The sexual strategy of the central long-toed salamander, Ambystoma macrodactylum columbianum, in south-eastern Washington

The sexual strategies of salamanders are largely'unobservable'owing to the low visibility typical of their preferred breeding habitats. We conducted both field and laboratory studies to investigate the sexual strategy of the North American central long-toed salamander, Ambystoma macrodactylum columbianum. Capture-recapture and cohort-marking of salamanders at an aquatic breeding site indicated that males arrive before females and remain in the water for longer periods of time (measured in weeks, rather than days as for females). We estimate that the overall duration of the breeding season is short, three weeks or less. Courtship between single males and females is characterized by repeated bouts of axillary amplexus, separated by multiple episodes of spermatophore deposition. Amplectant males provide females with considerable tactile stimulation in the form of head-rubbing. Unpaired males attempt to displace amplectant males from their partners by wrestling with them. In addition, unpaired males appear to interfere in ongoing courtship encounters during the spermatophore deposition stage, perhaps attempting to 'steal'inseminations. We suggest that explosive scramble-competition polygyny best describes the mating system of A. m. columbianum. Operational sex ratios are probably male-biased throughout the short breeding season, leading to intense competition among males for mates. Mate choice (by either sex) probably is of little importance as a determinant of mating success.     

Sexual Selection Under Scramble Competition: Mate Location and Mate Choice in the Eucalypt Leaf Beetle Chrysophtharta agricola (Chapuis) in the Field

For males of many species, the number of offspring sired can depend on the number of females mated. While pre- and postcopulatory choice by females can affect the outcome of potential mate encounters, mate location is a necessary prerequisite to any possible courtship and subsequent mating. Mate location of Chrysophtharta agricola in the field was examined using sticky traps baited with sexually receptive conspecific beetles. More beetles were caught on traps baited with conspecific beetles of either sex than on control traps that contained foliage only. Furthermore, 94% of beetles captured on control traps were males, indicating that the mating system of Chrysophtharta agricola can be labeled prolonged searching scramble competition polyandry, in which receptive females are evenly dispersed spatially and temporally, and males search competitively for them. Operational sex ratios were 1:1 throughout the season. By sampling paired and unpaired beetles in the field, we found that beetles generally did not select mates based on body size. Furthermore, neither sex mated preferentially with partners that were uninfected by parasitic mites or with beetles of the same generation. In the absence of postcopulatory female choice, the ability of males to locate females may therefore be the most important trait in determining male mating success.     

The effect of size and sex ratio experiences on reproductive competition in Nicrophorus vespilloides burying beetles in the wild

Male parents face a choice: should they invest more in caring for offspring or in attempting to mate with other females? The most profitable course depends on the intensity of competition for mates, which is likely to vary with the population sex ratio. However, the balance of pay‐offs may vary among individual males depending on their competitive prowess or attractiveness. We tested the prediction that sex ratio and size of the resource holding male provide cues regarding the level of mating competition prior to breeding and therefore influence the duration of a male's biparental caring in association with a female. Male burying beetles, Nicrophorus vespilloides were reared, post‐eclosion, in groups that differed in sex ratio. Experimental males were subsequently translocated to the wild, provided with a breeding resource (carcass) and filmed. We found no evidence that sex ratio cues prior to breeding affected future parental care behaviour but males that experienced male‐biased sex ratios took longer to attract wild mating partners. Smaller males attracted a higher proportion of females than did larger males, securing significantly more monogamous breeding associations as a result. Smaller males thus avoided competitive male–male encounters more often than larger males. This has potential benefits for their female partners who avoid both intrasexual competition and direct costs of higher mating frequency associated with competing males.     

