an emerging model for context-dependent vocal signaling and reception. Mouse ultrasonic vocalizations are robustly produced in social contexts. In adults, male vocalization during courtship has become a model of interest for signal-receiver interactions. These vocalizations can be grouped into syllable types that are consistently produced by different subspecies and strains of mice. Vocalizations are unique to individuals, vary across development, and depend on social housing conditions. The behavioral significance of different syllable types, including the contexts in which different vocalizations are made and the responses listeners have to different types of vocalizations, is not well understood. We examined the effect of female presence and estrous state on male vocalizations by exploring the use of syllable types and the parameters of syllables during courtship. We also explored correlations between vocalizations and other behaviors. These experimental manipulations produced four main findings: 1) vocalizations varied among males, 2) the production of USVs and an increase in the use of a specific syllable type were temporally related to mounting behavior, 3) the frequency (kHz), bandwidth, and duration of syllables produced by males were influenced by the estrous phase of female partners, and 4) syllable types changed when females were removed. These findings show that mouse ultrasonic courtship vocalizations are sensitive to changes in female phase and presence, further demonstrating the context-sensitivity of these calls.
The accessory gland proteins (Acps) that male Drosophila melanogaster produce
and transfer to females during copulation are key to male and female fitness. One Acp, the
sex peptide (SP), is largely responsible for a dramatic increase in female egg laying and
decrease in female receptivity after copulation. While genetic variation in male
SP expression levels correlate with refractory period duration in females, it
is unknown whether male SP expression influences female egg laying or if any
effect of SP is mediated by SP retention in the female reproductive tract. Here we
measured the amount of SP retained in the female reproductive tract after mating and
female egg laying after copulating with virgin males. We found no correlation between male
SP expression levels and egg laying, or the amount of SP in the female
reproductive tract after mating. Additionally, the amount of SP retained in the female did
not influence egg laying. These finding suggests that additional factors, such as
variation in other Acps, are important for the retention of SP in females and its
quantitative effects on egg laying. It also shows that egg laying and refractory period
response to SP is at least partially uncoupled.
; sex peptide; sexual conflict; sperm competition
Studies of mating preferences and pre-mating reproductive isolation have often focused on females, but the potential importance of male preferences is increasingly appreciated. We investigated male behavior in the context of reproductive isolation between divergent anadromous and stream-resident populations of threespine stickleback, Gasterosteus aculeatus, using size-manipulated females of both ecotypes. Specifically, we asked if male courtship preferences are present, and if they are based on relative body size, non-size aspects of ecotype, or other traits. Because male behaviors were correlated with each other, we conducted a principal components analysis on the correlations and ran subsequent analyses on the principal components. The two male ecotypes differed in overall behavioral frequencies, with stream-resident males exhibiting consistently more vigorous and positive courtship than anadromous males, and an otherwise aggressive behavior playing a more positive role in anadromous than stream-resident courtship. We observed more vigorous courtship toward smaller females by (relatively small) stream-resident males and the reverse pattern for (relatively large) anadromous males. Thus size-assortative male courtship preferences may contribute to reproductive isolation in this system, although preferences are far from absolute. We found little indication of males responding preferentially to females of their own ecotype independent of body size.
Long-term memory formation in Drosophila melanogaster is an important neuronal function shaping the insect’s behavioral repertoire by allowing an individual to modify behaviors on the basis of previous experiences. In conditioned courtship or courtship suppression, male flies that have been repeatedly rejected by mated females during courtship advances are less likely than naïve males to subsequently court another mated female. This long-term courtship suppression can last for several days after the initial rejection period. Although genes with known functions in many associative learning paradigms, including those that function in cyclic AMP signaling and RNA translocation, have been identified as playing critical roles in long-term conditioned courtship, it is clear that additional mechanisms also contribute. We have used RNA sequencing to identify differentially expressed genes and transcript isoforms between naïve males and males subjected to courtship-conditioning regimens that are sufficient for inducing long-term courtship suppression. Transcriptome analyses 24 hours after the training regimens revealed differentially expressed genes and transcript isoforms with predicted and known functions in nervous system development, chromatin biology, translation, cytoskeletal dynamics, and transcriptional regulation. A much larger number of differentially expressed transcript isoforms were identified, including genes previously implicated in associative memory and neuronal development, including fruitless, that may play functional roles in learning during courtship conditioning. Our results shed light on the complexity of the genetics that underlies this behavioral plasticity and reveal several new potential areas of inquiry for future studies.