Males,but not females,perform strategic mate searching movements between host plants in a leaf beetle with scramble competition polygyny

Mate searching is assumed to be performed mostly by males, but when females benefit from multiple mating or are under risk of failing to mate, they may also perform mate searching. This is especially important in scramble competition polygynies, in which mate searching is the main mechanism of mate competition. Typically, more mobile individuals are expected to achieve higher mating success because mobility increases their probability of finding mates. If we assume individual movements are mainly explained by mate searching in scramble competition polygynies, we can investigate searching strategies by asking when individuals should leave their location and where they should go. We hypothesize that individuals will leave their locations when mating opportunities are scarce and will seek spatially close sites with better mating opportunities. We tested these hypotheses for males and females of Leptinotarsa undecimlineata, a leaf beetle with scramble competition polygyny in which both sexes are promiscuous. Individuals mate and feed exclusively on Solanum plants, and thus, individual movements can be described as switches between plants. Females were less likely than males to leave isolated plants, and both males and females moved preferentially to neighboring plants. Males were more likely to leave when the local number of females was low, and the number of males was high. They moved to plants with more females, a behavior consistent with a mate searching strategy. Females were more likely to move to plants with fewer males and many females, a behavior consistent with male harassment avoidance. Strategic movement is widely considered in foraging context, but seldom in a mate searching context. Considering that selection to minimize searching costs, maximize mating success, and minimize harassment may be ubiquitous in nature, we argue that strategic movements by mate searching individuals are likely to occur in many species.     

Mate Switching in a Non-monogamous Species? The Case of the Common Quail (Coturnix coturnix)

We investigated the occutrence of mate switching in the common quail, a non-monogamous species with a temporal pair bond but without either male parental care or territoriality. The study was carried out throughout the breeding seasons of 1993–95 in Mas Esplugues (Catalonia, Spain) by monitoring 28 radio-tagged couples and 17 radio-tagged unpaired males. Agonistic interactions between males inside funnel traps containing a female were also recorded. Mate switching within the laying period occurred in 72% of the females, and new partners always had a higher body condition index than old partners. Agonistic interactions inside the funnel traps also showed that successful males had a better body condition index than losers. These results, along with the observed mate-guarding behaviour of females by their partners throughout the laying period, a highly male-biased sex ratio (five males per female), the lack of territoriality of males and the expected difficulties which unmated males experience in finding pairs, suggest that mate switching is not induced by paired males. Moreover, the constant inflow of new males observed throughout the fertile period of the female and the low costs stemming from mate change strongly support the idea that it is paired females who induce mate switching, in order to improve their fitness by mating with the best quality male available at every moment of their fertile period.     

Mutual mate choice by mountain pine beetles: size‐dependence but not size‐assortative mating

1. Mutual mate choice may be rare, occurring when both sexes invest heavily in reproduction, mating opportunities are abundant, and individuals differ in quality. 2. Mountain pine beetles, Dendroctonus ponderosae (Curculionidae: Scolytinae) appear to meet the conditions for mutual mate choice. We introduced males to females in breeding sites and observed the occurrence and speed of a male entering a female's gallery. We tested for consequences of mutual mate choice, namely condition‐dependent choosiness and assortative mating. 3. Males were more likely to enter a female's gallery when the gallery was in a smaller tree with less resin production and when the gallery was larger. Female body size and condition did not influence the probability of entry. Larger males were less likely to enter a gallery than were smaller males, probably because of size‐dependent choosiness rather than physical limitations. 4. Small males took longer to enter galleries of large females than of small females, whereas large males entered as quickly into galleries of large females as small females. This suggests size‐dependent choosiness by females. 5. No assortative mating by body size was detected, probably because males appeared to choose on the basis of female‐associated resources rather than on female traits.     

Early male reproductive advantage,multiple paternity and sperm storage in an amphibian aggregate breeder

We tested whether the order in which males encounter females affects reproductive fitness in spotted salamanders (Ambystoma maculatum). Using mating chambers in the field, we allowed one male access to a female before a second male. We then used four microsatellite markers in paternity analyses of the resulting larvae. First males sired a significantly larger number of offspring than second males, suggesting that male reproductive success is greatly enhanced by early arrival at breeding ponds. Multiple paternity was common among clutches, and frequently larvae were assigned to unidentified males that had not been in the chambers. Sperm from these males had either been stored by females for a year or obtained more recently at other breeding sites.     

The Starling Mating System as an Outcome of the Sexual Conflict

Many bird species demonstrate a variable mating system, with some males being monogamously mated and other males able to attract more than one mate. This variation in avian mating systems is often explained in terms of potential costs of sharing breeding partners and compensation for such costs. However, whenever there is a difference in the optimal mating system for males and females, a sexual conflict over the number of partners is expected. This paper contains a verbal model of how a conflict between male and female European starlings (Sturnus vulgaris),resulting from the fitness consequences of different mating systems for males and females differing over time, determines the mating system. We demonstrate that males and females have contrasting fitness interests regarding mating system, such that males gain from attracting additional mates whereas already mated females pay a cost in terms of reduced reproductive success if males are successful in attracting more mates. We demonstrate how this can be traced to the rules by which males allocate non-sharable care between different broods. Furthermore, we demonstrate that there exist male and female conflict behaviours with the potential to affect the mating system. For example, aggression from already mated females towards prospecting females can limit male mating success and males can circumvent this by spacing the nest-sites they defend. The realised mating system will emerge as a consequence of both the fitness value of the different mating systems for males and females, and the costs for males and females of intersexual competition. We discuss how this model can be developed and critically evaluated in the future.     

Differences in potential reproductive rates of male and female seahorses related to courtship roles

Dwarf seahorses, Hippocampus zosterae (Syngnathidae), are distinguished by extreme morphological specialization for paternal care, the formation of monogamous pair bonds and mating repeatedly over the course of a breeding season. To determine the potential reproductive rates of male and female dwarf seahorses, we measured (1) the maximum number of offspring produced per breeding cycle when sexually receptive mates were unlimited, and (2) the relative time each sex was unavailable for mating ('time out'). We paired sexually isolated males and females with sexually receptive partners and observed them from the day of introduction through to copulation, to determine the length of time it takes each sex to prepare to mate. We conducted additional experiments to determine the length of gestation, which when added to the time needed to prepare to mate and copulate gives an estimate of total reproductive cycle duration, T. We estimated potential reproductive rate by dividing the mean number of offspring produced per breeding cycle by the duration of the breeding cycle (T). We estimated reproductive 'time out' by identifying the period of time males and females were physiologically capable of mating ('time in', S) and subtracting time S from time T. When provided with sexually receptive partners, females took 2 days longer than males to complete courtship and copulation, but neither males nor females remated during gestation. Therefore, males could potentially produce 17% more offspring than females over the course of one breeding season. Females had reproductive 'times out' 1.2 times longer than did males, as they were only capable of mating during the 4 h directly preceding copulation. Thus, H. zosterae males have higher potential reproductive rates and shorter reproductive 'times out' compared with H. zosterae females. These results and previous work indicating that seahorses display traditional courtship roles support the prediction that the sex having the higher potential reproductive rate, or equivalently, the shorter 'time out', will compete more intensely for access to the opposite sex. Copyright 2000 The Association for the Study of Animal Behaviour.     

Familiarity and flexible mating strategies of a solitary rodent, Dipodomys ingens

Because mating is a product of individual reproductive strategies that may vary with changing conditions, we predicted variable mating behaviour in an arid-adapted, territorial rodent, the giant kangaroo rat, Dipodomys ingens. We also predicted that familiarity would facilitate nonaggressive courtship and mating in this solitary rodent. Through direct observations in the field, we found that mating varied from exclusive to multiple partners. Where densities were low, and on nights when multiple females were in oestrus, each animal mated with one member of the opposite sex. In conditions where the operational sex ratio was skewed towards multiple males, males footdrummed and competed for females. Males were able to mate with one or two females in adjacent territories, and they successfully competed for these same females throughout the breeding season. Females that mated exclusively with one male had more pups emerge from the burrow compared with females that experienced male competition. Females allowed nearest neighbour males to enter their burrows, and they engaged in more nonaggressive contact with close neighbours than with other males. Paired encounters in the field showed less aggression towards neighbours than strangers. In laboratory tests, females were less aggressive towards and allowed more contact with familiar than unfamiliar males. These results show that D. ingens can alter mating strategies as conditions change. Familiarity is an important factor in nonaggressive interactions between males and females and may be important to mate preferences in females during reproduction. The less aggressive behaviour to neighbours than to strangers (‘dear enemy’ phenomenon) is consistent with other solitary animals that defend multipurpose territories. Copyright 2002 The Association for the Study of Animal Behaviour. Published by Elsevier Science Ltd. All rights reserved.     