long-term memory; courtship conditioning; RNA seq; courtship behavior
Theory predicts that males will benefit when they bias their mating effort towards females of higher reproductive potential, and that this discrimination will increase as males become more resource limited. We conducted a series of experiments to test these predictions in a laboratory population of the fruitfly, Drosophila melanogaster. In this species, courtship and copulation have significant costs to males, and females vary greatly in fecundity, which is positively associated with body size. When given a simultaneous choice between small and large virgin females, males preferentially mated with larger, more fecund, females. Moreover, after males had recently mated they showed a stronger preference for larger females. These results suggest that male D. melanogaster adaptively allocate their mating effort in response to variation in female quality and provide some of the first support for the theoretical prediction that male stringency in mate choice increases as resources become more limiting.
male mate choice; mating costs; Drosophila melanogaster; fecundity
In many species, males can influence the amount of resources their mates invest in reproduction. Two favoured hypotheses for this observation are that females assess male quality during courtship or copulation and alter their investment in offspring accordingly, or that males manipulate females to invest heavily in offspring produced soon after mating. Here, we examined whether there is genetic variation for males to influence female short-term reproductive investment in Drosophila melanogaster, a species with strong sexual selection and substantial sexual conflict. We measured the fecundity and egg size of females mated to males from multiple isofemale lines collected from populations around the globe. Although these traits were not strongly influenced by the male's population of origin, we found that 22 per cent of the variation in female short-term reproductive investment was attributable to the genotype of her mate. This is the first direct evidence that male D. melanogaster vary genetically in their proximate influence on female fecundity, egg size and overall reproductive investment.
maternal investment; fecundity; egg size; differential allocation hypothesis; sexual conflict; Drosophila melanogaster
Male Drosophila fruit flies acquire and defend territories in order to attract females for reproduction. Both, male-directed agonistic behavior and female-directed courtship consist of series of recurrent stereotypical components. Various studies demonstrated the importance of species-specific sound patterns generated by wing vibration as being critical for male courtship success. In this study we analyzed the patterns and importance of sound signals generated during agonistic interactions of male Drosophila melanogaster. In contrast to acoustic courtship signals that consist of sine and pulse patterns and are generated by one extended wing, agonistic signals lack sine-like components and are generally produced by simultaneous movements of both wings. Though intra-pulse oscillation frequencies (carrier frequency) are identical, inter-pulse intervals are twice as long and more variable in aggression signals than in courtship songs, where their precise temporal pattern serves species recognition. Acoustic signals accompany male agonistic interactions over their entire course but occur particularly often after tapping behavior which is a major way to identify the gender of the interaction partner. Since similar wing movements may either be silent or generate sound and wing movements with sound have a greater impact on the subsequent behavior of a receiver, sound producing wing movements seem to be generated intentionally to serve as a specific signal during fruit fly agonistic encounters.
acoustic signals; temporal patterns; frequency analysis; agonistic behavior; courtship; ethogram
Traditional models of sexual selection posit that male courtship signals evolve as indicators of underlying male genetic quality. An alternative hypothesis is that sexual conflict over mating generates antagonistic coevolution between male courtship persistence and female resistance. In the scarabaeine dung beetle Onthophagus taurus, females are more likely to mate with males that have high courtship rates. Here, we examine the effects of exposing females to males with either high or low courtship rates on female lifetime productivity and offspring viability. Females exposed to males with high courtship rates mated more often and produced offspring with greater egg–adult viability. Female productivity and lifespan were unaffected by exposure to males with high courtship rates. The data are consistent with models of sexual selection based on indirect genetic benefits, and provide little evidence for sexual conflict in this system.
female choice; genetic benefits; sexual conflict; courtship rate
Interactions among reproductive season, testosterone (T) and female presence were investigated on the structure and function of forebrain and neuromuscular systems controlling courtship and copulation in the green anole lizard. Under breeding (BS) or non-breeding (NBS) environmental conditions, male green anoles were implanted with either T or blank capsules and exposed to one of three female stimulus conditions: physical, visual or no female contact. T and at least visual exposure to females increased courtship displays (extension of a throat fan, or dewlap), and these effects were greater during the BS than NBS. T also facilitated copulation, and did so to a greater extent in the BS. The hormone increased soma size in the preoptic area (POA) and amygdala (AMY), and in the AMY the effects were greater in the BS than NBS. Cross-sectional areas of copulatory organs and associated muscle fibers were enhanced by T, and more so in the BS than NBS. However, no effects on morphology of dewlap motoneurons or muscles or copulatory motoneurons were detected. Thus, (1) changes in behavior and neural and/or muscular morphology are not always parallel and (2) differences in responsiveness to T exist across seasons and among tissues.
Sexual behavior; Courtship; Lizard; Reptile; Forebrain; Neuromuscular
The adaptive significance of learning is supported by studies showing its positive effects on mating behaviour, but they rarely go beyond fertilization success. Here we studied how learning contributes to qualitative reproductive investment, by testing the hypothesis that mating in the context that predicts male appearance has positive effects on female reproductive investment compared with unsignalled mating. Using Japanese quail (Coturnix japonica), we found that effects of mating in the context predicting mating opportunity depend on female body condition and receptivity, while the outcome of unexpected mating depends on male behaviour. In particular, among females mated with the familiar male in the context predicting that he will appear, female condition positively affected the number of fertilized eggs and egg mass and more receptive females tended to produce more sons. Additionally, conditioned females laid heavier eggs for daughters than for sons. In contrast, in females that were mated unexpectedly and with a novel male, the number of fertilized eggs was highly dependent on male behaviour and was negatively related to maternal body condition. Egg mass was not related to body condition, and there were no indications of sex allocation. This is, to our knowledge, the first study demonstrating how female body condition and behaviour interact with the context of mating in shaping maternal reproductive investment.
Pavlovian conditioning; mate familiarity; body condition; maternal effects; sex allocation; fertilization success
The hypothesis that social stimulation, derived from the presence and activities of conspecifics, can hasten and synchronize breeding in colonies of birds was tested. A modified playback/recorder system was used to continuously exaggerate the amount of colony sound available to zebra finches throughout their courtship period. Males that heard ‘sound supplements’ generated from their own colony sang more than males in control colonies that did not receive playback; males that heard samples from a different colony, sang at an intermediate level. Females that were exposed to the vocalizations of their mate and playback from a colony other than their own, laid eggs earlier and more synchronously than females in control colonies. Females that heard the vocalizations of their mate along with playback samples generated from their own colony, laid eggs more synchronously but not earlier than control females. Both acoustic treatments caused females to lay larger clutches. Social stimulation influences the breeding schedule and clutch size in zebra finch colonies. If there are advantages associated with these effects, social stimulation may contribute to the maintenance of colonial breeding systems.
social stimulation; breeding schedules; clutch size; zebra finch; colony sound; synchrony
Although conspicuous courtship displays are an effective way of attracting the attention of receptive females, they could provide valuable information to rival males on the location of these females. In fiddler crabs, males that see a receptive female wave their single, greatly enlarged claw in a highly conspicuous courtship display. We test whether other males use this courtship display to alert them to the presence of receptive females that they cannot directly see. We show that male fiddler crabs (Uca mjoebergi) eavesdrop on the courtship displays of nearby males to detect mate-searching females. This allows males to begin waving before a female becomes visible. Furthermore, males appear to adjust their waving according to the information available: eavesdropping males wave 12 times faster than non-courting males but only 1.7 times slower than males in full visual contact with the female.