The Mating System of <Emphasis Type="Italic">Habropoda pallida</Emphasis> Timberlake (Anthophorinae: Apidae)

Males of Habropoda pallida searched for recently emerged females at a location in central Arizona where many females had nested in the preceding year. Patrolling males exhibited a variety of mate locating tactics ranging from inspecting existing emergence holes to patrolling flights around flowering creosote bush. These tactics appear to be under the control of a conditional strategy, given that male behavior shifted during the day. Moreover, some marked males mounted dead females placed on the ground in the emergence area as well as mounting dead females pinned to flowers in the emergence area and at a creosote bush some distance away. At least some males exhibited site fidelity in that numerous males marked within the nest site emergence area were recaptured there; others marked at a creosote bush were also seen again at that plant. The mating system of the bee involves scramble competition with individual males attempting to locate receptive emerging and flower-visiting females before rival males. In keeping with the scramble competition hypothesis, males are considerably smaller on average than females and are only aggressive when trying to push their way past rival males already mounted on a potential mate.     

Male mating competition,female choice and dominance in a free ranging group of Japanese macaques

Focal and ad libitum samples of high and middle ranked males in a group of free ranging Japanese macques were taken in order to examine rank related differences in male mating strategies. Males tended to have like ranked females as consort partners, with high rank males showing more consort activity, over all, than middle rank males. High rank males tended to interfere in consorts and middle rank males tended to have their consorts disrupted. Consorts involving high rank females were most subject to interference. With one exception, proximity between partners in consorts involving high rank males was due to male actions while proximity in consorts involving middle ranked males was due to female actions. The two highest ranked males were never observed copulating. Their mating failure may have been due to avoidance by females who had known them since immaturity. High rank males were somewhat more likely than middle rank males to have consorted with females during the period of likely conception. There was some evidence that frequent consort partners joined the same subgroup during a group fission. Males appeared to use the advantage conferred by high rank mainly in competition for high rank females. Females showed some indications of preference for mates likely to retain or attain high rank in the future.     

Determinants of breeding dispersal in the sand lizard, Lacerta agilis, (Reptilia, Squamata)

We studied determinants of breeding dispersal (the distance that an animal shifts its mean home range co-ordinate between reproductive events) in an individually marked population of sand lizards (Lacerta agilis) in south west Sweden during 1987–1991. Female breeding dispersal was not determined by age, size, body condition, or number of partners, and females and males that dispersed further did not experience a higher mortality. However, females with a low reproductive output dispersed further than females that reproduced more successfully, and males that lost in bodily condition dispersed further than males that better maintained body condition. We also looked for relationships between age-differences and band-sharing similarity (DNA fingerprinting = DFP) in three categories of lizards — all females, all males, and males and females — to establish whether males would be likely to mate with close kin. Age-difference was strongly correlated with band-sharing in only one category, males and females. When males were older than females this relationship was not significant. However, when females were older than males, age-difference was strongly correlated with band-sharing. Furthermore, females that were old enough to be the mothers of the courting males shared significantly more bands with these males than did the younger females, including the females of the same age as the males. Although parental-specific DFP bands necessary for establishing paternity among our adult lizards were inaccessible to us, we suggest that our circumstantial evidence strongly suggests that some males mate with their mothers. Males that were more closely related with their neighbours also moved further when we controlled statistically for age and mating success. We suggest that by mating with many partners males not only increase their mating success, but also increase the probability of mating with females with ‘good genes’: mean heterozygosity of parents (as revealed by micro-satellites) were strongly correlated with offspring survival.     