animal communication; eavesdropping; fiddler crabs; mate attraction; Uca mjoebergi
The sexual selection strategies of territorial Odonata that do not present courtship behavior is still not completely understood, especially the role of the females. Diastatops obscura Fabricius (Odonata: Libellulidae) females participate in mate selection in a passive manner, allowing copulation with the first male that captures them and afterwards choosing whether to oviposit or not. This study introduces the idea of female passive choice as an adaptative tactic in intersexual selection. Also discussed is the adaptative value of this tactic and its flexibility according to environmental conditions and reproductive strategies adopted by the males. A natural population of Diastatops obscura was observed in the Pitimbu River of northeast Brazil. Focal continuous and ad libitum techniques were used to record attempted copulation, copulation, and oviposition behavior, in addition to registering male territoriality. An estimate of individual reproductive success (IRS) was obtained by recording 187 reproductive events. Territorial males, mainly occupying areas near the river margin, achieved greater copulation and oviposition success (IRS = 0.371) than did satellite males (IRS = 0.028). Females that copulated with territorial males experienced, for the most part, only one copulation and oviposition event, while those that copulated with satellite males fled or performed a second copulation with a territorial male. Thus, the best tactic adopted by the D. obscura males was to occupy a territory providing the greatest access to females, while the females used passive choice for fitness optimization.
sexual selection; female choice; dragonfly; Pitimbu River
The spiny lizard Sceloporus grammicus (Squamata: Phrynosomatidae) is a small reptile from central México and the southern United States, occurring in a wide geographic area characterized by extensive variation in topographic and climatic regimes. Genetic variation among lineages from central México is substantial, though the extent to which this variation corresponds with life-history traits remains obscure. To address part of this puzzle, we studied a population of S. grammicus from Tepeapulco, Hidalgo, México. Male-biased sexual dimorphism was extensive in this population; males were larger than females overall, and expressed proportionately larger heads and longer limbs. Minimum size at sexual maturity was similar in the sexes (males: 43 mm; females: 42 mm). In contrast to other populations from the Central Plateau, reproductive activity of males and females was synchronous. Testicular recrudescence of adult males was initiated in October–November, and maximum testis size maintained from December to July. Female reproductive activity showed no clear seasonal pattern: females had vitellogenic follicles from October to July, and pregnant females were found throughout the year. Female body size was not related to litter size. Neither male nor female gonadal mass was correlated with any abiotic environmental variable examined. Differences in reproductive characteristics among populations of S. grammicus might be indicative of plasticity in response to local environmental conditions, local adaptation, or complex gene × environment interactions. We consider these results in the context of previously studied populations of S. grammicus from the Central Plateau and elsewhere, and propose directions for future research.
Interpopulation variation; life history; lizard; reproduction; sexual dimorphism
Activity patterns and social interactions play a key role in determining reproductive success, although this is poorly understood for species that lack overt social behaviour. We used genetic paternity analysis to quantify both multiple paternity and the relative roles of activity and social behaviour in determining reproductive success in a nondescript Australian lizard. During the breeding season we intensively followed and recorded the behaviour of a group of seven males and 13 females in a naturalistic outdoor enclosure to examine the relative roles of body size, activity and social interactions in determining male fertilization success. We found multiple paternity in 42% of clutches. No single behaviour was a significant predictor of male fertilization success in isolation, but male-female association, interactions and courtship explained 41% of the variation in male fertilization success. Males with the highest number of offspring sired invested heavily in interacting with females but spent very little time in interactions with males. These same males also sired offspring from more clutches. When taken collectively, an index of overall male activity, including locomotion and all social interactions, significantly explained 81% of the variation in the total number of offspring sired and 90% of the variation in the number of clutches in which males sired offspring. We suggest that the most successful male strategy is a form of endurance rivalry in which active mate searching and interactions with females have the greatest fitness benefits.