Extra‐group mating increases inbreeding risk in a cooperatively breeding bird

In many cooperatively breeding species, females mate extra‐group, the adaptive value of which remains poorly understood. One hypothesis posits that females employ extra‐group mating to access mates whose genotypes are more dissimilar to their own than their social mates, so as to increase offspring heterozygosity. We test this hypothesis using life history and genetic data from 36 cooperatively breeding white‐browed sparrow weaver (Plocepasser mahali) groups. Contrary to prediction, a dominant female's relatedness to her social mate did not drive extra‐group mating decisions and, moreover, extra‐group mating females were significantly more related to their extra‐group sires than their social mates. Instead, dominant females were substantially more likely to mate extra‐group when paired to a dominant male of low heterozygosity, and their extra‐group mates (typically dominants themselves) were significantly more heterozygous than the males they cuckolded. The combined effects of mating with extra‐group males of closer relatedness, but higher heterozygosity resulted in extra‐group‐sired offspring that were no more heterozygous than their within‐group‐sired half‐siblings. Our findings are consistent with a role for male–male competition in driving extra‐group mating and suggest that the local kin structure typical of cooperative breeders could counter potential benefits to females of mating extra‐group by exposing them to a risk of inbreeding.     

Role of Contests in the Scramble Competition Mating System of a Leaf Beetle

Male competition for mates can occur through contests or a scramble to locate females. We examined the significance of contests for mates in the leaf beetle Chrysomela aeneicollis, which experiences a short breeding season. During peak mating season, 18–52% of beetles are found in male-female pairs, and nearly half of these are copulating. Sex ratios do not differ from parity, females are larger than males, and positive size-assortative mating occurs. Males fight (2–4% of beetles) over access to females, and disruption of mating usually follows these contests. In the laboratory, we compared mating and fighting frequencies for males found in mating pairs (field-paired) and single males placed into an arena with a field-paired female. Mating pairs were switched in half of arenas (new male-female pairs) and maintained in the other half. For 2 days, each male was free to move about and fight; thereafter males were tethered to prevent contests. Mating frequencies were significantly greater for field-paired than single males in both situations. Male size was not related to mating frequency; however, large females received more matings than small ones. These data suggest that males fight for high quality females, but otherwise search for as many matings as possible.     

Correlations between body size, defensive behaviour and reproductive success in male Little Blue Penguins Eudyptula minor: implications for female choice

Field studies exploring relationships among body size, defensive behaviour (including vocalizations) and reproductive success in male Little Blue Penguins Eudyptula minor were conducted to identify behavioural and physical attributes that might provide females or opponents with information on male quality. Head length provided a simple estimator of body size in both sexes. Nests defended by large males produced eggs and chicks earlier in the breeding season, and contained chicks that grew more quickly than chicks in nests defended by small males. In contrast, female body size did not affect any of the reproductive parameters we measured. The pitch of male defensive calls varied significantly with male size, so females (or opponents) could potentially gauge male size by assessing the calls. Large males were also more likely to respond vocally to simulated nest intrusions. As a result, by 'keeping company' with prospective partners early in the breeding season, females can obtain information about males that would allow them to select better partners and improve their own reproductive potential.     

The male and female perspective in the link between male infant care and mating behaviour in Barbary macaques

Infant care from adult males is unexpected in species with high paternity uncertainty. Still, males of several polygynandrous primates engage in frequent affiliative interactions with infants. Two non‐exclusive hypotheses link male infant care to male mating strategies. The paternal investment hypothesis views infant care as a male strategy to maximize the survival of sired offspring, while the mating effort hypothesis predicts that females reward males who cared for their infant by preferably mating with them. Both hypotheses predict a positive relationship between infant care and matings with a particular female. However, the paternal investment hypothesis predicts that increased matings come before infant care whereas the mating effort hypothesis predicts that infant care precedes an increase in matings. Both hypotheses are usually tested from the perspective of the proportion of matings and care that individual females engage in and receive, rather than from the perspective of the care and mating behaviour of individual males. We tested the relationships between care and mating from both female and male perspectives in Barbary macaques. Mating predicted subsequent care and care predicted subsequent mating when viewed from the male but not the female perspective. Males mainly cared for infants of their main mating partners, but infants were not mainly cared for by their likely father. Males mated more with the mothers of their favourite infants, but females did not mate more with the main caretakers of their infants. We suggest that females do not choose their mating partners based on previous infant care, increasing paternity confusion. Males might try to increase paternal investment by distributing the care according to their own instead of female mating history. Further, males pursue females for mating opportunities based on previous care.     

Mate finding in the forest cockchafer, Melolontha hippocastani, mediated by volatiles from plants and females

The response of the forest cockchafer, Melolontha hippocastani F. (Coleoptera, Scarabaeidae), towards volatiles emitted by different host plants and conspecifics was tested in field experiments during the flight period at dusk. Funnel traps containing artificially damaged leaves from the host plants Carpinus betulus L. and Quercus rubra L., as well as from the non‐host plant Prunus serotina Ehrh. caught significantly more beetles than empty control traps. On the other hand, traps baited with undamaged leaves from Q. rubra did not catch significantly more beetles than empty controls. Leaves from C. betulus damaged by beetle feeding did not attract more beetles than artificially damaged leaves. By use of gas chromatography coupled with electroantennographic detection (GC‐EAD) electrophysiological responses of males and females were shown for 18 typical plant volatiles. A synthetic mixture of selected typical green plant volatiles was also highly attractive in the field. A total of 9982 beetles was caught during the field experiments, among them only 33 females. This suggests that attraction to damaged foliage during flight period at dusk is male‐specific. Field experiments testing the attractiveness of female M. hippocastani towards conspecific males by employing caged beetles and beetle extracts indicated that males of M. hippocastani use a female‐derived sex pheromone for mate location. On wired cages containing either unmated feeding females, or unmated females without access to foliage, or feeding males in combination with extracts from unmated females, significantly more males landed during the flight period than on comparable control cages containing feeding males or male extracts. A possible scenario of mate location in M. hippocastani involving feeding‐induced plant volatiles and a female‐derived sex pheromone is discussed.     

The role of male age, sperm age and mating history on fecundity and fertilization success in the hide beetle

Models of age-related mate choice predict female preference for older males as they have proven survival ability. However, these models rarely address differences in sperm age and male mating history when evaluating the potential benefits to females from older partners. We used a novel experimental design to assess simultaneously the relative importance of these three parameters in the hide beetle, Dermestes maculatus. In a two-part experiment we first explored age-related male mating success and subsequently examined the consequences of male age, sperm age and male mating history on female fecundity and fertilization success. In a competitive mating environment, intermediate-age males gained significantly higher mating success than younger or older males. To test the consequences for females of aged-related male mating success, a second set of females were mated to males varying in age (young, intermediate-age and old), in numbers of matings and in timing of the most recent mating. We found that male age had a significant impact on female fecundity and fertilization success. Females mated to intermediate-age males laid more eggs and attained consistently higher levels of fertilization success than females with young and old mates. A male's previous mating history determined his current reproductive effort; virgin males spent longer in copula than males with prior mating opportunities. However, differences in copulation duration did not translate into increased fecundity or fertilization success. There was also little evidence to suggest that fertilization success was dependent on the age of a male's sperm. The experiment highlights the potential direct benefits accrued by females through mating with particular aged males. Such benefits are largely ignored by traditional viability models of age-related male mating success.     

Sexual selection for male mobility in a giant insect with female-biased size dimorphism

Female-biased size dimorphism, in which females are larger than males, is prevalent in many animals. Several hypotheses have been developed to explain this pattern of dimorphism. One of these hypotheses, the mobility hypothesis, suggests that female-biased size dimorphism arises because smaller males are favored in scramble competition for mates. Using radiotelemetry, we assessed the mobility hypothesis in the Cook Strait giant weta (Deinacrida rugosa), a species with strong female-biased size dimorphism, and tested the prediction that male traits promoting mobility (i.e., longer legs, smaller bodies) are useful in scramble competition for mates and thus promote reproductive success. Our predictions were supported: males with longer legs and smaller bodies exhibited greater mobility (daily linear displacement when not mating), and more mobile males had greater insemination success. No phenotypic traits predicted female mobility or insemination success. In species with female-biased size dimorphism, sexual selection on males is often considered to be weak compared to species in which males are large or possess weaponry. We found that male giant weta experience sexual selection intensities on par with males of a closely related harem-defending polygynous species, likely because of strong scramble competition with other males.